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W.—Status of Acigona Hiibner (sensu Bleszynski) (Lepidoptera: Pyralidae-.Cram- binae) with changes in nomenclature ................. 0.000 e cece eee CBA is BRT 100 BAIMAI, V., R. G. ANDRE, B. A. HARRISON, U. KIJCHALAO, and L: PANTHUSIRI— Crossing and chromosomal evidence for two additional sibling species within the taxon Anopheles dirus Peyton and Harrison (Diptera: Culicidae) in Thailand ................. 157 BELLINGER, R. G. and R. L. PPENKOWSKI-—Life history observations on the grasshopper Appalachia hebardi Rehn and Rehn (Orthoptera: Acrididae: Melanoplinae) ............. 43 CLARKE, J. F. G.—The correct identity of Acleris inana (Robinson) (Lepidoptera: Tortricidae) 175 DIATLOFF, G. and W. A. PALMER—The host specificity of Neolasioptera lathami Gagné i (Diptera: Cecidomyiidae) with notes on its biology and phenology ..................... 122 HARRIS, S. C.—A new species of Agarodes (Trichoptera: Sericostomatidae) from southeastern WinttedStates nV Mi OME O TEE BEN UTE OAL eB a0 Ab A AAs Me Aa Re ae 74 HOFFMAN, K. M.—Earwigs (Dermaptera) of South Carolina, with a key to the eastern North American species and a checklist of the North American fauna .....................-. 1 KONDRATIEFF, B. C. and R. F. KIRCHNER — Additions, taxonomic corrections, and faunal affinities of the stoneflies (Plecoptera) of Virginia, USA ..............6.50..00002204-- 24 LOWRY, J. E. and H. D. BLOCKER—Two new species of Flexamia from the Nebraska Sand Hills\(Homoptera: Cicadellidae: Deltocephalinae))}.) 2. 0) P open Eee Fae Bioele ae 57 MARI MUTT, J. A.—A new Seira from the United States, with a redescription of Seira bi- punctata (Packard) and new records for Seira distincta Mari Mutt (Collembola: Entomobry- NELSON, C. R. and R. W. BAUMANN-—Scanning electron microscopy for the study of the winter stonefly genus Capnia (Plecoptera: Capniidae) ............ 2.00.2. ee eee eee eee 31 PALMER, W. A.—The phytophagous insect fauna associated with Baccharis halimifolia L. and B. neglecta Britton in Texas, Louisiana, and northern Mexico .............5-.-524.4005- (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ORGANIZED MARCH 12, 1884 OFFICERS FOR 1987 THOMAS E. WALLENMAIER, President MICHAEL J. RAupPP, Program Chairman F. EUGENE Woon, President-Elect GEOFFREY B. WHITE, Membership Chairman PAUL M. MArsH, Recording Secretary Victor L. BLACKBURN, Custodian RICHARD G. RosBINS, Corresponding Secretary MANYA B. STOETZEL, Delegate, Wash. Acad. Sci. NORMAN E. WoobpLey, 7reasurer RAYMOND J. GAGNE, Editor Publications Committee DAVID R. SMITH , THEODORE J. SPILMAN GEORGE C. STEYSKAL Honorary President C. F. W. MUESEBECK ; Honorary Members FREDERICK W. Poos ASHLEY B. GURNEY THEODORE L. BISSELL All correspondence concerning Society business should be mailed to the appropriate officer at the following address: Entomological Society of Washington, c/o 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. -— Published quarterly beginning with January by the Society at Washington, D.C. Members in good standing receive the Proceedings of the Entomological Society of Washington. Nonmember subscriptions are $35.00 per year, domestic, and $40.00 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. 730 of the July 1984 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 Wash- ington, c/o Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash- ington, D.C. 20560. Editor: Raymond J. Gagné, Systematic Entomology Laboratory, c/o U.S. National Museum NHB 168, Wash- ington, D.C. 20560. Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 30 January 1987 Second Class Postage Paid at Washington, D.C. and additional mailing office. PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA PROC. ENTOMOL. SOC. WASH. 89(1), 1987, pp. 1-14 EARWIGS (DERMAPTERA) OF SOUTH CAROLINA, WITH A KEY TO THE EASTERN NORTH AMERICAN SPECIES AND A CHECKLIST OF THE NORTH AMERICAN FAUNA KEVIN M. HOFFMAN Department of Entomology, Clemson University, Clemson, South Carolina 29634- 0365. Abstract. — Biological and distributional data are given for the nine species of Dermap- tera recorded from South Carolina. A key to the 17 eastern North American species is presented along with brief notes on the taxonomy and distribution of all 22 species occurring in North America north of Mexico. The most recent comprehensive checklist of North American Dermaptera was pub- lished over 35 years ago (Gurney, 1950), and the last key to the eastern North Amer- ican species was published by Blatchley (1920). In the intervening years new species have been introduced, and the taxonomic status of others has changed. This paper documents the species occurring in South Carolina, presents a revised checklist for the species occurring in North America north of Mexico, and provides a key to eastern North American species. A similar study of the Dermaptera of California provided a key to most of the known western species (Langston and Powell, 1975). The order Dermaptera comprises some 1200 species worldwide, occurring mostly in the tropics. Only 22 species are currently known from North America, and at least twelve of these were introduced from Eu- rope and the tropics. Seventeen species have been recorded from eastern North America, nine of which have been found in South Carolina. These nine are the most wide- spread species in the east; the other eight species are restricted to Florida and New Jersey. The remaining five North American species occur only in western United States. A brief summary of the North American records for non-South Carolina species fol- lows. There are eight eastern species not found in South Carolina. Pyragropsis buscki (Cau- dell) is a Caribbean species that has become established in the Miami, Florida area (Gur- ney, 1959). Euborellia ambigua (Borelli) was originally described from Costa Rica and has since been recorded from a mangrove swamp near Miami, Florida (Hebard, 1922). Euborellia caraibea Hebard, a species that is widespread in the Caribbean Islands, has been collected in sandy areas on Virginia Key, Florida (Brindle, 1971d). The char- acters originally used by Hebard (1922) to differentiate E. caraibea from E. ambigua have shown considerable variation when a larger series of E. caraibea was examined, making it difficult to distinguish the two species (Brindle, 1971d). This difficulty is reflected in the following key and awaits the collection of a larger series of E. ambigua before it can be resolved. Euborellia an- nulata (F.), a native of Indonesia, was re- corded by Townes (1946) from Miami, Florida (as Euborellia stali [Dohrn, 1864a], synonymized by Brindle, 1981). Specimens of this species are in the Florida State Col- lection of Arthropods from as far north as Daytona, Florida. Labia curvicauda (Motschulsky), origi- nally described from Ceylon, has been found on Long Key, Florida, occupying the bases of leaves in the tops of coconut palms (Rehn and Hebard, 1912). Labia rehni Hebard is known only from a single female found be- tween boards in a woodshed on Key West, Florida, on 7 July 1912 (Hebard, 1917; Brindle, 1971a). Doru davisi Rehn and He- bard is found only in southern Florida, es- pecially around Lake Okeechobee, and is usually associated with saw-grass (Gurney, 1972). Marava arachidis (Yersin) was de- scribed from southern Europe and has sub- sequently been introduced into southern Florida (Caudell, 1913). This species has been recorded from two localities in New Jersey by Townes (1946) (as Marava wal- lacei [Dohrn, 1864b], synonymized by Boeseman, 1954). It has also been recorded from Texas, Arizona, and California (He- bard, 1943; Langston and Powell, 1975). Five species occur only in western United States. Chelisoches morio (F.) is a south Pa- cific species that has been introduced into California (Langston and Powell, 1975). Euborellia cincticollis (Gerstaecker) is an African species that is now established in California and Arizona (Langston and Pow- ell, 1975). Euborellia femoralis (Dohrn), an Oriental species that is very similar to E. annulipes, has been recorded from Arizona and California (Steinmann, 1981). Vostox apicedentatus (Caudell) is a native species that has been recorded from California, Ar- izona, New Mexico, and Texas. It was listed in the genus Spongovostox by Langston and Powell (1975) but has been transferred to the genus Vostox by Brindle (1971b). Vos- tox excavatus Nutting and Gurney is known only from Arizona, New Mexico, and northwestern Mexico (Nutting and Gurney, 1961). Labia pilicornis (Motschulsky, 1863) was listed by Gurney (1950) as being established in the United States, but no North Ameri- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON can collection records were cited. A foot- note associated with the listing stated that Morgan Hebard considered Labia rehni to be ajunior synonym of L. pilicornis, so per- haps it was included in the list in antici- pation of this proposed synonymy. Because subsequent workers have maintained the two species as separate and no published Nearctic records have been found, it is doubtful that L. pilicornis occurs in North America and thus is omitted from the checklist below. In the Neotropical region, L. pilicornis is known only from Cuba (Brin- dle, 19714). Earwigs exhibit sexual dimorphism in both the number of abdominal segments and the shape of the forceps. Males have ten abdominal segments, whereas females have only eight apparent segments due to the fu- sion of the eighth and ninth segments with the tenth (Giles, 1963). The forceps of males are generally curved and widely separated at the base, and many species have one or more prominent teeth on the inner margins (Figs. 11-25). Female forceps are more or less straight, usually closer together basally than those of the male and lack prominent teeth on their inner margins (Figs. 26-31). Both sexes possess a divided plate called the pygidium, which is located between the bas- es of the forceps (Fig. 18). The pygidium is thought to be derived from the epiproct and is useful taxonomically, especially in males. Immature earwigs undergo five to six in- stars before reaching adulthood. Antennal segments are added at each molt, and wing pads begin to develop in the second or third instar. Immatures are lighter in color than adults and possess conspicuous ecdysial lines on the head and thoracic terga. Immatures have male-like, ten-segmented abdomens, but have female-like, straight forceps. Material for this study was examined from the following institutions: North Carolina State University, University of Georgia, Florida State Collection of Arthropods, Na- tional Museum of Natural History, and Clemson University. The body length of VOLUME 89, NUMBER 1 specimens was measured from the head to the tip of the forceps. A brief diagnosis and information on the synonymy, bionomics, world distribution, and South Carolina dis- tribution are given for those species record- ed from South Carolina. The synonymy giv- en for each South Carolina species is complete unless otherwise noted, in which case synonyms commonly used in previous literature on Nearctic species are provided, and a reference is cited which contains a more complete synonymy. South Carolina county records for each species include every county from which the species has been col- lected, but not necessarily every record from those counties. The known North American distribution of each species is included in the following checklist, with states desig- nated by the two-letter codens assigned by the United States Postal Service. The rec- ords used in compiling the checklist are con- sidered to represent established popula- tions, although it is possible that some may represent transient infestations that have since disappeared. The classification system used in the checklist is that of Popham (196Sa, b). CHECKLIST OF THE NORTH AMERICAN SPECIES OF DERMAPTERA Superfamily Pygidicranoidea Family Pygidicranidae Subfamily Pygidicraninae Pyragropsis buscki (Caudell, 1907); ge Superfamily Labioidea Family Carcinophoridae Subfamily Carcinophorinae Anisolabis maritima (Bonelli, 1832); widespread on sea coasts. Euborellia ambigua (Borelli, 1906); FL. Euborellia annulata (F., 1793); FL. Euborellia annulipes (Lucas, 1847): widespread. Euborellia caraibea Hebard, 1922; EL; Euborellia cincticollis (Gerstaecker, 1883); AZ, CA. Euborellia femoralis (Dohrn, 1863); AZ, CA. Family Labiidae Subfamily Labiinae Labia curvicauda (Motschulsky, 1863); FL. Labia minor (L., 1758); widespread. Labia rehni Hebard, 1917; FL. Subfamily Spongiphorinae Marava arachidis (Yersin, 1860); AZ, CA, EX, NIGEE. Marava pulchella (Audinet-Serville, 1839); southeastern U.S., TX. Vostox apicedentatus (Caudell, 1905); AZ, CA, NM, TX. Vostox brunneipennis (Audinet-Ser- ville, 1839); eastern U.S., TX, OK. Vostox excavatus Nutting and Gur- ney, 1961; AZ, NM. Superfamily Forficuloidea Family Labiduridae Subfamily Labidurinae Labidura riparia (Pallas, 1773); southeastern U.S., AZ, CA, TX, OK. Family Chelisochidae Subfamily Chelisochinae Chelisoches morio (F., 1775); CA. Family Forficulidae Subfamily Forficulinae Doru aculeatum (Scudder, 1876): eastern U.S., Ontario. Doru davisi Rehn and Hebard, 1914; FL. Doru taeniatum (Dohrn, 1862); southeastern U.S., AZ, CA, TX. Forficula auricularia L., 1758; wide- spread. KEY TO ADULTS OF THE EASTERN NORTH AMERICAN SPECIES OF DERMAPTERA Ie Tegminae absent or present only as rounded flaps that do not meet at the inner basal margins (Figs. | and 2); right forceps 4 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-10. Thoracic terga, metatarsi, and antennal bases of Dermaptera. 1, Euborellia annulata, thoracic terga, dorsal view. 2, E. ambigua, thoracic terga (redrawn from Hebard, 1922), dorsal view. 3, Marava pulchella, pronotum and wings of fully winged form, dorsal view. 4, M. arachidis, pronotum (redrawn from Brindle, 1971), dorsal view. 5, Doru aculeatum, right metatarsus, lateral view. 6, D. aculeatum, right metatarsus, dorsal view. 7, M. pulchella, right metatarsus, lateral view. 8, M. pulchella, right metatarsus, dorsal view. 9, Forficula auricularia, base of right antenna, dorsal view. 10, D. aculeatum, base of right antenna, dorsal view. a4 = fourth antennal segment, cl = tarsal claw, hw = hindwing, ms = mesonotum, mt = metanotum, pr = pronotum, te = tegmina, ts] = first tarsal segment, ts2 = second tarsal segment, ts3 = third tarsal segment. ZL): of male curved inward more strongly than left (Figs. 11-12) (Carcinophoridae) .... Tegminae normally developed and meet- ing along entire midline (Fig. 3); forceps of male symmetrical (Figs. 14-25) ..... Tegminae present as rounded lateral flaps on mesonotum (Figs. | and 2); Florida . Tegminae absent; widespread ......... Tegminal flaps widely separated on meso- notum (Fig. 1); legs with brown rings around femora and tibiae; left forceps of male curved only at apex (Fig. 12) ..... Se SEU Rtt, Arete ere Euborellia annulata (F.) 5(2). Tegminal flaps covering most of meso- notum (Fig. 2); legs unicolorous or femora darkened; left forceps of male curved al- most as much as right (Fig. 13) ....... Legs uniformly pale; tegminal flaps not always meeting at midline (Fig. 2); found only in mainland mangrove swamps ... PEAS PRE ee Euborellia ambigua (Borelli) Legs pale or with femora darkened; teg- minal flaps always meeting or overlapping at midline; found in various habitats on 1S] and Smee Euborellia caraibea Hebard Antennae entirely dark brown, 20-24 seg- VOLUME 89, NUMBER 1 Figs. 11-18. Male forceps of Dermaptera, dorsal views. 11, Anisolabis maritima. 12, Euborellia annulipes. 13, E. ambigua (redrawn from Hebard, 1922). 14, Labia minor. 15, L. curvicauda (redrawn from Brindle, 197 1a). 16, Marava arachidis (redrawn from Brindle, 1971b). 17, M. pulchella. 18, Vostox brunneipennis. a10 = tenth abdominal segment, fe = forceps, pg = pygidium. 6 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 22 23 Figs. 19-25. Male forceps of Dermaptera, dorsal views. 19, Doru taeniatum. 20, D. aculeatum. 21, D. davisi (redrawn from Gurney, 1972). 22, Pyragropsis buscki, arcuate form (redrawn from Gurney, 1959). 23, P. buscki, elongate form (redrawn from Gurney, 1959). 24, Labidura riparia. 25, Forficula auricularia. mented; femora and tibiae without encir- tibiae pale yellow with encircling dark cling dark bands; body length 20-25 mm bands; body length 12-18 mm ........ LEN eee eae Anisolabis maritima (Bonelli) oe .eeesee...... Euborellia annulipes (Lucas) - Antennae brown with 1-3 white subapical 6(1). Second tarsal segments cylindrical and not segments, 14-18 segmented; femora and wider than third, at most only slightly ex- VOLUME 89, NUMBER 1 Figs. 26-31. 28 Female forceps of Dermaptera, dorsal views. 26, Euborellia annulipes. 27, Labia minor. 28, L. rehni (redrawn from Hebard, 1917). 29, Marava pulchella. 30, Vostox brunneipennis. 31, Doru aculeatum. 7(6). 8(7). tended beneath third (Figs. 7 and 8) .... 7 Second tarsal segments dilated and much wider than third, extending conspicuously beneath third (Figs. 5 and 6) (Forficulidae) POR dekh ete meth: Chenidaiiiitnck ink heb iho teh te 14 Large pad-like arolium between tarsal claws; hindwings visible; body length 14— 19 mm; male forceps of two types, both forms curved strongly inward (Figs. 22 Andy?) legates Pyragropsis buscki (Caudell) No arolium between tarsal claws; hindw- ings visible or not; body length variable; male forceps not strongly incurved (Figs. 14-18) Antennae 25-30 segmented; pronotum light brown with two dark longitudinal stripes; body length 20-30 mm; male py- gidium reduced and not visible in dorsal view (Fig. 24) .... Labidura riparia (Pallas) Antennae 10-16 segmented; pronotum unicolorous; body length less than 20 mm; 9(8). 11(10). male pygidium prominent (Figs. 14-18) (albindae) =F, Sais) eae ace, Shame eante 9 Tegminae pubescent; body length less than 35000 00 Uap et eR re Nn ae ERE re adm er 10 Tegminae glabrous; body length 8-18 TNLIVDS 2 oye EAS ee rise 12 Tegminae as long as pronotum; visible portion of hindwings only half length of pronotum; inner margin of female forceps notched basally (Fig: 28) ......:....... Tegminae much longer than pronotum, visible portion of hindwings as long as pronotum; inner margin of female forceps not notched basally (Fig. 27) Head and thorax black; abdomen reddish; male pygidium quadrate; inner margin of male forceps curved (Fig. 15) ae ae Labia curvicauda (Motschulsky) Body yellowish-brown to brown; male py- gidium elongated medially; inner margin 8 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON of male forceps straight (Fig. 14) Sorat AO ri ts eee Labia minor (L.) Male pygidium large, produced medially (Fig. 18); hindwings always present; an- tennae entirely brown; body length 12-16 Vostox brunneipennis (Audinet-Serville) - Male pygidium not produced medially (Figs. 16-17); hindwings often absent or concealed; basal 2-3 antennal segments yellow, remainder brown; body length 8- 2 inne See, et es eee oe 13 Pronotum broader than long (Fig. 4); male pygidium pentagonal; male forceps with- out inner basal tooth (Fig. 16) ee ee ne Marava arachidis (Y ersin) - Pronotum as broad as long (Fig. 3); male pygidium quadrate; male forceps with in- ner basal tooth (Fig. 17) .. Marava pulchella (Audinet-Serville) Fourth antennal segment less than twice as long as broad (Fig. 9); tegminae dark brown; male pygidium truncate; male for- ceps broadened, almost touching basally (Figs 25) eis. fo5. ot Forficula auricularia L. - Fourth antennal segment more than twice as long as broad (Fig. 10); tegminae yellow with brown inner margins; male pygi- dium spine-like; male forceps widely sep- arated at base (Figs. 19-21) ........... IS} Hindwings visible beyond tegminae ... SE ete ror Roe acre Doru taeniatum (Dohrn) Hindwings not visible beyond tegminae 16 Male forceps with tooth near apex (Fig. 20); male pygidium distinctly shorter than length of last abdominal segment ...... rea Pe Doru aculeatum (Scudder) - Male forceps without tooth near apex (Fig. 21); male pygidium as long as length of last abdominal segment ....... Doru davisi Rehn and Hebard 129): 13(12). 14(6). 15(14). 16(15). DERMAPTERA OF SOUTH CAROLINA Anisolabis maritima (Bonelli, 1832) Fig. 11 Forficula maritima Bonelli, 1832, in Géné, Ann. Sci. Regn. Lomb.-Venet. 2: 224. Anisolabis maritima (Bonelli) Fieber, 1853, Lotoss22578 Steinmann (1984) gives a more complete synonymy. Anisolabis maritima can be distinguished from the other wingless species occurring in South Carolina by the 20-24 segmented an- tennae, lack of dark encircling bands on the legs, and 20-25 mm body length. This species has been cited as A. maritima (Géné, 1832) in earlier papers. Anisolabis maritima is usually found underneath litter and drift- wood along seashores (Langston, 1974). This earwig forages at night and is predaceous; its prey includes crickets, sandfleas, and smaller earwigs (Bennett, 1904). This species is essentially cosmopolitan. Introduced into North America, A. mari- tima now occurs locally along the Pacific coast from British Columbia south to Cal- ifornia, and along the Atlantic and Gulf coasts from Maine to Florida and westward to Texas (Langston and Powell, 1975). South Carolina Records.—Anisolabis maritima probably occurs locally along the entire South Carolina coast. Beaufort Co.: Beaufort, 14 June 1930, coll. O. L. Cart- wright, | 2; Bluffton, 2 April 1933, coll. D. Dunavan, | 4, 2 2, 2 immatures. Horry Co.: 30 August 1981, coll. S. Mudge, | 6. Pickens Co.: Clemson, 23 June 1980, under board in barn, coll. C. Lay, 19. Euborellia annulipes (Lucas, 1847) The Ringlegged Earwig Figs. 12, 26 Forficesila annulipes Lucas, 1847, Ann. Soc. Entomol. Fr. 5: 84. Anisolabis annulicornis Blanchard, 1851, in Gay, Hist. Fisica Pol. Chile, Zool. 6: 10. Euborellia annulipes (Lucas) Burr, 1915, J. R. Microsc. Soc. 35: 545. Sakai (1970a) gives a more complete syn- onymy. The ringlegged earwig can be distin- guished from the only other wingless species in South Carolina by its 14-18 segmented antennae, dark encircling bands on the fem- ora and tibiae, and 12-18 mm body length. The dark brown antennae generally have the third, fourth, and sometimes fifth subapical segments pale yellow to white, although a few specimens show only one or no pale segments. VOLUME 89, NUMBER 1 Euborellia annulipes is a general scav- enger that is commonly found on the ground underneath rocks, logs, and other debris. This earwig can cause minor damage to plants and stored foods when it enters greenhouses, nurseries, and warehouses (Bharadwaj, 1966). However, it is also pre- daceous and will attack other insect pests. The ringlegged earwig is an introduced species that occurs worldwide. It has been established in North America for at least 140 years and now occurs virtually through- out the United States and into British Co- lumbia (Langston and Powell, 1975; Scud- der and Vickery, 1985). South Carolina Records. — The ringlegged earwig is the most commonly encountered species in the state. Aiken Co.: Windsor, 25 November 1933, coll. O. L. Cartwright, 1 2. Anderson Co.: Pendleton, 31 October 1975, in cotton trash, coll. P. Zinsmelster, 1 6, 1 9, 2 immatures. Barnwell Co.: Elko, 30 July 1981, debris under pig feeding trough, coll. J. R. Brushwein, | 64, 2 2, 1 immature. Beaufort Co.: Beaufort, 30 Oc- tober 1925, under trash, coll. F. Sherman, 1 immature. Charleston Co.: Charleston, 5 May 1951, coll. D. Dunavan, | 2. Cherokee Co.: Gaffney, 15 August 1939, coll. L. M. Sparks, | ¢. Clarendon Co.: Summerton, 31 March 1929, coll. O. L. Cartwright, 1 im- mature. Colleton Co.: Bear Island, 30 Sep- tember 1979, grass, coll. B. Hendrix, | 9°. Darlington Co.: Darlington, 19 June 1929, coll. J. M. Napier, 1 ¢. Dorchester Co.: Sum- merville, 15 August 1931, F. Sherman, | é. Edgefield Co.: Trenton, 19 November 1959, trunk of peach tree, coll. T. E. Skelton, 1 immature. Florence Co.: Florence, 17 July 1981, leaf litter, coll. R. D. Simpson, 2 °. Lee Co.: Meredith, 22 June 1926, coll. O. L. Cartwright, 1 3, 1 2. Oconee Co.: Fairplay, 18 October 1984, under feathers in chicken house, coll. W. Barton, 2 immatures. Orangeburg Co.: Orangeburg, 22 July 1927, coll. F. Sherman, | ¢. Pickens Co.: Clemson, 1 December 1983, beneath rock in garden, coll. J. Joly, 1 2, 2 immatures. Richland Co.: Columbia, 21 February 1926, O. Cart- wright, 1 ¢. Spartanburg Co.: Greer, 31 Oc- tober 1976, coll. M. McClimon, | 9. Labia minor (L., 1758) Figs. 14, 27 Forficula minor Linnaeus, 1758, Syst. Nat. GOs. 423 nor. Labia minor (L.) Stephens, 1835, Ill. Brit. Entomol., Mand. 6: 8. Labia minuta Scudder, 1862, Boston J. Nat. Hist. 7: 415. Sakai (1970b) gives a more complete syn- onymy. Labia minor is distinguished from all oth- er earwigs in South Carolina by both the abundance of golden hair covering most of its body and its small body size (less than 8 mm long). It resembles a small rove beetle (Staphylinidae) and as such is often put into unsorted collections of these beetles. Labia minor is both a scavenger and a predator and can be found under various kinds of debris. It flies readily and is frequently at- tracted to lights. Labia minor is found in temperate and subtropical zones worldwide and is another species that has been intro- duced into the Nearctic region. Like E. an- nulipes, it is now distributed throughout North America (Langston and Powell, 1975; Scudder and Vickery, 1985). South Carolina Records. — Labia minor is presently known only from the extreme northwestern part of the state. It may well occur in other parts of the state, being com- monly overlooked because of its small size. Anderson Co.: Pendleton, 28 July 1972, in cottonseed, coll. R. P. Griffin, 1 °. Pickens Co.: Clemson, 23 June 1956, in light trap, coll. D. Dunavan, | 6; Clemson, 19 Septem- ber 1985, in cattle feed at Clemson Univ. dairy barn, coll. K. M. Hoffman, | imma- ture; Clemson, 5 October 1984, flying in field, coll. J. Barron, 1 4; Clemson, 20 April 1939, at light, coll. F. T. Arnold, | ¢. Marava pulchella (Audinet-Serville, 1839) Figs: 3, 7.8. 17, 29 Forficula pulchella Audinet-Serville, 1839, Hist. Nat. Ins. Orthop. p. 42. 10 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Labia guttata Scudder, 1876b, Proc. Boston Soc. Nat. Hist. 18: 265. Labia burgessi Scudder, 1876b, Proc. Bos- ton Soc. Nat. Hist. 18: 266. Labia melancholica Scudder, 1876b, Proc. Boston Soc. Nat. Hist. 18: 267. Prolabia pulchella (Audinet-Serville) He- bard, 1917, Entomol. News 28: 319. Laprobia_ pulchella (Audinet-Serville) Hincks, 1960, Proc. R. Entomol. Soc. Lond. (B) 29: 156. Marava pulchella (Audinet-Serville) Brin- dle 197 ta. J. Nat. Hist 53.557. Marava pulchella is the only earwig in South Carolina that exhibits both fully winged and brachypterous adults. When the tegminae and hindwings are fully devel- oped, the hindwings are visible, somewhat lighter in color basally, and the compound eyes are large. Brachypterous individuals have shorter tegminae, hindwings that are either absent or concealed, and smaller compound eyes. Factors governing the rel- ative proportions of these morphs in a pop- ulation are not known for M. pulchella, but temperature has been shown to be influ- ential for a different species (Knabke and Grigarick, 1971). Each branch of the male forceps generally has two teeth located api- cally and basally on the inner margin, al- though in some specimens the apical tooth may be missing. This earwig can be found under the bark of dead trees and in debris. Fully winged adults can be attracted to lights. Marava pulchella is native to southeastern United States, being found from North Carolina south to Florida and westward to Texas. It has also been recorded from Cuba (Alayo and Hernandez, 1980). South Carolina Records.—Marava pul- chella probably occurs statewide. Anderson Co.: Pendleton, 28 July 1972, in cotton seed, coll. R. P. Griffin, 1 6. Florence Co.: Flor- ence, 29 March 1930, coll. O. L. Cartwright, 1 6, 1 immature. Hampton Co.: Yemassee, 5 January 1928, coll. J. A. Berly, 1 6; Ye- massee, 28 December 1941, coll. O. L. Cart- wright, 1 6. Lee Co.: Meredith, 19 April 1928, coll. O. L. Cartwright, 1 4. Pickens Co.: Clemson, 7 March 1935, coll. J. G. Watts, 1 immature; Clemson, 20 October 1972; hickory log, coll/sR: PAGriihinyales: Isaqueena Forest, 2 August 1984, UV light trap, coll. K. M. Hoffman, 1 ?; Clemson, 15 August 1986, porchlight, coll. K. M. Hoff- man, | é. Richland Co.: Columbia, 20 Feb- ruary 1926, coll. O. Cartwright, 1 imma- ture. Vostox brunneipennis (Audinet-Serville, 1839) Figs. 18, 30 Psalidophora brunneipennis Audinet-Ser- ville, 1839, Hist. Nat. Ins. Orthop. p. 30. Vostox brunneipennis (Audinet-Serville) Burr, 1911, Genera Insect. 122: 51. This earwig is a reddish-brown to dark brown species with yellow hindwings that are bordered on their apical and inner mar- gins with brown. The male forceps usually have a single tooth located on the inner mar- gin near the midpoint, although some spec- imens possess a second tooth distally (Fig. 18). This species is most commonly found under the bark of dead trees, but adults can also be attracted to lights. Vostox brunneipennis, a native species, is found from Panama northward to the southern United States. The holotype is from Pennsylvania, and in the United States this species is known from Illinois, Indiana, and Virginia south to Florida and westward to Texas and Oklahoma (Brindle, 1971b; Ar- nold and Drew, 1979). South Carolina Records.— Vostox brun- neipennis probably occurs statewide. An- derson Co.: Anderson, 11 March 1982, coll. G. Jones, | 6. Greenwood Co.: Greenwood, 5 February 1977, coll. P. Ellenburg, 1 2. Lee Co.: Meredith, 12 February 1927, coll. O. L. Cartwright, 1 6, 3 2. Pickens Co.: Clem- son, 9 January 1986, under bark of dead standing oak tree, coll. J. R. Brushwein, 1 VOLUME 89, NUMBER 1 6; Clemson, 1 August 1956, in trap light, coll. D. Dunavan, | 9; Six Mile, 17 October 1973, under log, coll. R. Peigler, 1 imma- ture. Saluda Co.: Ridge Spring, 25 July 1984, under bark of dead oak tree on ground, coll. D. E. Scotten, 2 immatures. Labidura riparia (Pallas, 1773) The Striped Earwig Fig. 24 Forficula riparia Pallas, 1773, Reise Russ. Reiches: 2.727. Forficula bidens Olivier, 1791, Encycl. Method. 6: 466. Forficula erythrocephala Fabricius, 1793, Entomol. Syst. 2: 4. Labidura riparia (Pallas) Dohrn, 1863, Stett. Entomol. Zeit. 24: 313. For a more complete synonymy, see Stein- mann (1978). The striped earwig is most easily recog- nized by both its large size and the two dark bands running the length of the pronotum. Also, there is a broad, dark, medio-dorsal band extending the length of the abdomen. This species 1s primarily predaceous, ac- tively seeking its prey at night (Schlinger et al., 1959). It can be collected on the ground at night or under debris during the day. Labidura riparia is an introduced species that occurs worldwide in both tropical and temperate zones. In the United States this earwig is established in the southern third of the country, from North Carolina south to Florida and westward to Texas, Arizona, and California (Langston and Powell, 1975). South Carolina Records.—The striped earwig is found throughout the state. Barn- well Co.: Blackville, 24 October 1975, Ed- isto Expt. Sta., pitfall traps from soybeans, coll. J. F. Price, 5 6, 5 °. Beaufort Co.: Sea- brook, 18 October 1980, coll. M. K. Disney, 1 immature. Charleston Co.: Charleston, 7 February 1934, coll. J. A. Berly, 1 6. Edge- field Co.: Trenton, 19 November 1959, trunk of peach tree, coll. T. E. Skelton, 1 ¢. Greenville Co.: Greenville, 30 September 11 1933, coll He iabPoe, In iv6s Horry Cox Myrtle Beach, 27 December 1955, in dwell- ing of Joe C. Ivey, coll. D. Dunavan, 1 4, 1 2. Pickens Co.: Clemson, 10 October 1961, under wood slat, coll. R. E. O’Brien, | im- mature; Clemson, 26 September 1984, on cement beneath lights at night, coll. K. M. Hoffman, | °. Sumter Co.: Sumter, Febru- ary 1955, coll. unknown, | immature. Doru aculeatum Scudder, 1876 Figs. 5, 6, 10, 20, 31 Forficula aculeata (Scudder, 1876a, Proc. Boston Soc. Nat. Hist. 18: 262. Doru aculeatum (Scudder) Rehn and He- bard, 1914, J. N.Y. Entomol. Soc. 22: 93. Doru aculeatum is 12-18 mm long, and can be distinguished from the closely related species D. taeniatum by the lack of visible hindwings. This earwig is generally found on grasses and sedges growing near water (Hebard, 1934; Cantrell, 1968). Doru acu- leatum is native to eastern North America, ranging from southern Michigan and On- tario to Florida and westward to Illinois, Nebraska, and Louisiana. South Carolina Records.—Doru aculea- tum has only been found in the extreme northwestern part of the state and at Myrtle Beach. Anderson Co.: Portman Shoals, 24 November 1927, coll. F. Sherman, 2 4, 1 2. Horry Co.: Myrtle Beach, 1 March 1965, sedge, coll. V. M. Kirk, 2 6. Oconee Co.: Seneca, 10 September 1969, on jewel weed, coll. J. W. Van Duyn, | 6. Pickens Co.: Clemson, 14 October 1961, in corn ear, coll. J. A. DuRant, 3 6, 1 2; Pickens, 28 March 1978, coll. J. Keith, 1 ¢; Rocky Bottom, 22 May 1934, coll. O. L. Cartwright, 1 4; Six Mile, 26 October 1963, under dead willow bark, coll. D. G. Bottrell, 1 4, 1 2. Doru taeniatum (Dohrn, 1862) Fig. 19 Forficula taeniata Dohrn, 1862, Stett. Ento- mol; Zeits23: 230. 12 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Forficula californica Dohrn, 1865, Stett. Entomol. Zeit. 25: 85. Forficula exilis Scudder, 1876a, Proc. Bos- ton Soc. Nat. Hist. 18: 262. Doru exile (Scudder) Burr, 1911, Genera In- Sects 1225579" Doru lineare (Eschscholtz) Burr, 1911, Gen- era Insect. 122: 79 (partim). Doru taeniatum (Dohrn) Brindle, 1971Ic, Papeis Avulsos Zool. 23: 191. Doru taeniatum is distinguished from the closely related D. aculeatum by the presence of visible hindwings. The species Doru lin- eare (Eschschlotz, 1822) is listed by Blatch- ley (1920), Gurney (1950), and Langston and Powell (1975) as occurring in the United States. However, a revision of the genus Doru has shown the range of this species to be limited to Brazil, Argentina, and Para- guay (Brindle, 1971c). Records of this species in North America are now referrable instead to D. taeniatum (Gurney, 1972). Doru taeniatum ranges from Bolivia and Colombia northward to the southeastern United States (Brindle, 1971c). It has been found as far north as Maryland and as far west as Texas, Arizona, and California. It is unclear whether it is native to North America or was introduced from Central and South America (Gurney, 1972). South Carolina Records.—Doru taenia- tum probably occurs locally statewide, but seems to be more prevalent in the southern and eastern counties. Barnwell Co.: Black- ville, 18 October 1978, on soybeans, coll. G. Sanders, | ¢. Berkeley Co.: St. Stephen, 17 September 1983, coll. T. Johnson, 1 ¢. Charleston Co.: Charleston, 17 March 1980, coll. M. Wallace, 1 ¢. Dorchester Co.: Four Holes Swamp, 13 August 1976, Mellards Lake, coll. J. Morse and J. Chapin, | 9. Hampton Co.: | October 1983, coll. D. Ed- wards, | 6. Horry Co.: Conway, 20 March 1976, woods, coll. T. Thompson, | °. Pick- ens Co.: Clemson, 16 March 1976, on ground, coll. C. Wilson, | 3; Clemson, 10 ©ctober 1976, coll) 1 Currin; } adult: Clemson, 18 September 1978, coll. C. Whit- mire, 1 9. Forficula auricularia L., 1758 The European Earwig Figs. 9, 25 Forficula auricularia Linnaeus, 1758, Syst. Nats (10) 12423) not: Sakai (1973) gives a more complete syn- onymy. The European earwig is 15-20 mm in length and can be recognized by the basally broadened forceps of the male (Fig. 25). It is the most economically destructive ear- wig, occasionally causing substantial dam- age to cereals, fruits, seedling vegetables, and flowers. Unlike most earwigs, the European earwig does not survive well in warm cli- mates, preferring instead cool, moist re- gions (Crumb et al., 1941). The European earwig occurs in temperate regions around the world and has become widely estab- lished in North America. It has been re- corded locally across southern Canada southward to North Carolina and westward to Arizona and California (Langston and Powell, 1975; Scudder and Vickery, 1985). South Carolina Records.—The only re- cord of this species in South Carolina is of a female taken in Charleston. Due to this species’ preference for cool climates and the fact that Charleston is a major seaport, it is possible that this specimen is an adventive and not a member of an established popu- lation. Further collecting is necessary to de- termine whether the European earwig is es- tablished in South Carolina. Charleston Co.: Charleston, 30 October 1983, coll. B. F. Ce- lekeiGo: ACKNOWLEDGMENTS Iam grateful to the following for allowing me to examine material: Lewis Deitz and Carol Parron (North Carolina State Uni- versity), Cecil Smith (University of Geor- gia), Lionel Stange (Florida State Collection of Arthropods) and David Nickle (Agricul- VOLUME 89, NUMBER 1 tural Research Service, USDA, National Museum of Natural History). Gratitude is also extended to Peter H. Adler and John C. Morse (Clemson University) for review- ing the manuscript. This is Technical Con- tribution No. 2593 of the South Carolina Agricultural Experiment Station, Clemson University. LITERATURE CITED Alayo, D. and L. R. Hernandez. 1980. Introduccion al estudio del orden Dermaptera. Informe cient.- tec. Acad. Cienc. Cuba, No. 132: 1-30. Arnold, D. C. and W. A. Drew. 1979. Earwigs (Der- maptera) of Oklahoma. Proc. Okla. Acad. Sci. 59: 115-116. Audinet-Serville, J. G. 1839. Histoire naturelle des insectes. Orthopteres. Paris. 777 pp. Bennett, C. B. 1904. Earwigs (Anisolabis maritima Bon.). Psyche 11: 47-53. Bharadwaj, R. K. 1966. Observations on the bio- nomics of Euborellia annulipes (Dermaptera: La- biduridae). Ann. Entomol. Soc. Am. 59: 441-450. Blanchard, E. 1851. Orden IV. Orthopteros, pp. 1- 84. In Gay, C., Historia fisica y politica de Chile, Zoologica. Vol. 6. Paris. 152 pp. Blatchley, W. S. 1920. Orthoptera of Northeastern America with special reference to the faunas of Indiana and Florida. Nature Publ. Co., Indianap- olis. 784 pp. Boeseman, M. 1954. The Dermaptera in the Mu- seums at Leiden and Amsterdam. Zool. Verh. Lei- den 21: 1-122. Bonelli, F.-A. 1832. In Géné, G., Saggio di una monographie delle Forficule indigene. Ann. Sci. Nat. Regn. Lomb.-Venet. 2: 215-228. Borelli, A. 1906. Forficole di Costa Rica. Boll. Musei Zool. Anat. Torino 21: 1-19. Brindle, A. 1971la. A revision of the Labiidae (Der- maptera) of the Neo-tropical and Nearctic regions II. Geracinae and Labiinae. J. Nat. Hist. 5: 155- 182. 1971b. A revision of the Labiidae (Dermap- tera) of the Neotropical and Nearctic regions III. Spongiphorinae. J. Nat. Hist. 5: 521-568. 1971c. A revision of the genus Doru Burr (Dermaptera, Forficulidae). Papeis Avulsos Zool., Sao Paulo 23: 173-196. 1971d. The Dermaptera of the Caribbean. Stud. Fauna Curacao other Caribbean Islands, 131, 38: 1-75. . 1981. The types of Dermaptera described by Fabricius. Entomol. Rec. J. Var. 93: 14-16. 13 Burr, M. 1911. Dermaptera. Genera Insectorum 122: 1-112. 1915. On the male genital armature of the Dermaptera. J. R. Microsc. Soc. 35: 521-546. Cantrell, I. J. 1968. An annotated list of the Der- maptera, Dictyoptera, Phasmatoptera, and Or- thoptera of Michigan. Mich. Entomol. |: 299-346. Caudell, A. N. 1905. Ona collection of Orthoptera from southern Arizona, with descriptions of new species. Proc. U.S. Natl. Mus. 28: 461-477. 1907. On some Forficulidae of the United States and West Indies. J. N.Y. Entomol. Soc. 15: 166-170. 1913. Notes on Nearctic orthopterous in- sects: |—Nonsaltatorial forms. Proc. U.S. Natl. Mus. 44: 595-614. Crumb, S. E., P. M. Eide, and A. E. Bonn. 1941. The European earwig. U.S.D.A. Tech. Bull. No. 766, 76 pp. Dohrn, H. 1862. Die Dermaptera von Mexico. Stett. Entomol. Zeit. 23: 225-232. 1863. Versuch einer Monographie der Der- mapteren. Stett. Entomol. Zeit. 24: 309-322. 1864a. Versuch einer Monographie der Der- mapteren. Stett. Entomol. Zeit. 25: 285-296. . 1864b. Versuch einer Monographie der Der- mapteren. Stett. Entomol. Zeit. 25: 417-429. 1865. Versuch einer Monographie der Der- mapteren. Stett. Entomol. Zeit. 26: 68-99. Eschscholtz, J. F. 1822. Entomographien. G. Reimer, Berlin. 128 pp. Fabricius, J.C. 1775. Systema entomologiae, sistens insectorum classes, ordines, genera, species, ad- jectis synonymis, locis, descriptionibus, observa- tionibus. Flensburgi and Lipsiae. 832 pp. 1793. 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Reise durch verschiedene Prov- inzen des Russischen Reiches. St. Petersburg. 2: 701-744. Popham, E. J. 1965a. The functional morphology of the reproductive organs of the common earwig (Forficula auricularia) and other Dermaptera with reference to the natural classification of the order. J. Zool. 146: 1-43. 1965b. A key to Dermapteran subfamilies. Entomologist (Lond.) 98: 126-136. Rehn, J. A. G. and M. Hebard. 1912. On the Or- thoptera found on the Florida Keys and in extreme southern Florida. I. Proc. Acad. Nat. Sci. Phila. 64: 235-276. 1914. United States and Mexican records of species of the genus Doru. J. N.Y. Entomol. Soc. 22: 89-96. Sakai, S. 1970a. Dermapterorum catalogus praeli- minaris |. A basic survey for numerical taxonomy of the Carcinophoridae, Dermaptera of the world. Spec. Bull. Daito Bunka Univ. No. |: 1-91. 1970b. Dermapterorum catalogus praelimi- naris 2. A basic survey for numerical taxonomy of the Labiidae, the Dermaptera of the world. Spec. Bull. Daito Bunka Univ. No. 2: 1-177. 1973. Dermapterorum catalogus praelimi- naris 7. A basic survey for numerical taxonomy of the Forficulidae, the Dermaptera of the world. Spec. Bull. Daito Bunka Univ. No. 6: 1-357. Schlinger, E. I., R. V. Bosch, and E. J. Dietrick. 1959. Biological notes on the predaceous earwig Labi- dura riparia Pallas, a recent immigrant to Cali- fornia (Dermaptera, Labiduridae). J. Econ. Ento- mol. 52: 247-249. Scudder, G. G. E.and V. R. Vickery. 1985. A tabular check-list of Canadian orthopteroid insects. Notes Lyman Entomol. Mus., No. 13, 20 pp. Scudder, S. H. 1862. Materials for a monograph of the North American Orthoptera, including a cat- alogue of the known New England species. Boston J. Nat. Hist. 7: 409-480. . 1876a. Acentury of Orthoptera. Decade VI.— Forficulariae. Proc. Boston Soc. Nat. Hist. 18: 257- 264. . 1876b. Description of three species of Labia from the southern United States. Proc. Boston Soc. Nat. Hist. 18: 265-268. Steinmann, H. 1978. A systematic survey of the species belonging in the genus Labidura Leach, 1815 (Dermaptera). Acta Zool. Acad. Sci. Hung. 25: 415-423. 1981. A study of the circumtropical Der- maptera material in the “Instituut voor Taxon- omische Zoologie,”” Amsterdam. Acta Zool. Acad. Sci. Hung. 27: 187-210. 1984. Synopsis of Dermaptera of the world: 4. Family Carcinophoridae. Entomol. ABH (Dres.) 48: 63-82. Stephens, J. F. 1835. Illustrations of British Ento- mology. Vol. 6, Mandibulata. Baldwin and Cra- dock, London. 240 pp. Townes, H. K. 1946. Minutes of the 558th regular meeting of the Entomological Society of Wash- ington, Nov. 1, 1945. Proc. Entomol. Soc. Wash. 48: 47. Yersin, A. 1860. Note sur quelques orthoptéres nou- veaux ou peu connus d’Europe. Ann. Soc. Ento- mol. Fr. 8: 509-537. PROC. ENTOMOL. SOC. WASH. 89(1), 1987, pp. 15-23 OAXACACORIS, A NEW PLANT BUG GENUS AND THREE NEW SPECIES OF ORTHOTYLINI FROM MEXICO (HETEROPTERA: MIRIDAE) MICHAEL D. SCHWARTZ AND GARY M. STONEDAHL Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024. Abstract.— The new genus Oaxacacoris is diagnosed and described, including three new species: cygnus, pueblensis, schaffneri. All species occur in southcentral Mexico. The male genitalia of all species are illustrated, and a dorsal habitus provided for the male of cygnus. Scanning electron micrographs of the scale-like setae and pretarsus of schaffneri also are included. The idea for this paper originated during our revision of the genus Pseudopsallus (Stonedahl and Schwartz, 1986). Among the material on loan to us from Joseph C. Schaffner, Texas A&M University, College Station were specimens with a resemblance to Pseudopsallus, but which we now place in the new genus Oaxacacoris. The rela- tionship of Oaxacacoris to other orthotyline genera is not well understood. This is due to the limited knowledge of Mexican Miri- dae in general, as well as the poorly defined limits of many New World genera of Or- thotylini. In our revision of the western North American genus Pseudopsallus (Stonedahl and Schwartz, 1986), we selected Oaxaca- coris as the sister group of the former genus because these two genera shared derived characters of the male genitalia that have yet to be discovered in other genera of the tribe. Further, we suggested that Oaxaca- coris and Pseudopsallus belonged to a com- plex of genera united by similarities in type and fine structure of setae and by the general form of the male genitalia. Oaxacacoris can be distinguished from other genera of this group by the characters given in the generic diagnosis. The only known host association for the genus is Mimosa rhododactyla B. L. Robinson (Fabaceae). All holotypes and some paratypes are de- posited in the collection of the American Museum of Natural History, New York (AMNH). The remaining paratypes and ad- ditional specimens are deposited in the Tex- as A&M University collection (TAM). All measurements are in millimeters. Oaxacacoris Schwartz and Stonedahl, NEw GENUS Diagnosis.— Recognized by the green or yellow green, slightly shining general ap- pearance; dorsal vestiture with silvery and golden brown to black, narrow, scale-like setae with converging ridges (Fig. 2a); and male genitalia with the following character- istics: anterodorsal margin of aperture of genital capsule with three tergal processes, one right lateral and two left lateral of mid- line (Figs. 3, 10, 17); lateralmost pair of processes very large, slightly flattened, ex- tending well beyond margins of aperture; left medial tergal process small, simple, ori- ented ventrad into aperture; dorsal lobe of left paramere greatly expanded distally, with 16 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 4-9 strong spines (Figs. 6, 13, 19); right par- amere subquadrate in lateral view, with prominent, sometimes bifurcate, basal lobe (Figs. 8, 15, 22); and right spicula of vesica bisected to base, dorsal portion much small- er than ventral portion (Figs. 9, 16, 23). Description. — Male. Macropterous. Small to moderate subovate body form; length 3.50-—4.90 (apex of tylus to apices of hemel- ytra); width (across humeral angles of pronotum) 1.15-1.40; surface texture smooth; coloration dark to light green, vari- ably suffused or mottled with yellow, with frons, anterior portion of pronotum, em- bolium, scutellum, and cuneus sometimes extensively yellow; dorsal vestiture with re- cumbent, narrow, golden scale-like setae and long, suberect, golden simple setae, some- times also with recumbent, narrow, black scale-like setae and short to long, suberect, black simple setae. Head: Subtriangular; length of portion of head anterior of eye subequal to length of head posterior of an- terior margin of eye in dorsal or lateral view; posterior margin without carina; frons and vertex weakly convex, gently sloping anteri- ad, meeting tylus with weak suture; tem- poral area obsolete; eyes large, subovate, projecting slightly beyond anterolateral an- gles of pronotum in dorsal view, occupying six-sevenths of head height in lateral view, posterior margin sublinear, anterior margin slightly emarginate; antennal fossae small, contiguous with anterior margin of eye, ven- tral margin of fossae slightly to obviously dorsad of ventral margin of eye in lateral view; tylus smoothly curving to apex; juga triangular, flattened to slightly convex; lora rectangular, slightly swollen; gena present, width equal to diameter of antennal seg- ment I; gula small; length of buccular flange and cavity equal, not reaching posterior margin of head; labium reaching apices of metacoxae or sometimes to sixth sternite; antennal segment I with length equal to or slightly greater than width of vertex, linear, tapered basally, with a few erect bristle-like setae on distal and interior surface; seg- ments II-IV cylindrical, linear, diameter smaller than segment one, with densely dis- tributed, reclining, short, black simple setae. Pronotum: Trapezoidal, broader than long, slightly sloping transversely and longitudi- nally; lateral margins straight, smoothly curved at junction with propleura; posterior margin straight; anterior and posterior an- gles broadly rounded; anterior margin slightly arcuate medially; calli weakly con- vex, reaching anterior angles and lateral margin of pronotum, confluent anterome- dially, bordered posteriorly by faint to shal- low depression; mesoscutum narrowly ex- posed; scutellum weakly convex or flattened. Hemelytra: Subparallel-sided, widest me- dially; claval vein weakly elevated; radial vein elevated basally, obsolete distally; cu- neal incisure small; cuneal fracture angled slightly anteriad; cuneus longer than broad; membrane lightly to strongly suffused with uniform fuscous, light species with short spot or band of dark fuscous or black near in- tervein of inner cell; inner cell longer than cuneus, slightly narrowed distally; outer cell triangular. Legs: Coloration uniformly yel- low, or mixed yellow and yellow green with variable number of green spots; femora slightly flattened, tapered distally and ba- sally, with short, reclining, black or fuscous simple setae, sometimes with setae restrict- ed to anterior portion; meso- and meta- tibiae with several rows of minute, dark spinulae; tibiae with suberect, light, simple setae; tibial spines dark or light fuscous; api- ces of tibiae dark fuscous; apices of third tarsal segment black; tarsal segment one half as long as segment two and three; claws strongly curved, pulvilli connate to interior surface (at angle) of claw (Fig. 2b). Genitalia: Genital capsule: Large, subrectangular in ventral view, width greater than length, anterodorsal margin with three large var- iously shaped, posteriorly directed sclero- tized tergal processes; processes lateral of midline of capsule, and not supporting rec- tal opening; left processes consisting of sin- gle large, basally spinose process, originat- VOLUME 89, NUMBER 1 17 Fig. 1. Oaxacacoris cygnus, dorsal habitus of male. ing on posterior edge of anterodorsal margin (left tergal process, LP) and single small pro- cess originating on ventral surface of antero- dorsal margin, oriented ventrad into aper- ture (left ventral tergal process, LVP); right process solitary, large, usually orientated to left side of aperture (right tergal process, RP); posteroventral margin of capsule with deep paramere and phallotheca sockets, and without posteriorly directed processes; ap- 18 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 2. erture large, subcircular. Left paramere: Dorsal lobe (DLL) very large, flattened, strongly spinose, and usually expanded dis- tally; lateral lobe (LAL) narrow, rounded, and curved or sometimes notched distally. Right paramere: Subquadrate in lateral view; interior surface with prominent medial lobe (MIL), sometimes obsolete; dorsal lobe (DLR) usually spinose marginally and pro- jecting dorsad of medial lobe; basal lobe (BLR) solitary or strongly bifurcate, strong- ly spinose. Phallotheca: laterally com- pressed, open on dorsal and distal surfaces, opening narrow and linear dorsally, merg- ing with larger irregularly shaped distal por- tion; without secondary ventrobasal open- ing. Vesica: Ductus seminis simple, cylindrical, flexible with ribs; secondary gonopore horseshoe-shaped, base short and weakly sclerotized; two long posteriorly di- rected vesical spiculae narrowly attached right and left of midline of ductus basad of secondary gonopore, and practically sur- rounding ductus; right spicula bifurcate; dorsal portion (DS) short, simple or weakly serrate; ventral portion (RVS) with thick basal half, tapering to long, flattened or rounded, strongly and marginally serrate distal half; left spicula (LVS) with long, sometimes sinuate, rounded shaft, distal third recurved, flattened and strongly ser- rate marginally, base of recurved portion with fine serrations. Oaxacacoris schaffneri. 2a, Scale-like setae on dorsum. 2b, Pretarsal structures. Female. Macropterous. Similar to male in color, vestiture, and structure, except usually somewhat larger with slightly broader head and vertex. Genitalia: Follow- ing the terminology of Slater (1950). Scler- otized rings: Large, elongate, widely sepa- rated; lateral margin of ring moderately folded dorsally, not extending mesad. Pos- terior wall: K structures of moderate size, heart-shaped with bifid apex, well separated medially; L structure and portion of J struc- ture visible in anterior view. Etymology.— Named for the state of Oa- xaca in Mexico, the type locality of the type species; masculine. Type species. — Oaxacacoris cygnus, new species. Distribution. — Guerrero, Michoacan, Oaxaca, and Puebla, Mexico (Fig. 24). Discussion. —Identification of the species of Oaxacacoris is most easily accomplished by careful examination of the male genitalic structures. To faciliate identification we provide illustrations and descriptions of all the structures for each species. Oaxacacoris cygnus, NEW SPECIES Figs. 1, 3-9, 24 Diagnosis. — Recognized by the large size; labium reaching apices of metacoxae or slightly beyond; dorsal vestiture always with densely distributed, dark brown or black scale-like setae; and by the structure of the VOLUME 89, NUMBER 1 male genitalia, especially the shape of the tergal processes (Figs. 3, 4), the swan-shaped left paramere with 6-9 strong spines on dis- tal portion of dorsal lobe (Fig. 6), and basal lobe of right paramere bifurcate, with very large, laterally directed basalmost spine (Figs. 7, 8). Description. — Male (Fig. 1). Length 4.15- 5.20; mottled dark green general coloration; dorsal vestiture with recumbent, narrow, black scale-like setae and narrow, golden simple setae. Head: Width across eyes 0.85- 0.86, width of vertex 0.37-0.39; dark green, tinged with extensive yellow green or yellow areas overall; antennae yellow or yellow brown, segment IV and distal end of seg- ment III fuscous; length of segment I 0.35- 0.45, segment IT 1.51—1.65; labium reaching apices of metacoxae or slightly beyond. Pronotum: Posterior width 1.40-1.59: whit- ish green anterior of calli, calli and disk dark green, disk narrowly adjoining posterior margin of calli yellow, posterior angles faint green; mesoscutum yellow laterally and me- dially; scutellum with yellow apex. Hemel- ytra: Dark mottled green; cuneus, emboli- um, clavus and corium bordering embolium, and apex of clavus sometimes pale yellow or yellow green; membrane lightly suffused with fuscous and with band of dark fuscous bordering apex of inner cell; veins pale. Legs: Yellow, metafemora and sub-basal portion of metatibiae tinged with small green spots; femora with short, reclining black simple setae. Genitalia: RP very long, flattened, parallel-sided, distal fourth tapered and with several strong spines, or with only single large secondary serrate spine, apex pointed and directed to left; LP long, with bifurcate basal spine, or with simple basal spine, and recurved pointed apex; LVP cylindrical, sinuate with bifurcate pointed apex. Left paramere: DLL palmate with six to nine spines; LAL flattened, sinuate with pointed apex. Right paramere: MIL simple; DLR somewhat pointed distally with several small spines; BLR bifurcate, with long simple slightly recurved basalmost spine, distal- 19 most spine solitary, or weakly to deeply bi- furcate. Phallotheca: dorsodistal edge re- curved. Vesica: DS faintly serrate, shorter than RVS; RVS flattened and tapered, with strongly serrate margin, apex pointed; LVS with long, tapering recurved distal third, apex pointed. Female. Length 4.35-5.05; width across eyes 0.78-0.85, width of vertex 0.40-0.43: length of antennal segment I 0.43-0.44, seg- ment II 1.50-1.64; posterior width of pronotum 1.50-1.64. Etymology.—From the latin cygnus (swan), referring to the swan-like shape of the left paramere in lateral view. Distribution. — Figure 24. Holotype ¢6.—MEXICO: Oaxaca. 4 mi NE of Miltepec, July 21 1984, Carroll, Schaffner, Friedlander. Paratypes.— MEXICO: Oaxaca. 6 4, 5 2 same data as holotype. Additional specimens.—MEXICO: Guerrero. 13 8, 15 2 6 mi E of Xochipala, July 13, 1985, Jones, Schaffner. Puebla: 4, 24 mi SW of Acatepec, July 21, 1984, Car- roll, Schaffner, Friedlander; 4, 3 2 4.4 mi SW of Acatepec, July 9, 1977, Schaffner. Oaxacacoris schaffneri, New SPECIES Figs. 2a, 2b, 10-16, 24 Diagnosis.—Similar to cygnus, but dif- ferentiated by its smaller size; labium reach- ing well beyond apices of metacoxae, some- times to sixth abdominal sternite; dorsal vestiture without, or with only a few dark brown, scale-like setae; and by the structure of the male genitalia, especially the shape of the tergal processes (Figs. 10, 11) and dorsal lobe of left paramere with only four strong spines distally (Fig. 13). Description.— Male. Length 3.58—-4.05; mottled green general coloration; dorsal vestiture with recumbent, narrow golden or brown, scale-like setae and narrow, golden simple setae. Head: Width across eyes 0.72- 0.78, width of vertex 0.33-0.34; yellow or yellow green; antennae yellow or yellow brown, segments III and IV slightly fuscous; 20 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 3-16. Male genitalic structures. 3-9, Oaxacacoris cygnus. 3, 4, Tergal processes of genital capsule. 3, Dorsal view (LP, left process; LVP, left ventral process; RP, right process). 4, Posterior view. 5, 6, Left paramere. VOLUME 89, NUMBER 1 length of segment I 0.34—-0.35, segment II 1.19-1.26; labium surpassing apices of metacoxae, sometimes reaching sixth ster- nite. Pronotum: Posterior width 1.16-1.20; yellow or yellow green anteriad of posterior depression of calli, disk mottled green and yellow or sometimes entirely yellow green with margin of disk faint green; mesoscu- tum orange yellow; scutellum yellow with faint orange cast; area adjoining mesoscu- tum mostly orange. Hemelytra: Mottled green and yellow, or sometimes mostly mottled yellow; cuneus and embolium yel- low; membrane very lightly suffused with fuscous, more so peripherally, sometimes with small testaceous band near apex of in- ner cell; veins testaceous. Legs: Yellow; tarsi and apices of tibiae brownish yellow; fem- ora with short, reclining, yellow simple se- tae. Genitalia: RP long, flattened, parallel- sided, distal portion tapered and weakly serrate marginally, with apex pointed and directed to left; LP long, thick, with several large serrations basally, tapered and narrow distally, apex pointed; LVP cylindrical and simple. Left paramere: DLL palmate with four spines; LAL somewhat flattened, sin- uate with pointed apex. Right paramere: MIL simple; DLR with several small spines distally; BLR deeply bifurcate, with long simple basalmost spine, distalmost spine short with bifid apex. Vesica: DS serrate, shorter than RVS; RVS wide and flattened, with strongly serrate margin, apex truncate and serrate; LVS long, broad recurved distal third, medially tapering to narrow apex. Female. Length 3.81-—4.25; width across eyes 0.76—-0.80, width of vertex 0.38-0.40; length of antennal segment I 0.35-0.40, seg- — 21 s ment II 1.37-1.55; posterior width of pronotum 1.27-1.31. Etymology.—Named for Joseph C. Schaffner, who graciously provided all spec- imens for this study. Distribution. — Figure 24. Holotype 6.—MEXICO: Michoacan. 30 mi S of Nueva Italia, August 8, 1978, Plitt, Schaffner. Paratypes.— MEXICO: Michoacan. 5 6, 12 2 same data as holotype. Additional specimens.—MEXICO. Mi- choacan. 20 8, 30 2 28.5 mi S of Nueva Italia, July 9, 1985, Jones, Schaffner, ex Mi- mosa rhododactyla B. L. Robinson (Faba- ceae). Oaxacacoris pueblensis, NEw SPECIES Figs. 17-24 Diagnosis.—Readily distinguished from cygnus and schaffneri by the following char- acteristics of the male genitalia: left tergal process bulbous with numerous, tiny ser- rations (Figs. 17, 18); lateral lobe of left par- amere notched apically (Fig. 20); basal lobe of right paramere solitary, with large sec- ondary spines (Figs. 21, 22); and dorsal spic- ula of vesica small and nonserrate (Fig. 23). The scale-like setae of the dorsal vestiture are silvery to golden brown, never dark brown or black as in cygnus and sometimes schaffneri. Further distinguished from schaffneri by the shorter labium. Description.— Male. Length 3.43; mot- tled green and yellow general coloration; dorsal vestiture with recumbent, long, nar- row, golden scale-like setae and long, sub- erect, golden to nearly black simple setae; metacuneus, clavus and corium bordering 5, Posterior view (DLL, dorsal lobe; LAL, lateral lobe). 6, Lateral view (DLL, LAL; as in Fig. 5). 7, 8, Right paramere. 7, Dorsal view (BLR, basal lobe; DLR, dorsal lobe; MIL, medial interior lobe). 8, Inside lateral view (BLR, DLR, MIL; as in Fig. 7). 9, Spiculae of vesica (DS, dorsal; LVS, left ventral; RVS, right ventral). 10-16, Oaxacacoris schaffneri. 10, 11, Tergal processes of genital capsule. 10, Dorsal view. 11, Posterior view. 12, 13, Left paramere. 12, Posterior view. 13, Lateral view. 14, 15, Right paramere. 14, Dorsal view. 15, Inside lateral view. 16, Spiculae of vesica. 22 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON i9 Figs. 17-23. Male genitalic structures of Oaxacacoris pueblensis. 17, 18, Tergal processes of genital capsule. 17, Dorsal view. 18, Posterior view. 19, 20, Left paramere. 19, Lateral view. 20, Posterior view. 21, 22, Right paramere. 21, Dorsal view. 22, Inside lateral view. 23, Spiculae of vesica. claval suture sometimes with darker brown scale-like setae. Head: Width across eyes 0.73, width of vertex 0.34; yellow; antennae testaceous, segment IV and apex of segment III fuscous; length of segment I 0.38, seg- ment II 1.13; labium reaching apices of metacoxae. Pronotum: Posterior width 1.16; yellow anteriad of posterior depression of calli, calli somewhat darker, disk mottled green and yellow, peripheral border faint green; mesoscutum and scutellum orange yellow. Hemelytra: Mottled green and yel- low, interior of cuneus yellow; membrane darkly suffused with fuscous; veins fuscous, rufous adjacent to cuneus. Legs: Yellow; tarsi and apices of tibiae lightly fuscous, with long, suberect, yellow simple setae. Geni- talia: RP long, thick basally, narrowed me- dially, apex truncate with single large spine directed to left; LP bulbous, surface densely serrate, with single apical and basal spines; LVP cylindrical, short with pointed apex. Left paramere: DLL long, broad basally, ta- pering to pointed apex, proximal surface with five spines; LAL fairly broad, with notched apex. Right paramere: MIL obso- lete; DLR small, with several small spines on interiobasal surface; BLR large and sol- itary, with large spines, apex extending dis- tad of distal end of DLR, and with large basal spines. Vesica: DS very small, simple; RVS narrow with strong marginal serra- tions, much smaller than LVS; LVS with curved portion strongly hooked and mar- ginally serrate. Female. Length 4.01—4.32; width across VOLUME 89, NUMBER 1 ( Ss re Piel IN f = SS w Tt a ~ Nr \ Ly See aa { SIN we ( ay ca \. \ N a \ NS: \ EN J j vy NPS " \ Ne ( = ae Sta SS \ if iy \ Caray A eA A ( 4 es \ ae > a , } q 7, ‘ a: Sool a stake il 3 es ys > Be re ° Jo 4 fe Sy 7 4 Fa ee | ayy oe Le ee Fig. 24. Distribution of Oaxacacoris cygnus ®, Oaxacacoris schaffneri &, Oaxacacoris pueblensis @. eyes 0.76-0.77, width of vertex 0.39-0.40; length of antennal segment I 0.44—0.46, seg- ment II 1.40-1.65; posterior width of pronotum 1.31-1.38. Etymology.— Named for its occurrence in the state of Puebla, Mexico. Distribution. — Figure 24. Holotype ¢.—MEXICO: Puebla. 16 mi NW of Acatlan, July 14, 1974, Clark, Mur- ray, Ashe, Schaffner. Paratypes.—MEXICO: Puebla. 6, 15 2 same data as holotype. Additional specimens.—MEXICO: Pu- ebla: 42 5 mi SE of Izucar de Matamoros, July 20, 1974, Carroll, Schaffner, Friedlan- der; 2 7.3 mi SW Izucar de Matamoros, July 23 22, 1981, Bogar, Schaffner, Friedlander. In- dividuals of these populations are slightly greater in length (4.14—4.68) and darker than the holotype and paratypes but are indis- tinguishable from the later specimens with regard to external morphology and vesti- ture. We are tentatively recognizing them as conspecific with the type, pending the acquisition of male specimens from other areas for detailed comparison. All three lo- calities for this species are within a 35 mile area. ACKNOWLEDGMENTS We thank Kathleen Schmidt, Hillsdale, New York for the fine illustration of the adult male of Oaxacacoris cygnus, Lauren Duffy, Interdepartmental Laboratory, American Museum of Natural History, who assisted with the preparation of the scanning electron micrographs, and an anonymous reviewer for improving the manuscript. The authors contributed equally to all fac- ets of this paper. LITERATURE CITED Slater, J. A. 1950. An investigation of the female genitalia as taxonomic characters in the Miridae. Iowa St. J. Sci. 25: 1-81. Stonedahl, G. M. and M. D. Schwartz. 1986. Revi- sion of the plant bug genus Pseudopsallus Van Du- zee (Heteroptera: Miridae). Am. Mus. Novit. No. 2842, 58 pp. PROC. ENTOMOL. SOC. WASH. 89(1), 1987, pp. 24-30 ADDITIONS, TAXONOMIC CORRECTIONS, AND FAUNAL AFFINITIES OF THE STONEFLIES (PLECOPTERA) OF VIRGINIA, USA Boris C. KONDRATIEFF AND RALPH F. KIRCHNER! (BCK) Colorado State University, Department of Entomology, Fort Collins, Colorado 80523; (RFK) U.S. Army Corps of Engineers, Huntington District, Water Quality Section, 502 8th Street, Huntington, West Virginia 25701. Abstract.—One hundred and forty-nine species of stoneflies are recorded from Virginia. Of these, 80 species have distributions associated with the Appalachian Mountains, 57 are widely distributed east of the Rocky Mountains, 6 are associated with the Coastal Plain Physiographic Province, and 6 have ranges which include both the Appalachians and the Coastal Plain. There are apparently 7 regional endemic species. Twenty-one species are recorded from Virginia for the first time. Kondratieff and Voshell (1979) reported 116 species of Plecoptera known from Vir- ginia in their checklist of the stoneflies of Virginia. Since 1979, 21 state records have been added and seven new species described from Virginia. There also have been several recent important taxonomic changes: Pel- toperlidae (Stark and Stewart, 1981), Ptero- narcyidae (Stark and Szczytko, 1982), Per- lodidae (Stewart and Stark, 1984; Szczytko and Stewart, 1981), Chloroperlidae (Sur- dick, 1985), and Perlidae (Zwick, 1984). Three genera, Jsoperla by Szczytko, Sweltsa by Surdick, and Perlesta by Stark, are being revised and will add additional taxa to Vir- ginia. This species list is presented to provide a framework for future studies of the Plecop- tera of Virginia, including an identification manual. The 149 species reported herein from Virginia are the most recorded from any state or province from the Nearctic re- ' The views of the Author do not purport to reflect the position of the Department of the Army or the Department of Defense. gion. The following numbers of species have been reported from surrounding states: Kentucky—77 (Tarter et al., 1984), North Carolina—about 130 (Unzicker and Mc- Caskill, 1982), and West Virginia— 106 (Tarter and Kirchner, 1980). The richness of the stonefly fauna from Virginia is indicative of the range of topog- raphy which allows for heterogeneously di- verse habitats. Five major physiographic provinces of eastern North America (Coast- al Plain, Piedmont Plateau, Blue Ridge, Ridge and Valley, and the Appalachian Pla- teau) are represented in Virginia (Hoffman, 1969). Fig. 1 delimits the physiographic provinces of Virginia and the major river systems. Virginia has eight major river ba- sins, several of old geographical systems such as the New River (Hoffman, 1969). The to- pographic relief of Virginia ranges from sea level to 1743 m (Mt. Rogers, 5720 ft.). The distribution of Plecoptera of Virginia has three basic patterns. Eighty species are closely linked with the central Appalachians and foothills of the upper Piedmont Plateau region. This area has one of the most diverse VOLUME 89, NUMBER 1 RPA c a PA YS TA EA AFG \ ’ LG, Ark PEEDEE enn, POWELL CLINCH _NFK S.FK. Tenn Nw HOLSTON HOLSTON Fig. 1. 7 PH on Meal DISMAL swaMP N.C Physiographic provinces of Virginia (according to Hoffman, 1969). CP = Coastal Plain; PP = Piedmont Plateau; BR = Blue Ridge; RV = Ridge and Valley; AP = Appalachian Plateau. The symbol * identifies Mt. Rogers. fauna and flora assemblages of the North Temperate Zone. Immatures of these species usually occur in cooler streams or springs of the Appalachian Plateau, Ridge and Val- ley, Blue Ridge, and Piedmont Plateau provinces. Some of these species are pre- dominantly southeastern Appalachian, with southern Virginia being their northern limit of distribution. Included in this group are Allocapnia fumosa Ross, A. stannardi Ross, Alloperla nanina Banks, Megaleuctra wil- liamsae Hanson, Strophopteryx limata (Fri- son), Sweltsa mediana (Banks), S. urticae (Ricker), and Yugus arinus (Frison). The northern border of the southern Appala- chians is generally considered to be where the Roanoke River slices through the Blue Ridge Mountains (Fig. 1). Most the the species listed above may be collected from streams draining Virginia’s highest peaks, Mt. Rogers-White Top area in Grayson, Smyth, and Washington counties (Fig. 1). Several Appalachian species have strong northeastern affinities, and Virginia in- cludes their southern distribution limits. These species may have survived the ice ages in the extensive unglaciated areas of Canada. These likely northern emigrants in- clude: Allocapnia maria Hanson (most southern record, Smyth County), Alloperla banksi Frison, all four of the eastern species of Ostrocerca, and Sweltsa naica (Pro- vancher). Many species, however, are widespread, occurring all along the Appalachian Moun- tains from Canada to Georgia: Acroneuria carolinensis (Banks), Bolotoperla rossi (Fri- son), Hansonoperla applachia Nelson, Ma- lirekus hastatus (Banks), and Sweltsa onkos (Ricker). About 57 species may be considered to have widespread boreal distributions. These generally occur east of the Rocky Mountain region. In Virginia, these species are found in all five physiographic provinces. Good examples are Acroneuria abnormis (New- man), Allocapnia granulata (Claassen), A. recta Claassen, A. rickeri Frison, Alloperla caudata Frison, Amphinemura nigritta (Provancher), Attaneuria ruralis (Hagen), Clioperla clio (Newman), Haploperla brevis (Banks), Isoperla bilineata (Say), Leuctra ferruginea (Walker), Perlinella drymo (Newman), P. ephyre (Newman), Prostoia 26 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON completa (Walker), P. similis (Hagen), Neo- perla clymene (Newman), and Strophopte- ryx fasciata (Burmeister). Six species are generally restricted to the Coastal Plain physiographic province, and as Stark (1979) stated, occupy “‘a boomer- ang shaped range that extends northward along the Atlantic Coast and to the west along the Gulf Coast.” Acroneuria arenosa (Banks), Neoperla carlsoni Stark and Bau- mann, and Taeniopteryx lonicera Ricker and Ross are examples of this group. Nymphs of these species are most commonly col- lected on submerged woody substrate and among accumulated organic debris in the many shifting sand streams of the area. Adults of the perlids are often only collected in numbers using light traps. Several species, including Allocapnia virginiana Frison, A. wrayi Ross, Eccoptura xanthenes (New- man), Helopicus subvarians (Banks), and Paragnetina fumosa (Banks) range through- out the Coastal Plain but also occur in the Appalachians. There are apparently several regional en- demic species whose affinities are clearly with the Appalachian fauna. These include Diploperla morgani Kondratieff and Vosh- ell, D. kanawholensis Kirchner and Kon- dratieff, Al/operla biserrata Nelson and Kondratieff, Taeniopteryx nelsoni Kondra- tieff and Kirchner, Allocapnia harperi Kirchner, and Tallaperla lobata Stark. Significant range extensions are noted for several species. /sogenoides varians (Walsh) was collected from Bedford County (Big Ot- ter River). This species is known from II- linois, Indiana, Michigan, Mississippi, South Carolina, and Tennessee (Stewart and Stark, 1984). The Big Otter River has a diverse stonefly fauna. Diploperla kanawholensis was also collected from this river and rep- resents a new state record (Kirchner and Kondratieff, 1984). The eastern species of Ostrocerca have been considered rare. How- ever, the four eastern species often occur abundantly but locally in the Appalachian region of Virginia. In the list that follows, the 21 new state records are indicated by # and the seven species described since 1979 by +. In ad- dition, 21 other species that probably occur in Virginia are indicated by *, and the ad- jacent states where they have been reported are listed in parentheses. Each species in the following list is identified by its continental distribution: widespread boreal species (WB), Appalachian (AP), and Coastal Plain (CP), and Appalachian—Coastal Plain dis- tribution (AP-CP). We are following the classification of Zwick (1973). The follow- ing papers are necessary to supplement those listed by Kondratieff and Voshell (1979) to identify the stoneflies of Virginia and sur- rounding states: Harper and Stewart (1984), Kirchner (1980, 1982), Kirchner and Har- per (1983), Kirchner and Kondratieff (1984, 1985), Kondratieff and Kirchner (1982a, 1982b), Kondratieff and Voshell (1981, 1982), Kondratieff et al. (1981), Nelson (1979, 1982), Nelson and Kondratieff (1983), Stark (1983, 1985, 1986), Stark and Ray (1983), Stark and Stewart (1981), Stark and Szczytko (1981), Stewart and Stark (1984), Surdick (1985), and Szczytko and Stewart (1976). List OF STONEFLIES OF VIRGINIA ORDER PLECOPTERA SUBORDER ARCTOPERLARIA Group EUHOLOGNATHA SUPERFAMILY NEMOUROIDEA FAMILY Nemouridae SUBFAMILY Amphinemurinae Amphinemura delosa (Ricker) WB A. nigritta (Provancher) WB A. wui (Claassen) AP SUBFAMILY Nemourinae Ostrocerca albidipennis (Walker) AP # O. complexa (Claassen) AP # O. prolongata (Claassen) AP O. truncata (Claassen) AP Paranemoura perfecta (Walker) AP VOLUME 89, NUMBER 1 Prostoia completa (Walker) WB + P. hallasi Kondratieff & Kirchner CP P. similis (Hagen) WB Shipsa rotunda (Claassen) WB Soyedina carolinensis (Claassen) AP S. vallicularia (Wu) AP * Zapada chila (Ricker) (TN) AP FAMILY Taeniopterygidae SUBFAMILY Brachypterinae Bolotoperla rossi (Frison) AP Oemopteryx contorta (Needham & Claas- sen) AP * O. glacialis (Newport) (WV) WB Strophopteryx appalachia Ricker & Ross AP S. fasciata (Burmeister) WB S. limata (Frison) AP Taenionema atlanticum Ricker & Ross AP SUBFAMILY Taeniopteryginae Taeniopteryx burksi Ricker & Ross WB # T. lita Frison CP T. lonicera Ricker & Ross WB T. maura (Pictet) WB T. metequi Ricker & Ross WB + T. nelsoni Kondratieff & Kirchner AP T. parvula Banks WB T. ugola Ricker & Ross AP FAMILY Capniidae Allocapnia aurora Ricker AP * 4. brooksi Ross (TN) AP A. curiosa Frison AP * 4. forbesi Frison (KY, TN, WV) WB A. frisoni Ross & Ricker AP * 4. frumi Kirchner (WV) AP A. fumosa Ross AP A. granulata (Claassen) WB + A. harperi Kirchner AP . Illinoensis Frison WB . loshada Ricker AP . maria Hanson AP . mystica Frison AP . nivicola (Fitch) WB . pygmaea (Burmeister) WB . recta (Claassen) WB . rickeri Frison WB + A. simmonsi Kondratieff & Voshell AP Bmw BAA wD . stannardi Ross AP . virginiana Frison AP-CP . vivipara (Claassen) WB . wrayl Ross AP-CP . Zola Ricker AP Nemocapnia carolina Banks WB Paracapnia angulata Hanson AP I ed oe Br A me Ge a FAMILY Leuctridae SUBFAMILY Megaleuctrinae Megaleuctra flinti Baumann AP # M. williamsae Hanson AP SUBFAMILY Leuctrinae Leuctra alexanderi Hanson AP L. grandis Banks/biloba Claassen AP L. carolinensis Claassen AP L. duplicata Claassen AP L. ferruginea (Walker) WB L. mitchellensis Hanson AP L. monticola Hanson AP * L. nephophila Hanson (NC, TN) AP # L. rickeri James WB L. sibleyi Claassen AP # L. tenella Provancher WB L. tenuis (Pictet) WB L. triloba Claassen AP L. truncata Claassen AP L. variabilis Hanson AP Paraleuctra sara (Claassen) WB GROUP SYSTELLOGNATHA SUPERFAMILY PTERONARCYOIDEA FAMILY Pteronarcyidae Pteronarcys biloba Newman AP P. comstocki Smith AP P. proteus Newman AP P. scotti Ricker AP P. dorsata (Say) WB SUPERFAMILY PELTOPERLOIDEA FAMILY Peltoperlidae SUBFAMILY Peltoperlinae Peltoperla arcuata Needham AP P. tarteri Stark & Kondratieff AP 27 Tallaperla anna (Needham & Smith) AP # T. cornelia (Needham & Smith) AP # T. laurie (Ricker) AP 28 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON + T. lobata Stark AP T. maria (Needham & Smith) AP * Viehoperla ada (Needham & Smith) (NC, TN) AP SUPERFAMILY PERLOIDEA FAMILY Perlodidae SUBFAMILY Isoperlinae Clioperla clio (Newman) WB Isoperla bilineata (Say) WB * TJ. bellona Banks (NC, TN) AP I. burksi Frison WB * T. cotta Ricker (WV) WB # I. coushatta Szczytko & Stewart CP I. dicala Frison WB * T. distincta Nelson (NC, TN) AP * I. gibbsae Harper (WV) AP I. holochlora (Klapalek) AP I. lata Frison WB + I. major Nelson & Kondratieff AP I. marlynia Needham & Claassen WB I. montana (Banks) AP I. orata Frison AP * T. richardsoni Frison (WV) WB I. signata (Banks) WB I. similis (Hagen) AP I. slossonae (Banks) WB # I. transmarina (Newman) WB SUBFAMILY Perlodinae Cultus decisus (Walker) WB Diploperla duplicata (Banks) AP-CP # D. kanawholensis Kirchner & Kondratieff AP D. morgani Kondratieff & Voshell AP D. robusta Stark & Gaufin WB Helopicus subvarians (Banks) AP-CP Isogenoides hansoni (Ricker) AP # I. varians (Walsh) WB Malirekus hastatus (Banks) AP * Oconoperla innubila (Needham & Claas- sen) (NC, TN) AP Remenus bilobatus (Needham & Claassen) AP Yugus arinus (Frison) AP Y. bulbosus (Frison) AP FAMILY Chloroperlidae SUBFAMILY Paraperlinae * Utaperla gaspesiana Harper & Roy (WV) AP SUBFAMILY Chloroperlinae * Alloperla aracoma Harper & Kirchner (WV) AP . atlantica Baumann AP . banksi Frison WB . biserrata Nelson and Kondratieff AP . caudata Frison WB . chloris Frison AP . concolor Ricker AP . Idei (Ricker) AP . Imbecilla (Say) WB . nanina Banks AP * 4. neglecta Frison (NC, TN) AP A. usa Ricker AP Haploperla brevis (Banks) WB Suwallia marginata (Banks) AP Sweltsa lateralis (Banks) AP S. mediana (Banks) AP S. naica (Provancher) AP S. onkos (Ricker) WB S. urticae (Ricker) AP * Rasvena terna (Frison) (NC, TN, WV) AP Dawa AA RA DR FAMILY Perlidae SUBFAMILY Acroneuriinae Acroneuria abnormis (Newman) WB A. arenosa (Pictet) CP * 4. arida (Hagen) (TN) WB A. carolinensis (Banks) AP A. evoluta Klapalek WB A. filicis Frison WB A. flinti Stark & Gaufin AP # A. internata (Walker) WB A. lycorias (Newman) WB * 4, perplexa Frison (TN, WV) WB * 4. petersi Stark & Gaufin (NC, TN) AP Attaneuria ruralis (Hagen) WB * Beloneuria stewarti Stark & Szczytko (NC, TN) AP Eccoptura xanthenes (Newman) AP-CP # Hansonoperla appalachia Nelson AP VOLUME 89, NUMBER 1 Perlesta placida (Hagen) ““complex’” WB Perlinella drymo (Newman) WB P. ephyre (Newman) WB SUBFAMILY Perlinae # Neoperla carlsoni Stark & Baumann CP # N. catharae Stark & Baumann WB # N. clymene (Newman) WB * N. choctaw Stark & Baumann (WV) WB N. freytagi Stark & Baumann WB # N. stewarti Stark & Baumann WB Paragnetina fumosa (Banks) AP-CP P. immarginata (Say) AP # P. ichusa Stark & Szczytko AP P. media (Walker) AP # Agnetina annulipes (Hagen) CP A. capitata (Pictet) WB # A. flavescens (Walsh) WB ACKNOWLEDGMENTS We thank the following persons for mak- ing specimens available for study: M. I. Bass, OFS. Flint: Ir:* R«L: Hoffman; R. E. Jen- kins, M. Kosztarab, and J. R. Voshell, Jr. Rebecca F. Surdick kindly provided the Hansonoperla record. Stanley W. Szczytko allowed us to include several Jsoperla rec- ords from Virginia. Bill P. Stark verified several species determinations. The base map of Virginia was kindly provided by R. E. Jenkins, Roanoke College. It is a modi- fication of the U.S.G.S. state of Virginia, scale 1: 500,000 map, 1957 edition. LITERATURE CITED Harper, P. P. and K. W. Stewart. 1984. Plecoptera. Chapter 13, pp. 182-230. Jn Merritt, R. W. and K. W. Cummins, eds., An introduction to the aquatic insects of North America. 2nd Edition. Kendall/Hunt Publ. Co. Dubque, Iowa. Hoffman, R. L. 1969. The insects of Virginia. No. 1. Part II. The biotic regions of Virginia. Res. Div. Bull. 48: 23-62. VA Polytech. Inst. and State Univ. Kirchner, R. F. 1980. A new Allocapnia from Vir- ginia (Plecoptera: Capniidae). Entomol. News 91: 19-21. . 1982. A new Allocapnia from West Virginia (Plecoptera: Capniidae). Proc. Entomol. Soc. Wash. 84: 786-790. 29 Kirchner, R. F. and P. P. Harper. 1983. The nymph of Bolotoperla rossi (Frison) (Plecoptera: Taenio- pterygidae: Brachypterinae). J. Kans. Entomol. Soc. 56: 411-414. Kirchner, R. F. and B. C. Kondratieff. 1984. A new Diploperla from West Virginia (Plecoptera: Per- lodidae). Proc. Entomol. Soc. Wash. 86: 648-652. 1985. The nymph of Hansonoperla appala- chia Nelson (Plecoptera: Perlidae). Proc. Entomol. Soc. Wash. 87: 593-596. Kondratieff, B. C. and R. F. Kirchner. 1982a. Tae- niopteryx nelsoni, a new species of winter stonefly from Virginia (Plecoptera: Taeniopterygidae). J. Kans. Entomol. Soc. 55: 1-7. 1982b. Notes on the winter stonefly genus Allocapnia (Plecoptera: Capniidae). Proc. Ento- mol. Soc. Wash. 84: 240-244. Kondratieff, B. C. and J. R. Voshell, Jr. 1979. A checklist of the stoneflies (Plecoptera) of Virginia. Entomol. News. 90: 241-246. . 1981. Allocapnia simmonsi, a new species of winter stonefly (Plecoptera: Capniidae). Ann. Entomol. Soc. Am. 74: 58-59. . 1982. The Perlodinae of Virginia, USA (Ple- coptera: Perlodidae). Proc. Entomol. Soc. Wash. 84: 761-774. Kondratieff, B. C., R. F. Kirchner, and J. R. Voshell, Jr. 1981. Nymphs of Diploperla. Ann. Entomol. Soc. Am. 74: 428-430. Nelson, C. H. 1979. Hansonoperla appalachia, a new genus and a new species of eastern nearctic Ac- roneuriini (Plecoptera: Perlidae), with a phenetic analysis of the genera of the tribe. Ann. Entomol. Soc. Am. 72: 735-739. . 1982. Notes on the life histories of Stropho- pteryx limata (Frison) and Oemopteryx contorta (Needham and Claassen) (Plecoptera: Taeniop- terygidae) in Tennessee. J. Tenn. Acad. Sci. 57: 9-15. Nelson, C. H. and B. C. Kondratieff. 1983. Isoperla major, a new species of eastern nearctic Isoperli- nae (Plecoptera: Perlodidae). Ann Entomol. Soc. Am. 76: 270-273. Stark, B. P. 1979. The stoneflies (Plecoptera) of Mis- sissippi. J. Miss. Acad. Sci. 24: 109-122. . 1983. The Tallaperla maria complex of east- ern North America (Plecoptera: Peltoperlidae). J. Kans. Entomol. Soc. 56: 398-410. . 1985. Notes on Oconoperla (Plecoptera: Per- lodidae). Entomol. News. 96: 151-155. 1986. Review of Agnetina (Plecoptera: Per- lidae). J. Kans. Entomol. Soc. 59: 437-445. Stark, B. P. and B. C. Kondratieff. A new species of Peltoperla from eastern North America (Plecop- tera: Peltoperlidae). Proc. Entomol. Soc. Wash. (In press). 30 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Stark, B. P. and D. H. Ray. 1983. A revision of the genus Helopicus (Plecoptera: Perlodidae). Fresh- wat. Invertrbr. Biol. 2: 16-27. Stark, B. P. and K. W. Stewart. 1981. The nearctic genera of Peltoperlidae (Plecoptera). J. Kans. Entomol. Soc. 54: 285-311. Stark, B. P. and S. W. Szczytko. 1981. Contributions to the systematics of Paragnetina (Plecoptera: Per- lidae). J. Kans. Entomol. Soc. 54: 625-648. 1982. Egg morphology and phylogeny of Pteronarcyidae (Plecoptera). Ann. Entomol. Soc. Am. 75: 519-529. Stewart, K. W. and B. P. Stark. 1984. Nymphs of North American Perlodinae genera (Plecoptera: Perlodidae). Great Basin Nat. 44: 373-415. Surdick, R. F. 1985. Nearctic genera of Chloroper- linae (Plecoptera: Chloroperlidae). Ill. Biol. Mon. 54: 1-146. Szczytko, S. W. and K. W. Stewart. 1976. Three new species of nearctic Jsoperla (Plecoptera). Great Ba- sin Nat. 36: 211-220. 1981. Reevaluation of the genus Clioperla. Ann. Entomol. Soc. Am. 74: 563-569. Tarter, D. C. and R. F. Kirchner. 1980. List of the stoneflies (Plecoptera) of West Virginia. Entomol. News 91: 49-53. Tarter, D.C., D. A. Adkins, and C. V. Covell, Jr. 1984. A checklist of the stoneflies (Plecoptera) of Ken- tucky. Entomol. News 95: 113-116. Unzicker, J. D. and V. M. McCaskill. 1982. Plecop- tera, pp. 5.1-5.50. Jn A. R. Brigham, W. U. Brig- ham, and A. Gnilka, eds., Aquatic insects and oli- gochaetes of North and South Carolina. Midwest Aquatic Enterprises, Mahomet, Illinois. Zwick, P. 1973. Insecta: Plecoptera. Phylogenetisches system und Katalog. Das Tierreich 94. Walter de Gruyter and Co. Berlin. 465 pp. . 1984. Notes on the genus Agnetina (= Phas- ganophora) (Plecoptera: Perlidae). Aquat. Insects 6: 71-79. PROC. ENTOMOL. SOC. WASH. 89(1), 1987, pp. 31-42 TAXONOMY AND IDENTIFICATION OF THE EGG PARASITES (HYMENOPTERA: PLATYGASTRIDAE, TRICHOGRAMMATIDAE, MYMARIDAE, AND EULOPHIDAE) OF CITRUS WEEVILS (COLEOPTERA: CURCULIONIDAE) MICHAEL E. SCHAUFF Systematic Entomology Laboratory, BBII, Agricultural Research Service, USDA, % U.S. National Museum, NHB 168, Washington, D.C. 20560. Abstract.—The egg parasitoids of citrus weevils are reviewed, and an illustrated key is presented. Nine species of Hymenoptera in two superfamilies (Chalcidoidea: Eulophidae, Mymaridae, and Trichogrammatidae and Proctotrupoidea: Platygastridae) are now known to be primary or secondary parasites in these weevil egg masses. Two new species, Ho- rismenus bennetti and Tetrastichus fennahi, both eulophids, are described and figured. Additional data on the known hosts and distributions of the parasitoids are given. Weevils in the genera Pachneus, Exop- thalmus, Artipus, and Diaprepes (Curcu- lionidae) feed on a variety of plants. Adults are leaf feeders and larvae feed on and bore into the roots of the host plants. When these weevils attack citrus trees, they are collec- tively called citrus weevils and are capable of causing substantial economic damage. Larvae can even kill citrus by girdling the roots. Since 1964, when specimens of Diaprepes abbreviatus (L.) were discovered in citrus groves in the vicinity of Apopka, Florida (Woodruff, 1964), several attempts to es- tablish effective biological control programs have been made (e.g. Sutton et al., 1972; Beavers and Selhime, 1975). Unfortunately, these efforts have met with limited success. The weevil is native to Puerto Rico and the West Indies (Woodruff, 1968), but the exact route by which it became introduced into continental United States is unknown. In Puerto Rico, where D. abbreviatus has been a pest of sugarcane (commonly known as the sugar-cane root-stalk borer weevil), it has been effectively suppressed by an egg parasite, Tetrastichus haitiensis Gahan (Wolcott, 1948). This parasite has been the subject of most of the efforts to establish a biological control agent in the U.S. How- ever, several other parasites have been reared from the eggs of citrus weevils and the inability to identify and correctly assign specific names to these species has ham- pered efforts at control (R. Woodruff, pers. comm.). In addition, it is now apparent that some of these species are undescribed. I take this opportunity to clarify the tax- onomy of these species and provide a key to enable field workers to identify them. Much information about the biology and interrelationships of these species remains to be discovered and a sounder taxonomy should enable that work to proceed. Listed hosts for the parasites refer only to known species of citrus weevils and, in the case of hyperparasites, to hosts associated with cit- rus weevils. The majority of specimens used in this study were collected by R. Woodruff, E. E. Grissell and F. D. Bennett. They and the types are deposited in either the Florida State 32 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Collection of Arthropods at Gainesville (FSCA), the British Museum of Natural History (BMNH) or the U.S. National Mu- seum, Washington, D.C. (USNM). Morphological terminology follows that of Graham (1959), except that I use the term metasoma to refer to that part of the ab- domen past the propodeum (often called the gaster) and metanotum rather than meta- scutellum. Terms for sculpturing follow Harris (1979). KEY TO THE PARASITES OF CITRUS WEEVIL EGGS 1. Forewings nearly parallel sided (Fig. 5) and with marginal cilia longer than width of wing; fe- male antennae with 6 funicle articles and club entire (Fig. 14); male antennae with 13 articles BP ESTERS CORE ee OIA CLO et LAE Cleruchus sp. — Forewings not parallel sided (Figs. 1-4) and with marginal cilia much shorter than wing width; female antennae with fewer than 6 fu- nicle articles and club when present with 2 or 3 articles (Figs. 6-11); male antennae with few- erathanwllstanticlest ss een ee aa eee 2 Tarsi 5-segmented; forewing with only sub- marginal vein (Fig. 1); antenna with 4 small funicular articles (Fig. 6) and a 3-segmented club; scutellum without paired setae (Fig. 15) ESR Sten meee See Platystasius citri Nixon — Tarsi 3 or 4-segmented; forewing with sub- marginal, marginal and stigmal veins (Figs. 2— 4); antennae with fewer than 4 funicular arti- cles or club not 3-segmented (Figs. 7-11); scu- tellum with | or two pairs of setae (Figs. 16, Zs ()) Pete eR MRE af To RN teeter ere Per sy reve atc 3 3. Tarsi 3-segmented; forewings with obvious se- tal tracts radiating from stigmal vein and other wing veins as in Fig. 2; female antenna as in Fig. 10; body length about 0.8 mm SES RO ee See Brachyufens osborni (Dozier) — Tarsi 4-segmented; forewings without setal tracts, wing veins as in Figs. 3, 4; antennae as in Figs. 7-9,.11; body length greater than 0.9 mm 4. Scutellum with 2 longitudinal submedian grooves and 2 pairs of setae (Figs. 16, 17); ax- illae advanced forward of anterior edge of scu- tellum; clypeus bilobed (Fig. 24); propodeum narrowed medially, only about as wide as metanotum, with a simple median carina which may be incomplete (Fig. 23); metasoma broad- ly joined to propodeum, petiole not apparent bo — Scutellum without grooves or with only a single median groove (Figs. 18-20), with only a single pair of setae; axillae not advanced past anterior edge of scutellum, clypeal margin straight; pro- podeum wider than metanotum; metasoma petiolate 5. Scutellum with median longitudinal groove (Figs. 18, 19); propodeum with median raised area flanked by submedian carinae (Fig. 22) .. 6 — Scutellum without median groove (Fig. 20), propodeum with a pair of parallel submedian carinae (Fig. 21) ... Pediobius irregularis Kerrich 6. Notaulices complete, well defined (Fig. 18); first metasomal tergum with narrow reticulate band of sculpture posteriorly (Fig. 12); body metallic green; male femora and scape yellow ....... BANE Sewer e sd Horismenus cupreus (Ashmead) — Notaulices faint, incomplete (Fig. 19); first metasomal tergum nearly smooth or with faint open crack-like sculpture on posterior half (Fig. 13); body black; male femora and scape brown tONblaCkKeee nee: Horismenus bennetti sp. nov. 7. Most of head and metasoma yellow, rest of body often with extensive areas of yellow, may be nearly entirely yellow; male flagellar articles with long whorled setae (Fig. 11) i Fee One e nae eer Tetrastichus gala (Walker) — Body dark brown or blackish (except legs yel- low); male flagellar articles without long whorled Setaci(Hig"S) memes end Lene ts ee ee 8 8. Submarginal vein with a single seta (rarely 2); mesoscutum midlobe with single row of setae (Fig. 17); body without metallic greenish lustre; base of metasoma yellow; propodeum smooth and without carinae arising from nucha CS Rae oe Tetrastichus haitiensis Gahan — Submarginal vein with 3 or 4 setae (Fig. 4); mesoscutum with an irregular second row of setae (Fig. 16); body with metallic greenish lustre; propodeum sculptured and with carinae arising from nucha (Fig; 2)... o.5...... see te FONE eR Tetrastichus fennahi sp. nov. TRICHOGRAMMATIDAE Brachyufens osborni (Dozier) Figs. 2, 10 Ufens osborni Dozier, 1932: 36. Notes.— This species was described from specimens reared from Diaprepes abbrevi- atus eggs in Puerto Rico. In his original de- scription, Dozier noted that osborni was somewhat different than other species of Ufens, but decided that it still fell within the limits of that genus. Doutt and Viggiani (1968) noted several differences between the VOLUME 89, NUMBER 1 \ \ aN \ X ; Ss QA \ \ \\ Neat \\\ // / Vy) VW) WAH Sf le / fy) / LG IL Vay fifeies dita Figs. 1-5. fennahi. 5, Cleruchus sp. Scale line equals 0.1 mm. other species of Ufens and osborni and transferred it to their new genus Brachyu- fens as the type species. B. osborni has been reared from Pachnaeus litus eggs as well as species of Diaprepes. It has been reported to be quite common at times and has parasi- tized as many as 81% of the eggs of P. /itus (Baranowski, 1960). In view of the known biology of other trichogrammatids, it is 33 Forewings. 1, Platystasius citri. 2, Brachyufens osborni. 3, Horismenus bennetti. 4, Tetrastichus doubtful that reports of this species as a possible secondary parasite (Burks, 1979) are correct. Diagnosis.—As the only trichogramma- tid reared from citrus weevil eggs, this species is quite easily separated from the other parasites by the following characters: tarsi 3-segmented (4 or 5 segmented in oth- ers), forewings with distinct setal tracts (no 34 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON SSS SS = — SS —— = Ze, O Figs. 6-14. 6-11, Antennae. 6, Platystasius citri, female. 7, Horismenus bennetti, female. 8, Tetrastichus haitiensis, male. 9, Tetrastichus fennahi female. 10, Brachyufens osborni, female. 11, Tetrastichus gala, male. 12, 13, Metasomas. 12, Horismenus cupreus. 13, Horismenus bennetti. 14, Cleruchus sp., female antenna. Scale line equals 0.1 mm. tracts in other species) and venation as in Fig. 2, length about 0.8 mm and female an- tennae as in Fig. 10 (other antennae as in Figs. 6-9). Hosts. — Pachneus litus (Germar), P. opa- lus (Olivier), Diaprepes abbreviatus (L.). Specimens examined.—Dominican Re- public: San Cristobal Prov., San Cristobal. Puerto Rico: Mayaguez, University Cam- pus; Maricao. Andros Island, San Andros. Montserrat, Plymouth. United States. Flor- ida: Dade Co., Homestead; Palm Beach Co., West Palm Beach; Indian River Co., Vero Beach; St. Lucie Co.; Ft. Pierce, Hardee Co., Ft. Green. Collection dates range from 14 June through 3 October. EULOPHIDAE Tetrastichus haitiensis Gahan Figs: 8, 17 Tetrastichus haitiensis Gahan, 1929: 17. Notes. — This species was described from a series of specimens reared from Exop- thalmus quadrivittatus eggs at Port-au- Prince, Haiti. It is a primary parasite of the VOLUME 89, NUMBER 1 35 Figs. 15-20. SEMs. Thorax, dorsal view. 15, Platystasius citri. 16, Tetrastichus fennahi. 17, Tetrastichus haitiensis. 18, Horismenus cupreus. 19, Horismenus bennetti. 20, Pediobius irregularis. eggs and has also been reared from Dia- prepres abbreviatus and Pachnaeus litus. It is apparently the commonest parasite of cit- rus weevil eggs in this area and at times may kill up to 100% of the eggs and egg masses (Van Whervin, 1968). As a result, it has received the most attention from biological control workers and has been imported and released into Florida from the West Indies on several occasions (Sutton et al., 1972; Beavers and Selheime, 1975). A few spec- imens have recently been recovered from citrus weevil eggs in Central Florida (C. Tar- rant, pers. comm.). Diagnosis.—This species is most easily confused with Tetrastichus gala or T. fen- nahi. It differs in the following: the body is generally dark brown without a greenish 36 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tinge and with the legs and base of the metasoma yellow (body extensively yellow in gala and body with greenish tinge and base of metasoma not yellow in fennahi); submarginal vein usually with only | seta (submarginal with 3 or more in others); se- tae on male funicles about 2 as long as width of segment (Fig. 8) (shorter in fennahi and much longer and whorled in gala, Fig. 11); propodeum smooth and without para- spiracular carinae (propodeum sculptured in others and with paraspiracular carinae in fennahi),; mid lobe of scutum with only a single row of setae laterally (Fig. 17) (7. fen- nahi with partial second row, Fig. 16). Variability.—Females may occasionally have some yellow or very light brown areas around the scrobes, occelli, and notauli. There may also be some light brown on the femora. A few males have been observed with much shorter setae on the funicular articles than is typical for this species and the expanded ridge on the anterior surface of the scape is present although not as no- ticeable as in males with elongated funicular setae. Hosts.— Diaprepes abbreviatus (L.), Ex- opthalmus quadrivittatus (Olivier), Pach- neus litus (Germar), and P. opalus (Olivier). Specimens examined.—Dominican Re- public: La Romana Prov., Cacata; San Cris- tobal Prov., San Cristobal; Republic Prov., Monte Cristi, 4 Km. N. Villa Vasquez. Puerto Rico: Isabela; Mayaguez, University Campus; Ponce, Fortuna Fruit Experiment Station. Andros Island, San Andros. Ja- maica: St. Catherine Parish, Charlton nr. Ewarton; Red Hill. Cuba, Santiago, D. I. Vegas. Haiti, Port-au-Prince. United States. Florida, Hardee Co., Ft. Green. Collection dates range from March through October. Tetrastichus gala (Walker) Figs. 11, 24 Tetrastichus gala Walker, 1847: 28. Notes.—This species has been misiden- tified as T. marylandensis Girault (1916) (Z. Boucek, pers. comm.) and the exact rela- tionship of these two species remains in doubt. I can find little difference between specimens of the two species, but the host ranges indicate that two species may indeed be involved. 7. marylandensis is recorded as a parasite of lepidopterous larvae, aphids, and midges as well as from the eggs of citrus weevils. When associated with citrus wee- vils, 7. gala (cited as marylandensis) has been cited as an‘‘egg predator” feeding ex- ternally in the weevil egg masses (Van Whervin, 1968). I have studied specimens identified as both marylandensis and gala reared from various hosts, but many are in such poor condition that definitive identi- fication to species is not possible. Since this paper is not revisionary in scope, I am re- luctant to propose the synonymy of mary- landensis with gala. Until further specimens from the various hosts can be obtained or a comprehensive revision is undertaken, I believe it better to leave the two names as they are. Diagnosis.—This species is most easily confused with Tetrastichus fennahi and T. haitiensis. It can be separated by the follow- ing: submarginal vein with 3-5 setae (sub- marginal with only one seta or rarely 2 in T. haitiensis); body with at least some yel- low markings on the face, venter of thorax, and metasoma and often nearly entirely yel- low (body entirely dark brown or blackish except for the legs or base of the metasoma in other Tetrastichus species); male funic- ular articles with greatly lengthened, whorled setae (Fig. 11) (funicular setae much shorter in other Tetrastichus as in Fig. 8). Variability.—This species shows marked variation in coloration. Some specimens are almost entirely yellow with only a little brown on the edges of the metasomal terga, posterior margin of the propodeum, ante- rior edge of the pronotum, and around the foramen of the head. At the other extreme a series of specimens from Jamaica has al- most the entire dorsum of the thorax dark brown, and has large brown areas on the VOLUME 89, NUMBER | 37 Figs. 21-24. SEMs. 21-23, Propodeums. 21, Pediobius irregularis. 22, Horismenus bennetti. 23, Tetrastichus fennahi. 24, Face of Tetrastichus gala. metasoma, side of the thorax, and head. Variation in color ranges almost the whole spectrum between these two extremes. The face and frons are apparently always light colored, as is the venter of the thorax. Hosts.—Diaprepes abbreviatus (L.), D. famelicus (Olivier). Specimens examined.—Jamaica, St. Catherine Parish, Charlton nr. Ewarton. Montserrat, Plymouth. Guadeloupe, Do- main Duclos. Puerto Rico, Isabela, Maya- quez. Dominica. Collection dates range from December through June. Tetrastichus fennahi Schauff, NEw SPECIES Figs. 4, 9, 16, 23 Holotype female.— Length 1.2 mm. Col- or as follows: body dark brown with me- tallic green reflections; basal half of hind coxae light brown; rest of legs, tegulae, and antennae yellow. Head slightly wider than thorax, about as high as wide; face, frons, occiput with scattered silvery setae; frons with scattered small setigerous punctures near eye margins; occiput lightly imbricate; POL 2.5 x OOL; malar space 2 eye height; malar suture complete, curved; antenna in- serted on line with ventral margin of eye; scape not quite reaching level of tip of eye; antenna as in Fig. 9; scutum (Fig. 16) im- bricate, median longitudinal groove com- plete; midlobe with 2 irregular lines of setae; slightly longer than scutellum (18:15); scu- tellum as long as wide, propodeum (Fig. 23) imbricate, with several irregular small cari- nae projecting anteriorly from nucha, me- dially only about as wide as metanotum, paraspiracular carinae complete, with only a single seta laterad of spiracle; metasoma slightly longer than thorax; first tergum 38 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON smooth medially, becoming imbricate lat- erally, other terga imbricate; ovipositor sheaths cylindrical, barely protruding past tip of metasoma; forewing slightly more than 2x as long as wide (43:20) (Fig. 4); sub- marginal vein with 3 setae; ratio submar- ginal : marginal : stigmal 15:25:7; tip of hindwing rounded, marginal fringe 3 width of hindwing at hamulus. Male.—The only available male speci- men is very badly shriveled and no obvious differences from the female (except for the genitalia) can be discerned. Diagnosis.—This species is most easily confused with T. haitiensis and gala. It can be differentiated from them and from other Tetrastichus species by the following: body dark brown with greenish tinge (at least face and venter of thorax yellow in 7. gala, no greenish tinge in 7. haitiensis or base of metasoma yellowish); mid lobe of scutum with partial second row of setae (Fig. 16) (mid lobe with only a single row of setae in other species (as in Fig. 17); submarginal vein with 3—5 setae (submarginal with only 1 or rarely 2 setae in 7. haitiensis); propo- deum imbricate and with carinae arising from nucha (Fig. 23), paraspiracular carina present (other species with propodeum mostly smooth, without carinae, paraspi- racular carina absent); male flagellum with setae only about as long as width of segment (male flagellar setae about 2x as long as width of segment in haitiensis (Fig. 8), much longer and whorled in T. gala (Fig. 11). Variability.— Very little variation was observed in the specimens available for study. The body length ranges from |.0-1.3 mm. Eye color varies from bright red to silver or grayish. In a few specimens, the hind coxae are nearly entirely yellow. The majority of the specimens are badly shriv- eled and the propodeum is collapsed mak- ing it difficult to see if the paraspiracular carinae are as evident as in the type female and in a specimen that was photographed with the scanning electron microscope. In some of these poor specimens it appears that the carina is absent and caution should be used when assessing this character. Types.— Holotype 2 on point with data: St. Lucia, B.W.I., 1937, R. G. Fennah, Ex. Diaprepes abbreviatus eggs. Forewing and antenna slide mounted. Deposited in U.S. National Museum of Natural History. Para- types: 17 2, 1 6 same data as holotype; 1 Barbados, W. I., ex. eggs of Diaprepes ab- breviatus on citrus; 5 2 Machourie, Domin- ica, B.W.I., June, 1954, Coll. F. D. Bennett, ex. ova Diaprepes on legume; | 2 same data as above except collected at Grand Savan- nah [Grande Savanne], July, 1954; 2 2 Ja- maica, St. Catherine Parish, Charlton nr. Ewarton, 19-VI-1975, em. 29-VI-1975. E. E. Grissell, R. E. Woodruff. ex. Exopthal- mus or Pachneus eggs. Paratypes deposited in the USNM except for 2 2 each deposited in British Museum, Canadian National Col- lection, and Florida State collection of Ar- thropods. Hosts. —Diaprepes abbreviatus (L.). Etymology.—This species is named in honor of the collector of the holotype, R. G. Fennah. Pediobius irregularis Kerrich Figs. 20, 21 Pediobius irregularis Kerrich, 1973: 190. Notes.— This species was described from specimens reared from the egg mass of Ex- opthalmus viticollis Champion on citrus by L. W. Van Whervin taken in Belize (British Honduras). Little else is known of this species and it apparently has not been col- lected in the West Indies where most of the collecting by biological control workers has taken place. Kerrich listed this species as a primary parasite of the weevil eggs. Diagnosis.—This species is most easily confused with the two species of Horisme- nus. It can be separated by the following: scutellum without median groove (Fig. 20) (groove present in Horismenus, Fig. 19); propodeum with a pair of parallel subme- dian carinae (Fig. 21) (propodeum medially VOLUME 89, NUMBER 1 with a shiny raised area in Horismenus, Fig. DD). Hosts.—Exopthalmus vitticollis Cham- pion. Specimens examined.—Paratypes of P. irregularis and an additional short series of specimens collected a year earlier by the same collector (L. W. Van Whervin) who collected the types series (from the same locality). This species is known only from the type locality (Belize). Horismenus cupreus (Ashmead) Figs. 12, 18 Holcopelte cupreus Ashmead, 1894: 171. Notes.—The types of this species were collected on St. Vincent from unknown host (specimens deposited in BMNH). There are additional specimens in the USNM labelled as reared from eggs of Diaprepes famelicus esuriens on Montserrat. Whether it is a pri- mary or secondary parasite of the eggs is unknown. Diagnosis.—This species is most easily confused with H. bennetti or Pediobius ir- regularis. It is easily separated from Pe- diobius by the presence of a longitudinal groove on the scutellum and the raised smooth median area of the propodeum (as in Figs. 19, 22) (scutellar groove absent in Pediobius and median propodeum with paired carinae, Figs. 20, 21). H. cupreus dif- fers from bennetti by having complete and well defined notauli (Fig. 18), the first meta- somal tergum with a narrow sculptured band (Fig. 12), the body is black, and the male scape and femora are yellow (in bennetti, the notauli are not well defined (Fig. 19), the first tergum usually has only minute cracklike sculpture (Fig. 13), the body is black, and the male femora and scape are brown or black). Hosts.—Diaprepes famelicus esuriens (Gyllenhal). Specimens examined.—Type of H. cu- preus and other specimens from St. Vincent, and Montserrat. 39 Horismenus bennetti Schauff, NEw Species Figsa3i1 361 9:522 Holotype female.—Length 1.6 mm. Col- or black except the following: scape, legs past coxae yellow; last tarsomere brownish apically. Antennae as in Fig. 7; apex of scape even with arms of frontal forks at margin of eye; face below toruli lightly imbricate; area laterad of scrobes and below frontal grooves more strongly sculptured, nearly al- veolate; between scrobes smooth; frons me- dially above frontal forks very lightly im- bricate, nearly smooth; vertex imbricate; genae smooth to very faintly strigate, oc- ciput alveolate; POL 3x OOL; pronotum imbricate except smooth along posterior margin; scutum and scutellum imbricate as in Fig. 19; notaulices fading anteriorly; me- dian scutellar groove nearly reaching pos- terior margin of scutellum; with row of small alveolae extending posteriorly from scutel- lar setae and curving inwards near margin; metanotum smooth; propodeum (Fig. 22) smooth except at posterior edge and laterad of nucha; petiole in dorsal view slightly longer than wide (13:10), rugulose; prepec- tus imbricate, mesopleuron smooth; meta- soma equal in length to thorax, nearly 2 x as long as wide (60:35); first tergum covering 7/3; length, smooth except for small postero- medial patch of very fine crack-like acicu- lations (Fig. 13); forewing as in Fig. 3. Male.—Similar to female except the fol- lowing: length about 1.1 mm; scape and femora brown to black; basal half of tibiae occasionally light brown; scape 3 x as long as wide, funicular articles covered by nu- merous white setae; petiole 2 as long as wide; metasoma ovate, only about as wide as long. Diagnosis.—This species is most easily confused with Pediobius irregularis and H. cupreus. It can be separated from Pediobius by the presence of a longitudinal groove on the scutellum and the raised smooth median area of the propodeum (as in Figs. 19, 22) (scutellar groove absent in Pediobius and 40 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON median propodeum with paired carinae, Figs. 20, 21). It differs from H. cupreus by: the black body color (cupreus and many oth- er species are metallic green); the notauilces are faint and incomplete (Fig. 19) (notau- lices well defined in cupreus, Fig. 18); the first metasomal tergum nearly smooth, with only faint crack-like sculpture (Fig. 13) (nar- row reticulate band in cupreus, Fig. 12); male femora and scape brown to black (male scape and femora of cupreus yellow). Additional characters which help to separate this species from other Horismenus are: propodeum mostly smooth and without reticulation near nucha (many other species have some re- ticulate sculpture on the propodeum); legs of the female yellow past coxae (several species have the femora and/or tibiae dark colored); small row of alveolae adjacent to scutellar setae and scutellar surface nearly completely smooth (alveolae lacking and scutellar surface sculptured in some other species). Variability.—There is some variation in the appearance of the sculptured area on the first metasomal tergum. In one specimen, it appears as more of a reticulated pattern somewhat similar in appearance to that in cupreus. However, in this specimen, the band of sculpture was quite wide (about half as wide as long), while in cupreus the band is very narrow (only about a sixth as wide as long). Types.— Holotype 2 on point with data: Jamaica, Red Hill, May 1956. Curculionid eggs on citrus. Coll. by F. D. Bennett. De- posited in the U.S. National Museum of Natural History. Paratypes: 3 2; Puerto Rico, Isabela, VI-16-1932, G. N. Wollcott, ex. eggs Diaprepes abbreviatus; 1 °, 1 8, Jamaica, B.W.I., July 1954, Ova Prepodes, coll. by Simmonds; | ¢?, Jamaica, Mona, Dec. 1, 1967, ex. eggs of Tetrastichus parasite on citrus weevil eggs, coll. by Van Whervin; 5 6 and 4 2, Dominican Republic, San Cris- tobal, 23-VI-1976, on citrus, emerged 24- VI-1976; 8 2 and 3 4, Jamaica, Parish of St. Catherine, Charlton. Exp. Sta., 19-VI-1975. Grissell & Woodruff. Ex. Exopthalmus eggs on citrus. em. 25-VI-1975. One 6 and | 2 paratype deposited in BMNH and FSCA, rest in USNM. Hosts.—The species is apparently a hy- perparasite. Its most likely host is Tetra- stichus haitiensis, although it may also para- sitize other Tetrastichus. Etymology.—This species is named in honor of F. D. Bennett who collected many of the specimens used in this study. PLATYGASTRIDAE Platystasius citri Nixon Rigs 6. 15 Platystasius citri Nixon, 1969: 447. Notes.—This species is a primary endo- parasite (Van Whervin, 1968) of citrus wee- vil eggs (Exopthalmus or Pachneus sp. ac- cording to label data). It is known from Jamaica, although it is possible that it oc- curs on other islands in the West Indies. Although originally described in the genus Platystasius, this species is now considered better placed in Fidiobia (L. Masner, pers. comm.). A formal change in generic place- ment is being proposed in a manuscript on platygastrid taxonomy that is currently in preparation but has not yet been published. Therefore, I have used the currently pub- lished combination. Diagnosis. — This species is the only proc- totrupoid that has been associated with cit- rus weevil eggs. It can be identified by the following: tarsi 5-segmented (3 or 4 seg- mented in other species); forewing with only a submarginal vein (Fig. 1) (obvious sub- marginal and marginal and stigmal veins in other species, except Cleruchus sp. (see Figs. 2-4); antenna with four small funicles and large 3-segmented club (Fig. 6) (antennae of other species as in Figs. 7-11, 14); scutellum without obvious paired setae (Fig. 15) (scu- tellum with one or two pairs of large setae in others, Figs. 16, 17, 20, 22). Hosts.— Pachneus or Exopthalmus sp. Specimens examined.—Jamaica, St. VOLUME 89, NUMBER 1 Catherine Parish, Worth Park and Charlton nr. Ewarton; Manchester. Mona Island. Collected in June, August, and September. MyYMARIDAE Cleruchus sp. Notes.— This species was recently reared from eggs of Artipus floridanus Horn in the vicinity of Wabasso, Florida (Indian River Co.). Unfortunately, only 4 specimens have been collected, and these are in poor con- dition. I would place them in the genus Cle- ruchus (sensu Schauff, 1984). They are very similar to C. brevipennis Ogloblin (1940). This species is almost certainly unde- scribed, but without additional specimens I am reluctant to name it at this time. Diagnosis.—This species is the only member of the family Mymaridae yet re- corded from citrus weevil eggs. It can be identified by the following characters: tarsi 4-segmented; forewings nearly parallel-sid- ed and with marginal cilia much longer than wing width (Fig. 5) (other species with fore- wings much broader and not parallel-sided and marginal cilia much shorter than wing width); female antenna with 6 funicular ar- ticles and a single segmented club (Fig. 14); male antennae with 13 segments (other species with male antennae with fewer than 13 segments). ACKNOWLEDGMENTS I give special thanks to E. E. Grissell whose preliminary work on the taxonomy of these parasites made possible the timely comple- tion of this study. I thank also R. Woodruff, who together with Dr. Grissell collected much of the material on which this study has been based. R. E. White, E. E. Grissell, D. Wahl, and F. D. Bennett made many valuable comments on the manuscript. Iam grateful to J. S. Noyes and Z. Bouéek for the loan of specimens from the British Mu- seum and L. Stange for material from the Florida State collection of Arthropods. Iam grateful to L. Masner (Biosystematic Re- 41 search Institute, Agriculture Canada, Ot- tawa, Ontario), for his help with determin- ing the platygastrid and D. Whitehead (Systematic Entomology Lab, USDA, Washington, D.C.) for help with the weevil names. C. Tarrent (University of Florida) provided valuable specimens and _ infor- mation on Brachyufens osborni and the species of Cleruchus. LITERATURE CITED Ashmead, W. H. 1894. Report upon the Parasitic Hymenoptera of the Island of St. Vincent. J. Lin- nean Soc. 25: 56-254. Baranowski, R. M. 1960. Notes on a parasite of the citrus root weevil Pachnaeus litus (Germ.). Fla. Entomol. 43: 197. Beavers, J. B. and A. G. Selhime. 1975. Further at- tempts to establish the weevil egg parasite Je- trastichus haiteinsis in Florida. Fla. Entomol. 58: 29-31. Burks, B. D. 1979. Family Trichogrammatidae, pp. 1033-1043. In Krombein, K. V. et al., eds., Cat- alog of Hymenoptera in America north of Mexico, Vol. 1. Symphyta and Apocrita (parasitica). Smithsonian Institution Press, Washington, D.C., 1198 pp. Dozier, H. L. 1932. Descriptions of new trichogram- matid (Hymenoptera) egg parasites from the West Indies. Proc. Entomol. Soc. Wash. 34: 29-37. Doutt, R. L.and G. Viggiani. 1968. The classification of the Trichogrammatidae (Hymenoptera: Chal- cidoidea). Proc. Calif. Acad. Sci. (4th ser.) 35: 477- 586. Gahan, A. B. 1929. Description of an egg-parasite of Exopthalmus quadrivitattus (Olivier). Proc. Ento- mol. Soc. Wash. 31: 17-18. Girault, A. A. 1916. New miscellaneous chalcidoid Hymenoptera with notes on described species. Ann. Entomol. Soc. Am. 9: 291-308. Graham, M. W. R. DeV. 1959. Keys to the British genera and species of Elachertinae, Eulophinae, Entedontinae, and Euderinae (Hym., Chalcidoi- dea). Trans. Soc. British Entomol. 13: 169-204. Harris, R. A. 1979. A glossary of surface sculpturing. Occasional papers of laboratory services/ Ento- mology no. 28. California Department of Food and Agriculture, Sacramento, California, 31 pp. Kerrich, G. J. 1973. A revision of the tropical and subtropical species of the eulophid genus Pedio- bius Walker (Hymenoptera: Chalcidoidea). Bull. Br. Mus. (Nat. Hist.). 29: 115-199. Nixon, G.E. J. 1969. Two new species of Platystasius Nixon with a note on the generic relationship be- 42 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tween Platystasius and Fidiobia Ashmead. Proc. Entomol. Soc. Wash. 71: 445-449. Ogloblin, A. A. 1940. Dos Mymaridae nuevos de Misiones (Hym.). Rev. de Entomol. Rio de Ja- neiro 11: 597-603. Schauff, M. E. 1984. The Holarctic genera of My- maridae (Hymenoptera: Chalcidoidea). Mem. Entomol. Soc. Wash. 12: 67 pp. Sutton, R. A., A. G. Selhime, and W. McCloud. 1972. Colonization and release of Tetrastichus haitiensis as a biological control agent for citrus root weevils. J. Econ. Entomol. 65: 184-185. Van Whervin, L. W. 1968. The citrus weevils of Ja- maica and some of their parasites. Univ. West Indies, St. Augustine, Trinidad Tech. Bull. 1: 1-11, 1-23. Walker, F. 1847. Characters of undescribed Chalci- dites collected in North America by E. Doubleday, and now in the British Museum. Ann. Mag. Nat. Hist. 20: 19-29. Wolcott, G. N. 1948. The insects of Puerto Rico; Coleoptera. J. Agr. Univ. Puerto Rico. 32: 225- 416. Woodruff, R. 1964. A Puerto Rican weevil new to the United States. Fla. Dept. Agr., Div. Plant Ind. Ent. Circ. 30: 1-2. 1968. The present status of a West Indian weevil (Diaprepes abbreviata (L.)) in Florida (Co- leoptera: Curculionidae). Fla. Dept. Agr., Div. Plant. Ind. Ent. Circ. 77: 1-4. PROC. ENTOMOL. SOC. WASH. 89(1), 1987, pp. 43-46 LIFE HISTORY OBSERVATIONS ON THE GRASSHOPPER APPALACHIA HEBARDI REHN AND REHN (ORTHOPTERA: ACRIDIDAE: MELANOPLINAE) ROBERT G. BELLINGER AND ROBERT L. PIENKOWSKI Department of Entomology, Virginia Polytechnic Institute and State University, Blacks- burg, Virginia 24061. Abstract. —The brachytperous melanopline grasshopper, Appalachia hebardi Rehn and Rehn, is a late season, univoltine species that occurs in the Appalachian ridges of West Virginia, Virginia, and Pennsylvania, U.S.A. Data on this species were taken from a population near Mathias, West Virginia. A. hebardi feeds on understory vegetation and prefers older foliage. Both sexes went through five instars to become adults. Body size was within previously published ranges. Mean population ovariole number was 11.44 + 1.13 (Bellinger and Pienkowski, 1985), and mean pod size was 9.67 + 2.08. Maximum pod size is estimated to be 14. Some females can lay at least two pods during the season. Our observations corroborate previous reports and extend the life history information on A. hebardi. Appalachia hebardi Rehn and Rehn is a brachypterous grasshopper from the Ap- palachian Mountains of West Virginia, Vir- ginia, and Pennsylvania (Rehn and Rehn, 1936, 1939). The genus contains one other species, A. arcana Hubbell and Cantrall, known from Michigan (Hubbell and Can- trall, 1938). Little 1s known of their biology. This paper provides new information, in- cluding food plants in the field and labo- ratory, development, fecundity, and sea- sonal occurrence of A. hebardi. We previously published femur lengths and ovariole numbers for this species (Bellinger and Pienkowski, 1985), but include these data here for completeness. MATERIALS AND METHODS Grasshoppers were collected by the senior author (RGB) from the undergrowth around a cabin on the eastern slope of Big Ridge, east of the boundary of Lost River State Park, Mathias, Hardy Co., West Virginia (Bellinger and Pienkowski, 1985). The ele- vation of the site is ca. 700 meters. The species had been seen feeding in small num- bers on native and cultivated plants during 1981 and 1982, but were less abundant in 1983 (F. E. Wood, Dept. of Entomology, University of Maryland, personal commu- nication). The collection site was charac- terized by an overstory dominated by chest- nut oak, with an understory of black gum (Nyssa sylvatica Marsh.), dogwood, and sucker growth of American chestnut (Cas- tanea dentata (Marsh.) Borkh.). Under- growth was predominately mountain laurel (Kalmia latifolia L.), and Vaccinium spp. Grasshoppers were all on the ground when collected. Collecting was difficult because of the scarcity of individuals and because when disturbed they jumped long distances, even in the dense undergrowth. One immature and one adult male, and one immature and two adult females were collected on 12 Sep- tember, 1983. One adult male and six adult 44 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON females were collected in mid-October, 1983. Individuals from each collection were returned to the laboratory, at Blacksburg, VA. The initial intent of the collections was to identify the species and determine ovar- iole number in the females. The species was tentatively identified by RGB as A. hebardi. Grasshoppers were maintained at 30°C in a rearing chamber, with 2-3 individuals per 0.45 L paper carton with screened top and bottom. Initially, grasshoppers were pro- vided with fresh romaine lettuce (Lactuca sativa L. cv. ‘Romana’), ground dry dog food (25% crude protein), and misted with water twice daily. Because the grasshoppers did not noticeably feed on the lettuce, other plants were tried as food, including fescue grass (Poa pretensis L.), leaves of dandelion, (Taraxacum officinale Weber), Crimson clover (Trifolium incarnatum L.), three species of dogwood (Cornus florida L. and C. spp.), and red, white, and chestnut oak (Quercus rubra L., Q. alba L., and Q. prinus L.). Chestnut oak leaves were brought from the collection site. All other plants were col- lected near the laboratory. Grasshoppers from each collection were held for three to four days, or until all in- dividuals had become adults. All individ- uals were sacrificed at that time. Females were dissected to determine ovariole num- ber (Bellinger and Pienkowski, 1985) and their ovipositional status (Launois-Luong, 1978). Measurements of total body length, hind femur, and tegmen were made on three males and nine females, and of the length and width of the pronotum of the females. Because development of the grasshoppers was not observed in the field or in the lab- oratory, the number of antennal segments in the adults were counted to determine number of instars required to reach the adult (Shotwell, 1941). RESULTS The identification of this species as Ap- palachia hebardi Rehn and Rehn was con- firmed by Irving J. Cantrall, Museum of Zo- ology, University of Michigan, Ann Arbor (retired). Habitat and food plants.— Feeding pre- viously had been noted on native mountain laurel, sucker growth on cut chestnut oak stumps, several herbaceous weeds, and on cultivated rose of sharon (Hibiscus sp.). In the laboratory the grasshoppers ate ground dry dog food but not romaine lettuce. Grass- hoppers ate old leaves from lower parts of the three dogwood species but not newer leaves from branch ends. Similarly, only older leaves of dandelion were eaten. Grass- hoppers did not feed on foliage from the three species of oak, or the grass, or the clover. Body size.— Body measurements (mean + SD (mm)) were: total body length, males— 20.87 + 1.16, females—25.27 + 2.06, hind femur, males—10.9 + 0.35, females— 12.28 + 0.52, tegmen, males—4.33 + 0.57, females—4.59 + 0.38, pronotal length, fe- males—5.09 + 0.43, pronotal width— 4.10 + 0.38. Antennal segments and instar number. — Two of the males had 23 antennal segments, and one had 24 segments. Three of the fe- males had 23 segments, and the remaining six had 24. There was no apparent difference in segment number because of collection dates. Antennal segment counts showed that all individuals in each sex had gone through five developmental stadia to reach the adult. Antennae were long for the number of seg- ments. Individual segments were visibly longer than in the genus Me/anoplus Stal. Fecundity.—Appalachia hebardi mated readily in the laboratory. No eggs were laid in the laboratory, probably because a suit- able ovipositional media was not provided. Mean ovariole number per female was 11.44 + 1.13 SD, and ranged from 10 to 13 (Bellinger and Pienkowski, 1985). There were five to seven ovarioles per ovary, so there could be up to 14 ovarioles per female. Dissections of females, collected as adults in the field, showed that the two adults col- lected in September each had 12 ovarioles. VOLUME 89, NUMBER 1 One showed little development of the ova- ries, and the other had laid one pod with 12 eggs but had not yet begun development of a second pod. Adult females collected in October showed a range of ovarian devel- opment. One female had oviposited twice and was developing a third pod. This female had 11 ovarioles and had laid eight eggs in the first pod and nine eggs in the second pod. Seven oocytes were being developed, indicating a maximum of seven eggs in the potential third pod. None of the others had Oviposited, nor had they begun oocyte de- velopment, but two had 12 ovarioles and 12 well developed oocytes each. Average pod size was thus 9.67 + 2.08 (n = 3 pods). While color of the chorion in grasshopper eggs varies among species, developing oo- cytes of this species were bright yellow, the same color observed in other Melanoplinae, and in most Oedipodinae and Gomphocer- inae (unpublished). We did not observe any post-ovipositional eggs. No parasites were noted in any dissection. DISCUSSION Observations made on a small population of A. hebardi from West Virginia confirm some observations previously made on the species. The collection site was a habitat similar to those described by Rehn and Rehn (1936). The elevation in West Virginia was within the 550 to 1220 m range of elevations given for the species (Rehn and Rehn, 1939). We found as did Rehn and Rehn (1936) that the species jumped long distances to escape capture. Plant associations but no food plants were given by Rehn and Rehn (1936). Our field observations and laboratory feed- ings, although somewhat limited, suggested that the species feeds on older foliage of undergrowth species. Individuals from the Mathias, WV pop- ulation in 1983 went through five stadia. This is acommon number of stadia for ac- ridid grasshoppers. Brachypterous acridid grasshoppers frequently go through a re- duced number of stadia (four or five), and 45 as a result have small body sizes (Mason, 1954; Uvarov, 1966, 1977). Species from woodlands and higher altitudes are fre- quently brachytperous. Body size measure- ments for the Mathias, WV population fell within the range reported by Rehn and Rehn (1936), except for the length of the tegmen in males, which was longer. Appalachia herbardi has small ovaries (low number of ovarioles), a condition which is related to its small body size (Bellinger and Pienkowski, 1985). Our counts of ovar- ioles indicate potential pod sizes of 10 to 14 eggs per pod. Our data showed that pod sizes in the field averaged 9.67 eggs. Rehn (1938) reported “‘approximately ten eggs” from a single pod. Females in the Mathias population are capable of laying at least two, and possibly three pods per female, based on one female. Thus, this species may lay up to ca. 42 eggs per female, on the basis of these limited data. The oviposition sub- strate for this species remains unknown. While most acridid grasshoppers oviposit in the soil, some woodland species oviposit in holes in dead wood (Blatchley, 1920). Rehn and Rehn (1936) reported that A. hebardi occurred as early as the first week of July, to as late as early September, over its range. Our collections were made in mid- September, when we collected immatures and adults, and mid-October. The repro- ductive status of adult females showed that the population began oviposition before mid-September, but that as late as mid-Oc- tober some females had yet to oviposit, or even begin to develop their first pod. At the elevation of the Mathias site, the growing season 1s over by mid-October, and unless this species 1s adapted to cooler tempera- tures and senescent food plants, some fe- males in the population may never oviposit. ACKNOWLEDGMENTS We thank F. E. Wood for informing us of the study population and allowing us ac- cess to it. We thank Irving J. Cantrall for confirming the identification of Appalachia 46 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON hebardi and encouraging our studies of grasshopper biology. We thank also David A. Nickle, Systematic Entomology Labo- ratory, Agricultural Research Service, c/o U.S. National Museum, Washington, D.C. for providing us with literature on Appa- lachia. LITERATURE CITED Bellinger, R. G. and R. L. Pienkowski. 1985. Inter- specific variation in ovariole number in melanop- line grasshoppers (Orthoptera: Acrididae). Ann. Entomol. Soc. Am. 78: 127-130. Blatchley, W. S. 1920. Orthoptera of northeastern America. The Nature Publishing Co. Indianapolis. 784 pp. Hubbell, T. H. and I. J. Cantrall. 1938. A new species of Appalachia from Michigan (Orthoptera, Acri- didae, Cyrtacanthacridinae). Univ. Michigan Mus. Zool. Occ. Paper No. 389: 1-21. Launois-Luong, M. H. 1978. Methode pratique d’in- terpretation de l’etat des ovaires des acridiens du Sahel. Ann. Zool. Ecol. Anim. 10: 569-587. Mason, J. B. 1954. Number of antennal segments in adult Acrididae (Orthoptera). Proc. R. Entomol. Soc. Lond. B 23: 228-238. Rehn, J. A. G. and J. W. H. Rehn. 1936. On new or redefined genera of nearctic Melanopli (Orthop- tera: Acrididae, Cyrtacanthacridinae). Trans. Am. Entomol. Soc. 62: 1-56. 1939. Studies of certain cyrtacanthacridoid genera (Orthoptera: Acrididae). Part I. The Podis- ma complex. Trans. Am. Entomol. Soc. 65: 61- 95. Rehn, J. W. H. 1938. Notes on the eggpods of Ap- palachia hebardi and Dendrotettix quercus (Or- thopter: Acridiae; Cyrtacanthacridinae). Entomol. News 49: 259-260. Shotwell, R. L. 1941. Life histories and habits of some grasshoppers of economic importance on the Great Plains. U.S. Dept. Agric. Tech. Bull. No. 774. Uvarov, B. 1966. Grasshoppers and locusts: a hand- book of general acridology, vol. 1. Cambridge Uni- versity Press, Cambridge. 481 pp. . 1977. Grasshoppers and locusts: a handbook of general acridology, vol. 2. Centre for Overseas Pest Research, London. 613 pp. PROC. ENTOMOL. SOC. WASH. 89(1), 1987, pp. 47-50 AGATHIS THOMPSONI N. SP., A NEARCTIC SPECIES OF AGATHIDINAE (HYMENOPTERA: BRACONIDAE) PARASITIC ON GREYA SUBALBA (BRAUN) (LEPIDOPTERA: INCURVARIIDAE) MICHAEL J. SHARKEY Biosystematics Research Centre, Agriculture Canada, Ottawa, Ontario K1A 0C6, Canada. Abstract. — Adults of Agathis thompsoni n. sp. (Braconidae: Agathidinae) from north- western U.S.A. are described and illustrated. Larvae are parasitic on Greya subalba (Braun), which feeds on the schizocarps of Lomatium spp. (Umbelliferae). Females of the new species are differentiated from those of both European and North American species. Among the many undescribed species of Agathis Latreille in North America is one that has been studied by J. N. Thompson (Thompson, 1986). It is to compliment his research on oviposition behaviour and searching efficiency that I describe this new species. Agathis thompsoni Sharkey, NEw SPECIES Diagnosis. —Agathis thompsoni is distin- guished from other species of Agathis by the following combination of character states: basal flagellomere 1.6 x longer than follow- ing flagellomere; malar space 0.6 x greatest diameter of eye; ovipositor 1.2 x as long as metasoma when fully extended. Description, holotype ¢.—(Intraspecific variation is given in parentheses). Color: Black except metasoma slightly paler lat- erally and yellowish-orange as follows: mandible, all femora over distal 0.7, fore and middle tibiae, hind tibia except for bas- al and apical melanic bands, basal 0.2 of all basitarsomeres; fore wing hyaline. Head (Fig. 1): Antenna with 26 (23-26) flagello- meres; basal flagellomere 1.6 x longer than following flagellomere; head subrostiform, malar space 0.6 x greatest diameter of eye; weak, V-shaped depression anterior to me- dian ocellus; smooth, longitudinal ridge from near median ocellus to level of anten- nal insertion; galea 2.2 x longer than max- imum width. Mesosoma (Figs. 2, 3): Notauli deeply impressed, pitted, scutellar groove with numerous longitudinal ridges; prono- tum smooth except for crenulae along pos- terior margin; sternaulus 0.7 length of mesopleuron and complete to posterior margin; metapleuron rugose over ventral 0.2 (0.2-0.3); propodeum with transverse, an- terolateral ridge and with 3 longitudinal ridges, medial ridge weak; (propodeum may be somewhat rougher than in Fig. 2); mid tibia with 3 (2-3) preapical spines; hind tib- ia with 6 (4-6) apical spines; hind tarsal claw with strong basal tooth. Metasoma (Fig. 4): First tergum as long as wide, with pair of weak longitudinal ridges and weak striae over anterior 0.5; tergum 2 + 3 mostly smooth with basal swelling; ovipositor 1.2 x longer than metasoma when fully extended; Ovipositor sheaths slightly shorter (0.9 x) than metasoma. Length: 3.7 (3.5—4.3) mm. Allotype 6.—As for the holotype except antenna with 23 flagellomeres. (Left hind leg missing after coxa). This species is named after John N. Thompson. Material examined. — Holotype 2, U.S.A., Washington, Whitman Co., Smoot Hill Biol. 48 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON > Ne Sian Figs. 1, 2. Agathis thompsoni. 1, Head, lateral. 2, Metanotum and propodeum, dorsal. Pres. nr. Albion, ex. Greya subalba (Braun), 5.VI.1985, J. N. Thompson, (United States National Museum). Allotype 6, same data as holotype except date, 17.VI.80. Para- types: 38 2, same data as holotype, (Cana- dian National Collection, United States Na- tional Museum). | 4, same data as allotype, (Canadian National Collection). Discussion.—In a recent paper (Sharkey, 1985) I defined my concept of Agathis La- treille. The following Nearctic and Holarc- tic species belong to this genus: 4. brevicor- nis (Muesebeck), A. cupressi Muesebeck and Walkley, 4. gibbosa (Say), A. malvacearum Latr., A. pumila (Ratzburg), A. rubripes Cresson, A. terminata Cresson, A. thomp- soni Sharkey, A. tibiator Provancher. Females of A. thompsoni differ from those of most other North American species of Agathis by their short ovipositor, which is only 1.2 as long as the metasoma when fully extended. Other species have the ovi- positor fully as long as the body except A. pumila, a Holarctic species. Unlike 4. thompsoni, A. pumila has the two most bas- al antennal flagellomeres subequal in length. In Nixon’s (1986) key to the European females of Agathis, A. thompsoni keys to couplet number 30. Females differ from those of five of the six species that key through this couplet by their short ovipos- itor. Females of A. melpomene Nixon, which also have a short ovipositor, differ in that the first tergum of the metasoma is mostly smooth, though sometimes with weak ru- gosity medially. Females of A. thompsoni have striae in the anterior 0.6 of the first metasomal tergum. ACKNOWLEDGMENTS I thank G. Gibson and J. Huber for their reviews, B. Jinkinson for the scanning elec- VOLUME 89, NUMBER 1 Figs. 3, 4. tron photomicrographs and the Electron Microscope Centre, Agriculture Canada, for the use of its facilities. LITERATURE CITED Muesebeck, C. F.W. 1927. A revision of the parasitic wasps of the subfamily Braconinae occurring in OUR pod ~ “ AZ 2 ) rata Agathis thompsoni. 3, Mesosoma, lateral. 4, Metasoma, dorsal. America north of Mexico. Proc. U.S. Natl. Mus. 69: 1-73. Nixon, G. E. J. 1986. A revision of the European Agathidinae (Hymenoptera: Braconidae). Bull. Br. Mus. (Nat. Hist.) Entomol. 52: 183-242. Sharkey, M. J. 1985. Notes on the genera Bassus Fabricius and Agathis Latreille, with a description 49 50 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON of Bassus arthurellus n. sp. Can. Entomol. 117: searching efficiency in a natural population of a 1497-1502. braconid parasitoid. J. Anim. Ecol. 55: 351-360. Thompson, J. N. 1986. Oviposition behaviour and ANNOUNCEMENT With this issue the Proceedings begins a two-column format not including titles and abstracts. This is a cost-saving measure that the Publications and Executive Committees believe does not sacrifice utility or beauty. The new format should allow for some growth in the number of articles without the bound volume becoming too large. Space saving comes mainly from fitting narrow tables and plates into only one column rather than having them extend across a whole page. There is also a gain of 4, inch width of print per page. — Editor. PROC. ENTOMOL. SOC. WASH. 89(1), 1987, pp. 51-56 SCANNING ELECTRON MICROSCOPY FOR THE STUDY OF THE WINTER STONEFLY GENUS CAPNIA (PLECOPTERA: CAPNIIDAE) C. RrLEY NELSON AND RICHARD W. BAUMANN Department of Zoology, Brigham Young University, Provo, Utah 84602. Abstract. —Scanning electron micrographs are produced for nine species of Capnia: C. barberi, C. cheama, C. coloradensis, C. decepta, C. elevata, C. fibula, C. manitoba, C. melia, and C. uintahi. Structural details not previously reported such as spines and sensilla may prove useful in constructing future phylogenies. The value of scanning electron microscopy for species identification and systematic work is discussed. Scanning electron microscopy (SEM) has been used to examine structural details of Plecoptera eggs (Baumann, 1973; Stark and Baumann, 1978; Szczytko and Stewart, 1979; Stark and Stewart, 1981; Stark and Szczytko, 198la, b; Stark and Stewart, 1982c; Stark and Ray, 1983; Stark, 1983; Stark and Szczytko, 1982; Stark and Szczyt- ko, 1984), mouthparts (Baumann, 1973; Stark and Stewart, 1981; Stark and Stewart, 1982a-c; Stark and Ray, 1983), and sensilla (Stark and Stewart, 1981; Stark and Stewart, 1982a; Stark and Ray, 1983; Stark, 1983; and Kapoor and Zachariah, 1984). These studies have helped in constructing phylog- enies such as those of Nelson (1984). The use of SEM to illustrate male genitalia of stoneflies has been limited to the species description of Prostoia hallasi (Kondratieff and Kirchner, 1984). A study of the male genitalia of Capnia was undertaken to ex- amine the feasibility of using SEM as a tool to differentiate species and species groups in this large genus. MATERIALS AND METHODS Specimens of North American Capnia were examined and compared to holotypes (except that of C. fibula Claassen, which was missing). The term genitalia refers to the epiproct of the male and associated sclerites while terminalia refers to all the appendages of the abdominal segments including tergal knobs, genitalia, and cerci. Collection data for the specimens used in this study are: Capnia barberi Claassen—CALIFOR- NIA, Nevada County, Sagehen Creek, 26 February 1965, Sheldon and Hawthorne. Capnia cheama Ricker—MONTANA, Lincoln County, Kootenai River, 19 March 1970, R. L. Newell. Capnia coloradensis Claassen—COLO- RADO, Routt County, Fish Creek, Hwy 40, Steamboat Springs, 6 April 1963, A. R. Gaufin. Capnia decepta Banks—ARIZONA, Cochise County, Huachuca Mountains, Ramsey Canyon, 17 January 1984, Bau- mann and Nelson. Capnia elevata Frison—OREGON, Was- co County, creek, Hwy 30, 4 mi E Rowena, 3 March 1984, Baumann and Nelson. Capnia fibula Claassen— ARIZONA, Co- conino County, West Fork Oak Creek, 6 February 1984, M. W. Sanderson. Capnia manitoba Claassen— MAINE, sta. 2 Rt. 5 Saco, 3 April 1964, C. H. Nelson. Capnia melia Frison—OREGON, Clack- 52 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON amas County, Salmon River at headwaters, jet. Hwys 26 & 35, 2 March 1984, Bau- mann, Nelson, Fiala. Capnia uintahi Gaufin—UTAH, Davis County, Farmington Canyon at Fork Bridge, 5 April 1966, R. W. Baumann. In preparation for study with the scanning electron microscope, specimens were seri- ally dehydrated from their storage fluid of 70% alcohol for more than 10 minutes in 95% ethanol, 20 minutes in absolute (100%) ethanol over calcium sulfate, and stored for more than 20 minutes in acetone previously kept over calcium sulfate. The specimens were then transferred to acetone-immersed holding baskets in a Sorvall critical point drying apparatus attached to a carbon diox- ide tank. The dried abdomens of the spec- imens were then placed on aluminum stubs covered with double-sided tape. The spec- imen-bearing stubs were gold coated using a Polaron gold-coating apparatus. Coated specimens were then examined using an AMRay 1000 scanning electron microscope at 20 kV. RESULTS This study reveals that male terminalia of Capnia are well suited for examination using SEM. The epiproct, which is used in species determinations, is heavily sclero- tized and thus resistant to deformation dur- ing drying and other preparation procedures necessary for viewing with a scanning elec- tron microscope. This structural stability in the Capnia is uncommon in stoneflies since important taxonomic characters in genera in other families are usually on softer body parts that are susceptible to deformation during specimen preparation. Detailed structures not previously noted in species descriptions were seen with the SEM. A few of these features that indicate phylogenetic relationships are outlined be- low. In C. cheama (Fig. 1) and C. fibula (Figs. 4 and 12) details of the dorsal abdominal hump are shown. Both trichoid and cam- paniform sensilla are present. The paucity of trichoid sensilla on the knob of tergum seven in C. fibula compared with the nu- merous sensilla of the knob of tergum eight in C. cheama distinguishes two groups of species in the genus. In C. excavata and C. uintahi the setation of the tergal knob re- sembles that of C. cheama and the setation of C. fibula is similar to that of C. venosa, C. elevata, C. wanica, and C. manitoba. In C. fibula (Fig. 3) spines are asymmet- rical along the lateral margins of the epi- proct when seen in dorsal view. These spines are directed cephalad and would seemingly hinder insertion of the epiproct into the fe- male. Similar spines occur in several species groups of Capnia and take on a diversity of placements including the stout, tightly bunched configuration of C. decepta (Figs. 11, 13 and 15) and a scattered pattern along the anterior margin of the epiproctal bulb of C. melia (Figs. 14 and 16). Other species such as C. barberi (Fig. 9) show no indica- tion of these spines. The two-limbed epiproct of C. manitoba (Fig. 8) probably represents a plesiomorphic condition for at least one group of Capnia. The presence of a lower epiproctal limb was first noted using SEM in C. elevata (Fig. 10) and later noted using a light microscope in C. fibula, C. venosa, and C. wanica. These species show a reduced lower process and a cline featuring a gradual reduction in the length of this character as one moves away from the range of C. manitoba. No other named species of Capnia in North America has a similar two-limbed epiproct or hint of a reduced lower limb. A relatively simple tube epiproct such as that of C. coloradensis (Fig. 7) is shared by several species in the genus, but only one other, C. petila, has the apex of the epiproct drooping ventrally. The tube epiproct of some Capnia represents the most apo- morphic condition of the epiproct and is shared by several species, including C. con- VOLUME 89, NUMBER 1 53 Figs. 1-6. Male terminalia of Capnia spp. 1, C. cheama Ricker, epiproct and tergal knob, 225 x (dorsal). 2, C. cheama Ricker, tip of epiproct, 650 (anterolateral). 3, C. fibula Claassen, epiproct, 180 (dorsal). 4, C. fibula Claassen, epiproct and tergal knob, 75 x (lateral). 5, C. uintahi Gaufin, epiproct and tergal knob, 105 x (dorsal). 6, C. unitahi Gaufin, epiproct and tergal knob, 110 (lateral). fusa, C. elongata, C. gracilaria, C. lacustra, and C. uintahi demonstrate the value of C. lineata, C. promota, C. vernalis, and C. SEM in illustrating characters useful for de- zukeli. terminations at the specific level. The dis- Two closely related species, C. cheama tribution of the two kinds of sensilla on the 54 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 7-12. Male terminalia of Capnia spp. 7, C. coloradensis Claassen, epiproct and tergal knob, 105 x (lateral). 8, C. manitoba Claassen, epiproct and tergal knob 120 x (lateral). 9, C. barberi Claassen, epiproct, 90 x (lateral). 10, C. elevata Frison, epiproct, 105 (lateral). 11, C. decepta Frison, epiproct, 100 (lateral). 12, C. fibula Claassen, tergal knob, 700 = (dorsal). tergal knobs (Figs. 1 and 5) are useful in separating these two species. Higher mag- nification of the tip of the epiproct (Figs. 2 and 6) shows a dorsal appendage near the apex in C. cheama which is not present in C. uintahi. These structures are not readily apparent using a dissecting microscope and phylogenetically important information given by these characters has been unavail- able using classical techniques. VOLUME 89, NUMBER 1 14, C. melia Frison, epiproct, 150 x (dorsal). 15, C. decepta Frison, epiproct, 85 x (dorsal). 16, C. melia Frison, epiproct, 800 x (dorsal). 56 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON DISCUSSION The higher magnification and greater depth of field available make SEM a valu- able tool for investigating evolutionary and taxonomic relationships within Capnia and the potential for greater use is apparent. Comparison of minute details between taxa can aid systematists in understanding species variation and constructing accurate phylog- enies. The expense of SEM work and the often destructive manipulation of speci- mens, though, limit usefulness of SEM for routine species determination. These con- straints could result in overconfidence based on the characters observed from a single specimen rather than from a series of spec- imens. However, once a new character 1s observed using SEM more specimens may be examined using a dissecting microscope to confirm the character from a series of individuals. Although relatively low magnification was used during this preliminary study, other structures of probable taxonomic worth, such as epiproctal spines, sensilla, and se- tation patterns were observed. Once the mi- nute characters are examined from series of specimens and relationships established, routine species determinations can be per- formed using characters visible with an or- dinary dissecting microscope. ACKNOWLEDGMENTS We thank Wilford M. Hess, James V. Al- len, and Connie Swensen of the Electron Optics Laboratory at Brigham Young Uni- versity for their help with technical and crit- ical aspects of the scanning electron mi- croscopy. LITERATURE CITED Baumann, R. W. 1973. Studies on Utah stoneflies (Plecoptera). Great Basin Nat. 33: 91-108. Kapoor, N. N. and K. Zachariah. 1984. Scanning and transmission electron microscopy of the devel- opmental stages of the flower-shape sensillum of the stonefly nymph Thaumatoperla alpina Burns and Neboiss (Plecoptera: Eusthentidae). J. Insect Morphol. Embryol. 13: 177-189. Kondratieff, B. C. and R. F. Kirchner. 1984. A new species of Nemouridae (Plecoptera) from the Great Dismal Swamp, Virginia, USA. Proc. Entomol. Soc. Wash. 86: 578-581. Nelson, C. H. 1984. Numerical cladistic analysis of phylogenetic relationships in Plecoptera. Ann. Entomol. Soc. Am. 77: 466-473. Stark, B. P. 1983. A review of the genus Soliperla (Plecoptera: Peltoperlidae). Great Basin Nat. 43: 30-44. Stark, B. P. and R. W. Baumann. 1978. New species of nearctic Neoperla (Plecoptera: Perlidae), with notes on the genus. Great Basin Nat. 38: 97-114. Stark, B. P. and D. H. Ray. 1983. A revision of the genus Helopicus (Plecoptera: Perlodidae). Fresh- wat. Invertebr. Biol. 2: 16-27. Stark, B. P. and K. W. Stewart. 198i. The nearctic genera of Peltoperlidae (Plecoptera). J. Kans. Entomol. Soc. 54: 285-311. 1982a. Oconoperla, a new genus of North American Perlodinae (Plecoptera: Perlodidae). Proc. Entomol. Soc. Wash. 84: 746-752. 1982b. The nymph of Viehoperla ada (Ple- coptera: Peltoperlidae). J. Kans. Entomol. Soc. 55: 494-498. . 1982c. Notes of Perlinodes aurea (Plecoptera: Perlodidae). Ann. Entomol. Soc. Am. 75: 84-88. Stark, B. P. and S. W. Szczytko. 1981la. Contributions to the systematics of Paragnetina (Plecoptera: Per- lidae). J. Kans. Entomol. Soc. 54: 625-648. . 1981b. Skwala brevis (Kopenen) from Japan (Plecoptera: Perlodidae). Aquat. Insects 3: 61-63. 1982. Egg morphology and phylogeny in Pteronarcyidae (Plecoptera). Ann. Entomol. Soc. Am. 75: 519-529. 1984. Egg morphology and classification of Perlodinae (Plecoptera: Perlodidae). Ann. Limnol. 20: 99-104. Szczytko, S. W. and K. W. Stewart. 1979. The genus Isoperla (Plecoptera) of western North America; Holomorphology and systematics, and a new stonefly genus Cascadoperla. Mem. Am. Entomol. Soc. 32: 1-120. PROC. ENTOMOL. SOC. WASH. 89(1), 1987, pp. 57-60 TWO NEW SPECIES OF FLEXAMIA FROM THE NEBRASKA SAND HILLS (HOMOPTERA: CICADELLIDAE: DELTOCEPHALINAE)! J. E. Lowry AND H. DERRICK BLOCKER (JEL) Insect Pathology Laboratory, Agricultural Research Service, USDA, Beltsville, Maryland 20705; (HDB) Department of Entomology, Kansas State University, Manhat- tan, Kansas 66506. Abstract.—Two new species of Flexamia from perennial chloridoid grasses in the Ne- braska Sand Hills are described and illustrated. These are F. celata and F. arenicola. The previously unknown female of F. dakota is also described. The genus Flexamia is one of the most important and conspicuous groups of grass- land cicadellids. In addition to species rec- ognized by Young and Beirne (1958) in their revision, Hamilton and Ross (1975) de- scribed F. satilla, and Ross and Cooley (1969) added F. delongi. Although the latter form may be a geographic variation of san- dersi, its description raises to 36 the number of described species. In this report we de- scribed two new species and the female of dakota. The new species, F. arenicola and F. ce- lata, were discovered in sand blowouts in the Nebraska Sand Hills; they are closely related to F. flexulosa and F. stylata re- spectively, but differ in male genitalic struc- tures. F. dakota, whose female had been previously unknown, was collected from Texas to North Dakota and on west-facing slopes in the Loess hills of western Iowa. All collections of dakota were from Schi- zachyrium scoparium (Michx.) Nash. The cicadellid species we describe are named from material collected by R. F. Whitcomb. ' Contribution No. 85-535-J, Department of Ento- mology, Kansas Agricultural Experiment Station, Kan- sas State University, Manhattan, Kansas 66506. Type material is deposited in the U.S. Na- tional Museum of Natural History (USNM) and Kansas State University (KSU). Para- types are deposited also in the Grassland Cicadellid Collection of the Insect Pathol- ogy Laboratory, ARS, USDA, Beltsville Ag- ricultural Research Center-East (BARC-E), Beltsville, Maryland. Flexamia celata Lowry and Blocker, NEW SPECIES Fig. 1 Description. — Length of 6 4.4 mm, 2 4.4— 4.7 mm; head width of 4 1.3 mm, ? 1.4 mm; pronotal width of ¢ 1.2 mm, 2 1.3 mm; in- terocular width of 6 0.6 mm, 2 0.7 mm; vertex length of 6 0.7 mm, ? 0.8 mm; pro- notal length of ¢ 0.5 mm, ? 0.6 mm. Vertex not produced; median length of vertex ap- proximately 2 head width and slightly long- er than interocular width. Color pale stramineous with irregular dark markings on dorsum and forewings; vertex with black interocular line; head and prono- tum with faint tan markings as stylata; ven- ter and legs with irregular fuscous markings; 2 sternum VII with pair of conspicuous fus- cous stripes. Male pygofer with central margin strongly 58 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON D Figs 1. G Flexamia celata. A, aedeagus and connective, lateral aspect. B, same, ventral aspect. C, right style, dorsal aspect. D, male plates. E, male pygofer, lateral aspect. F, female seventh sternum. G, bases of first valvulae of female. produced to form spinelike process as sty- lata; plates extend to approximately 7 length of pygofer, fused mesally for 2 length, nar- rowed apically to rounded lateral lobe, api- ces of the two plates meeting in v-shaped notch; connective in lateral view with dorsal keels broad, approximately '/ height of dor- sal apodeme; apodemal processes straight, unmodified, in dorsal aspect parallel to ae- deagal shaft; style digitate in apical ', preapical lobe pronounced; aedeagus sym- metrical, shaft slender, not conspicuously curved, not expanded apically, central and paired apical processes approximately equal in length and less than '4 length of shaft, gonopore apical on caudodorsal surface. Female sternum VII with posterior mar- gin produced medially, with slight medial notch; ovipositor with basal processes of first valvulae recurved and extending laterad be- yond lateral margin. Types.—Holotype ¢: Garden Co., Ne- braska, Crescent Lake National Wildlife Refuge, 27-VI-1984, R. F. Whitcomb. De- posited in USNM. Paratypes: | 2, Sheridan Co., Nebraska, Lakeside, 28-VI-1984; 2 4, holotype locality, 28-VI-1984. Deposited at KSU and BARC-E. Remarks. — Flexamia celata keys to cou- plet 9 in Young and Beirne’s key to males and is apparently closely related to stylata. It can be distinguished by the shaft of the aedeagus which is slender, not conspicu- ously curved, not expanded apically, and which has shorter apical appendages. The processes at the base of the first valvulae of the ovipositor are diagnostic. Additional records: Most specimens were VOLUME 89, NUMBER 1 D 59 E G Fig. 2. Flexamia arenicola. A, aedeagus and connective, lateral aspect. B, same, ventral aspect. C, right style, dorsal aspect. D, male plates. E, male pygofer, lateral aspect. F, female seventh sternum. G, bases of first valvulae of female. taken from Redfieldia flexuosa (Thurb.) Va- sey. NEBRASKA: Box Butte, Cherry, and Hooker counties June-August. The name celata is an adjective meaning “hidden.” Flexamia arenicola Lowry and Blocker, NEw SPECIES Fig. 2 Description. — Length of 4 3.7-3.8 mm, 2 3.8—4.1 mm; head width of ¢ 1.1-1.2 mm, 2 1.2-1.3 mm; pronotal width of 4 1.0-1.1 mm, 2 1.1—1.2 mm; interocular width of 3 0.5-0.6 mm, ? 0.6-0.7 mm; vertex length of ¢ 0.6-0.7 mm, 2 0.7-0.8 mm; pronotal length of ¢ 0.4-0.5 mm, 2 0.5-0.6 mm. Me- dian length of vertex approximately 7 head width and 1'4 times interocular width. Color pale stramineous with irregular dark brown markings on forewings and apex of vertex; faint stripes on vertex and prono- tum; basal interocular lines fuscous to black; venter and legs with irregular fuscous mark- ings; 2 sternum VII with pair of fuscous spots on hind margin. Male pygofer with posterior lobe pro- duced anteroventrally, caudal margin fuscous, with or without a small acute projection apically; plates extending to ap- proximately % length of pygofer, mesal mar- gins touching for '2 length then forming con- spicuous v-shaped notch apically; style digitate apically; connective in lateral aspect with dorsal keels 2 height of dorsal apo- deme; aedeagus symmetrical, apodemal processes divergent apically, nearly attain- ing tip of shaft, gonopore apical on caudo- ventral surface, with pair of lateral apical processes approximately '4 length of shaft and curved laterodorsally, with unpaired ventral process of approximate same length directed anteriorly. 60 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON A B Fig. 3. Flexamia dakota, female. A, seventh ster- num. B, bases of first valvulae. Female sternum VII with posterior mar- gin produced medially, with slight median notch; ovipositor with recurved processes at base of first valvula not exceeding lateral margin, processes extending caudad, sin- uate and digitate apically. Types.—Holotype ¢: Garden Co., Ne- braska, Crescent Lake National Wildlife Refuge, 27-VI-1984. Deposited at USNM. Paratypes: 40 4, 15 2, same data as holotype; 1 2, Sheridan Co., Nebraska, 28-VI-1984: 1 6, Cherry Co., Nebraska, Brownlee, 9-VIII- 1977. Deposited at KSU, BARC-E. Flexamia arenicola is apparently closely related to flexulosa but can be distinguished by the expanded dorsal keels of the con- nective, the more robust aedeagal shaft and the longer aedeagal processes that are pres- ent in arenicola. It keys to couplet 17 in Young and Beirne’s key to males. It can be distinguished from modica and texana by its larger size, and its nearly symmetrical aedeagus with lateral apical processes that are dorsally curved. The processes at the base of the first valvulae of the ovipositor are diagnostic. Additional records. — All specimens were taken from Muhlenbergia pungens Thurb. NEBRASKA: Lincoln, McPherson, and Box Butte counties; COLORADO: Morgan Co., June-August. The name arenicola is a noun in appo- sition meaning “‘sand inhabitant.” Flexamia dakota Young and Beirne Fig. 3 Description of 2.—Length 3.1-3.2 mm; head width 0.9-1.0 mm; pronotal width 0.8- 0.9 mm; interocular width 0.4—0.5 mm; ver- tex length 0.6—0.7 mm; pronotal length 0.3- 0.4 mm. Median length of vertex approxi- mately 73 head width and 1'/ times inter- ocular width. Sternum VII with posterior margin produced with slight medial con- cavity; Ovipositor without basal processes. Remarks.— Young and Beirne did not separate the female of dakota from that of sandersi (see also delongi of Ross and Coo- ley, 1969). The females described here were found in association with males of dakota. Additional records.— All specimens tak- en from Schizachyrium scoparius. IOWA: Harrison, Monona, Pottawattomie, and Woodbury counties; NEBRASKA: Cherry, Lincoln and McPherson counties; WYO- MING: Campbell, Platte, and Weston counties; TEXAS: Roberts and Travis counties; OKLAHOMA: Alfalfa Co., June- August. ACKNOWLEDGMENTS The authors acknowledge the assistance of R. F. Whitcomb who collected the ma- terial used in this paper and initiated this collaboration. We also thank W. E. Steiner and A. L. Hicks for advice and assistance. LITERATURE CITED Hamilton, K. G. A. and H. H. Ross. 1975. New species of grass-feeding deltocephaline leafhoppers with keys to the Nearctic species of Palus and Rosenus (Rhynchota: Homoptera: Cicadellidae). Can. Entomol. 107: 601-611. Ross, H. H. and T. A. Cooley. 1969. A new Nearctic leafhopper of the genus Flexamia (Hemiptera: Ci- cadellidae). Entomol. News 80: 246-248. Young, D. A. and B. P. Beirne. 1958. A taxonomic revision of the leafhopper genus Flexamia and a new related genus (Homoptera: Cicadellidae). USDA Tech. Bull. 1173, 53 pp. PROC. ENTOMOL. SOC. WASH. 89(1), 1987, pp. 61-73 CHANGES IN NOMENCLATURE AND CLASSIFICATION OF SOME OPIINE BRACONIDAE (HYMENOPTERA)! ROBERT A. WHARTON Department of Entomology, Texas A&M University, College Station, Texas 77843. Abstract. —Lectotypes are designated for 13 species of opiine Braconidae and a neotype is designated for Bracon carbonarius Nees von Esenbeck, 1834. Three genera are trans- ferred to the Opiinae: Baeocentrum Schulz and Coeloreuteus Roman from the Rogadinae, and Neodiospilus Szépligeti from the Helconinae. All are synonyms of Opius Wesmael. Diachasmimorpha Viereck and Psyttalia Walker are elevated to generic rank in the Opi- inae, and Austroopius Szépligeti is placed as a subgenus of Psyttalia. Rhynchosteres (Fopius), New Subgenus, is described for several Ethiopian species previously referred to Biosteres. Four species of Opius are renamed, one new species, Rhynchosteres silvestrii, parasitic on Tephritidae is described, and 12 taxa are newly synonymized. Repositories are listed for the types of three species previously treated as nomina nuda or nomina dubia. Fischer (1963a, 1966, 1972, 1977) re- vised the opiine Braconidae from all major geographic regions of the World, and pub- lished a catalog of the known species (Fisch- er, 1971). Over the past 30 years, Fischer has also redescribed almost all of the older opiine species, thus providing a uniform format for comparison. His works are of particular value in pinpointing repositories for type specimens, and in clarifying the sta- tus of many uncertain species. A few prob- lems still remain, however (e. g. Fischer, 1971, pp. 130-132), and the purpose of the present work is to solve some of these. The following abbreviations are used for museums: IRSN, Institut Royal des Sci- ences Naturelles de Belgique, Bruxelles (Wesmael Collection); TAMU, Texas A&M University, College Station; ZMHB, Zoo- logisches Museum der Humboldt Univer- sitaet, East Berlin (Foerster Collection); ' Approved as TA #21733 by the Texas Agricultural Experiment Station. BMNH, British Museum (Natural History), London; CNC, Canadian National Collec- tion, Ottawa; NRS, Naturhistoriska Riks- museet, Stockholm; WIEN, Naturhistorish- es Museum, Vienna; USNM, U.S. National Museum of Natural History, Washington, D.C. All of the lectotypes designated herein have been so labelled by me. Baeocentrum Schulz, 1911 Szépligeti (1907) established the mono- typic genus Brachycentrus for a species col- lected by M. Rothschild in British East Af- rica. Szépligeti placed this genus in the Rogadinae. Schulz (1911), noting that Bra- chycentrus was preoccupied, renamed Szé- pligeti’s genus as Baeocentrum. The type species, minutus Szépligeti, 1907, was de- scribed from two specimens. One of these, bearing Rothschild’s blue locality label, a printed type label, and Szépligeti’s hand- written type label, is in the Muséum Na- tional d’Histoire Naturelle, Paris. I hereby designate this specimen as lectotype. The specimen matches Szépligeti’s original de- 62 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON scription, except that the specimen is a male and Szépligeti’s description is of a female. The male genitalia are strongly protruding in this specimen, however, and this feature apparently misled Szépligeti. The paralec- totype (with metasoma missing) is conspe- cific, and located in the Hungarian Natural History Museum, Budapest. This species is clearly a member of the genus Opius s. L., as characterized by Fischer (1972). I there- fore consider Baeocentrum to be a junior synonym of Opius, New Synonym. Opius minutus (Szépligeti), New Combination, (not minutus Granger, 1949, see below) fits the characterization of the subgenus Phlebose- ma Fischer, 1972. In Fischer’s (1972) clas- sification, Phlebosema thus becomes a ju- nior subjective synonym of Baeocentrum. Coeloreuteus Roman, 1910 This genus was established by Roman (1910) for Atoreuteus africanus Szépligeti, 1908, and placed in the Exothecinae. Shene- felt (1975) retained it in the Rogadinae. I have examined the holotype of africanus (in the NRS), and it belongs in Opius s. 1. (as characterized by Fischer, 1972). The ab- sence of the mid-dorsal portion of the oc- cipital carina, the flat, exposed labrum, and the fore wing venation all support this place- ment. I therefore consider Coeloreuteus to be a junior synonym of Opius, New Syn- onym. I have not examined the other species listed under Coe/oreuteus (Shenefelt, 1975) to determine their placement. Diachasmimorpha Viereck, 1913 Gahan (1915) synonymized Diachasmi- morpha with Opius. Fullaway (1951) was the first to subsequently recognize this dis- tinctive group of tephritid parasitoids, which he called the Opius longicaudatus group (or group 1). Fullaway (1951) characterized this group by the presence of a postnervellus in the hind wing and the deep but unsculptured notauli. Wharton and Gilstrap (1983) re- ferred to this as the ¢ryoni species group, transferred several species to it, and treated some of those included by Fullaway (1951) as synonyms of longicaudatus Ashmead. Wharton and Gilstrap (1983) further char- acterized this group on the basis of the api- cally sinuate ovipositor. Diachasmimorpha tryoni (Cameron), New Combination, is clearly a member of this group based on the shape of the ovipositor, wing venation, body sculpture, and shape of the clypeus, despite the almost complete absence of an occipital carina. Fischer (1967a) used the absence of an occipital carina as the basis for his genus Parasteres. Since the type species of Par- asteres is a synonym of tryoni (Wharton and Marsh, 1978), however, Parasteres becomes a junior subjective synonym of Diachas- mimorpha, New Synonym. The genus Di- achasmimorpha is very similar to the Neo- tropical genus Doryctobracon Enderlein, 1920, based on similarities in clypeal mor- phology (lower margin of clypeus more or less sinuate, ventral-lateral margin of clyp- eus completely separated from face ventrad of the anterior tentorial pit). Doryctobracon lacks the sinuate ovipositor of Diachasmi- morpha, and generally has the recurrent vein antefurcal to interstitial rather than the postfurcal insertion of Diachasmimorpha. The clypeal morphology and fore wing ve- nation are, in my opinion, sufficient for re- moving Diachasmimorpha from Biosteres, where it has been placed by all recent au- thors (e.g. Fischer, 1972; Wharton and Marsh, 1978). Hexaulax Cameron, 1910 Fischer (1971, p. 130) was unable to place this monotypic genus, stating only that it belonged ‘to the Opiinae according to Muesebeck.’’ Muesebeck (1967, p. 48), however, treated Hexaulax as a synonym of Opius on the authority of Fischer (1964). A clearly labelled specimen of Hexaulax ruf- iceps Cameron, 1910 (type species by monotypy of Hexaulax), collected by Fruhstorfer and from the type locality in VOLUME 89, NUMBER 1 Java, and bearing a co-type label, is now in the BMNH. This specimen belongs to Opius s. 1, as interpreted by Fischer (1972), but cannot readily be assigned to a subgenus because a pin through the mesonotum makes it impossible to determine whether or not a midpit is present. Cameron stated in his original description that the occiput was not margined. The apparent absence of an oc- cipital carina is an artifact, however, due to the head being pinched in the occipital re- gion. Traces of the carina can still be seen on close inspection. Because of this occipital feature, the matching locality label, and the lack of evidence in the original description that more than one specimen was involved, this specimen may be the holotype. How- ever, in order to alleviate any confusion in the future, I am treating it as a lectotype, and hereby designate it as such. Lytacra Foerster, 1862 Fischer (1959) stated that the type of Ly- tacra stygia Foerster, the type species of Ly- tacra, was lost. Fischer (1971) repeated this statement and placed stygia in Ademon Hal- iday, 1833 based on Foerster’s original de- scription. A single female from Foerster’s collection, labelled Lytacra stygia, is now present in the ZMHB; and I believe this to be the specimen on which Foerster (1862) based his description. The tips of the wings are missing and Foerster’s characterization of the radial cell incompletely closed (and thus similar to the condition in Ademon) is therefore misleading (and in fact incorrect). Foerster’s specimen is identical to the lec- totype and paralectotype of Diachasma caf- fer (Wesmael) in the IRSN. I therefore con- sider stygia to be a junior synonym of caffer, New Synonym, and Lytacra thus becomes a synonym of Diachasma. Neodiospilus Szepligeti, 1911 Szépligeti (1911) included two species (terebrator Szépligeti, 1911 and flavipes Szé- pligeti, 1911) in his original description of Neodiospilus. Brues (1926) subsequently 63 designated flavipes as the type species. Shenefelt (1970), following Szépligeti (1911), placed Neodiospilus in the Diospilini, and listed the holotype of flavipes as being in the NRS. A search for this specimen in Stock- holm in 1984 was unsuccessful. However, a female collected by Fuelleborn from the type locality in Langenburg on Lake Nyasa is now in the ZMHB. It bears a red type label, and a determination label in Szépli- geti’s handwriting which reads “Neodio- spilus luteipes m”’; and it matches Szépli- geti’s original description of flavipes. In the drawer near this specimen is an old note saying the label = luteipes but the MS = flavipes. In the same drawer are the male and female syntypes of Neodiospilus tere- brator, both with Szépligeti determination labels. I have no doubt that these are the specimens used by Szépligeti as the basis for his description of Neodiospilus, and consid- er the specimen labelled as /uteipes to be the holotype of flavipes. Both flavipes and ter- ebrator are opiines, and belong in Opius s. /. as interpreted by Fischer (1972). Neo- diospilus thus becomes a junior synonym of Opius, New Synonym. There are pins ob- literating the mesonotal regions of the /fla- vipes and terebrator specimens. The two species thus cannot readily be assigned to a subgenus, though it is clear from the clypeal morphology and wing venation that they should belong to different subgenera in Fischer’s (1972) classification. Szépligeti (1914), in an apparent lapsus, described another genus with the name Neo- diospilus. The type series for the two in- cluded species, baumanni Szépligeti and zenkeri Szépligeti, are also in the ZMHB. Shenefelt (1970) renamed Neodiospilus Szé- pligeti, 1914 as Repetiodiospilus. It is cor- rectly placed in the Diospilini (as presently understood). Psyttalia Walker, 1860 Muesebeck (1931) synonymized Psyttalia with Opius, and renamed the type species walkeri Muesebeck. Fischer (1971) listed 64 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Psyttalia under Opius, but inadvertently omitted walkeri from his species catalog. Fischer (1972) treated Psyttalia as a sub- genus of Opius. Psyttalia walkeri belongs to the concolor species group as characterized by Wharton and Gilstrap (1983). As noted by Wharton and Gilstrap (1983), the con- color and fletcheri species groups share a number of characters suggestive of a close relationship. In addition to characters noted by Fischer (1972), these include reduced mesonotal sculpture, large, attenuate hy- popygium, short clypeus, exposed labrum with reduced setation, and identical hind wing venation. I believe these characters are sufficient to elevate Psyttalia to generic rank in the Opiinae, with Austroopius Szépligeti, 1900 included as a subgenus. Members of the subgenus Psyttalia (= concolor species group) generally have the gaster weakly sculptured, at least basally, while in the sub- genus Austroopius (= the fletcheri species group) the recurrent vein is somewhat bowed. I do not consider the thickening of the first cubital cross-vein adequate for sep- arating the species formerly included in Austroopius (e. g. Fischer, 1963b, 1966) from closely related species, such as fletcheri Sil- vestri and incisi Silvestri, which lack this thickening. Opius africanus Szepligeti, 1910 With the transfer of Coeloreuteus to Opius (see above), Opius africanus (Szépligeti, 1908) becomes a senior secondary hom- onym of Opius africanus Szépligeti, 1910. The latter is therefore renamed Opius teph- ritivorus, New Name. Opius tephritivorus 1s a member of the subgenus Utefes Foerster, as characterized by Fischer (1972), and is at least subgenerically different from afri- canus. Pachythecus albobalteatus Cameron, 1912 This species was transferred to Biosteres by Fischer (1967b). The sculpture and gen- eral color pattern of the male holotype in the BMNH (BM type # Hym. 3.c.711) are the same as that of angaleti Fullaway, 1952. Although the mid coxae tend to be darker in most specimens of angaleti that I have seen, I can find no other differences, and therefore treat angaleti as a junior subjec- tive synonym of albobalteatus, New Syn- onym. The New Combination is Diachas- mimorpha albobalteatus (Cameron). The type locality of albobalteatus is “Borneo,” that of angaleti North Borneo (= Sabah). Diachasma brevistyli Paoli, 1934 Fischer (1971) treated brevistyli as a no- men nudum, referring only to a subsequent note on brevistyli by Martelli (1937). Through the courtesy of the librarians at the Museum “‘La Specola’’ in Firenze, Italy, I was able to locate Paoli’s (1934) orginal description, and therefore consider it (as well as Phanerotoma somalica Paoli, 1934) to be a validly described species. The type se- ries of brevistyli is in the Museum “‘La Spe- cola.”” This species is almost identical to Biosteres carinatus Szépligeti, 1910; and the generic placement of these two species is currently under investigation. Opius caffer Wesmael, 1835 Wesmael (1835) described caffer on the basis of two specimens, a male and a female. Both are in excellent condition in the IRSN. The female is hereby designated as lecto- type. It bears the following labels: top label: Coll. Wesmael (printed); 2nd label: 1865 (printed); 3rd label: Opius caffer mihi (hand- written) dét. C. Wesmael (printed); 4th la- bel: Type (printed in red). Bracon carbonarius Nees von Esenbeck, 1834 See procerus Wesmael (below). Biosteres carinatus Szepligeti, 1910 I have examined the holotype male in the NRS and am unable to separate this species from Hedylus giffardii Silvestri (as charac- VOLUME 89, NUMBER 1 terized by Wharton and Gilstrap, 1983). I therefore consider giffardii to be a junior subjective synonym of carinatus, New Syn- onym. See additional comments under brevistyli (above). Opius cingulatus Wesmael, 1835 This is the type species (by original des- ignation) of Nosopoea Foerster, 1862. Wes- mael (1835) described cingulatus on the ba- sis of 14 specimens. Eleven specimens, bearing identical type and determination la- bels, but representing at least three species, are now present in the IRSN. I hereby des- ignate as lectotype one of four females most closely matching the original description, and bearing the following labels: top label: 1832 (printed); 2nd label: Coll. Wesmael (printed); 3rd label: Type (printed in red); 4th label: 6 Opius 2 cingulatus. mihi (hand- written) dét. C. Wesmael (printed). The lec- totype agrees with Fischer’s (1972) defini- tion of cingulatus; and also fits the concept of cingulatus used by Haliday (1837), Foers- ter (1862), Marshall (1891) and Thomson (1895). The lectotype has the hypostomal and occipital carinae widely separated at the mandible, and the mandible bears a carinate ridge basally on its ventral border. Paralec- totypes which are not conspecific have the oral and occipital carinae meeting near the base of the mandible, the sternaulus at least weakly sculptured, and/or a broad basal lobe ventrally on the mandible (as in O. pallipes Wesmael or O. crassipes Wesmael). Opius coffeae Fischer, 1962 I have been unable to locate the holotype of coffeae, but a paratype in the Naturhis- torisches Museum, Basel is identical to the lectotype of Biosteres caudatus Szépligeti, 1913. I therefore treat coffeae as a junior subjective synonym of caudatus, New Syn- onym. If the holotype is ever located, it should be examined to confirm this syn- onymy. This species is characterized by the narrow patch of deep punctures between eye 65 and ocelli (Wharton and Gilstrap, 1983, Fig. 12). Generic placement 1s discussed below under si/vestrii, n. sp. Opius comatus Wesmael, 1835 This is the type species (by original des- ignation) of Holconotus Foerster, 1862. Ashmead (1900) later proposed the name Aulonotus for Foerster’s Holconotus, which was preoccupied. Wesmael (1835) de- scribed comatus on the basis of 11 speci- mens. There are now eight specimens in IRSN, two of which fit the description of Wesmael’s var. |. I hereby designate as lec- totype of comatus the female bearing R. Koenig’s 1968 lectotype label. Koenig un- fortunately never published a lectotype des- ignation for this species. The specimen has the following additional labels: top label: Coll. Wesmael (printed); 2nd label: 1860 (printed); 3rd label: ¢ Opius 2 comatus. mihi (hand-written) dét. C. Wesmael (printed); 4th label: Type (printed in red); Sth label: R. I: Se: N: .B. 1: ¢::3.3173 6th label: LEC- TOTYPE Opius comatus Wesmael 2 dé- signé par R. Koénig 1968. There are two more females bearing type and comatus determination labels (includ- ing the additional specimen of var. 1) than listed by Wesmael (1835) in the original de- scription. It is not possible at present to determine whether specimens were added after the original description, or Wesmael erred in listing the sexes. Labelling of para- lectotypes has therefore been conservative. Opius crassipes Wesmael, 1835 This is the type species (by original des- ignation) of Hypocynodus Foerster, 1862. Wesmael (1835) described crassipes on the basis of a single female, and also indicated that he had a male which possibly belonged to this species. In the Wesmael Collection there are now two females plus a third spec- imen lacking an abdomen. All bear Opius crassipes type labels. The two intact females are not conspecific, however, and I have 66 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON labelled the one that fits Wesmael’s original description as the holotype. Neodiospilus flavipes Szepligeti, 1911 Since flavipes is preoccupied in Opius by flavipes Szépligeti, 1898, it becomes nec- essary to rename the junior homonym fol- lowing transfer of the type species of Neo- diospilus to Opius (see above). It is therefore renamed Opius sepivalfus, New Name. The specific epithet is an arbitrary combination of letters. Opius irregularis Wesmael, 1835 This is the type species of A//otypus Foers- ter, 1862. Wesmael (1835) described Opius irregularis on the basis of one male and five females. Six specimens (2 4, 4 2) matching his original description and bearing iden- tical type and determination labels are in the IRSN. These specimens also bear the lectotype and paralectotype labels of R. Koenig. Unfortunately, Koenig’s lectotype designation has never been published. Koe- nig labelled a male as lectotype, but I believe the female offers better characters for species discrimination. I therefore designate as lec- totype a female with the following labels: top label: Coll. Wesmael (printed); 2nd la- bel: 1847 (printed); 3rd label: ¢ Opius ¢ ir- regularis mihi (hand-written) dét. C. Wes- mael (printed); 4th label: Type (printed in red); 5th label: PARALECTOTYPE Opius irregularis Wesmael ? désigné par R. K6nig 1968. Opius levis Wesmael, 1835 This is the type species of the subgenus Opiothorax Fischer, 1972. Wesmael (1835) described Opius levis on the basis of seven specimens. There are now six specimens in the IRSN. I hereby designate as lectotype the female labelled by Koenig in 1968 as lectotype (but never published). The spec- imen has the following additional labels: top label: Coll. Wesmael (printed); 2nd label: 1835 (printed); 3rd label: Opius levis mihi. 6. 2 (hand-written) dét. C. Wesmael (print- ed); 4th label: Type (printed in red); Sth label: R. I. Sc. N. B. I. G. 3.317; 6th label: LECTOTYPE Opius levis Wesmael ¢ dé- signé par R. K6nig 1968. Fischer (1972) rec- ords this as a color-variable species, but specimens with dark hind coxae will not run to /evis in his key (Fischer 1972, p. 441). Wesmael (1835) specifically mentions the dark coxae in his original description; all members of the type series have this feature. Rhogadopsis miniacea Brethes, 1913 This is the type species (by monotypy) of Rhogadopsis Bréthes, 1913. It was trans- ferred to Opius by de Santis (1967). I des- ignate as lectotype a female in the Museo ‘Bernardino Rivadavia’’ (Buenos Aires) with a single label in Bréthes’ handwriting which reads Rhogadopsis miniacea Br. The species runs to the tucumanus-group of Lis- sosema in Fischer (1977). Rhogadopsis thus becomes a senior synonym of the subgenus Lissosema Fischer, 1972, New Synonym. Opius minutus Granger, 1949 It becomes necessary to rename the junior homonym following transfer of Brachycen- trus minutus Szépligeti, 1907 to Opius (see above). Opius minutus Granger is therfore renamed Opius gregnar. The specific epithet is an arbitrary rearrangement of the name Granger. Opius pallipes Wesmael, 1835 This is the type species (by subsequent designation of Muesebeck and Walkley, 1951), of Opius Wesmael, 1835. Wesmael (1835) described this species on the basis of 15 females and five males. There are now 19 specimens in the IRSN. I designate as lectotype a female with abdomen mounted beneath the rest of the specimen on a sep- arate card, and characterized by an excep- tionally smooth petiole and propodeum, 24 segmented antenna (right side, left side bro- ken), and pale clypeus. This characteriza- tion fits just within the range of variation given for this species by Wesmael. The lec- VOLUME 89, NUMBER 1 totype was selected specifically on the basis of the sculptural characters of the petiole and propodeum so as to more clearly define this species. It should be pointed out, how- ever, that studies on intraspecific variation in sculptural features have not been con- ducted for this species and are desperately needed. The diverse array of host records for pallipes (e. g. Fischer, 1971) suggests that some confusion has existed in the past re- garding the application of this name. It 1s hoped that restriction of the definition of pallipes by means of this lectotype desig- nation will enable future workers to more accurately identify this species; and that this in turn will lead to a re-examination of pub- lished host records. The species 1s of poten- tial use in biological control of Liriomyza (Hendrikse, 1980). The lectotype bears the following labels: top label: Coll. Wesmael (printed); 2nd label: 1831 (printed); 3rd la- bel: 6 Opius 2 pallipes mihi (hand-written) dét. C. Wesmael (printed); 4th label: Type (printed in red). Opius procerus Wesmael, 1835 Wesmael (1835) described this species from two individuals. There are still two specimens (both male) of this species in the IRSN labelled as types. I hereby designate as lectotype the male bearing R. Koenig’s Opius carbonarius (Nees) determination la- bel. It has the following additional labels: top label: Coll. Wesmael (printed); 2nd la- bel: 1872 (printed); 3rd label: Opius pro- cerus. mihi ¢ (hand-written) dét. C. Wes- mael (printed); 4th label: Type (printed in red). I designate this specimen as also the neotype of Bracon carbonarius Nees von Esenbeck, 1834. The vast majority of Nees von Esenbeck’s collection, including the type(s) of carbonarius, have been destroyed (e. g. Shenefelt, 1970; Papp, 1985; and Van Achterberg, pers. comm.). Haliday (1837) treated procerus Wesmael as a synonym of carbonarius Nees, and I consider Haliday as the first reviser relative to the establish- ment of a fixed concept for carbonarius. Al- 67 though this concept is in agreement with the latest treatment of carbonarius (Fischer, 1977), at least two major authors (Wesmael, 1835; Thomson, 1895) have interpreted carbonarius differently. Bracon carbonarius is the type species of Biosteres Foerster, 1862. The subgeneric concepts used by Fischer (1972) for Biosteres are based on Haliday’s interpretation of carbonarius, and are therefore in conflict with the interpre- tations of Wesmael and Thomson. A neo- type designation for carbonarius will resolve this conflict. Fischer (1972, 1977) has pre- sented keys for the separation of carbon- arius from closely related species. Opius ruficeps Wesmael, 1835 This is the type species (by original des- ignation) of Therobolus Foerster, 1862. Wesmael (1835) described this species from two specimens. There are now three speci- mens in the Wesmael Collection, all bearing the identical type and determination labels noted above for other Wesmael species. I hereby designate as lectotype a female hav- ing an additional label with the hand-writ- ten number 3 on it. Hexaulax ruficeps Cameron, 1910 Transfer of the type species of Hexaulax to Opius results in a secondary homonym. I therefore rename ruficeps Cameron (not ruficeps Wesmael, 1835) as Opius indenta- tus, New Name, in reference to the damaged lectotype and misleading original descrip- tion (Cameron, 1910) resulting therefrom. Celiestiella testaceipes Cameron, 1903 This is the type species (by monotypy) of Celiestiella Cameron, 1903. Cameron (1903) mentioned only the male sex in his ex- tremely brief description of this species. However, the clearly labelled type series in the BMNH (BM type # Hym 3.c.713) con- sists of two males and one female. Of the two males, one is damaged by the pin. I therefore designate the other male as lec- totype. Fischer (1967b) redescribed this 68 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON species from the female, and transferred it to Biosteres. I agree with this placement. Opius testaceus Wesmael, 1838 This is the type species (by original des- ignation) of Utetes Foerster, 1862. Wesmael (1838) described this species from four fe- males and one male, but there are now only three specimens in the IRSN. I hereby des- ignate as lectotype the remaining intact fe- male. It bears the following labels: top label: 119. (hand-written); 2nd label: Coll. Wes- mael (printed); 3rd label: Type (printed in red); 4th label: O. testaceus. mihi (hand- written) dét. C. Wesmael (printed); 5th la- bel: cf. Opius (hand-written) Rév. J. A. Mar- shall (printed) 1887 (hand-written). Biosteres testaceus Szepligeti, 1914 I have examined the holotype female in the ZMHB, and am unable to separate this species form Biosteres fullawayi Silvestri, 1913. I therefore treat testaceus Szépligeti as a junior subjective synonym of fullawayi, New Synonym. This species was recently diagnosed by Wharton and Gilstrap (1983). It’s generic placement is currently under 1n- vestigation. DESCRIPTION OF NEw TAXA Most of the Ethiopian and Indo-Pacific species currently placed in Biosteres are not readily accommodated by the existing op1ine classification. Most authors, following Fischer (1963a, 1966), have placed the in- cluded species in Biosteres on the basis of the short second cubital cell. However, this feature is inadequate for characterizing opiine genera. Closer examination of the wing venation shows that there are signifi- cant differences between Biosteres s. s. and these Ethiopian and Indo-Pacific species. The most obvious difference is in the shape of the stigma. It is longer and narrower in Biosteres s. s., with the first radial segment arising basad of the midpoint. The post- nervellus is also greatly reduced in Biosteres s. s., and is often only barely visible as a weakly infumate crease. Other major dif- ferences are in the shape of the mandible, which has a basal tooth in Biosteres s. S.; and in the host preferences. Some of the Indo-Pacific species formerly placed in Biosteres have now been trans- ferred to Diachasmimorpha (see above). Based on the differences enumerated in the preceding paragraph, I believe most of the remaining Indo-Pacific and Ethiopian species are more closely related to Diachas- mimorpha and Rhynchosteres Fischer, 1965 than to Biosteres s. s. 1 prefer to retain Di- achasmimorpha as restricted by the defi- nition given earlier in this paper. Since the new species described below does not fit this restricted definition, I have placed it in Rhynchosteres. Rhynchosteres Fischer, 1965 Fopius Wharton, NEw SUBGENUS Type species: Rhynchosteres silvestrii, new species. Occipital carina absent dorsally, but well- developed laterally; oral and hypostomal carinae widely spearated at mandible. La- bral setae sparse, confined to ventral and lateral margins; labrum concealed by large, somewhat hemispherical clypeus with con- vex to nearly truncate lower margin; margin of clypeus extending ventrally from anterior tentorial pit somewhat reflected, and sepa- rated from gena by a sharp groove. Man- dibles gradually and evenly tapering distal- ly, without basal tooth or lobe, not deflected ventrally; ventral tooth shorter than dorsal tooth. Apical antennal segment with spine- like tip. Genal sulcus shallow, often ob- scured by facial sculpture. Second radial segment shorter than Ist cubital cross-vein; recurrent vein antefurcal to interstitial; first radial segment arising from or distad of mid- stigma; stigma discrete, not merging im- perceptibly with metacarpus distally; post- nervellus long, distinctly sclerotized and pigmented; radiella absent at base, often in- dicated by fold or infumate trace distally. VOLUME 89, NUMBER 1 Pronotum largely unsculptured and without median pit. Sternaulus crenulate; notauli complete to midpit and crenulate through- out. Postpectal carina usually present an- teriorad mid coxae. Claws simple, with broad base. Petiole with deep glymma; dor- sope present or absent. Spiracles of terga 2 + 3 near lateral margins (tergum 3 spi- racle often not visible in dorsal view due to postmotem collapse of tergum). Hypopyg- ium attenuate. Known species are parasit- oids of tephritids, and have the ovipositor at least 1.5 times length of mesosoma. Diagnosis.—Rhynchosteres differs from other large tropical opiines with short sec- ond cubital cell and broad, discrete stigma as follows: the occipital carina is absent in Doryctobracon, the ovipositor is sinuate in Diachasmimorpha, and the mandible has a well-developed flange along the ventral bor- der; the pronotum has a large median pit in Pseudorhinoplus Fischer, 1972. Rhynchos- teres was described for species with a dis- tinctively protruding and medially reflected clypeus (Fischer, 1965; Van Achterberg, 1983). In Rhynchosteres (Fopius), the clyp- eus does not protrude in this fashion. Nevertheless, some of the described species have the clypeus thickened ventral-medi- ally, and I believe this is the initial step in the transformation to the condition found in Rhynchosteres s. s. Once the Afrotropical fauna becomes better known, I suspect the distinction between Rhynchosteres s. s. and R. (Fopius) will be less obvious. The per- sulcatus species group (Fullaway, 1951; Wharton and Gilstrap, 1983) is somewhat intermediate between Diachasmimorpha and R. (Fopius); and needs more detailed study before it can be accurately placed. I include the following species in Rhyn- chosteres (Fopius): bevisi (Brues), New Combination, caudatus (Szépligeti), New Combination, desideratus (Bridwell), New Combination, niger (Szépligeti), New Com- bination, ottotomoanus (Fullaway), New Combination, and pyknothorax (Fischer), New Combination. At least three species 69 groups are evident, based on clypeal mor- phology and sculpture of the petiole and frons. I transfer also c/ypeatus (Bridwell) to Rhynchosteres s. s., New Combination. The subgeneric name is masculine and is a contraction formed from the words Fisch- er and Opius. The type species has been misidentified in the past as Biosteres caudatus, and has recently been referred to as caudatus auct. (Wharton and Gilstrap, 1983; Steck et al., 1986). I have been unable to find an avail- able name for this species, and it is therefore described as new. Rhynchosteres (Fopius) silvestrii Wharton, NEw SPECIES Figs., 1.2 Female.— Head: 1.66 + 0.06 times broader than long; 1.35 + 0.05 times broad- er than mesonotum; face with deep punc- tures and well-developed midridge, the lat- ter extending between antennal bases as a low, sharp ridge; frons varying from nearly unsculptured in small individuals to exten- sively rugosopunctate (variation as in Wharton and Gilstrap, 1983; Figs. 10, 11); ocellar triangle margined, at least in part, and often completely, by a crenulate sulcus. Ventral margin of clypeus weakly convex, not noticeably thickened in middle. Eyes bare or apparently so, large, 2.36 + 0.43 times longer than temples; temples receding in dorsal view. Antenna roughly 3.4 times longer than mesosoma; Ist flagellomere subequal to 2nd. Mesosoma: 1.28 + 0.05 times longer than high, 1.67 + 0.04 times longer than broad. Median mesonotal lobe with 2 parallel, ru- gosopunctate grooves anteriorly; lateral me- sonotal lobes hairy, and usually with scat- tered, deep punctures; with setae confined to margins in smaller species; notauli meet- ing in a narrow midpit, midpit varying from discrete furrow bordered anteriorly by striae to a small rugosostriate patch of sculpture. Propodeum rugose, sculpture variable, but often with obvious transverse elements on 70 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. anterior face, sculpture on posterior, decli- vous face more uniform and slightly weak- er; flange-like midridge present anteriorly. Sternaulus broad, deep, crenulate, but usu- ally absent over posterior one-third; cren- ulate sculpture extending dorsally along an- terior border of mesopleuron through subalar depression; posterior margin of me- sopleuron noticeably crenulate ventrad of speculum, suture unsculptured or nearly so dorsad of speculum; mesopleural disc hairy; postpectal carina usually well-developed mid-ventrally. Wing: 2nd radial segment 2.23 + 0.29 times longer than Ist; 1st cubital cross-vein 1.22 + 0.09 times longer than 2nd radial segment; 3rd radial segment extending near- ly to wing tip; cubitus arising from about anterior 0.15 of basal vein, with Ist segment Fore and hind wing of Rhynchosteres (Fopius) silvestrii. sinuate; nervulus inclivous, usually post- furcal by less than its length, more rarely interstitial; junction between nervulus and basal vein thickened; recurrent vein ante- furcal to interstitial; parallel vein arising be- low middle of brachial cell. Postnervellus reclivous, though usually weakly recurved posteriorly, extending nearly to posterior margin as a well-developed, deeply im- pressed crease which is usually weakly pig- mented. Metasoma: Petiole 1.00 + 0.05 times longer than apical width, apex 2.04 + 0.14 times wider than base; finely striate, with a stronger median longitudinal carina some- times evident; dorsal carinae well-devel- oped basally, weak over posterior half, but usually extending to posterior margin, very weakly converging (at posterior margin, dis- VOLUME 89, NUMBER 1 tance between carinae usually equal to or slightly less than distance to lateral margin); dorsope well-developed. Metasoma un- sculptured beyond petiole. Ovipositor tip not narrowed, with barely developed dorsal notch and weak ventral serrations; 2.56 + 0.15 times longer than mesosoma; ovipos- itor sheath dorsally with 2 rows of setae, each row with about 35 setae, setae shorter and about twice the density ventrally. Color: Quite variable, generally reddish- brown to black; palps white to pale yellow; scape, pedicel, mandibles (except tips), all legs, and terga 2 + 3 yellow (remaining terga usually darker); clypeus, gena, orbits, pronotum, and antenna variable, but usu- ally at least partly yellow; wings hyaline. Male.—Somewhat darker than female, with all metasomal terga brown to black (rarely pale at extreme base of tergum 2). Otherwise essentially as in female, but with mesonotum slightly narrower, midpit often weakly developed to more or less absent, and petiole 1.21 + 0.07 times longer than apical width (and thus distinctly longer and narrower than in female). Length (exclusive of ovipositor): 2.0-3.5 mm. Material examined.— Holotype 2, CAM- EROON: Centre-Sud Province, Nkolbis- son, 22.VIII.1982, G. Steck, reared ex ripe coffee berries (USNM). Paratypes (TAMU, CNC, WIEN): Same locality, host plant, and collector as holotype: IV.18.1982 (1 9), Wel982 (1 3), 22.Vill.1982 (9. 2,,,4 <3); 17.1X.1982 (2 9, 12), 26.1X.1982 (2 9, 1 3), IX.1982 (5 9, 2 6), 3.X.1982 4 9, 2 8), HOPETOS2- (67 2, 6S); 17x. 1982’ G Sd): 26.X.1982 (4 2, 1 8), X.1982 (10 2, 46 38), EXTEII S25 (200-35), Oe 98.2.(2:. 9,16); 15.X1.1982 (4 2, 10 4), 22.X1.1982 (4 9, 13 6), X1.1982 (3 2, 2 4); CAMEROON: Centre- Sud Prov., Akonolinga, VI.1982 (1 @, 1 3); DA VINLLOS2 (1 2); 28.VHI.1982 (1 6), WATT 982 (5:9, 13's); LX.1982 G2): X.1982 (2 2), G. Steck, reared from Coffea robusta berries, with Trirhithrum coffeae (Bezzi) (Tephritidae) as probable host; Ouest Prov., 71 Ba Fig. 2. Ovipositor tips. A, Rhynchosteres (Fopius) silvestrii. B, R. (F.) caudatus (Szépligeti). Bafoussam, 30.VI.1982 (1 9), 13.VIH.1982 (1 9), G. Steck, reared from Coffea arabica berries; Centre-Sud Prov., Etoug-Ebe, 26.V.1982, G. Steck, reared from Dacus on squash (1 ¢?); Yaounde, Agr. Res. Inst., 2.V1.1980, F. E. Gilstrap, W. G. Hart, & D. Perkins, from coffee berries (12 2). COSTA RICA: Turrialba, 12.XI.1982, M. Fischel, recovered from release site in coffee plan- tation (1 4); shipped to Costa Rica from Cameroon, via Texas, IX.23.1982 (3 9), X1.1982 (3 2). TOGO: Kpime-Seva, 16.XII.1981, G. Steck, from Coffea robusta berries: (19). The material collected in 1982 was shipped through the Texas A&M Univer- sity (TAMU) quarantine as #s T82001, T8200S9> “S8Z013;5" P8203) maT 82033, T82034, T82036, 182039, T82040, T82041, T82044, T82046, and 182047. Further details are provided by Steck et al. (1986). Diagnosis.—This species has frequently been reared with R. (F.) caudatus from a variety of tephritids in West Africa; and has often been confused with the latter (Whar- ton and Gilstrap, 1983; Steck et al., 1986). In caudatus, the dorsope is not developed, the clypeus tends to be thickened medially along the ventral margin, the ovipositor is slightly shorter (usually less than twice length of mesosoma), the frons bears a very dis- tinctive transverse row of deep, close-set punctures on an otherwise unsculptured backgorund (frons weakly to heavily rugose in silvestrii), and the median mesonotal lobe has only a single longitudinal groove me- dially. The two species probably attack dif- ferent immature stages of their host tephri- tids since the ovipositor is exceptionally 72 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON narrow at the tip in caudatus (Fig. 2). A similar situation was described in detail by Van den Bosch and Haramoto (1951) for members of the persulcatus species group. A much closer relationship exists between silvestrii and pyknothorax, and these two species are difficult to separate. However, the latter, a South African species, lacks punctures and setae on the lateral lobes of the mesonotum, and the clypeus is slightly thickened mid-ventrally. Discussion.— This species is named in honor of the Italian entomologist Filippo Silvestri, who did much to further our knowledge of opiine biology through his bi- ological control efforts. The above mea- surements (given as a mean + standard de- viation) are based on 20 specimens each. ACKNOWLEDGMENTS I am most grateful to the following cu- rators and assistants, without whose help this work could not have been accom- plished: A. Bachmann (Buenos Aires), P. Dessart (IRSN), E. Haeselbarth (Munich), T. Huddleston (BMNH), F. Koch (ZMHB), P. Marsh (USDA, USNM), S. Mascherini (Firenze), J. Papp (Budapest) and A. Sten- mark (NRS). I thank also M. Fischer (Wien), C. Van Achterberg (Leiden), J. Schaffner (College Station), and F. Bin (Perugia) for assistance and fruitful discussions, and W. Hart, F. Gilstrap, and G. Steck for making available the material on which the new species was based. Finally, I thank R. Koe- nig for graciously allowing me to publish the lectotype designations for Wesmael’s Opius species. LITERATURE CITED Ashmead, W.H. 1900. Some changes in generic names in the Hymenoptera. Can. Entomol. 32: 368. Brues, C. T. 1926. Studies on Ethiopian Braconidae with a catalogue of the African species. Proc. Am. Acad. Arts Sci. 61: 205-436. Cameron, P. 1903. Descriptions of new genera and species of Hymenoptera from India (continua- tion). Z. Hym. Dipt. 3: 9-16, 177-184, 298-304, 337-344. . 1910. Onsome Asiatic species of the subfam- ilies Braconinae and Exothecinae in the Royal Ber- lin Museum. Soc. Entomol. 25: 11-12, 14-16, 19- 20, 22-23, 25-26. De Santis, L. 1967. Catalogo de los Himenopteros argentinos de la serie Parasitica, incluyendo Be- thyloidea. Com. Invest. Cient. Prov. Buenos Aires, La Plata, pp. 33-34. Fischer, M. 1959. Die européischen Opiinae (Hy- menoptera, Braconidae). Acta Entomol. Mus. Nat. Pragae 33: 241-263. . 1963a. Die Opiinae der athiopischen Region. Beitr. Entomol. 13: 194-221, 662-747. 1963b. Das Genus Austroopius Szépligeti (Hymenoptera, Braconidae, Opiinae). Mitt. Zool. Mus. Berlin 39: 173-186. 1964. Die Opiinae der nearktischen Region (Hymenoptera, Braconidae). I. Teil. Polski Pismo Entomol. 34: 197-530. 1965. Zwei neue Opiinen—Gattungen aus dem Kongo (1. Beitrag ueber die Opiinae des Mu- see Royal de l’Afrique Centrale) (Hymenoptera, Braconidae). Rev. Zool. Bot. Afr. 71: 309-323. 1966. Revision der indo-australischen Opi- inae. Dr. W. Junk B. V., The Hague. 167 pp. . 1967a. Zusammenfassung der neotropischen Opiinae mit Ausschluss der Gattung Opius Wesm. (Hymenoptera, Braconidae). Beitr. Neotrop. Fau- na. 5: 1-21. . 1967b. Redeskriptionen von einigen Opiinen (Hymenoptera, Braconidae). Arbeitsgemeinschaft Oesterr. Entomol. 19: 59-69. . 1971. Hym. Braconidae. Index of World Opi- inae. Index of entomophagous insects. Le Fran- cois, Paris. 189 pp. 1972. Hymenoptera Braconidae (Opiinae I). Das Tierreich 91: 1-620. . 1977. Hymenoptera Braconidae (Opuinae II— Amerika). Das Tierreich 96: 1-1001. Foerster, A. 1862. Synopsis der Familien und Gat- tungen der Braconen. Verh. naturh. Ver. preuss. Rheinl. 19: 225-288. Fullaway, D. T. 1951. Review of the Indo-Austra- lasian parasites of the fruit flies (Tephritidae). Proc. Haw. Entomol. Soc. 14: 243-250. Gahan, A.B. 1915. A revision of the North American ichneumon-flies of the subfamily Opiinae. Proc. U.S. Natl. Mus. 49: 63-95. Haliday, A. H. 1837. Essay on Parasitic Hymenop- tera. Entomol. Mag. 4: 203-221. Hendrikse, A. 1980. A method for mass rearing two braconid parasites (Dacnusa sibirica and Opius pallipes) of the tomato leafminer (Liromyza by- roniae). Meded Fac. Landbou. Rijksuniv. Gent 45: 563-571. Marshall, T. A. 1891. A monograph of the British Braconidae. Part IV. X XIII. Opiides. Trans. Ento- mol. Soc. Lond. 1891: 7-61. Martelli, G. M. 1937. Contributo alla conoscenza VOLUME 89, NUMBER 1 biologica del Dacus oleae Rossi e dei suoi parassiti in Tripolitania. Agric. Colon. 31(A): 149-155. Muesebeck, C. F. W. 1931. Descriptions of a new genus and eight new species of ichneumon-flies with taxonomic notes. Proc. U.S. Natl. Mus. 79: 1-16. 1967. Family Braconidae, pp. 27-60. In Krombein, K. V. and B. D. Burks. Hymenoptera of America north of Mexico, synoptic catalog. Sec- ond Supplement. USDA Agric. Mon. 2. 584 pp. Muesebeck, C. F. W. and L. M. Walkley. 1951. Fam- ily Braconidae, pp. 90-184. Jn Muesebeck, C. F. W., K. V. Krombein, and H. K. Townes. Hyme- noptera of America north of Mexico, synoptic cat- alog. USDA Agric. Mon. 2. 1420 pp. Paoli, G. 1934. Diachasma brevistyli Paoli, pp. 349- 351. In Prodromo di entomologia agraria della Somalia Italiana. Firenze, Istituto Agricolo Co- loniale Italiano. Papp, J. 1985. Taxonomical and faunistical novelties of the Opiinae from the Old World tropics (Hy- menoptera: Braconidae). Acta Zool. Hung. 31: 185- 216. Roman, A. 1910. Notizen zur Schlupfwespensamm- lung des schwedischen Reichsmuseums. Entomol. Tidsk. 31: 109-196. Schulz, W. A. 1911. Zweihundert alte Hymenopter- en. Zool. Ann. 4: 1-220. Shenefelt, R. D. 1970. Pars 5. Braconidae 2. Hel- coninae, Calyptinae, Mimagathidinae, Triaspinae. In Ferriere, Ch. and J. van der Vecht, eds., Hy- menopterorum catalogus (nova editio). Dr. W. Junk B. V., The Hague, pp. 177-306. 1975. Pars 12. Braconidae 8. Exothecinae Rogadinae. /n van der Vecht, J. and R. D. Shene- felt, eds., Hymenopterorum catalogus (nova edi- tio). Dr. W. Junk B. V., The Hague, pp. 1115- 1262. 73 Steck, G. J., F. E. Gilstrap, R. A. Wharton, and W. G. Hart. 1986. Braconid parasitoids of Tephritidae (Diptera) infesting coffee and other fruits in West- Central Africa. Entomophaga 31: 59-67. Szépligeti,G. 1907. Collections faites par M. le Baron Maurice de Rothschild dans |l’Afrique Orientale. Bull. Mus. Hist. Nat. Paris 1907: 34-36. 1911. Braconidae der I. Zentral-Afrika-Ex- pedition. Wiss. Ergebn. dtsch. Zentr. Afr. Exped. 3: 393-418. 1914. Afrikanische Braconiden des Koenig. Zoologischen Museums in Berlin. Mitt. zool. Mus. Berlin 7: 153-230. Thomson, C.G. 1895. LIl. Bidrag till Braconidernas kaénnedom. Opusc. Entomol. 20: 2141-2339. Van Achterberg, C. 1983. A new species of Rhyn- chosteres Fischer from Zaire (Hymenoptera, Bra- conidae). Zool. Meded. 57: 91-95. Van den Bosch, R. and F. H. Haramoto. 1951. Opius oophilus Fullaway, an egg-larval parasite of the Oriental Fruit Fly discovered in Hawaii. Proc. Haw. Entomol. Soc. 14: 251-255. Wesmael, C. 1835. Monographie des Braconides de Belgique. Nouv. Mém. Acad. R. Bruxelles 9: 1- 252. 1838. Monographie des Braconides de Bel- gique. Nouv. Mém. Acad. R. Bruxelles 11: 5-166. Wharton, R. A. and F. E. Gilstrap. 1983. Key to and status of opiine braconid (Hymenoptera) parasit- oids used in biological control of Ceratitis and Dacus s. |. (Diptera: Tephritidae). Ann. Entomol. Soc. Am. 76: 721-742. Wharton, R. A. and P. M. Marsh. 1978. New World Opiinae (Hymenoptera: Braconidae) parasitic on Tephritidae (Diptera). J. Wash. Acad. Sci. 68: 147- 167. PROC. ENTOMOL. SOC. WASH. 89(1), 1987, pp. 74-76 A NEW SPECIES OF AGARODES (TRICHOPTERA: SERICOSTOMATIDAE) FROM SOUTHEASTERN UNITED STATES'! S. C. HARRIS Aquatic Biology Program, Department of Biology, University of Alabama, Tuscaloosa, Alabama 35487. Abstract.—A new species, Agarodes alabamensis, from coastal Alabama is described and illustrated. The genus now totals ten species, all restricted to eastern North America. The genus Agarodes is restricted to and widespread throughout eastern North America from Canada to Florida (Wiggins, 1977). Nine species are recognized in the genus, with all but Agarodes distinctus Ul- mer and Agarodes griseus Banks restricted to southeastern United States (Ross and Scott, 1974). In Alabama, four species oc- cur: Agarodes crassicornis (Walker) and Agarodes libalis Ross and Scott on the Coastal Plain; A. griseus Banks at scattered localities in northern Alabama; and Aga- rodes stannardi which 1s restricted to a small area in the northwestern portion of the state. To this list is added an apparently rare new species, Agarodes alabamensis, known only from the type locality on the Coastal Plain. As is typical for the genus, this new species was collected with a black-light along a small sand-bottom stream. Type material will be deposited at the Na- tional Museum of Natural History, Smith- sonian Institution. Terminology follows that of Schmid (1980) and Ross and Scott (1974). Agarodes alabamensis Harris, NEW SPECIES Fig. 1 Diagnosis. —In many respects this species resembles Agarodes tetron (Ross) but differs ' Contribution No. 93 from the Aquatic Biology Pro- gram, The University of Alabama. in the rounded apex of segment X and in the elongate mesal branch on the basomesal process of the inferior appendage. In addi- tion, A. alabamensis is a Coastal Plain in- habitant, while A. tetron occurs in the mon- tane regions of northern Georgia and North Carolina. Male.— Length 17-19 mm. Body, legs and head light brown to yellow. Antennae brown, with 43 segments, the scape small and trap- ezoidal. Labial palpus 1.3 mm long; max- illary palpus 0.8 mm in length, oblong, bear- ing 2 narrow inner processes. Wings light brown; forewing with extensive peg-like se- tae in anal region and posteriorly along M and Cu veins. Abdominal segment IX an- nular. Segment X elongate and narrow in dorsal view, divided into lateral plates di- verging apically; in lateral view sinuate dor- sally and lightly sclerotized, distally round- ed to blunt apex, lacking ventral projection. Preanal appendage narrow and elongate, ex- tending nearly half length tergum X, setose distal portion curving dorsad in lateral view; in dorsal view somewhat spatulate distally and curving mesad. Inferior appendages slender anteriorly, widening posteriorly in lateral view; in ventral view united basally, slender with irregular margins, basomesal processes divided basally. Mesal processes over *%4 length of inferior appendages, slen- der and narrowing to acute apex, distally serrate and diverging; lateral processes thin VOLUME 89, NUMBER 1 Agarodes alabamensis / Figs 1 Agarodes alabamensis n. sp., male genitalia. A, Lateral view. B, Dorsal view. C, Ventral view. 75 76 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON and nearly equal in length to mesal pro- cesses. Phallus long and generally cylindri- cal, membranous apically with acute apex, ventral notch subapically. Female. — Unknown. Immatures.— Unknown. Etymology.—Latin: of Alabama. Holotype.—Alabama, Clarke County, Fishers Creek at unmarked Co. Rd., | mile SW Whatley (R3E T8N S25), 27 April 1985, S.C, Harts, Paratype.— Alabama, same as above, | 4. Distribution.—Agarodes alabamensis is known only from the type locality and ap- pears to be rare in coastal Alabama. Exten- sive black-light trapping throughout Clarke County and adjacent counties has not yield- ed any additional specimens. Repeated black-light collecting along Fishers Creek at several locations in May 1985, early and late April 1986, and May 1986 found no additional material. As well, on four occa- sions an extensive search for Agarodes lar- vae in Fishers Creek and its tributaries was unsuccessful. Discussion. —Agarodes alabamensis is a member of the subgenus Agarodes Banks (Ross and Wallace, 1974) based on the small antennal scape and slender mesal lobe of the maxillary palp. The key to Agarodes males in Ross and Scott (1974) is amended as follows to accommodate the new species. 7. Mesal branch of basomesal process short, sit- uated apically, much as in Fig.3 ...... griseus — Mesal branch of basomesal process long, sit- uated basally 7A. Mesal branch of basomesal process about half as long as lateral branch, apex of seqment X with ventral projection (Ross, 1948, Fig. 4) FO RO case le ces, shes Ne ty oA ECT tetron — Mesal branch of basomesal process over half as long as lateral branch, apex of segment X without ventral projection alabamensis ACKNOWLEDGMENTS The Geological Survey of Alabama pro- vided equipment and facilities during the study and is gratefully acknowledged. John Unzicker of the Illinois Natural History Survey provided specimens of Agarodes te- tron for comparison. I thank also Patrick O’Neil for his help in the collections, Ruth Turner for photographing the figures, and Irene Thompson for typing the manuscript. LITERATURE CITED Ross, H. H. and D. C. Scott. 1974. A review of the caddisfly genus Agarodes, with descriptions of new species (Trichoptera: Sericostomatidae). J. Ga. Entomol. Soc. 9: 147-155. Ross, H. H. and J. B. Wallace. 1974. The North American genera of the family Sericostomatidae (Trichoptera). J. Ga. Entomol. Soc. 9: 42-48. Schmid, F. 1980. Genera des Trichoptéres du Canada et des Etats adjacents, pt. 7, 296 pp. /n Les insectes et arachnides du Canada. Agric. Canada, Ottawa. Wiggins, G. B. 1977. Larvae of the North American caddisfly genera (Trichoptera). Univ. Toronto Press, Toronto. 401 pp. PROC. ENTOMOL. SOC. WASH. 89(1), 1987, pp. 77-94 MORPHOMETRIC ANALYSIS OF UNIPARENTAL 4APHYTIS REARED FROM CHAFF SCALE, PARLATORIA PERGANDIT COMSTOCK, ON TEXAS CITRUS (HYMENOPTERA: APHELINIDAE; HOMOPTERA: DIASPIDIDAE) JAMES B. WOOLLEY AND HAROLD W. BROWNING (JBW) Department of Entomology, Texas A&M University, College Station, Texas 77843; (HWB) Department of Entomology, Agricultural Research and Extension Center, Texas A&M University, 2415 East Highway 83, Weslaco, Texas 78596. Abstract.—The literature on parasites of chaff scale, Parlatoria pergandii Comstock, is briefly reviewed with emphasis on chaff scale in Texas. A survey of the natural enemies of chaff scale in Texas citrus showed two thelytokous and closely related (cryptic) species, Aphytis hispanicus (Mercet) and Aphytis comperei DeBach and Rosen, to be the most common parasites. Since these species are reported in the literature to be sympatric in many localities, and since individuals with an apparently intermediate morphology were found, we tested the hypothesis that the concepts of A. hispanicus and A. comperei rep- resent two points in a continuous distribution of phenotypes. A morphometric study of the material was conducted to determine if two distinct morphs corresponding to A. hispanicus and A. comperei occur in Texas citrus, and if so, to identify useful morphological characters to distinguish between them. Sixteen measurements of anatomical structures, Six meristic characters, and two qualitative characters were scored for 146 specimens reared from isolated chaff scale. The measurement data were analyzed using principal component and canonical variates analyses. Principal component analysis of the raw and log-transformed data showed that two distinct morphs exist which correspond to A. hispanicus and A. comperei. In addition, a third group of individuals, designated as A. ?hispanicus, was found. These individuals are close to, but somewhat distinct from, A. hispanicus. Principal component analysis and canonical variates analysis suggest that the A. ?hispanicus group consisted of small specimens of A. hispanicus. Canonical variates analysis also showed that 6 of the 17 characters used were useful in discriminating between A. comperei and A. hispanicus. Two meristic characters showed strong discontinuities between A. comperei and A. hispanicus. We conclude that two species, A. comperei and A, hispanicus, are the common parasites of chaff scale in Texas citrus. in Texas (Dean, 1955; Dean et al., 1983), Israel (Gerson, 1977; Harpaz, 1961), Spain This paper is the first in a series reporting the results of a survey of the parasites of armored scale on citrus in south Texas. Here we discuss the status of uniparental (thely- tokous) Aphytis (Hymenoptera: Aphelini- dae) reared from chaff scale, Parlatoria per- gandii Comstock (Homoptera: Diaspididae). Chaff scale is a recurring problem on citrus (Limon et al., 1977; Carrero, 1980), and in other locations worldwide (Talhouk, 1975) Parlatoria pergandii occurs sympatrically in Israel with Parlatoria cinerea Doane and Hadden, the tropical grey chaff scale (Ger- son, 1967a, b). 78 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON The most common parasites we encoun- tered on chaff scale were referable to one of two species in the proclia group of Aphytis: A. hispanicus (Mercet) and A. comperei DeBach and Rosen. Both species are uni- parental (Rosen and DeBach, 1979) and males referable to this species group were quite rare in our collections. The similarities between these two species and the occur- rence of apparent intermediate forms prompted this study. Prior to the work of DeBach and Rosen (summarized in Rosen and DeBach, 1979) species identification in Aphytis was ex- tremely difficult and often controversial. Dean (1965) expressed some frustration at the different species names various special- ists provided for Aphytis reared from chaff scale on Texas citrus—variously A. proclia (Walker), A. diaspidis Howard, and A. his- panicus. All of these species are 1n the pro- clia group, as are A. maculicornis (Masi) and A, paramaculicornis DeBach and Rosen which were imported and released against the olive scale, Parlatoria oleae (Colvée), in California and elsewhere (Rosen and DeBach, 1979). Dean (1965), Dean and Hoelscher (1967) and Dean and Bailey (1960) refer to an Aphytis ““complex”’ as the dominant parasites of chaff scale in Texas. Later, DeBach and Rosen (1976) deter- mined that Dean’s chaff scale parasite ma- terial contained two very similar Aphytis species: A. hispanicus and a second species they described as A. comperei. Our examination of early correspondence and unpublished reports indicate that chaff scale was present early in the history of the citrus industry in Texas. A complete review of all parasites of chaff scale in Texas is forthcoming. Introduction of other Aphytis species have been made for chaff scale con- trol in Texas, but establishments of exotic species have not been documented (Dean and Bailey, 1960). In 1968 a strain of A. paramaculicornis originally obtained from chaff scale on citrus at Escondido, Califor- nia was shipped from Riverside to Texas and approximately 35 adults were released on chaff scale at Weslaco. This somewhat enigmatic Aphytis was originally thought to be the “Iran” strain of A. paramaculicornis, but Rosen and DeBach (1979) regard it as distinct and apparently indigenous to Cal- ifornia. No recoveries of this Aphytis have been made. In fact, we have not reared ma- terial from chaff scale in Texas referable to any introduced species. All Aphytis found on chaff scale in Texas can therefore be con- sidered to be indigenous, or exotic species which moved with chaff scale when it was introduced to Texas. The holotype of Aphytis hispanicus Mer- cet (1912) was from material reared from chaff scale on citrus at Valencia, Spain. The species was most recently redescribed by Rosen and DeBach (1979), whose list of lo- calities for this species includes Spain, Italy, Turkey, Israel, the Caucasus, Morocco, Tai- wan, Brazil, Trinidad, Mexico, Florida, Cal- ifornia, and notably, Texas. Most of the specimens were reportedly reared from P. pergandii, and to a less extent from P. ci- nerea, P. oleae, Aspidiotus nerii Bouché, In- sulaspis pallida (Green), Lopholeucaspis Japonica (Cockerell), and Mytilaspis conch- iformis (Gmelin). Records from Acutaspis scutiformis (Cockerell), Aonidiella aurantii (Maskell) and Chrysomphalus dictyospermi (Morgan) were regarded as questionable. Aphytis hispanicus has also been reported attacking chaff scale in Morocco (Abassi, 1975), Spain (Limon et al., 1976, 1977) and Florida (Muma, 1971). Crouzel (1973) list- ed A. hispanicus and Aphytis argentinus Bréthes as parasites of P. cinerea and P. pergandii, respectively, in Argentina. How- ever, Gerson (1977) stated that the chaff scale parasite which she listed as A. argen- tinus is probably a synonym of A. comperei or A. hispanicus. Rosen and DeBach (1979) describe A. hispanicus as a uniparental, sol- itary parasite of Parlatoria species which attacks second instars, male scale, and adult VOLUME 89, NUMBER 1 79 Table 1. Diagnostic characters for Aphytis comperei and A. hispanicus from Rosen and DeBach (1979). Information taken from key to species and species redescriptions. A. hispanicus A. comperei Genal sutures to eye Antennal club Pedicel, funicle, base of club Antennal club Crenulae heavily sclerotized, infuscate from oral margin to about ¥4 distance length/width = 2.5-3.0 uniformly and strongly infuscate apical '4 blackish (infuscate) 6-8 per side, elongate, non-over- less heavily sclerotized, faintly in- fuscate shorter, thicker, length/width about 2.3 paler tip with conspicuous black spot 3-5 per side, wider, distinctly black- lapping, faintly infuscated ish Ratio of lengths of ovipositor/ less than or = 1.33 middle tibia Forewing, length/width Setae in delta region of fore- wing LMC forewing/width forewing 2.75-3.00 0.17-0.33 59-161 in 9-12 rows longer, up to 2.00 2.5—2.66 51-96 in 7-9 rows 0.20 female scale, preferring the latter. Gerson (1967b, 1968), Rosen (1965, 1967, 1969) and Rivnay (1968) provide information on the biology of A. hispanicus in Israel. DeBach and Rosen (1976) described A. comperei and provided diagnostic charac- ters to distinguish it from A. hispanicus and A, proclia. The holotype female was reared from ‘“‘Aonidiella aurantii material” on cit- rus in McAllen, Texas; however, Rosen and DeBach (1979) regarded the California red scale record for the holotype as question- able. They point out that California red scale, chaff scale, and other scale species are often found mixed together on citrus and cross contamination of rearing samples is com- mon. Most of the records listed for A. com- perei are from chaff scale, and records from A. aurantii, Chrysomphalus aonidum (L.), and Cornuaspis beckii (Newman) were re- garded as questionable (Rosen and DeBach, 1979). The distribution of A. compere in- cludes Texas, Mexico, Florida, Jamaica, South Africa, Hong Kong and Canton, China. Little additional information is available on the biology of A. comperei be- yond the observation of Rosen and DeBach (1979) that this species is uniparental. Rosen and DeBach (1979) provide a number of characters to distinguish com- perei and hispanicus in their key to Aphytis species and in the redescriptions of the two species. We list their criteria in Table 1 and provide figures of typical character states for each species. Of particular interest are the shape and coloration of the antennal seg- ments, the conformation of the crenulae, and aspects of the forewings. In the next few paragraphs, all discussion of the diagnostic characters for the two species refers to the criteria of Rosen and DeBach (1979). The antennal club of 4. hispanicus (Fig. 1, Table 1) 1s characterized by a blackish, infuscate region in about the distal third. The antennal club of A. comperei (Fig. 2, Table 1) is shorter, thicker, and the infuscate area 1s confined to a black spot at the tip. The propodeum of Aphytis typically bears several posterior, lamellate projections called crenulae. Variations in the size, shape, number, and color of the crenulae have been used to distinguish between many Aphytis species (Rosen and DeBach, 1979). These authors state that 12-16 crenulae are found on A. hispanicus (Table 1), and further, that the crenulae in hispanicus are elongate, pale 80 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-5. Photomicrographs of Aphytis species. 1, Antennae of A. hispanicus, medial aspect. 2, Antennae of A. comperei, medial aspect. 3, Propodeum of A. hispanicus. 4, Propodeum of A. comperei. 5, Eighth abdominal tergum of A. hispanicus. (cr = crenulae, Is = lateral setae, ms = medial setae). in color, and do not overlap one another (Fig. 3). In contrast, A. comperei is char- acterized by 6-10 crenulae (Table 1), and the crenulae are wider, overlapping, and distinctly darker in color (Fig. 4). The forewing of A. hispanicus is longer and narrower than that of A. compere (Ta- ble 1, Figs. 6 and 7), but the differences are not striking. The oblique, bare streak in the AN & aphelinid wing is properly termed the linea calva (Hayat, 1983). In Aphytis the region of the forewing basal to the linea calva is called the delta region (Rosen and DeBach, 1979). As noted in Table 1, specimens of A. hispanicus are said to commonly bear more setae (microtrichiae) in the delta region (59- 161) than specimens of 4A. comperei (51-96 setae). Finally, the ratio of the longest mar- VOLUME 89, NUMBER 1 81 Ss Lee fe Ore is oo 2 oo a = = = — ~Ne rae ——— Ss TP Es amo SSS eS eS vy a S rateve:9 — aS Sas = wee af Cire (he ae pees ee - ies Seip = Sve-_- --->S sry Ae EG EAC Ge Le ree LI Te A Gite NIE a Na ep Ga ; = ROE Ce eel Oh ie age i TEC. ar eg ee IN Se : ws Ce pe ae - ts Wp Sane OE I Fe = ad J Ao aT = aaa VELL. “> oS oF Sa ios ae Came —_—— Loe eae ce Li RO == ai lon > —_—— —_ = <= eae re ne ee a lee ee ie ca — -_— wee em ee Ona PN a ee oc ge Se eg eS ge = —— ~” ie -—S ae ae TEES Tt rh ile te —— wae Z SIAR aoe eae Ge ae aa ae AS ae SS Shee / Vaaf Ug Et Gea SS aR SSS aneen EL; AEH Age ge ae Rie a SNe ae EN Cina cf wa 7 ee ec RI om coo —— a rane NS eS a ye pe EE GAS i ae ER Sas Ns SS GEG AL AE EE Se eg oe ne Ea NOS Se ties = fe =< = a el ie If LG ae GTA eC ao ENN = LSS eee aL a (eS SENN De oee aS ‘N= =~ = - ~ See pied hae Fes SN \ RS Sea AS ~Ne ee us NY AN << REN CS — — OR PTR SE RON Se Ue ae ESN == Ss N See are PNGNT Ney > = NN mS =~ NENTS =~ SAR ~ SEBEL ee EAL ee AGE OS a EN = _ St eo “Aas eae ae. Fa o— so Pa Cae Na ~ a SED Pim Vt Aa Ee — a he =f US Bi me el a c oo WESC, i i LES BE aSoaoe = rT (ss 3 Pia) fi A oe pe y, Ae A iA “ol BR aN NS = = : Gee Se eo LE ON by Gra ee GEE SR SEO SSS ING US eae Ss hf Ye Ya — rs Goal / LE EE EE SE Oe ES 2. = J v FE all 7 ys ee >N ea ss Re kad d 7. wee — / ( / CF pale Se Se eee wey re oe ae ee ees EEA EO en ree NSCS SS me erty AL Me aed g, NP pee a ee NE - 1 4 or _ om ter ey NL N ~ on Bae eae ft GEL Le ‘ LES AAS tee woe Soran oe DR EMME Tcl tile Ue ee See Pea ki aee COME Be Pa DE Ge ryt Ci ie aie SSN eS SU pment ht POS Sas Se Ny alee ie Ua! / = SS Sa Sw ft , ( oo Reece se ~ a AOE RE Ma = Oe ae SEN De Ee - = = LGBT Zen a hana = NEY = => - 0.15, Kolmogorov test). For this rea- son, the 95% confidence ellipse for comperei was constructed without the score for the outlier observation. RESULTS Principal component analysis.—A_ pro- jection of the individual specimens on the first two principal components computed from the variance/covariance matrix from the original data is presented as Fig. 8. To- gether, the first two principal components account for 93.0% of the original variance. The remaining principal components rep- resent increasingly miniscule proportions of the total variance: from 2.1% for the third to 0.03% for the seventeenth. The unitized eigenvectors associated with the first two principal components are shown in Table 3. The elements of each vector have been scaled so that the sum of the squares of all the elements in each vector is unity. Thus, the elements represent weights, and the val- ue for each element squared represents the proportion of variance 1n the principal com- ponent which each variable contributes, as- suming that PCA has produced uncorrelat- ed linear transformations of the original variables. Two distinct clusters of points represent- ing A. comperei and A. hispanicus individ- uals were found (Fig. 8). The individual ten- tatively determined as 4. ?comperei lies well within the cluster formed by A. comperei individuals. The clusters formed by A. com- perei and A. hispanicus are distinct with re- spect to the second principal component only. Thus, by examining the weights 1n Ta- ble 3 for the second principal component, one can gauge the contribution of individual variables to the location of individuals on this axis. For example, from Fig. 8 and Ta- ble 3 it is noted that A. hispanicus individ- uals tend to have longer and narrower fore- wings with long marginal cilia, a longer 85 Table 3. Eigenvalues and weights for the first two principal components, computed from the covariance matrix from the untransformed data. The vectors are scaled so that the sum of the squares of the elements in each vector is unity. The rows have been sorted on the elements for the second principal component, from numerically highest to lowest. Variable or Quantity PGI PC Il Eigenvalue 3243.92 555.63 Proportion of Variance 0.794 0.136 14) Setae in delta region, forewing 0.20 0.56 5) Length of infuscate area on club 0.03 0.27 13) Length of LMC on fore- wing —0.04 0.26 11) Length of forewing 0.79 0.16 15) Length of middle tibia 0.23 0.12 9) Length of propodeum 0.08 0.10 17) Length of midtibial spur 0.07 0.09 16) Length of basitarsus 0.10 0.07 4) Length of club 0.10 0.03 8) Length of metanotum 0.02 0.01 7) Length of scutellum 0.13 —0.004 1) Length of scape 0.12 —=(: 01 6) Length of mesoscutum 0.15 —0.02 3) Length of apical funicle segment 0.04 —0.02 2) Length of pedicel 0.04 —0.03 2) Width of forewing 0.30 —0.28 ) Length of ovipositor 0.32 —=().62 infuscate area on the antennal club and more setae in the delta region of the forewing. Aphytis comperei individuals tend to have wider forewings and longer ovipositors. The first principal component in Fig. 8 fits some of the criteria (Blackith and Rey- ment, 1971; Jolicoeur and Mosimann, 1960) commonly used to identify a size vector, or a principal component which reflects pri- marily variation in overall size of the spec- imens. The weights for the first principal component in Table 3 are all positive except for character 13. The weights are not, how- ever, of uniform magnitude. The heavily weighted characters (14, 11, 15, 12, and 10) are measurements of comparatively large structures with correspondingly large vari- 86 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON compere ? comperel hispanicus ? hispanicus IT (28%) I (54%) Fig. 9. Observations plotted on the first two principal components computed from the covariance matrix from the log-transformed data. The first principal component contains 54% of the sample variance, the second principal component contains 28%. ance. We note also that the clusters of points for each species in Fig. 8 are roughly parallel to the first principal component axis, lend- ing further support to the interpretation of this principal component as a size vector. If the first principal component simply ap- proximates overall size, then the five indi- viduals originally determined as A. ?hispan- icus would appear to be small specimens of A, hispanicus. Figure 9 is a projection of the individuals on the first two principal components com- puted from a variance/covariance matrix derived from the log-transformed data. This transformation should reduce the overall ef- fect of the large variances associated with variables with numerically large values. Ta- ble 4 shows the eigenvalues and eigenvec- tors for the first two principal components, as discussed above for Table 3. That the tranformation was successful can be seen in Table 4, in that the first principal compo- nent now accounts for only 53.8% of the total variance, with a greater proportion, 27.7%, now contained in the second prin- cipal component. The remaining principal VOLUME 89, NUMBER 1 components now account for somewhat greater, but still relatively small proportions of the total variance: 4.7% for the third and 0.1% for the seventeenth. The weights for the first principal component (Table 4) are all positive, although not of uniform mag- nitude, suggesting that the first principal component retains some variance associ- ated with overall size of the specimens. Again, two distinct clusters of points in Fig. 9 correspond with original determina- tions of either 4. comperei or A. hispanicus and A. ?hispanicus. However, the clusters in Fig. 9 are at oblique angles to two prin- cipal component axes, suggesting that now the first principal component expresses variance associated with shape differences in addition to size differences. Also, the in- clusion of a point in one or another cluster is now determined by a contribution from both principal components. The contribu- tion of individual variables to the clustering of individuals when projected on the first two principal components can be assessed by examining the relative weights of vari- ables on each component. The length of the infuscate portion of the antennal club is the only variable which makes a strong contri- bution to both the first and second principal component, accounting for just over 25% of the variance of each. The number of setae in the delta region of the forewing accounts for slightly over 25% of the variance in the first principal component, and the length of the marginal cilia on the forewing for slight- ly over 25% of the variance represented by the second principal component. Other variables with high weights on the first prin- cipal component are the length of the pro- podeum and the length of the basitarsus. The width of the forewing, length of the pedicel and third funicle segment on the antenna, lengths of the mesoscutum and scutellum, and length of the ovipositor all have relatively high weights on the second principal component. The A. ?hispanicus individuals are again concentrated at one end of the distribution 87 Table 4. Eigenvalues and weights for the first two principal components, computed from the covariance matrix from the log-transformed data. The vectors are scaled so that the sum of the squares of the elements in each vector is unity. PCI Variable or Quantity PC Il Eigenvalue 0.0341 0.0176 Proportion of Variance 0.538 0.277 1) Length of scape 0.13 0.17 2) Length of pedicel 0.07 0.26 3) Length of apical funicle segment 0.10 0.20 4) Length of club 0.14 0.12 5) Length of infuscate area on club 0.57 0:51 6) Length of mesoscutum 0.15 0.20 7) Length of scutellum 0.18 0.20 8) Length of metanotum Oy 0.09 9) Length of propodeum 0.25 0.11 10) Length of ovipositor 0.05 0.26 11) Length of forewing 0.15 0.15 12) Width of forewing 0.11 0.26 13) Length of LMC on fore- wing 0.09 0:52 14) Setae in delta region, forewing 0.53 =—0:01 15) Length of middle tibia 0.19 O35 16) Length of basitarsus 0.26 0.18 17) Length of midtibial spur 0.20 0.08 of A. hispanicus individuals in Fig. 9. How- ever, the effect now appears to be spread between the first and second principal com- ponents. Furthermore, the points now over- lap with A. hispanicus more with respect to their location on the second principal com- ponent. In this case, more than a simple size effect seems to be involved, as the location of points with respect to either component is not simply size related. It appears that these individuals are intermediate with re- spect to some aspects of morphology, al- though more similar to A. hispanicus than to A. comperei. Aphytis ?hispanicus was re- tained as a distinct a priori class in the ca- nonical variates analysis for this reason. The A.?comperei specimen was treated as a member of the 4. comperei class for canon- ical variates analysis because the point rep- resenting this specimen fell in the middle of 88 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON © comperei © hispanicus Oo ? hispanicus -10 UJ Fig. 10. Observations plotted on the two canonical variate axes. The ellipses around each cluster of points represent 95% prediction regions (Owen and Chmielewski, 1985). the points for the A. comperei specimens when projected against either set of prin- cipal components (Figs. 8 and 9). Canonical variates analysis.—Homoge- neity of the covariance matrices for the three classes used in canonical variates analysis was tested using the SAS DISCRIM pro- cedure. The covariance matrix for the ?his- panicus class was not of full rank due to the small number of observations, and there- fore could not be tested against the other two. However, the tests did show that the null hypothesis that the covariance matrices of the comperei and hispanicus classes were homogeneous could not be rejected (P = 1.000, likelihood ratio test). We assumed that the covariance matrix for the ?hispan- icus class was also homogeneous with the other two, since we had no reason to assume otherwise. Figure 10 is a plot of the projection of the individuals on the first two canonical vari- ates. The first canonical variate contains 97.6% of the between-groups variance, and since with three groups only two canonical variates can be constructed, the second ca- nonical variate contains 2.4% of the be- tween-groups variance. One would expect VOLUME 89, NUMBER 1 89 Table 5. Standardized coefficients and total canonical structure for the canonical variates analysis. The standardized coefficients are the amount that the canonical variate score will change for a change in the original variable of one standard deviation. The total canonical structure values are the total-sample correlations between the original variables and the canonical structure scores. The rows have been sorted by the elements of the vector of coefficients for the first canonical variate. Standardized Coefficients Variable CVI 5) Length of club infuscation 1.45 14) Setae in delta region, forewing enly/ 9) Length of propodeum 0.91 13) Length of LMC on forewing 0.80 17) Length of midtibial spur 0.35 15) Length of middle tibia 0.33 11) Length of forewing 0.18 8) Length of metanotum 0.14 16) Length of basitarsus 0.09 4) Length of antennal club 0.09 2) Length of pedicel —=(102 6) Length of mesoscutum SOY 1) Length of scape =0:35 12) Width of forewing —0.41 7) Length of scutellum —0.50 3) Length apical funicle segment —0.58 10) Length of ovipositor —0.87 Total Canonical Structure CV Il CVI CV Il =0:23 0.91 0.15 0.75 0.73 0.57 0.40 0.46 0.62 =0:39 0.74 =0:35 0.37 0.45 0.65 —0.45 0.24 0.71 0.33 0.10 0.76 =O: 0.26 0.40 =0:26 0.30 0.67 0.25 0.14 0.66 —0.04 =(0:22 0.47 —0.67 =—0:02 0.68 0.18 —0.01 0.70 0.14 =0:30 0.64 —0.38 0.04 0.66 0.83 =O:19 0.88 0.11 —0.54 0.58 the very tight clusters of points representing the comperei and hispanicus individuals (Fig. 10), because canonical variates are con- structed to maximize between-group co- variance relative to within-group covari- ance. The 95% confidence ellipse for the comperei specimens (with one outlier re- moved, as discussed above) is well separat- ed from the ellipses for both the hispanicus and ?hispanicus groups, while the ellipses for hispanicus and ?hispanicus are broadly overlapping. We note also that the ?hispan- icus ellipse differs in size, shape, and ori- entation from the other two ellipses, an in- dication that the covariance matrix for the this class may not be equal (Owen and Chmielewski, 1985). Clearly, the first canonical variate un- ambiguously discriminates the comperei group from the hispanicus plus ?hispanicus groups (Fig. 10). The standardized canoni- cal coefficients for the canonical variates (Table 5) are the products of the canonical vector coefficients and the pooled within- group standard deviations for each variable. They represent the amount that the canon- ical variate score (e.g. in Fig. 10) will change for each change of the original variable by one standard deviation. A large absolute value for a standardized coefficient gener- ally indicates a variable which will be useful in discrimination (but see Campbell and Atchley, 1981, for a discussion of potential problems with this interpretation). The strong positive standardized coefficient scores for the first canonical variate for vari- ables 5, 9, 13, and 14 in Table 5 indicate that hispanicus individuals tend to have a longer infuscate area on the antennal club, a longer propodeum, longer marginal cilia on the forewing, and more setae in the delta region of the forewing, respectively. The strong negative score for variable 10 on the first canonical variate indicates that com- perei individuals have longer ovipositors because the comperei cluster in Fig. 10 is in the negative range of the first canonical vari- ate. Weaker negative scores on the first ca- nonical variate in Table 5 for characters 7 and 3 indicate that comperei individuals tend 90 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON specimens observed 6 7! 8 2) Fig. 11. Oe tl 12 3 | als number of crenulae Number of specimens with observed number of crenulae. (Black bars represent 4. comperei, white bars represent A. hispanicus, shaded bars represent A. ?hispanicus specimens.) to have a longer scutellum and a longer api- cal funicle segment. The same pattern is evident in the total canonical structure values in Table 5. These values represent the total-sample correla- tions between the original variables and the canonical structure scores. As with the stan- dardized coefficients, variables 5, 9, 13, and 14 have strong positive correlations on the first canonical variate. Variable 17, the length of the midtibial spur, also shows a strong positive correlation on the first ca- nonical variate, but its standardized coef- ficient is relatively low. Variable 10, the length of the ovipositor, has a strong neg- ative correlation on the first canonical vari- ate, with the same implication as discussed above. Meristic characters.— At least two of the meristic characters for which we recorded data show marked differences between these species. As noted by Rosen and DeBach (1979), hispanicus individuals tend to have more crenulae that comperei individuals. Figure 11 is a histogram of the number of specimens observed with a particular num- ber of crenulae. Most comperei specimens had 8-10 crenulae (see Fig. 4), most his- panicus specimens had 11-15 (see Fig. 3), while the ?/ispanicus specimens had an in- termediate number. Another useful meristic character is the number of medial setae on the eighth abdominal tergum (Fig. 5). As can be seen in Fig. 12, most comperei spec- imens had two such setae, rarely | or 3, while most hispanicus specimens had 4, or 5, rarely 3 or 6. The ?hispanicus specimens were again intermediate with 2 or 3 setae in this location. DISCUSSION The ?hispanicus individuals do not ap- pear to represent a morph distinct from the hispanicus individuals. In all plots (Figs. 8, 9, 10) the ?hispanicus observations cluster at one end or the other of the distributions of hispanicus observations. In fact, the ?his- panicus specimens appear to be simply small hispanicus individuals. This can be seen clearly in Fig. 10 and Table 5. The ?hispan- VOLUME 89, NUMBER 1 specimens observed Fig. 12. 91 3 4 5 6 medial setae on eighth tergum Number of specimens with observed number of medial setae on the eighth tergum. (Black bars represent A. comperei, white bars represent A. hispanicus, shaded bars represent A. ?hispanicus specimens.) icus individuals lie below the hispanicus cluster in Fig. 10, with virtually all discrim- ination between these two groups on the second canonical variate. In Table 5, the total canonical structure values show strong positive correlations between the scores on the second canonical variate and the origi- nal variables for all variables except 5 and 13. Therefore, since ?hispanicus individuals tend to have numerically lower (more neg- ative) scores on the second canonical vari- ate, they tend to have numerically smaller values for all variables except 5 and 13. This trend is also apparent in Table 2, in which the means for ?hispanicus individuals for all variables except 13 are lower than the means for hispanicus individuals. The meristic characters are intermediate for the ?hispan- icus Class (Figs. 11 and 12), but again, if the ?hispanicus group consists of small hispan- icus individuals, they would be expected to have fewer crenulae and fewer medial setae on the eighth tergum. The comperei individuals (except the one outlier) are morphologically distinct from the hispanicus specimens (including the ?hispanicus specimens). In all of the results, the following patterns are consistent. Aphy- tis comperei individuals tend to have a long- er Ovipositor than hispanicus individuals. Aphytis hispanicus individuals tend to have a longer infuscate area on the antennal club, more setae in the delta region of the fore- wing, longer marginal cilia on the forewing, and a longer propodeum than comperei in- dividuals, and to some extent, a longer mid- dle tibia with a longer apical spur. Of the diagnostic characters used by DeBach and Rosen (1976) and Rosen and DeBach (1979) (Table 1), most are well supported by our results. However, we did not find A. com- perei individuals to have consistently short- er clubs (Tables 3-5) as they stated, nor did we find consistently shorter or wider fore- wings in A. comperei (Tables 3-5). In Table 2 we have tabulated the means, 95% confidence intervals for the means, and observed ranges for the quantitative vari- ables. The means for characters 5, 10, 12, 13, and 14 for comperei and hispanicus are 92 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON well separated (as indicated by the confi- dence intervals), but the ranges for these variables are strongly overlapping, with the exception of character 5 for which the ranges are contiguous. Therefore, while there is in- formation in many of these characters, any particular variable for a single specimen will not necessarily be discriminating. In fact, the covariation of traits leads to the dis- tinctly different morphologies of comperei and hispanicus. This points out the dangers in comparison of mean values for characters (e.g. with f-tests) or of the ranges of char- acters in making taxonomic decisions. In this case, comparison of means only would overstate the differences between comperei and hispanicus, while comparison of ranges only would not reveal trends which do occur in these data. Multivariate techniques such as PCA and CVA explicitly represent the covariation between many characters, thereby providing a method to assess trends which occur in several characters simulta- neously. For an excellent discussion of this general problem, see Albrecht (1980). CONCLUSIONS Our results strongly support the conclu- sions of Rosen and DeBach (1979) that A. comperei and A. hispanicus are two distinct, but closely related species. Of course, with thelytokous forms such as these, notions of reproductive isolation do not apply, and the boundaries of species are necessarily arbi- trary. We have found rare male specimens in our material referable to one of the two species. The results of R6ssler and DeBach (1972, 1973) suggest that the rare males in Aphytis species are functional, at least in some species. Therefore, the possibility ex- ists that mating occurs in the field, and that gene flow occurs between clones and even between species as we now recognize them. More likely, however, is the situation ob- served with uniparental strains of A. ma- culicornis and biparental strains of A. par- amaculicornis in which laboratory studies have indicated that these forms are com- pletely reproductively isolated (Rosen and DeBach, 1979). The possible role of males in comperei and/or hispanicus is a matter for further investigation. Nevertheless, we have found two distinct morphs corresponding almost exactly to the concepts of A. comperei and A. hispanicus presented by DeBach and Rosen (1976). The results from the principal components anal- ysis are the strongest evidence for the dis- tinctness of the two forms, as this technique does not utilize any a priori grouping cri- teria in dimension reduction. The meristic characters provide further evidence for the discontinuity in the morphologies of the two species. The A. ?hispanicus specimens re- main somewhat problematic, as both the meristic characters and the multivariate analyses indicate that these individuals are intermediate in form. However, the multi- variate analyses also suggest that these spec- imens are smaller A. hispanicus individuals and meristic data are consistent with that hypothesis. We do not, therefore, support recognizing these as a third, distinct morph. Given that A. comperei and A. hispanicus are morphologically distinct, how can one best identify individual specimens? The cri- teria of Rosen and DeBach (1979) (Table 1) will generally be useful, although we would not recommend using the length/width ra- tios of either the antennal clubs or the fore- wings. One new meristic character, the number of medial setae on the eighth ter- gum will be useful in most, but not all cases. However, as noted above, it is the covaria- tion in traits which makes the two species morphologically different, and no single characteristic will provide a reliable crite- rion for identification in all cases. Discrim- inant analysis provides a statistical method to identify unknowns to known groups in just such a situation. Discriminant analysis produces a single linear transformation of the original variables to optimally discrim- inate between a set of predefined groups. Canonical variates analysis, which we have used, is essentially the multi-group exten- VOLUME 89, NUMBER 1 sion of discriminant analysis (Albrecht, 1980), and the special case of CVA of two groups 1s equivalent to discriminant anal- ysis (Neff and Marcus, 1980). Once a set of diagnostic characters and a discriminant function has been developed for a pair (or group) of cryptic species, the information can be distributed to persons who need to make routine identifications, but who do not have extensive experience with the species and with their subtle diagnostic characters. This person could input a series of measurements for each unknown speci- men. The discriminant function would then provide the a posteriori probability of an unknown belonging to one of the known species. However, one important caveat ex- ists, all possible species to which the un- known might be referable must be included in the discriminant function for the tech- nique to be valid. ACKNOWLEDGMENTS We thank Penny Wilkinson for slide- mounting the specimens used in this study, and for much of the data entry on the com- puter. We thank also Willy Melton for his assistance with the field work and with many other matters. Fred Hendricks, John Her- aty, William Smith, and Tom Unruh re- viewed the manuscript and each made many valuable suggestions and comments. This paper is Technical Article No. 21841 of the Texas Agricultural Experiment Station. The specimens discussed above are deposited as Voucher Series No. 902 in the Insect Col- lection, Department of Entomology, Texas A&M University. LITERATURE CITED Abassi, M. 1975. Notes bio-écologiques sur Parla- toria pergandei Comstock (Homopt. Coccidae) au Maroc. Fruits 30: 179-184. Albrecht, G. H. 1980. Multivariate analysis and the study of form, with special reference to canonical variate analysis. Amer. Zool. 20: 679-693. Blackith, R. E. and R. A. Reyment. 1971. Multi- variate morphometrics. Academic Press, London and New York. Campbell, N. A. and W. R. Atchley. 1981. The ge- 93 ometry of canonical variate analysis. Syst. Zool. 30: 268-280. Carrero, J. M. 1980. Estado actual de la lucha biol- ogica contra las cochenillas de los agrios en Val- encia (Espana). Fruits 35: 625-631. Crouzel, I. S. de. 1973. Estudios sobre control biol- ogico de cochinillas Diaspididae que atacan citri- cos en la Republica Argentina. Idia 304: 15-39. Dean, H.A. 1955. Factors affecting biological control of scale insects in Texas citrus. J. Econ. Entomol. 48: 444-447. 1965. An Aphytis complex (Hymenoptera: Eulophidae) of chaff scale. Ann. Entomol. Soc. Am. 58: 142-145. Dean, H. A. and J. C. Bailey. 1960. Introduction of beneficial insects for the control of citrus scale insects and mites. J. Rio Grande Valley Hort. Soc. 14: 40-46. Dean, H. A. and C. E. Hoelscher. 1967. Chaff scale parasite complex as affected by carbaryl. J. Econ. Entomol. 60: 729-730. Dean, H. A., J. V. French, and D. Meyerdirk. 1983. Development of integrated pest management in Texas citrus. Texas Agric. Exp. Stn. Bull. 1434. DeBach, P. and D. Rosen. 1976. Twenty new species of Aphytis (Hymenoptera: Aphelinidae) with notes and new combinations. Ann. Entomol. Soc. Am. 69: 541-545. Gerson, U. 1967a. Studies of the chaff scale on citrus in Israel. J. Econ. Entomol. 60: 1145-1151. . 1967b. The natural enemies of the chaff scale, Parlatoria pergandii Comstock, in Israel. Ento- mophaga 12: 97-109. 1968. The comparative biologies of two hy- menopterous parasites of the chaff scale, Parla- toria pergandii. Entomophaga 13: 163-173. 1977. Statut actuel de Parlatoria pergandii Comstock en Israél. Fruits 32: 407-411. Harpaz,I. 1961. Coccoidea, pp. 126-175. In Avidov, A., Pests of the cultivated plants of Israel. The Magnes Press, Jersualem. [In Hebrew. ] Hayat, M. 1983. The genera of Aphelinidae (Hy- menoptera) of the world. Syst. Entomol. 8: 63- 102. Johnson, R. A. and D. W. Wichern. 1982. Applied multivariate statistical methods. Prentice-Hall, Engelwood Cliffs. Jolicoeur, P. and J. E. Mosimann. 1960. Size and shape variation in the painted turtle. A principal component analysis. Growth 24: 339-354. Limon, F., A. Melia, J. Blaso, and P. Moner. 1976. Estudio de la distribucion, nivel de ataque y par- asitos de las cochinillas diaspinas Chrysomphalus dictyospermi Morgan y Parlatoria pergandii Comst. en citricos de la provincia de Castellon. Bol. Serv. Plagas 2: 73-87. 1977. Contribution a l'étude de la distribu- 94 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tion, du niveau d’attaque et des parasites des Cochenilles diaspines (Chrysomphalus dictyosper- mi Morgan et Parlatoria pergandii Comst.) des Citrus de la Province de Castellon. Fruits 32: 354- 359. Mercet, R. G. 1912. Los enemigos de los parasitos de las plantas. Trab. Mus. Cienc. Nat. Madrid, 10, 306 pp. Muma, M.H. 1971. Parasitism of armored scale in- sects in commercial Florida citrus groves. Fla. Entomol. 54: 139-150. Neff, N. A. and L. F. Marcus. 1980. A survey of multivariate methods for systematics. New York: privately published. Owen, J. G. and Chmielewski, M. A. 1985. On ca- nonical variates analysis and the construction of confidence ellipses in systematic studies. Syst. Zool. 34: 366-374. Rivnay, E. 1968. Biological control of pests in Israel (a review 1905-1965). Israel J. Entomol. 3: 1-156. Rosen, D. 1965. The hymenopterous parasites of cit- rus armored scales in Israel (Hymenoptera: Chal- cidoidea). Ann. Entomol. Soc. Am. 58: 388-396. 1967. Biological and integrated control of citrus pests in Israel. J. Econ. Entomol. 60: 1422- 1427. 1969. The parasites of coccids, aphids, and aleyrodids of citrus in Israel: some zoogeograph- ical considerations. Israel J. Entomol. 4: 45-53. Rosen, D. and P. DeBach. 1979. Species of Aphytis of the world (Hymenoptera: Aphelinidae). Series Entomologica V. 17, Dr. W. Junk BV, The Hague. Réssler, Y. and P. DeBach. 1972. The biosystematic relations between a thelytokous and an arrheno- tokous form of Aphytis mytilaspidis (Le Baron) (Hymenoptera: Aphelinidae) |. The reproductive relations. Entomophaga 17: 391-423. 1973. Genetic variability in a thelytokous form of Aphytis mytilaspidis (Le Baron) (Hyme- noptera: Aphelinidae). Hilgardia 42: 149-176. SAS Institute. 1982a. SAS user’s guide: basics. SAS Institute Inc., Cary, North Carolina. 1982b. SAS user’s guide: statistics. SAS In- stitute Inc., Cary, North Carolina. Talhouk, A. S. 1975. Citrus pests throughout the world, pp. 21-23. Jn Hafliger, Ernst, Citrus, Tech- nical Monograph No. 4, Ciba-Geigy Ltd., Swit- zerland. PROC. ENTOMOL. SOC. WASH. 89(1), 1987, pp. 95-99 NEW GENERA OF BAETIDAE (EPHEMEROPTERA) FROM AFRICA R. D. WALTZ AND W. P. MCCAFFERTY Purdue University, Department of Entomology, West Lafayette, Indiana 47907. Abstract. —Three new genera of Ephemeroptera are described for Afrotropical species of Baetidae with highly distinctive and unusual larvae. They are Pseudopannota Waltz and McCafferty, n. gen., including Pseudopannota bertrandi (Demoulin), n. comb. (type species) and Pseudopannota vinckei (Demoulin), n. comb.; Ophelmatostoma Waltz and McCafferty, n. gen., including Ophelmatostoma kimminsi Waltz and McCafferty, n. sp. (= Pseudocloeon sp. A. Kimmins); and Acanthiops Waltz and McCafferty, n. gen., including Acanthiops marlieri (Demoulin), n. comb. Pseudopannota and Ophelmatostoma have highly modified filtering/sweeping mouthparts. Pseudopannota also has fused mesotho- racic wingpads. Acanthiops is one of several genera of baetids with prominent dorsal tubercles. A largely undescribed fauna of mayflies in the Afrotropics provides a rich source of interesting and phyletically important species. We assign the species treated herein to three new genera as a part of our con- tinuing study of baetid systematics. All three of the new genera are striking as larvae be- cause of variously exaggerated morphology not typically found among Baetidae. Pseudopannota Waltz and McCafferty, NEw GENUS Larva.—Labrum (Demoulin, 1967, Fig. 2a; 1973, Fig. 3b) notched medially on an- terior margin and with labral shelf. Man- dibles (Demoulin, 1967, Fig. 2b, c; 1973, Fig. 3c, d) with incisors fused to apex; bases of prosthecae appearing recessed into man- dibular margin. Thumb of left mandible broad based and slightly elevated above plane of incisor bases. Maxillae (Demoulin, 1967, Fig. 2d; 1973, Fig. 3e) with elongate three-segmented palps that exceed length of galealacinia; segment 3 of palp much elon- gated and swollen relative to two basal seg- ments; galealacinia with several apical den- ticles and adjoining row of bristles as in most baetids. Labium (Demoulin, 1967, Fig. 2f; 1973, Fig. 3g) with glossae and para- glossae subtruncate apically, subparallel, and with long, fine setae apically; paraglossae exceed glossae in length; palps two seg- mented and with second segment swollen or expanded laterally and bearing long, fine setae. Legs with or without row of long, fine setae on foretibiae and foretarsi (Demoulin, 1967, Fig. la; 1973, Fig. 3h). Ventral fem- oral patch absent. Claws (Demoulin, 1967, Fig. 2g; 1973, Fig. 31) with numerically re- duced but prominent denticles and without subapical bristles. Wingpads of mesothorax fused medially and nearly to apices (Fig. 1 and Demoulin, 1967, Fig. 1a; 1973, Fig. 3a). Abdominal terga with scales and fine se- tae. Gills broadly rounded, thickened along anterior margin, with distinctly spiculate surfaces and spinous margins (Demoulin, 1967, Fig. 1b; 1973, Fig. 3n). Median ter- minal filament subequal to cerci. 96 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON igaale fax Pseudopannota bertrandi larva, mesotho- Type species.—Pseudocloeon bertrandi Demoulin, 1967: 227 [= Pseudopannota bertrandi (Demoulin), New Combination]. Included species.—Pseudopannota _ber- trandi (Demoulin) and Pseudopannota vinckei (Demoulin) n. comb. Distribution.—Ivory Coast, Madagascar, Senegal (River Gambia). Etymology.—Feminine Latin: pseudo— false, pan—fused, and nota—back, in ref- erence to the fused back condition of this schistonote mayfly. It thus appears falsely to be a member of the mayfly suborder Pan- nota sensu McCafferty and Edmunds (1979). Remarks. — Adults are unknown present- ly; however, the larvae of Pseudopannota are distinct from all other described Baeti- dae on the basis of the medially fused meso- thoracic wingpads, unique mouthparts, and uniquely denticulate claws. The unnamed larva described by Crass (1947: 64) as Pseu- docloeon sp. appears to approximate very closely mouthpart, claw, and gill characters described for known species of Pseudopan- nota. Although we suspect that this species will prove to be a member of Pseudopan- nota, we must wait to confirm this because we have thus far been unable to study any of Crass’s (1947) material. Ophelmatostoma Waltz and McCafferty, NEw GENUS Larva.—Labrum (Kimmins, 1955, Fig. 3b) notched medially on anterior margin and with labral shelf. Mandibles (Kimmins, 1955, Fig. 3c, d) with incisors fused to apex; prosthecae prominent and bases appearing recessed into mandibular margin; spiculae present between bases of incisors and molar area. Thumb of left mandible broad based, curved, and slightly elevated above plane of incisor bases. Maxillae (Kimmins, 1955, Fig. 3f) with slender two-segmented palps and subequal in length to galealacinia; galeala- cinia with prominent falcate denticle api- cally in addition to two adjoining rows of bristles apically. Labium (Kimmins, 1955, Fig. 3g) with glossae and paraglossae greatly elongated, slender, and apically with long, fine setae; paraglossae subequal to glossae in length; palps apparently two segmented (segments 2 and 3 fused) and swollen api- cally, basal segment abruptly angled toward interior. Legs with specialized ventral row of long, stout bristles on forelegs (Kimmins, 1955, Fig. 3h). Ventral femoral patch absent. Claws with single row of denticles and without subapical bristles. Abdominal terga with scales and fine se- tae. Gills broadly rounded and with serra- tions on distal anterior margins. Median terminal filament subequal to cerci. Type species.—Ophelmatostoma kim- minsi Waltz and McCafferty n. sp. Distribution.— Malawi [= Nyasaland (Kimmins, 1955)], South Africa (Trans- vaal), Zimbabwe [= Southern Rhodesia (Agnew, 1963)], and Senegal (River Gam- bia). Etymology.— Neuter Greek: ophelma- to— broom, stoma—mouth, in reference to the distinctive filtering/sweeping-type mouthparts. VOLUME 89, NUMBER 1 Remarks.—Adults of Ophelmatostoma are presently unknown. This genus, viz. the larvae, is clearly distinct from all other gen- era of Baetidae on the basis of the mouth- part characters (Fig. 2) described above. The described genera of baetids known to have filtering/sweeping mouthparts, i.e. Guaji- rolis Flowers (1985) and now Pseudopan- nota, and Ophelmatostoma, have several similarly adapted but independently de- rived characters of the mouthparts. These include the profusion of long bristles and fine setae, prosthecae that appear to be re- cessed in the mandibular margin, and very long, thin apical denticles of the galealaci- nia. Specific character states of the mouth- parts, tergum, claws, gills, and available knowledge of the adult stages indicate that these genera represent independently de- rived lineages. Ophelmatostoma kimminsi Waltz and McCafferty, New SPEcIES Pseudocloeon sp. A, Kimmins, 1955: 866. Larva.—Characterized as per Kimmins (1955) and as supplemented by the above generic description. Type materials.— Holotype, larva, in al- cohol, Malawi (=Nyasaland), Tengadzi Stream, 22.vii1.1952, L. Berner, deposited British Museum (Natural History), London. Paratypes: 4 larvae; 3 paratypes in alcohol and | paratype slidemounted in euparal (solvent: absolute alcohol), Senegal (River Gambia), Kedougau, 28.vi.1981, M. T. Gil- lies, deposited Purdue University Ento- mological Research Collection, West Lafa- yette, Indiana, USA. Etymology.—The specific epithet is for D. E. Kimmins, the British ephemeropterist who first studied the species. Acanthiops Waltz and McCafferty, NEw GENUS Larva.—Labrum (Demoulin, 1967, Fig. 4a) notched medially on anterior margin and with labral shelf. Mandibles (Demoulin, 97 Fig. 2. Ophelmatostoma kimminsi larva, head cap- sule (lateral left side). 1967, Fig. 4b, c) with patch of setae between base of united incisors and molar region. Left mandible with incisors fused to apex; prostheca stouter than prostheca of right mandible and not appearing recessed into mandibular margin; thumb of molar region stout and distinctly elevated above plane of incisor bases. Right mandibular incisors apically separated. Maxillae (Demoulin, 1967, Fig. 4d) with palps two segmented and subequal to or less than length of gal- ealacinia. Labium (Demoulin, 1967, Fig. 4f) with glossae and paraglossae slightly ta- pered apically; paraglossae subequal to glos- sae in length; palps three segmented; lateral margins of segment 2 subparallel; segment 3 short and narrower at base than apex of segment 2. Legs elongate and without highly modi- fied setal areas (Demoulin, 1967, Fig. 4a). Ventral femoral patch absent. Claws (De- moulin, 1967, Fig. 4g) with two rows of denticles and one pair of subapical ante- riorly directed bristles. Prothorax and me- sothorax with more or less prominent dorsal tubercles (Demoulin, 1967, Fig. 3a, b). Abdominal terga with scales and fine se- 98 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tae in addition to more or less prominent dorsal, medioposterior tubercles (Demou- lin, 1967, Fig. 3a, b; 1964: 288). Gills asym- metric (Demoulin, 1967, Fig. 41, 1-p), with spinous margins anteriorly, and numerous fine tracheae. Median terminal filament greatly reduced relative to cercl. Type species.—Centroptilum marlieri Demoulin, 1967: 230 [= Acanthiops mar- lieri (Demoulin), New Combination]. Distribution. — Zaire [= Belgian Congo], ?Kenya. Etymology.— Masculine Greek: akan- thos —thorn, i6ps—small fish, an allusion to this genus being a tuberculate or spiny min- now-like mayfly. Remarks.—The abdominal tubercles of Acanthiops are very highly developed, more so than in other baetids. The few other tu- bercled baetid genera, e.g. Jubabaetis Miil- ler-Liebenau (1980), Neobaetiella Miiller- Liebenau (1985), and Baetodes Needham and Murphy (1924), are clearly separable from Acanthiops on the basis of mouthpart, tergal, gill, and claw characters. Although adults of Acanthiops are unknown at the present time, Demoulin (1964) provided il- lustrations of a subimago associated with the larva he described as Centroptilum No. 3 from Kenya, a species which is probably congeneric with 4. marlieri and differing from this latter species by lacking gill 1. The hindwing of this subimago possesses a bi- furcate median costal process. This peculiar character state is thus far only associated with certain Afrotropical baetids, i.e. Cen- troptiliodes Lestage, and some Centropti- lum. We suspect several Afrotropical species, which are clearly provisional with respect to their present generic placement, will eventually prove to belong to Acan- thiops. These provisional species, whose lar- vae remain unknown at the present time, include Centroptilum biarcuatum Kolpelke (1980), C. boettgeri Kolpelke (1980), C. di- centrum Demoulin (1956), C. montanum Kimmins (1960), and C. sudafricanum Les- tage (1924). ACKNOWLEDGMENTS We thank P. C. Barnard, British Museum (Natural History), London; P. Grootaert, Institut Royal des Sciences Naturelles de Belgique, Brussels; and H. André, Musée Royale de l’Afrique Centrale, Tervuren, for loan of material. We also thank M. T. Gil- lies, Sussex, England, for the gift of speci- mens now deposited in PERC, and A. V. Provonsha for illustrations. This paper has been assigned Purdue University Experi- ment Station Journal Number 10,676. LITERATURE CITED Agnew, J. D. 1963. New South African records of imperfectly known Baetidae (Ephem.). Hydro- biol., Acta Hydrobiol. Hydrograph. Protist. 22: 41-45. Crass, R.S. 1947. The may-flies (Ephemeroptera) of Natal and the eastern Cape. Ann. Natal Mus. 11: 37-110. Demoulin, G. 1956. Quelques Ephéméroptéres du Kivu. Bull. Ann. Soc. R. Entomol. Belgique 92: 277-284. 1964. Mission H. Loffler en Afrique Orien- tale Ephemeroptera. Bull. Ann. Soc. R. Entomol. Belgique 100: 279-294. 1967. Description de deux larves atypiques de Baetidae (Ins. Ephemeroptera). Bull. Ann. Soc. R. Entomol. Belgique 103: 226-232. 1973. Ephéméroptéres de Madagascar. III. Bull. Inst. R. Sci. Nat. Belgique, Entomol. 49: 1- 20. Flowers, R. W. 1985. Guajirolis, anew genus of Neo- tropical Baetidae (Ephemeroptera). Stud. Neotr. Fauna Envirn. 20: 27-31. Kimmins, D. E. 1955. Ephemeroptera from Nyasa- land, with descriptions of three new species and some interesting nymphal forms. Ann. Mag. Nat. Hist., Ser. 12 8: 859-880. . 1960. Notes on East African Ephemeroptera, with descriptions of new species. Bull. Brit. Mus. Nat. Hist. (Entomology) IH: 23-34. Kolpelke, J-P. 1980. Ephemeroptera aus der Emer- genz des zentralafrikanischen Berbaches Kalengo (Zaire). Teil 1: Baetidae. Entomol. Abh. Staat. Mus. f. Tierk. Dresden 43: 99-129. Lestage, J. A. 1924. Les Ephéméres de l’Afrique du Sud. Catalogue critique et systématique des es- péces connues et description de trois genres nou- veaux et de sept espéces nouvelles. Revue Zool. Africaine 12: 316-351. McCafferty, W. P. and G. F. Edmunds, Jr. 1979. The higher classification of the Ephemeroptera and its VOLUME 89, NUMBER 1 99 1985. Baetidae from Taiwan with remarks evolutionary basis. Ann. Entomol. Soc. Am. 72: 5122 on Baetiella Ueno, 1931 (Insecta, Ephemerop- Miiller-Liebenau, I. 1980. Jubabaetis gen. n. and tera). Arch. f. Hydrobiol. 104: 93-110. Platybaetis gen. n., two new genera of the family Needham, J. G. and H. Murphy. 1924. Neotropical Baetidae from the Oriental Region, pp. 103-114. mayflies. Bull. Lloyd Library, 24, Entomol. Ser. In J. F. Flannagan and K. E. Marshall eds., Ad- 4: 1-79. vances in ephemeroptera biology. Plenum Pub- lishing Corporation, New York. ANNOUNCEMENT Erratum: In the article by Wheeler, G. C. and J. Wheeler, 1986, Proc. Entomol. Soc. Wash. 88(4), p. 685, couplet 3b, please change “‘separate”’ to “paper.” PROC. ENTOMOL. SOC. WASH. 89(1), 1987, pp. 100-102 STATUS OF ACIGONA HUBNER (SENSU BLESZYNSKI) (LEPIDOPTERA: PYRALIDAE: CRAMBINAE) WITH CHANGES IN NOMENCLATURE CHARLES W. AGNEW Department of Entomology, Texas A&M University, College Station, Texas 77843- 2475. Abstract. —Acigona Hibner has been incorrectly used as a generic name in the Cram- binae due to an earlier type designation of a noctuid species. Acigona (sensu Bleszynski) is composed of genus level taxa which should not be considered congeneric. Friedlanderia n. name is proposed as a replacement name for Chiloides Bleszynski 1963, a homonym of Chiloides Butler 1881. The type, Tinea cicatricella Hiibner, had been incorrectly des- ignated the type of Acigona by Bleszynski. Eoreuma morbidella (Dyar) n. comb. and Coniesta forsteri (Bleszynski) n. comb. are transferred from Acigona. Some unfortunate taxonomic errors have resulted in much confusion regarding the use of the generic name Acigona within the subfamily Crambinae (Lepidoptera: Pyral- idae). The incorrect usage of this name for crambine species is especially widespread in the non-systematic literature as the larvae of several species have been reported to in- fest agronomically-important Poaceae. Al- though many of the problems cannot be remedied until a revision of Acigona (sensu Bleszynski, 1967) is completed, some con- tributions toward nomenclatural stability can be made at this time. The genus Acigona was proposed by Hiib- ner ({1826]: 342) for two species without a type species being designated. Hampson (1926) selected one of these, Phalaena man- to Cramer (Noctuidae), as the type of Aci- gona and treated the genus as a synonym of Euclystis Hiibner. Bleszynski and Collins (1962), apparently unaware of this action, designated the other included species, Tinea cicatricella Hiibner (Pyralidae) as the type of Acigona in the Crambinae. Bleszynski (1963) also designated 7. cicatricella as the type of Chiloides Amsel 1949. Amsel (1949) originally included two species, 7. cicatri- cella and Chilo hederalis Amsel in Chi- loides. Bleszynski (1963) decided Amsel had not properly designated a type species as required by The International Code of Zoo- logical Nomenclature (Article, 13b), which made the generic name unavailable. Al- though Amsel appeared to indicate T. ci- catricella was to be the type, the wording 1s somewhat ambiguous and probably fails to satisfy the Code. In either case, Bleszynski intended Chiloides to become a junior ob- jective synonym of Acigona. Bleszynski (1965, 1967) expanded his concept of Acigona to encompass species of the genera Coniesta Hampson, Eoreuma Ely, Haimbachia Dyar, Achilo Amsel, Do- nacoscaptes Zeller, Girdhiara Kapur and Xubida Schaus, all of which share a con- dition of the female genitalia where a nar- rowed extension or “bridge” from the an- terior margin of the eighth tergite extends down onto the ostium bursa. Bleszynski (1967) also transferred several species to Acigona from Chilo, Erupa, and Eufernal- VOLUME 89, NUMBER 1 dia and described one species, 4. forsteri, in Acigona (Bleszynski, 1965). The mono- typic genus Occidentalia was included un- der Acigona by Klots (1970, 1983), but not by Bleszynski. Because Hampson’s (1926) type desig- nation places Acigona in the Noctuidae, there is no name for Bleszynski’s concept in the Pyralidae. Donacoscaptes Zeller is the oldest available name, but Bleszynski’s ge- neric concept, regardless of the name ap- plied to it, is unsatisfactory. Even before the problem with the erroneous type species designation for Acigona was brought to light (Nye, 1975; Fletcher and Nye, 1984), work- ers had begun to recognize the validity of some of the genera synonymized by Ble- szynski (Klots, 1970; Gaskin, 1973). I also believe that Acigona (sensu Bleszynsk1) con- tains several good genera as well as species for which new genera probably should be proposed. For example, Coniesta, Eo- reuma, and Haimbachia are closely related, but can be separated by genitalic characters of both sexes, especially the females. Several tropical species of Eoreuma and Haimba- chia are incorrectly assigned. After an ex- amination of the male type and an associ- ated female, I am transferring one former member of Acigona, Chilo morbidellus Dyar, to Eoreuma as E. morbidella (Dyar) n. comb. This South American species has the typical uncus and gnathos of Eoreuma and most closely resembles E. /oftini (Dyar) and E. evae Klots in the shape of the costal processes of the valvae. The female genitalia are also similar to those of E. /oftini. No single action can correct the problems with the name and the concept of Acigona (sensu Bleszynski). A replacement name for the genus could come from designating the type species of the senior synonym, Dona- coscaptes validus Zeller, as a new type for the generic concept to which Bleszynski ap- plied the name Acigona, but this would still result in the synonymization of good genera. In removing the genera from synonymy, we do not resolve the problem of the species 101 which do not belong in one of eight valid genera once included under Acigona. Be- cause 7. cicatricella, the ‘type’ of Acigona (sensu Bleszynski), is also the type of Chi- loides, it appears that there is an available generic name for this species. However, Chiloides Bleszynski 1963, is a junior hom- onym of Chiloides Butler 1881, (Lepidop- tera: Tortricidae) and thus unavailable. I therefore propose the following replace- ment name for Chiloides Bleszynski. Friedlanderia Agnew, NEw NAME Chiloides Amsel 1949, nomen nudum. Chiloides Bleszynski 1963, preoccupied by Chiloides Butler 1881 (Lepidoptera: Tor- tricidae). Acigona: Bleszynski and Collins 1962 (not Hiibner 1816 [1826]), incorrect type des- ignation. Type species: Jinea cicatricella Hubner [1823]-[1824]: pl. 68, fig. 455. Diagnosis of genus.—Same as for the type cicatricella, provided by Bleszynski (1965) who figured both sexes (Plate 5, Figs. 67-,.;), their genitalia (6, Plate 44, Fig. 67; 2, Plate 94, Fig. 67), and the larva (Figs. 67,;.,). The heavily sclerotized ovipositor of the female is unusually shaped and distin- guishes Friedlanderia cicatricella (Hiibner) N. Comp. from species in other genera. The basally lobed uncus can be used to distin- guish the male from species in related gen- era. The degree of sexual dimorphism in the wing pattern of F. cicatricella is much more marked than in most Crambinae. The genus is named for Timothy P. Friedlander, whose knowledge of Lepidoptera provided me with a helpful introduction to the Crambinae. At this time, only the type species, F. ci- catricella, belongs in the genus as currently defined. The other species included by Am- sel (1949), Chiloides hederalis (Amsel), 1s a synonym of Thopeutis galleriella (Ragonot) (Bleszynski and Collins, 1962). No useful purpose would be served by transferring from Acigona to Friedlanderia 102 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON those species originally described in Chilo, Erupa, and Eufernaldia. For the present, these misplaced species should revert to those genera until their generic affinities can be determined. Species from the genera once synonymized under Acigona have valid combinations available. One species with- out a valid combination, Acigona forsteri from China, is transferred to the Old World genus Coniesta to become Coniesta forsteri (Bleszynski) N. Comp. based on an illustra- tion of the male genitalia by Bleszynski (1965). Further study may necessitate plac- ing it elsewhere, but I feel that the present lack of a valid combination should be rec- tified for nomenclatural purposes. ACKNOWLEDGMENTS I thank Horace Burke, Timothy Fried- lander, Robert Wharton, and James Wool- ley of Texas A&M University for taxonom- ic and editorial advice on this manuscript; Frederick Rindge of the American Museum of Natural History for the loan of material; Douglas Ferguson of the Systematic Ento- mology Laboratory, Agricultural Research Service, for the loan of material and com- ments on the manuscript; and J. W. Smith, Jr. of Texas A&M University for providing support for this study. Approved by the Texas Agricultural Experiment Station as TA #21788. Research supported in part by Hatch project 6796 and a grant from the Rio Grande Valley Sugar Growers, Inc. LITERATURE CITED Amsel, H. G. 1949. On the Microlepidoptera col- lected by E. P. Wiltshire in Irak and Iran in the years 1935 to 1938. Bull. Soc. Fouad I Entomol. 33: 271-351. Bleszynski, S. 1963. Studies on the Crambidae (Lep- idoptera). Review of the genera of the family Crambidae with data on their synonyms and types. Acta Zool. Cracov. 8: 91-132. . 1965. Crambinae. Jn Amsel, H. G., Reisser, H., and Gregor, F. eds., Microlepidoptera Pa- laearctica 1. Verlag Fromme, Wien. 553 pp. 1967. Studies of the Crambinae (Lepidop- tera). Part 44. New neotropical genera and species. Preliminary checklist of neotropical Crambinae. Acta Zool. Cracov. 12: 39-110. Bleszynski, S. and R. J. Collins. 1962. A short cat- alogue of the world species of the family Cram- bidae (Lepidoptera). Acta Zool. Cracov. 7: 197- 389. Butler, A.G. 1881. Onacollection of nocturnal Lep- idoptera from the Hawaiian Islands. Ann. Mag. Nat. Hist. (5) 7: 392-408. Fletcher, D. S. and I. W. B. Nye. 1984. The Generic Names of Moths of the World. Volume 5 [Pyra- loidea]. Brit. Mus. (Nat. Hist.). London. 185 pp. Gaskin, D. E. 1973. A revision of New Zealand Chi- lonini (Lepidoptera: Pyralidae) and redescription of some Australian species. N. Z. J. Sci. 16: 435- 463. Hampson, G. F. 1926. Descriptions of new genera and species of Lepidoptera Phalaenae of the subfamily Noctuinae (Noctuidae) in the British Museum (Natural History). British Mus. (Nat. Hist.), London. 641 pp. Hiibner, J. 1796 [1836]. Sammlung europdischer Schmetterlinge 8: (Tineae). 78 pp., 71 pls. Augs- burg. 1816 [1826]. Verzeichniss bekannter Schmettlinge [sic], pp. [1]—431. Augsburg. Klots, A. B. 1970. North American Crambinae: Notes on the tribe Chiloini and a revision of the genera Eoreuma Ely and Xubida Schaus (Lepidoptera: Pyralidae). N.Y. Entomol. Soc. 78: 100-120. 1983. Crambinae. Jn Hodges, R. W. et al, eds., Check List of the Lepidoptera of America North of Mexico. E. W. Classey Ltd. & Wedge Entomol. Res. Found. London. 284 pp. Nye, I. W. B. 1975. The Generic Names of Moths of the World. Volume | [Noctuoidea, part]. Brit. Mus. (Nat. Hist.). London. 568 pp. PROC. ENTOMOL. SOC. WASH. 89(1), 1987, pp. 103-121 THE AFROTROPICAL PACHYGASTRINE GENERA ASHANTINA KERTESZ AND MERISTOMERINGINA JAMES, WITH TWO NEW GENERIC SYNONYMS (DIPTERA: STRATIOMYIDAE) NORMAN E. WoOoDLEY Systematic Entomology Laboratory, BBII, Agricultural Research Service, USDA, c/o U.S. National Museum of Natural History NHB 168, Washington, D.C. 20560. Abstract.—The two Afrotropical pachygastrine genera Ashantina Kertész and Meris- tomeringina James are reviewed. Two new generic synonymies are proposed: Neosolva Séguy is a new junior synonym of Ashantina (based on type species synonymy, N. dubia Séguy is a new junior synonym of Ashantina antennata Kertész) and Agnathomyia Lindner is a new junior synonym of Meristomeringina. Lectotypes are designated for Ashantina antennata Kertész and Neosolva dubia Séguy. A key to the six known species of Meris- tomeringina is provided as well as illustrations of male genitalia of all species, four of which are new (country of type locality in parentheses): M. aka (Zaire), M. cholo (Malawi), M. kontagora (Nigeria), and M. praestigiator (Uganda). The Pachygastrinae is a large subfamily of stratiomyid flies. Their extreme struc- tural diversity has made them a difficult group to work with and has resulted in the description of a large number of genera, each with few included species. Of the 62 Afro- tropical genera recognized by James (1980), 38 are monotypic, only seven have five or more included species, and only eight occur outside of the Afrotropical Region. The only recent key to genera from the region is that of Lindner (1966), which excludes those found only in Madagascar (15 genera), and nine other genera subsequently described or discovered from mainland Africa. Conse- quently, even generic determinations within the subfamily can be difficult to make. The character states used to define the Pachygastrinae, loss of vein M, beyond the discal cell and the abdomen composed of five major segments (i.e. those beyond five much reduced), are found in other stratio- myids. Thus the monophyly of the subfam- ily has not been adequately demonstrated. Based on their studies of female terminalia of representative stratiomyid taxa, Naga- tomi and Iwata (1978) noted that the ‘“‘Cli- tellariinae and Pachygasterinae may be het- erogenous.” This paper reviews two genera included by James (1952) in the tribe Meristomerin- gini, a tribe not maintained by him in sub- sequent publications. None of the seven character states mentioned by James (1952) is unique to the group. While the Afrotrop- ical genera included in the tribe by James are quite similar in general appearance, the possible monophyly of this suprageneric grouping requires much further documen- tation. The two genera treated in this paper may be identified using Lindner’s (1966) ge- neric key. The male genitalia for all known species in both genera are illustrated, which should allow for detection of additional species if they exist. Genus ASHANTINA Kertesz Ashantina Kertész, 1914: 539. Type species, A. antennata Kertész, by original desig- nation. 104 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Neosolva Séguy, 1953: 152. Type species, N. dubia Séguy, by original designation. NEW SYNONYM Y. Diagnosis. — Ashantina 1s most easily rec- ognized by the structure of its antenna (Fig. 1), in particular the eighth flagellomere, which is thick basally but quickly tapers to an arista-like apex. Only the basal portion of the eighth flagellomere is haired. Also, the rounded but protruding face, which ex- tends well below the lower eye margin (Fig. 1), and which is squared when viewed an- teriorly, is characteristic. Both of these char- acter states are likely autapomorphic for the genus; I am aware of no other pachygas- trines with similar features. Other features exhibited by Ashantina, al- though found in other pachygastrines, are useful in characterizing the genus. They in- clude the elongate antennal flagellum, the spineless scutellum, vein R,,, arising well beyond crossvein r-m, and the elongate ab- domen. Remarks.—The synonymy of Neosolva with Ashantina has been confirmed by ex- amination of type material for each of the included species. Séguy’s (1953) original de- scription of Neosolva was brief, not partic- ularly useful, and has made subsequent rec- Loy 4 (4 Ze Ashantina antennata Kertész, left lateral view of male head. ognition of his taxon difficult. Neoso/lva was excluded from the Xylomyidae by Naga- tomi and Tanaka (1971), who did not ex- amine the types, but guessed that it might be a pachygastrine. James (1980: 274) treat- ed Neosolva as an unplaced genus of Stra- tiomyidae. Ashantina runs rather ambiguously through paragraph 19 of Lindner’s generic key. It should probably be traced through the alternate, paragraph 14, as the last flagel- lomere is “bristle-form,” with only the base thickened and haired. Ashantina antennata Kertesz Ashantina antennata Kertész, 1914: 539. Neosolva dubia Séguy, 1953: 152. NEW SYNONYMY. Type material.—One female syntype of A. antennata, hereby designated as the lec- totype, still exists in the British Museum (Natural History), London. The other syn- types, which were in the Hungarian Natural History Museum, were destroyed in 1956. The lectotype female is labeled: ““Caught on leaf./Dunkwa, Ashanti, W. Africa. 2.VII.1907. Dr. W. M. Graham. 1908-245./ Ashantina @! antennata Kert det. Kertész/ LECTOTYPE Ashantina antennata Ker- VOLUME 89, NUMBER 1 tész, 1914: 539 des. N. E. Woodley 1981.” The specimen is in reasonably good con- dition but is missing the left antennal fla- gellum, the entire right antenna, the last two tarsomeres of the left foreleg, and the right hindleg beyond the trochanter. Four syntypes of N. dubia Séguy are pres- ent in the Muséum national d’Histoire na- turelle, Paris. I have examined one of these, a female, that is hereby designated as the lectotype. It is labeled: ‘““Thio/MUSEUM PARIS Nimba (Guinée) M. Lamotte II.VI.42/Neosolva dubia Typ. Seguy @/LECTOTYPE Neosolva dubia Séguy, 1953: 152 des. N. E. Woodley 1981/Ash- antina antennata Kertész, 1914: 539 Det. N. E. Woodley 1981.” The lectotype is in excellent condition, lacking only the last four tarsomeres of the left hindleg. Diagnosis.—As Ashantina is monotypic, A. antennata may be recognized by the ge- neric characters discussed above. Figs. 2, 4, and 5 illustrate the male terminalia, which should allow determination of this species with certainty and allow detection of further undescribed species if they exist. Distribution.—James (1980) states that A. antennata 1s found from Liberia to Zaire. Its distribution 1s shown in Fig. 6. Material examined.—_CAMEROUN: 1 8, Dept. Nyong-Sanaga, Nkolbisson, Septem- ber 963,91. G. Sesers (MRAG): 2 ¢; Yaounde, 2600 ft., 29-30 May 1936, Van Zwaluwenberg & McGough (USNM). CEN- TRAL AFRICAN REPUBLIC: 10 4, 3 9, Dept. Lobaye, La Maboke, sur Whitfieldia longifolia, 31 August and 4 October 1970, L. Matile (MNHN). GHANA: | 2 (lecto- type) Ashanti, Dunkwa, caught on leaf, 2 July 1907, W. M. Graham (BMNH). GUINEA: (1 2 (lectotype of N. dubia), Nim- ba [Mountains], Thio, February—April 1942, M. Lamotte (MNHN). LIBERIA: 4 2, | 8, Robertsport, Bendu, 26 February to | April 1943, F. M. Snyder (AMNH). NIGERIA: 1 6, Ile-Ife, 20 June 1970, J. T.°Medler (CNC); 3 4, 4 2, Ile-Ife, 26 January 1975 and 16 March 1975, J. T. Medler (USNM). 105 ZAIRE: 6 6, Luebo, 27 April 1958, F. J. Francoise (IRNSB); | 4, 1 2¢, Mbanza-Ngun- gu [as Thysville], 15°0’E, 5°30’S (AMNH); 1 6, Parc National Albert, W. Ruwenzori (1200-1500 m), March 1937, Hackars (MRAC); 1 4, Pare National Albert, Kivu, Rutshuru (riv. Rutshuru) 1000 m, 3 July 1935, G. F. de Witte (MRAC). Remarks. — Ashantina antennata is a dis- tinctive Afrotropical species and is fairly common in collections. Lindner (1938, 1955, 1966, 1970) gives numerous records. The sexes are quite similar in habitus. The frons and face in females are slightly wider than in males. Females are generally some- what darker in coloration, which is espe- cially noticeable on the thoracic pleura and dorsum of the abdomen. The species is quite variable in size; specimens examined range from 5.5 to 9.0 mm in length. The end of the abdomen in both sexes is unusual as the fifth tergite is shorter and narrower than the sternite, and it is appar- ently movable and capable of closing the end of the abdomen (Fig. 3). The aedeagal complex is complicated in structure and is somewhat simplified in my illustration (Fig. 5). In profile it is nearly straight, and is not illustrated in this view because it was dif- ficult to delimit structures which were ob- scured by membrane. The homologies are difficult to determine, but the aedeagus is apparently strongly fused to the aedeagal valves (the lateral processes in the normal trifid aedeagal complex of stratiomyids). Genus MERISTOMERINGINA James Meristomeringina James, 1952: 38. Type species, M/. mimetes James, by original designation. Agnathomyia Lindner, 1958: 124. Type species, 4. combinata Lindner, by mono- typy. NEW SYNONYMY. Diagnosis. — Meristomeringina is the only genus of Afrotropical Pachygastrinae that has a combination of a simple antennal fla- gellum and a scutellum with two spines. The 106 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON S) Figs. 2-5. Features of Ashantina antennata Kertész. 2, Male genital capsule, dorsal view. 3, End of female abdomen, dorsal view. 4, Male epandrium and post-genital segments, dorsal view. 5, Male aedeagal complex, dorsal view. only other genus on mainland Africa with ent habitus. Several pachygastrines from two scutellar spines is Goetghebueromyia Madagascar also have two scutellar spines Lindner, but this genus has the last antennal but are also very different in general form. flagellomere aristate and has a much differ- Remarks. — Meristomeringina is difficult VOLUME 89, NUMBER 1 107 Fig. 6. to define with a conclusively autapomorph- ic character state. The shape of the antennal flagellum is characteristic but is of the more general, plesiomorphic form, i.e. without any fusion of flagellomeres or modification of the terminal ones to form a style or arista (Figs. 7-9). The antennae of the pachygas- trines are so variable in structure that char- acter polarities are difficult to determine in many cases. Presence of scutellar spines is also plesiomorphic within the subfamily, al- though the character state of having only two is rare in pachygastrines. The aedeagal Distribution of Ashantina antennata Kertész. complex is similar in all species included in the genus, and the presence of toothlike pro- cesses on the aedeagal valves may be aut- apomorphic for the genus. The six species now known of Meristo- meringina are extremely similar in mor- phology. Several species from central Africa can be identified with certainty only by ex- amining the male genitalia. Females are vir- tually identical, and the descriptions are based on specimens associated with males that have identical collecting data. The fe- male genitalia are not distinctly different. 108 A 65 >400 Baccharis neglecta Britt. >150 >550 Solidago altissima L. 0 0 Chrysothamnus nauseosus (Pall.) Britt. 0 0 Aster novae-angliae L. 0 0 Helianthus annuus L. 0) 0 Dahlia pinnata Cav. 0) 0 mortality factor. It was then quite common to fail to recover a single fly from a collec- tion of galls. Only occasional galls were seen in Texas during the remainder of the year. In Texas, galls were never very abundant and only rarely were more than 10-20 galls found on the one plant. At these population levels the insect had little effect on the growth of mature plants. However, in Florida seed- ling plants only a few inches high were seen with as many as five large galls and these caused severe stunting. Host SPECIFICITY TESTING Oviposition. — Because the adult fly is the only mobile stage the species could be con- sidered monophagous if specificity of ovi- position could be satisfactorily demonstrat- ed. Two series of tests were therefore conducted to determine the specificity of Oviposition. A smaller test using plants closely related to Baccharis was conducted in Texas. In each of two 35 x 27 x 17 cm clear plastic cages two bouquets of foliage and growing tips of each of seven plants were randomly placed. Into one cage were placed galls col- lected from B. halimifolia near Conroe, Texas and into the other were placed galls collected from B. neglecta near Temple, Texas. After sufficient exposure to the emerging flies, the bouquets were examined 124 Table 2. List of plant species against which N. /a- thami was tested in order to obtain permission to re- lease it in Australia. APIACEAE: Daucus carota L.; Pastinaca sativa L. ANACARDIACEAE: Mangifera indica L. ASTERACEAE: Baccharis halimifolia L.; Carthamus tinctorius L.; Chrysanthemum sp.; Dahlia sp.; He- lianthus annuus L.; Lactuca sativa L. BRASSICACEAE: Brassica oleraceae (L.) Alef.; Bras- sica rapa L. BROMELIACEAE: Ananas comosus (L.) Merr. CARICACEAE: Carica papaya L. CHENOPODIACEAE: Beta vulgaris L. CONVOLVULACEAE: Ipomoea batatas (L.) Lam. CUCURBITACEAE: Cucumis melo L.; Cucumis sa- tivus L.; Curcubita maxima Duch. FABIACEAE: Arachis hypogaea L.,; Centrosema pu- bescens Benth.; Desmodium canum (Gmel.); Glycine wightii (R. Grah. ex Wight & Arn.) Verdc.; Glycine max L. Merr.; Medicago sativa L.; Phaseolus atro- purpureus DC.; Phaseolis vulgaris L.,; Pisum sativum L.; Stizolobium sp.; Stylosanthes gracilis, Trifolium repens L.; Vigna catjang V. LINACEAE: Linum usitatissimum L. MALVACEAE: Gossypium hirsutum L. MIMOSACEAE: Leucaena leucocephala (Lam.) de Wit. MUSACEAE: Musa sapientum M. PASSIFLORACEAE: Passiflora edulis Sims PINACEAE: Pinus radiata D. Don.; Pinus taeda L. POACEAE: Avena sativa L.; Digitaria decumbens Stent.; Pannicum maximum Jacq.; Paspalum dilatatum Poir.; Pennisetum clandestinum Chiov.; Saccharum officinarum L.; Sorghum vulgare L.; Triticum aes- tevum L.; Zea mays L. PROTEACEAE: Macadamia integrifolia Maid & Betche ROSACEAE: Fragaria vesca L.; Malus sylvestris Mill.; Prunus domestica L.; Prunus persica (L.) Batch.; Py- rus communis L.; Rosa sp. RUTACEAE: Citrus limon (L.) Burm. F.; Citrus par- adisi Macfady.; Citrus reticulata Blanco; Citrus sin- sensis (L.) SAPINDACEAE: Litchi chinensis Sonn. SOLANACEAE: Capsicum annuum L.; Lycopersicum esculentum Miller; Nicotiana tabacum L.; Solanum tuberosum L. VITACEAE: Vitis vinifera L. ZINGIBERACEAE: Zingiber officinale Roscoe. under a binocular microscope and any eggs deposited were counted. The results (Table 1) indicated that both populations of flies oviposited on both species of Baccharis but not on any other plant. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON A more comprehensive testing program, designed to satisfy the Australian Depart- ment of Health’s requirements for intro- duction into Australia, was conducted using 65 species of plants (Table 2). This list of plants included the most important agri- cultural species. Bouquets of the growing tips of 8 plant species from the list, together with a bouquet of B. halimifolia were placed into glass and stainless steel aquaria (60 x 36 x 40cm). Twenty-five pairs of flies from galls collected in Florida were placed in each cage. Each test was duplicated. Eggs were counted after a three day duration. Many eggs were deposited on all the B. halimifolia controls but none were found on any other plant. Gall development tests.—A series of tests was carried out in an insectary to determine on which plants galls would develop. Each group of well-developed potted plants was infested with 25 pairs of flies collected in Florida. Each test was duplicated. After three days the plants were placed in a glasshouse. Hatching, movement of larvae and gall for- mation were observed and recorded. Eggs were found only on the B. halimifolia plants. After three to five days, larvae emerged and entered the young tips. There was no move- ment of these larvae onto other plants. Galls were formed in two weeks and the life cycle was completed in six to eight weeks. DISCUSSION The experimental work supported the previous host records from the field by 1n- dicating that N. /athami is host specific to those species of Baccharis on which it is found in the field (i.e. B. halimifolia and B. neglecta). Permission was therefore sought and obtained to import and release this in- sect for the biological control of B. halimifo- lia in Australia. Although this fly does not cause dramatic control in its native habitat, this does not preclude it from being successful in a new environment. The fly has a high reproduc- tion rate and a short life cycle. The fact that VOLUME 89, NUMBER 1 it is usually at low population densities is due primarily to the high rate of attack by parasites. Before being released in Australia parasites will be very carefully eliminated from the population in a quarantine facility. Without these parasites a much higher pop- ulation of the fly may develop in Australia, perhaps to a level where significant effect on the population of the weed may occur. ACKNOWLEDGMENTS We thank Raymond J. Gagné of the Sys- tematic Entomology Laboratory, Agricul- tural Research Service, USDA, Washing- ton, D.C. for authoritative identifications of our material and other information and help. LITERATURE CITED Gagné, R. J. 1971. Two new species of North Amer- ican Neolasioptera from Baccharis (Diptera: Ce- cidomylidae—Compositae). Proc. Entomol. Soc. Wash. 73: 153-157. McFadyen, P. J. In press. Introduction of the gall fly Rhopalomyia californica from the USA into Aus- tralia for the control of the weed Baccharis hali- mifolia. Proc. VI Int. Symp. Biol. Contr. Weeds, Vancouver, 1984. PROC. ENTOMOL. SOC. WASH. 89(1), 1987, pp. 126-131 A NEW SEIRA FROM THE UNITED STATES, WITH A REDESCRIPTION OF SEIRA BIPUNCTATA (PACKARD) AND NEW RECORDS FOR SEIRA DISTINCTA MARI MUTT (COLLEMBOLA: ENTOMOBRYIDAE) JosE A. MARI Mutt Department of Biology, University of Puerto Rico, Mayagiiez, Puerto Rico 00708. Abstract. —Seira cryptica n. sp. is described from specimens collected in Florida and Arizona. This species was previously confused with S. bipunctata (Packard), which is redescribed from type material and specimens from Nebraska. Seira distincta Mari Mutt is recorded for the first time from the United States and Mexico. During a recent review of the Puerto R1- can species of Seira (Mari Mutt, 1987) I considered one local species as possibly S. bipunctata (Packard), described in 1873 from Texas and redescribed by Christiansen and Bellinger (1980) from specimens col- lected in various localities of the United States. These authors reported a degree of variation in coloration and chaetotaxy that suggested their samples contained more than one species. Christiansen and Bellinger had considered this possibility but preferred to list all their records under one species. Thanks to Kenneth Christiansen, who has placed at my disposal his collection of S. bipunctata, and to the Museum of Com- parative Zoology of Harvard University, which lent the necessary type material, I have been able to reinterpret the North American material of S. bipunctata. Some of the specimens in the Christiansen collec- tion belong to Packard’s species while oth- ers belong to a new species that is described below. I received also from Dr. Christiansen some specimens from Massachusetts and Mexico that represent new records for Seira distincta Mari Mutt (1986), known previ- ously only from Puerto Rico. The type series of Seira bipunctata and the holotype of Seira cryptica n. sp. are de- posited in the Museum of Comparative Zo- ology, Cambridge, Massachusetts. The re- maining material is deposited in the collection of Dr. Christiansen, Department of Biology, Grinnell College, Iowa. In the species descriptions and discussions, Ant. 2, Th. 2, Abd. 2, etc. means second antennal segment, second thoracic segment, etc. Seira cryptica Mari Mutt, NEw SPECIES Seira bipunctata (Packard): Christiansen and Bellinger 1980: 925-926, Fig. 756 B, E, F (Florida), misidentification. Description.— Length from front of head to end of Abd. 6 up to 1.8 mm. Body with a lateral band of pigment along its length, pigment extending dorsally but with dimin- ishing intensity, leaving dorsum of body un- pigmented (Fig. 12, see also Christiansen and Bellinger, 1980: 926, Fig. B). Some lighter specimens almost white, darker specimens with pigment extending almost to midline of body. Antennae and legs light- ly pigmented throughout length. Manu- brium with some pigment basally. Anterior margin of mesonotum rounded, not pro- jecting over head. Apex of Ant. 4 without proturberance or pin seta but with bilobed papilla (Fig. 13). Head rounded, distribu- Figs. 1-7. Seira bipunctata. 1, Distribution of head macrochaetae. 2, Distribution of body macrochaetae, bothriotricha (wavy lines) and pseudopores () (setae signalled by arrows are absent in some specimens—see text. On Th. 2, a broken line surrounds the external posterior group of macrochaetae). 3, Mucro. 4, Metathoracic claws. 5, Apex of Ant. 4. 6, External labial papilla. 7, Labral papillae. 128 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VOLUME 89, NUMBER 1 tion of macrochaetae as in Fig. 14. Eyes G and H reduced, not visible in cleared spec- imens. Outer labral papillae rounded, inner papillae conical and not bifurcated (Fig. 13). Body macrochaetotaxy as in Fig. 11; 2 + 2 setae on Abd. 1. Other characters described under S. bipunctata identical in this species. Comments.— The specimen from locality 2425 has three outer macrochaetae on both sides of Abd. 3; all other specimens possess two setae. I have examined two specimens from a cave in Puebla, Mexico (coll. no. 4067) which possess the typical coloration of this species but their guts are filled with so much food that the abdominal chaeto- taxy cannot be studied. Diagnosis. — Seira cryptica is very similar to S. distincta Mari Mutt (1986), described from Puerto Rico and now reported in this paper from the United States (Massachu- setts) and Mexico. These species differ in coloration, the pigment in S. distincta is re- stricted to the sides of Th. 3 to Abd. 2 (Mari Mutt, 1986: Fig. 21). In addition, S. dis- tincta lacks the macrochaeta inserted above the pseudopore of Abd. 4 and consistently possesses three outer macrochaetae on Abd. 3 (two in S. cryptica). Seira cryptica can be distinguished from S. bipunctata by the presence of 2 + 2 mac- rochaetae on Abd. | (4 + 4 in S. bipunc- tata) and also by the coloration, although well pigmented specimens of S. cryptica may closely resemble specimens of S. bipunc- tata. Specimens of S. bipunctata possess a somewhat conical mesonotum which pro- jects over an elongated head. In S. cryptica the mesonotum does not project over the head and the head is rounded. Seira bi- punctata has an apical protuberance on Ant. 4 (absent in S. cryptica) and always has 129 three outer macrochaetae on Abd. 3 (usually two in S. cryptica). Finally, the posterior outer group of setae on Th. 2 is usually com- posed of six setae in S. bipunctata and of five in S. cryptica. Material examined.— Florida, Monroe County, on several small mangrove islands, 4.VII.1969 to 16.VII.1970, collection num- bers: 2425,.243259243352506.2827-.283. 2834; D. Simberloff, holotype and 17 para- types on slides. Arizona, nr. Phoenix, McDowell Mt. Park, on dry stream bed, 19. 111.1986, coll. no. 6620, K. Christiansen, 1 specimen on slide. Seira bipunctata (Packard) Lepidocyrtus bipunctatus Packard 1873: 37 (Texas). Seira bipunctata—Christiansen & Bellinger 1980: 925-926, Fig. 756 A, C (Nebraska and New Mexico). Description.— Length from front of head to end of Abd. 6 up to 2.4 mm. Coloration variable (see comments). Anterior margin of mesonotum conical, projecting over head. Distribution of scales: head and body, dor- sum of Ant. | and Ant. 2, dorsal proximal half of Ant. 3, all leg segments, and furcula. Ant. 4 slightly annulated; its apex (Fig. 5) with distinct protuberance and bilobed pa- pilla but without pin seta. Head elongate, macrochaetae distributed as in Fig. 1. Eyes G and H not greatly reduced. Interocular chaetotaxy consists of a small ciliated seta external to eye D, a similar seta between eyes E and F, and 5 longer setae in area between eyes C to H (arrangement identical in S. caheni Jacquemart— Mari Mutt, 1986: Fig. 10). Prelabral setae ciliated, labral setae smooth. Labral papillae well developed, sometimes bifurcated apically —_ Figs. 8,9. Seira bipunctata. 8, Distribution of violet pigment, lectotype. 9, Ventral manubrial chaetotaxy, basal area of manubrium at top of page. Figs. 10-14. Seira cryptica. 10, Labral papillae. 11, Distribution of body macrochaetae, bothriotricha (wavy lines) and pseudopores (x). 12, Distribution of violet pigment. 13, Apex of Ant. 4. 14, Distribution of head macrochaetae. 130 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (Fig. 7). Labial chaetotaxy: al—a5,M1M2r (reduced)EL1L2. One long ciliated seta on each side of ventral groove near posterior margin of head. One to 3 pairs of scales inserted between this seta and setae of post- labial quadrangle. Setae of maxillary palpus smooth, subequal in length and shape. Dif- ferentiated seta of outer labial papilla clearly surpassing apex of papilla (Fig. 6). Body macrochaetotaxy as in Fig. 2; 4 + 4 setae on Abd. 1. Detailed chaetotaxy of Abd. 2 and Abd. 3 and distribution of setae asso- ciated with anterior bothriotricha of Abd. 4 asin S. caheni (Mari Mutt, 1986: Figs. 16- 18). Trochanteral organ with up to 30 slen- der smooth setae. Structure of claws as in Fig. 4. Ventral manubrial chaetotaxy as in Fig. 9. Mucro as in Fig. 3. Male genital plate circinate, with 11 smooth setae in a circle around genital pore and at east 1+1 setae near pore. Comments.—The types of this species possess a small patch of pigment along the anterior margin of the mesothorax and a larger patch on the lateral-posterior margin of Abd. 4 (Fig. 8). On the other hand, the specimens from Nebraska are almost com- pletely pigmented as illustrated by Chris- tiansen and Bellinger (1980: 926, Fig. 756C). The first two antennal segments of the latter specimens are violet but Ant. 3 and Ant. 4 are pale. Legs are violet to the apical 4 of femur, remainder of leg is pale yellow. The specimens from New Mexico have all the antennal segments, head, and thorax pig- mented but the abdomen lacks color. No anatomical differences were detected among these populations. Setae signalled by arrows on Th. 2 and Abd. 4 in Fig. 2 are absent on one side of the body of one specimen from Nebraska. The signalled seta on Abd. 3 is absent on both sides of the body of one specimen from the same locality. Diagnosis.— This species is similar to S. caheni Jacquemart (1976), described from the Galapagos Islands and reported from Cuba (Gruia, 1983) and Puerto Rico (Mari Mutt, 1986). Both species possess identical head and body macrochaetotaxy except that S. bipunctata has six setae on the outer pos- terior group of Th. 2 (Fig. 2) while most specimens of Jacquemart’s species have five setae (some specimens have six setae). Seira caheni lacks an apical protuberance on Ant. 4 and has a pair of setae between the apical pair on the manubrium of S. bipunctata (cf. Fig. 9 and Mari Mutt, 1986: Fig. 11). These species also differ in typical coloration. Material examined.—Texas, McLennan County, Waco, Lectotype (here designated), 4 paralectotypes on slides and 35 paralec- totypes in alcohol; Nebraska, Thomas County, Halsey, 13.11.1957, coll. no. 461x, Henzlik, col., 4 specimens on slides; New Mexico, Eddy County, Carlsbad Caverns National Park, bat cave site no. 4, 25.1V.1975, coll. no. 3668, 3 ona slide with specimens of an Entomobrya. Seira distincta Mari Mutt Seira distincta Mari Mutt, in press. Described originally from Puerto Rico, this species is reported for the first time from the United States and Mexico. Seira dis- tincta is very similar to S. cryptica; the dif- ferences between both are listed in the lat- ter’s diagnosis. Material examined.—USA, Massachu- setts, Cambridge, Biological Laboratories, 26.11.1948, K. Christiansen, coll. no. 5985, 4 specimens on slides; Mexico, Veracruz, Cueva del Rio, 3 km E of Atoyac, 6.1I.1957, coll. no. 4093, J. Reddell, 1 specimen on slide. LITERATURE CITED Christiansen, K. and P. Bellinger. 1980. The Collem- bola of North America north of the Rio Grande, a taxonomic analysis. Part 3, Family Entomobryi- dae. Grinnell College, lowa, pp. 785-1042. Gruia, M. 1983. Collemboles arthropléones de Cuba récoltés par les expeditions cubano-roumaines en 1969-1973, II. Result. Exped. Biospeleol. Cu- bano-Roumaines Cuba 4: 191-205. Jacquemart, S. 1976. Collemboles nouveaux des Iles Galapagos. Mission zoologique belge aux Iles Ga- lapagos et en Ecuador (N. et J. Leleup, 1964-1965) 3: 137-157. VOLUME 89, NUMBER 1 Mari Mutt, J. A. In press. Puerto Rican species of Seira. Carib. J. Sci. 22(3-4). Packard, A. S. 1873. Synopsis of the Thysanura of 131 Essex County, Mass., with descriptions of a few extralimital forms. Rep. Peabody Acad. Sci. 5: 23- Sil: PROC. ENTOMOL. SOC. WASH. SOQ) 1987 sips 131 NOTE Brachymeria discretoidea, a new junior synonym of Brachymeria discreta (Hymenoptera: Chalcididae) Brachymeria (Gahanula) discreta Gahan and Brachymeria (Gahanula) discretoidea Gahan were described in 1942 (Proc. U.S. Natl. Mus. 92: 43-44) and considered to be similar to one another. Brachymeria dis- creta was based on specimens from Mexico, discretoidea on specimens from Panama. Brachymeria discreta was later recorded from Texas, Arizona, Southern California, and Hawaii, and discretoidea from Arizona, Texas, and Mexico (Burks, 1979. Chalcid- idae, pp. 860-874. Jn Krombein, K. V. et al., eds., Catalog of Hymenoptera in Amer- ica North of Mexico. Vol. I. Smithson. Inst. Press., Wash., D.C.). Brachymeria discreta has been reared from the nest of Polistes instabilis Sauss. (Hymenoptera: Vespidae), probably from a pyraustid moth that in- fested the wasp nest, and from a species of Tinea (Lepidoptera: Tineidae) breeding in chicken droppings. Brachymeria discretoi- dea has been reared from the nest of Trigona amalthea (Oliv.) (Hymenoptera: Apidae) infested by an unidentified moth (Gahan, ibid.; Burks, 1960, Trans. Am. Entomol. Soc. 86: 225-273). Burks (1960) indicated difficulty in de- termining material as either discreta or dis- cretoidea and that the differences noted by Gahan tended to intergrade. He chose not to synonomize the two species and indicat- ed that, except for the characters in his key (‘frontal carina always well developed ver- sus weak-virtually absent; ocellocular line %, versus *% as long as diameter of lateral ocellus; marginal vein 4 versus 3! times as long as postmarginal; propodeum with an elongate-median areola versus lacking’’), the two species were alike in color and struc- ture. Upon examining the types of both species (in the U.S. Museum of Natural History, Washington, D.C., Nos. 55149 and 55150) and additional material from various lo- calities, I have concluded that the material represents one species, that the color differ- ences and intergrading charcters of Gahan and Burks merely represent intraspecific variation. I therefore consider discretoidea a junior synonym of discreta. I thank D. J. Burdick, Department of Bi- ology, California State University Fresno, Fresno; N. J. Smith, Fresno County Agri- cultural Commissioner’s Office, Fresno, California; and R. D. Haines, Tulare Coun- ty Agricultural Commissioner’s Office, Vi- salia, California for editorial comments on this paper. I thank also the following insti- tutions and their personnel for the oppor- tunity to have examined their material of B. discreta: University of California, Davis; California Department of Food and Agri- culture, Sacramento; Florida Department of Agriculture and Consumer Affairs, Gaines- ville; Bishop Museum, Honolulu, Hawaii; National Museum of Natural History, Washington, D.C.; Systematic Entomology Laboratory, USDA, Washington, D.C.; and the Fresno County Agricultural Commis- sioner’s Office, Fresno, California. Jeffrey A. Halstead, 2/710 N. Hayes, Fres- no, California 93722. PROC. ENTOMOL. SOC. WASH. 89(1), 1987, pp. 132-136 HOST RANGE AND NEW HOST RECORDS FOR THE PLUME MOTH PLATYPTILIA CARDUIDACTYLA (LEPIDOPTERA: PTEROPHORIDAE) FROM CALIFORNIA THISTLES (ASTERACEAE) CHARLES E. TURNER, ROBERT W. PEMBERTON, AND SARA S. ROSENTHAL Biological Control of Weeds, U.S. Department of Agriculture, Agricultural Research Service, Western Regional Research Center, Albany, California 94710. Abstract. —Seventeen new host records are reported for the artichoke plume moth Pla- typtilia carduidactyla (Lepidoptera, Pterophoridae) from native and naturalized thistles (Asteraceae, Cardueae) in California. All are adult rearing records from field populations of the host thistles. The number of moths reared and the amount of host material collected for rearing are reported. The new records include the naturalized species Arctium minus and Carduus pycnocephalus, and 15 native Cirsium species: Cirsium andersonii, C. an- drewsii, C. brevistylum, C. californicum, C. campylon, C. ciliolatum, C. crassicaule, C. cymosum, C. douglasii, C. fontinale obispoense, C. hydrophilum vaseyi, C. loncholepis, C. pastoris, C. rhothophilum, and C. tioganum. Seven of the newly reported Cirsium hosts are considered rare. Previous host records are also summarized. With one exception, all known hosts are in the thistle subtribe Carduinae. The so-called artichoke plume moth, Pla- typtilia carduidactyla (Riley) (Lepidoptera, Pterophoridae) is the most serious pest of introduced cultivated globe artichoke (Cyn- ara scolymus L., Asteraceae) in California (Haynes et al., 1981; Lange, 1941; Lange et al., 1954). This moth, which is native and widespread in North America, is muitivol- tine and feeds on leaf, stem, and flowerhead tissue of its host plants (Lange, 1950). The thistle tribe (Asteraceae, Cardueae) is an im- portant group of native species, naturalized weeds, and the cultivated crops safflower (Carthamus tinctorius L.) and globe arti- choke (Moore and Frankton, 1974; Ownbey et al., 1975; Reed, 1970; Robbins et al., 1970; USDA, SCS, 1982). Our study of the host pattern of the plume moth 1s part of a broader entomofaunal study of the thistle tribe in California (Pemberton et al., 1985; Turner et al., in press) as it relates to bio- logical control of weedy naturalized thistles. The previously reported host records for P. carduidactyla are as follows: Centaurea melitensis L. (Lange, 1941, 1950); Cirsium arvense (L.) Scop. (Lange, 1950); C. calli- lepis (Greene) Jeps. (reported as C. ameri- canum Daniels var. callilepis Jeps.) (Lange, 1950): C. discolor (Muhl. ex Willd.) Spreng. (Marcovitch, 1916); C. edule Nutt. (possi- bly confused with C. brevistylum Cronq.) (Lange, 1941, 1950); C. occidentale (Nutt.) Jeps. (Lange, 1941, 1950); C. proteanum J. T. Howell (reported as C. occidentale var. venustum Jeps.) (Lange, 1941, 1950); C. quercetorum (Gray) Jeps. (Lange, 1941, 1950); C. undulatum (Nutt.) Spreng. (Lange, 1941, 1950); C. vulgare (Savi) Tenore (re- ported as C. /anceolatum (L.) Scop.) (Riley, 1869; Lange, 1941, 1950); Cynara cardun- culus L. (Lange, 1941, 1950); C. scolymus L. (Essig, 1922; Lange, 1941, 1950); Sily- bum marianum (L.) Gaertn. (Essig, 1922). Where we use a different host name from VOLUME 89, NUMBER 1 that originally reported, this was done in accordance with more recent treatments of the genus Cirsium (Jepson, 1925; Munz, 1973). There has been confusion between C. brevistylum and C. edule, which are closely related and similar (Moore and Frankton, 1962). It is doubtful that true C. edule occurs in California (Moore and Frankton, 1962; Munz, 1973). Of these pre- viously reported hosts, explicit mention of emerged adults from host material is only made for C. discolor, C. vulgare, and C. scolymus. Of the previously reported hosts, C. callilepis, C. discolor, C. edule, C. occi- dentale, C. proteanum, C. quercetorum, and C. undulatum are native to North America (WSDAY SCS, 1982). All the new host records reported here are from material collected and reared by the authors except for the records prior to 1982, which are from material collected and reared by K. E. Frick and R. B. Hawkes, formerly affiliated with our Laboratory. Host mate- rial collected in 1982 consisted of periodic samples of ten whole plants from the same field populations of native and naturalized Carduus and Cirsium species. In 1983, 1984, and 1985 we sampled the flowerheads only of field populations of native and intro- duced species of Arctium, Carduus, Cartha- mus, Centaurea, Cirsium, Cynara, Onopor- dum, Saussurea, and Silybum. We collected lateral and terminal flowerheads at a stage between flowering and seed dissemination. All of the introduced species sampled were from naturalized populations except for samples from planted fields of artichoke and safflower. The host material was brought back to the lab for the rearing of adult plume moths and other thistle insects (Pemberton et al. 1985: Turner etal, in press). D. C. Ferguson and W. H. Lange, Jr. de- termined the plume moths. The authors, G. D. Barbe, T. C. Fuller, and G. B. Ownbey determined the thistles. Voucher specimens of most plume moths and host thistles are retained in our lab. We report here 17 new host records for 133 P. carduidactyla from native and natural- ized thistle species, including information on host location and sample date, number of flowerheads sampled, and the number of emerged adults. The new host records are from the relatively common native thistles Cirsium andersonii (Gray) Petrak, C. an- drewsii (Gray) Jeps., C. brevistylum Cronq., C. californicum Gray, C. cvmosum (Greene) J. T. Howell, C. douglasii DC., C. pastoris J. T. Howell, and C. tioganum Congd.; and from the relatively rare (Federal Register, 1985) native thistles C. campylon H. Sharsm., C. ciliolatum (L. Henders.) J. T. Howell, C. crassicaule (Greene) Jeps., C. fontinale (Greene) Jeps. var. obispoense J. T. Howell, C. hydrophilum (Greene) Jeps. var. vaseyi (Gray) J. T. Howell, C. loncho- lepis Petrak, and C. rhothophilum Blake. New host records also are from the natu- ralized thistles Arctium minus Bernh. and Carduus pycnocephalus L. These newly reported host species occur in a wide variety of habitats located throughout cismontane California (Munz, 1973). The number of emerged adult plume moths was generally smaller than the num- ber of larvae and pupae associated with the host plants. This difference in number be- tween adult versus immature plume moths could have been due to insect parasitoids (Lange, 1950) or to our rearing conditions. We also reared adult plume moths from C. occidentale, C. proteanum, C. quercetorum, C. undulatum, and C. vulgare among the hosts previously reported. Native Cirsium species are the only known native North American hosts for P. car- duidactyla. In North America Cirsium con- tains ca. 130 native species (Moore and Frankton, 1974; Ownbey et al., 1975), with ca. 30 species native to California (Munz, 1973). The moth is now known from 21 Cirsium species (excluding C. edule) native to North America, and 19 species native to California. Ironically, the plume moth was first described from specimens reared from naturalized Cirsium vulgare in Missouri 134 (Riley, 1869). Riley (1869) used the perhaps more appropriate colloquial name, thistle plume moth, for P. carduidactyla. Globe artichoke has been grown com- mercially in California at least since 1900 (Stokdyk, 1932). P. carduidactyla has been a pest of globe artichoke in California at least since 1922 (Essig, 1922; Lange, 1941). Both Cirsium and Cynara are in the sub- tribe Carduinae (Dittrich, 1977), which in- dicates that they are relatively closely re- lated. The plume moth is a native stenophagous insect that is capable of host utilization of an introduced crop plant that is relatively closely related to its native hosts. All known host taxa are in the thistle tribe Cardueae; and all known hosts are in the subtribe Carduinae with the exception of Centaurea melitensis, which is in the sub- tribe Centaureinae (Dittrich, 1977). Lange (1950) reported a larval feeding record of P. carduidactyla from a C. melitensis pop- ulation adjacent to a planting of globe ar- tichoke that had the moth. We did not rear P. carduidactyla from five samples of C. melitensis, or from any of the 14 samples from 3 other species of Centaurea that we studied (unpublished data). New Host RECORDS Arctium minus. Dimmick State Park, Men- docino Co., Calif.: host plants coll. 6-[X- 62 by K. E. Frick, 9 adults emerged by 27-IX-62. Carduus pycnocephalus. 2.5 km N. of Napa, Napa Co., Calif.: host plants coll. 15-V- 63 by R. B. Hawkes, 1 adult emerged on 17-VI-63. Cirsium andersonii. Gumboot Rd., 19.0 km from Castle Lake Rd., Siskiyou Co., Cal- if.: 108 host heads coll. 22-VIII-84, 1 adult emerged by 25-IX-84. Cirsium andrewsii. Abbott’s Lagoon, Pt. Reyes, Marin Co., Calif.: host plants coll. 11-V-82, 2 adults emerged by 5-VI-82; host plants coll. 1-VI-82, 1 adult emerged from head by 22-VI-82; host plants coll. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 13-VII-82, 5 adults emerged from heads by 17-XI-82; host plants coll. 3-VII-82, 3 adults emerged from heads by 11-VI- 83; 100 host heads coll. 22-VI-85, 1 adult emerged by 5-XI-85. Cirsium brevistylum. ca. 14 km W. of Wil- lits, Mendocino Co., Calif.: host plants coll. 25-VII-61 by K. E. Frick, 2 adults emerged by 4-VIII-61; host plants coll. 2-VII-62 by K. E. Frick, 20 adults emerged by 23-VII-62. Crescent City, Del Norte Co., Calif.: host plants coll. 16- VIII-61 by K. E. Frick, 5 adults emerged by 5-IX-61. Mt. Tamalpais, Marin Co., Calif.: host plants coll. 5-VII-62 by K. E. Frick, 3 adults emerged. (Additional later host records from Del Norte and Men- docino counties coll. by K. E. Frick avail- able from the authors.) Abbott’s Lagoon, Pt. Reyes, Marin Co., Calif.: 100 host heads coll. 17-V-84, 3 adults emerged by 1-VIII-84; 100 host heads coll. 5-VII-84, 3 adults emerged by 14-VIII-84. MacKerricher Beach, Mendocino Co., Calif.: 89 host heads coll. 10-VII-84, 17 adults emerged by 14-VIII-84. Cirsium californicum. Hwy. 49 between Downieville and Sierra City, Sierra Co., Calif.: 100 host heads coll. 21-VI-84, 1 adult emerged by 31-VII-84. Cirsium campylon. Blackbird Valley off Mines Rd., Santa Clara Co., Calif.: host plants coll. 8-VI-82, 1 adult emerged from head by 30-VI-82; host plants coll. 1-VII- 82, 1 adult emerged from head by 21- VII-82; host plants coll. 31-VIII-82, 2 adults emerged from head by 17-XI-82. Mines Rd., 11.6 km S. of Alameda—San- ta Clara county line, Santa Clara Co., Calif.: 100 host heads coll. 30-VI-85, 1 adult emerged by 5-XI-85. Cirsium ciliolatum. Y ork Rd. at Mulloy Rd., Siskiyou Co., Calif.: 100 host heads coll. 27-VI-84, 1 adult emerged by 30-VII-84. Cirsium crassicaule. Kern Nat. Wildlife Refuge, Kern Co., Calif.: 60 host heads coll. 23-VI-85, 1 adult emerged by 5-VII- 85. VOLUME 89, NUMBER 1 Cirsium cymosum. Callahan—Cecilville Rd., 8.0 km from Callahan, Siskiyou Co., Calif.: 67 host heads coll. 5-VIII-83, 1 adult emerged by 22-XII-83. Cirsium douglasii. Callahan—Cecilville Rd., 8 km from Callahan, Siskiyou Co., Calif.: host plants coll. 27-VII-82, 1 adult emerged from heads by 18-VIII-82; host plants coll. 17-VIII-82, 1 adult emerged from heads by 17-XI-82. Hwy. 3, 3.9 km S. of Scott Mt. Pass, Trinity Co., Calif:: 100 host heads coll. 4-VIII-84, 2 adults emerged by 3-IX-84. Cirsium fontinale obispoense. San Simeon Cr. Rd., 8.4 km from Hwy. 1, San Luis Obispo Co., Calif.: 100 host heads coll. 26-V-84, | adult emerged by 31-VII-84. Cirsium hydrophilum vaseyi. West Point Inn trail, Mt. Tamalpais, Marin Co., Calif.: 100 host heads coll. 22-VI-85, 1 adult emerged by 5-XI-85. Cirsium loncholepis. Guadalupe Dunes, San Luis Obispo Co., Calif: 60 host heads coll. 14-VI-83, 1 adult emerged by 22- XII-83. Cirsium pastoris. Callahan—Cecilville Rd., 2-8 km from Callahan, Siskiyou Co., Calif.: host plants coll. 7-VII-82, 1 adult emerged from stem by 21-VII-82; 100 host heads coll. 9-VIII-84, 1 adult emerged by 4-X-84. Slough Rd., S. of Louie Rd. exit from I-5, Siskiyou Co., Calif.: 100 host heads coll. 29-VI-83, 3 adults emerged by 1-X-83. Edgewood exit from I-5, 5.6 km N. of Weed, Siskiyou Co., Calif.: 100 heads coll. 9-VIII-84, 3 adults emerged by 4-X-84. Cirsium rhothophilum. Guadalupe Dunes nr. Oso Flaco Lake, San Luis Obispo Co., Calif.: 42 host heads coll. 15-VI-83, 2 adults emerged by 5-VII-84. Cirsium tioganum. Willow Cr. Rd., Siski- you Co., Calif.: 82 host heads coll. 12- VII-84, 1 adult emerged by 30-VII-84. Dorris Brownell Rd., nr. jct. with Willow Cr. Rd., Siskiyou Co., Calif.: 100 host heads coll. 18-VII-85, 9 adults emerged by 6-XI-85. 135 ACKNOWLEDGMENTS K. F. Haynes and D. MacNeill reviewed the manuscript. D. C. Ferguson (Systematic Entomology Laboratory, Agricultural Re- search Service, U.S. Department of Agri- culture) and W. H. Lange, Jr., helped de- termine the P. carduidactyla. G. D. Barbe, T. C. Fuller, and G. B. Ownbey helped de- termine the thistle species. R. Colville and M. T. Johnson, Jr., helped with the rearing and processing of insects, and the latter also helped with the sampling of thistleheads. LITERATURE CITED Dittrich, M. 1977. Cynaraea-systematic review, pp. 999-1038. In Heywood, V. H., J. B. Harborne, and B. L. Turner, eds., The biology and chemistry of the Compositae, 2 vols. Academic, New York. Essig, E.O. 1922. The artichoke plume moth. Calif. Dept. of Agric. Monthly Bull. 11: 454-456. Federal Register. 1985. Endangered and threatened wildlife and plants; review of plant taxa for listing as endangered or threatened species; notice of re- view. Federal Register 50: 39526-39584. Haynes, K. F., M. C. Birch, and J. A. Klun. 1981. Sex pheromone offers promise for control of ar- tichoke plume moth. Calif. Agric. 35: 13-14. Jepson, W.L. 1925. A manual of the flowering plants of California. Univ. of California, Berkeley. Lange, W. H., Jr. 1941. The artichoke plume moth and other pests injurious to the globe artichoke. Calif. Agr. Exp. Sta. Bull. 653. . 1950. Biology and systematics of plume moths of the genus Platyptilia in California. Hilgardia 19: 561-668. Lange, W. H., R. H. Sciaroni, and A. S. Greathead. 1954. Artichoke plume moth damage. Calif. Agric. 8(7): 7-8, 12. Marcovitch, S. 1916. Insects attacking weeds in Min- nesota. Minn. State Entomol. 16th Report: 135- 152. Moore, R. J. and C. Frankton. 1962. Cytotaxonomy and Canadian distribution of Cirsium edule and Cirsium brevistylum. Can. J. Bot. 40: 1187-1196. 1974. The thistles of Canada. Research Branch, Canada Dept. Agriculture, Monograph No. 10. Munz, P.A. 1973. A California flora and supplement. Univ. of California, Berkeley. Ownbey, G. B., P. H. Raven, and D. W. Kyhos. 1975. Chromosome numbers of some North American species of the genus Cirsium. III. Western United States, Mexico, and Guatemala. Brittonia 27: 297- 304. 136 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Pemberton, R. W., C. E. Turner, and S. S. Rosenthal. 1985. New host records for tephritid flies from Cirsium and Saussurea thistles in California. Proc. Entomol. Soc. Wash. 87: 790-794. Reed, C. F. 1970. Selected weeds of the United States. U.S. Dept. of Agric., Agric. Hdbk. No. 366. Riley, C. V. 1869. First annual report on the noxious, beneficial, and other insects of the state of Mis- souri. pp. 180-181. State of Missouri, Jefferson City. Robbins, W. W., M. K. Bellue, and W. S. Ball. 1970. Weeds of California. Calif. Dept. Agric., Sacra- mento. Stodyk, E. A. 1932. Marketing globe artichokes. Cal- if. Agric. Exp. Sta. Bull. 524. Turner, C. E., R. W. Pemberton, and S. S. Rosenthal. Host utilization of native Cirsium thistles (Aster- aceae) by the introduced weevil Rhinocyllus coni- cus (Coleoptera: Curculionidae) in California. En- viron. Entomol. (In press.) U.S. Dept. Agriculture, Soil Conservation Service. 1982. National list of scientific plant names. Vol. 1. List of plant names. PROC. ENTOMOL. SOC. WASH. 89(1), 1987, pp. 137-140 A NEW NEARCTIC DIXA4A (DIPTERA: DIXIDAE) FROM PENNSYLVANIA T. MICHAEL PETERS Professor and Curator, Department of Entomology, University of Massachusetts, Am- herst, Massachusetts 01003. Abstract.— Dixa adleri, new species, from Pennsylvania is described with illustrations of the mature larva, male genitalia, and female bursa copulatrix. Adults are compared with other known Nearctic dixids. Larval habitats of dixid genera are compared. Larval Dixidae live in or just above the meniscus in a wide variety of freshwater habitats, from stagnant bogs to rushing brooks. The three Nearctic genera have been associated in the literature with distinct types of freshwater environments, differences that have been correlated with unique anatom- ical characteristics of each of the genera. However, comparison of environmental data available for a new species of Dixa found in eastern United States with those of my previous collections, revealed some anomalies. Following the description of the new species is a discussion of overlap in larval habitats of the two most widely dis- tributed genera; Dixa Meigen and Dixella Dyar and Shannon. Dixa adleri Peters, NEw SPECIES Figs. 1-4 Adult.— Head: Medium brown in speci- mens preserved in alcohol; without micro- trichia; a line of 14 long setae along periph- ery of compound eye from dorsum of vertex to venter of eye; frontoclypeus with 2 or 3 setae near ventral edge; antennae concol- orous with head, first flagellomere subcylin- drical, width:length 1:10. Thorax: Medium brown in mature specimens, but with dis- tinctly darker vittae on scutum, vittae in- distinguishable in teneral individuals; an- terior pronotum of male with 6-8 setae (9- 11 in ) as long as width of sclerite; 1 seta near dorsal suture on posterior pronotum; scutellum of males with a transverse row of 11 setae, the central one with another pos- terior to it, scutellum of females with an anterior transverse row of 13 setae, followed by a transverse row of 3 setae and followed by a single mesal seta. Wing: Clear, without pigmented areas, length 3.25-3.69 mm (3)(3.69-4.08 in 9); in 6 M3 + 4:M1 + 2 as 1:1.4-1.6, M3 + 4:Mst as 1:1.5-2.5; R2 + 3:R3 as 1:2.1-3.1; in 9, vein ratios are within same range as those of 6. Crossvein m-cu broken. Halter: Hyaline. Legs: Distal spiniform seta on 3rd tarsomere of foreleg (on tarsomeres 3-4 of 1 2), on 1-3 of midleg and on 1-4 of hindleg; basal recurved spi- niform seta on tarsomere 5 of fore and mid- leg of 4; claws simple in 2, in ¢ hind claw simple, fore and midclaws with 5-6 ventral teeth (3-4 long, | short, 1 long); femur:tibia: tarsus length ratios of forelegs as 1:0.96—- 1.00:1.50-1.90 in 4, in 2 1:0.96—1.00:1.40- 1.48; midleg 1:0.92—1.05:1.13-1.74 in 4, in 2 1:1.00-1.05:1.32-1.41; hindleg 1:1.00- 1.05:1.54-1.74 in 6, in 2 1:1.14—-1.21:1.67- 1.95. Abdomen: Mottled brown and grey, mixed lighter and darker areas; in 4, 9th sternite widest laterally, wider than the more lightly sclerotized tergite 9; tergite 10 with non-segmented cerci (Fig. 1); gonocoxite and gonostylus as in Figs. | and 2; ejaculatory 138 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-4. Dixa adleri. 1, Lateral view of male terminalia. 2, Dorsal view of same. 3, Bursa copulatrix. 4, Posterior abdominal structures of fourth instar larva; left half dorsal, night half ventral. duct very short and very lightly sclerotized; Fourth instar larva.— Length 6.6—9.0 mm claspette and penis valves not discernable (n = 8); coloration of specimens available is in cleared glycerin mount, even under in- _ bleached out, probably due to preservation terference microscopy; ? with three tufts of | techniques. Terminology follows Peters and setae in bursa copulatrix (Fig. 3). Adamski (1982). Head: Seta 10 about 2 VOLUME 89, NUMBER 1 length of 9, length of 7 intermediate between them; head seta | long, wide; seta 2 slender but longer than 1, 3 very short and wide, widest at middle. Antenna: Tip with several sharp spines surrounding a larger, pear- shaped, thin-walled sensillum. Thorax: An- terior ventral setae of prothorax not ex- tending beyond mouth brushes, arranged as 1-1-4-1-1-4-1-1, each group of 4 without common sclerotized base. Abdomen: Pro- legs of segments | and 2 with uniordinal, biserial crochets of subequal length, anterior proleg preceded by row of spicules subequal in length to crochets, 25 crochets in first row, second row thickest, with 24 crochets; posterior proleg with 21 crochets per row, posterior row thickest, followed by an un- even line of thin spicules which are subequal in length to crochets; corona of branched spicules on dorsum of segments 2-7; am- bulatory combs on venter of segments 5—7 each with 9-10 spines on either side of a medial subrectangular plate, spines in 2 dis- tinct rows of alternating heavy and more slender spines, heavier ones projecting at less acute angle from surface of segment; segment 8 with only 6 slender setae on ven- ter, shorter than length of segment; paraspi- racular setae, postspiracular process, and metaspiracular plate as in Fig. 4; segment 9 with fringe of setae continuous from pos- terolateral process anterior to postspiracu- lar process, median plate as in Fig. 4, pecten of anterolateral plate simple, as in Fig. 4; ventrolateral plate of segment 10 with a thick seta directed posteriorly, another arising near its base, is much more slender and directed laterally, anterior to the plate is another more dorsal hair; caudal hairs of postanal process over 2x length of process. Specimens examined.— Holotype é, Slab Cabin Run, Pine Grove Mills, Centre Co., Pennsylvania, 18-IX-81, collected by Peter H. Adler with Malaise trap over stream; deposited in the Peters dixid collection, De- partment of Entomology, University of Massachusetts, Amherst, Massachusetts. Eighteen paratype 7 6 and 11 2, same data as holotype, with associated immature skins. 139 Additional sites, all in Pennsylvania: 29 specimens from Scott Road (Slab Cabin Run), State College, and Briesly. DISCUSSION If keyed using Peters and Cook (1966), mature D. adleri males come out as Dixa jraterna Garrett. They may be separated on the basis of the ejaculatory duct: it is large and heavily sclerotized in D. fraterna, but scarcely visible in D. adleri. Teneral males without distinctly darker scutal vittae key to D. inextricata but have more than 5 setae on the anterior pronotum. Females key to D. fluvica but may be separated by details of the bursa copulatrix. The three tufts of setae each possesses are subequal in length in D. fluvica, with the median group con- sisting of 3-4 setae. In D. adleri the median group 1s distinctly shorter than the other two and consists of 5-6 setae. All setae in the bursa copulatrix of D. adleri are very stout. They are slender in D. fluvica. Collection sites for Dixa adleri are char- acterized in Adler et al. (1983) and Adler and Kim (1984). Adults were collected with a Malaise trap erected over the stream. Lar- vae were collected with drift nets. Several adults with associated larval and/or pupal skins were reared by Dr. Adler after whom the species 1s named. In his lengthly discussion of larval habi- tat, Nowell (1952) observed that Dixa is a fast water form “always found out in the center of streams where the water 1s swiftest ..., Meringodixa inhabits ‘“‘quieter, smoother streams. . . congregating in quiet niches, or quiet pools .. .”” while Dixella is “in two habitats [it] may be in swiftly run- ning streams or in very still waters.” He continued, “This is the only one of the groups which is represented in the quiet pools.”’ These observations were based on California dixids and fit well the anatomical differences among larvae of each of the gen- era. Dixa has coronae of branched hydro- fuge hairs on abdominal segments 2-7, while Dixella has no flotation structures other than those on the posterior that surround the spi- 140 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON racles in all known dixid larvae. Meringo- dixa has only the posterior flotation struc- ture, but it has long lateral tufts of hairs on abdominal segments 2-6 that are reported to aid in flotation (Nowell, 1951). While my own observations of Dixa and Dixella from other parts of the U.S. gen- erally agree with Nowell’s, the actual situ- ation is more complex. Average current for my field collections is measured by timing a half-filled plastic vial over a measured dis- tance. Dixa was collected from moving waters in 8 of 12 instances, with average current of 1.83 ft/s (range = 0.67-3.5 ft/s). Dixella was collected from predominantly still water— ponds, bogs, pools—in 14 of 23 instances, with average current in the other collections of 0.94 ft/s (range of 0.1-3.0 ft/ s). An additional difference is that Dixa sometimes (4 of 12 instances) occurs where “emergent” vegetation is lacking, but Dix- ella, with one exception noted below, al- ways (23 of 23 instances) is found where vegetation emerges from the water or vege- tation on the bank (grasses, roots) hangs into the water. In six collections, where both Dixa and Dixella were present, four of six sites were still water, but current in the other two ranged from 1.0-2.6 ft/s. In one site, both genera were taken from rocky banks; all oth- ers had ‘‘emergent”’ vegetation. Larval habitats of Dixa and Dixella ap- parently overlap to a great extent, with Dixa predominant in rapidly moving water and Dixella more common in still water. Dixa with its 6 coronae of branched, hydrofuge spicules should more effectively stay afloat in rapid water if caught in the current or undergoing a diel drift (Waters, 1962). Thus, Dixa has an anatomical advantage 1n rap- idly moving water, but is not restricted to this environment. Dixella occurs in both moving and still water, but usually is found in areas with less rapid current than Dixa. They do occur together in both habitats, suggesting that larval habitat is controlled by two factors, maternal selection of ovi- position site and sites available to the ovi- positing female. ACKNOWLEDGMENT This research was partially supported by Hatch project #541 of the Massachusetts Agricultural Station. LITERATURE CITED Adler, P. H. and K. C. Kim. 1984. Ecological char- acterization of two sibling species, HIL-1 and IS- 7, in the Simulium vittatum complex (Diptera: Simuliidae). Can J. Zool. 62: 1308-1315. Adler, P. H., R. W. Light, and K. C. Kim. 1983. The aquatic drift patterns of black flies (Diptera: Sim- uliidae). Hydrobiologica 107: 183-191. Nowell, W. R. 1951. The dipterous family Dixidae in Western North America (Insecta:Diptera). Mi- croentomol. 16: 187-270. Peters, T. M. and D. Adamski. 1982. A description of the larva of Dixella nova (Walker)(Diptera:Dix- idae). Proc. Entomol. Soc. Wash. 84: 521-528. Peters, T. M. and E. F. Cook. 1966. The Nearctic Dixidae (Diptera). Misc. Publ. Entomol. Soc. Am. 5: 231-278. Waters, T. F. 1962. Diurnal periodicity in the drift of stream invertebrates. Ecology 43: 316-320. PROC. ENTOMOL. SOC. WASH. 89(1), 1987, pp. 141-146 A NEW SPECIES OF PELTOPERLA FROM EASTERN NORTH AMERICA (PLECOPTERA: PELTOPERLIDAE) B. P. STARK AND B. C. KONDRATIEFF (BPS) Department of Biology, Mississippi College, Clinton, Mississippi 39058; (BCK) Department of Entomology, Colorado State University, Fort Collins, Colorado 80523. Abstract.—A new species, Peltoperla tarteri Stark and Kondratieff, from Virginia and West Virginia is described. This new species was previously confused with P. arcuata Needham, but males lack the peculiar, basally curved cerci of that species. Both species inhabit springs and springbrooks of the central Appalachians but there are no known co- occurences. The Nearctic genus Pe/toperla presently includes a single eastern species, P. arcuata Needham (Stark and Stewart, 1981). This species was described from a female adult collected in Ithaca, New York (Needham, 1905). Twenty years later Needham and Claassen (1925) described the male from two specimens taken in Ithaca and at Pres- ident (Venango County), Pennsylvania. The nymph of this species was reared and de- scribed by Claassen (1931), also from Ith- aca. Frison (1942) redescribed the male in order to note the unusual cercal shape, pre- viously not emphasized. During a study of Nearctic peltoperlid genera by Stark and Stewart (1981), speci- mens lacking this peculiar cercal character were examined from Virginia and West Vir- ginia. These specimens were used to illus- trate the male genitalia for the genus Pel- toperla in the above paper. A study of these and additional material indicates that these specimens actually represent an unde- scribed species. Terminology used in this paper follows Stark and Stewart (1981). The following museums, institutions and individuals provided material for study: R. W. Baumann, Brigham Young University; O.S. Flint, United States National Museum of Natural History; R. F. Kirchner, Hun- tington, West Virginia; J. K. Liebherr, Cor- nell University; E. C. Masteller, Behrend College; R. F. Surdick, Front Royal, Vir- ginia; and J. R. Voshell, Virginia Polytech- nic Institute and State University. Peltoperla arcuata Needham Needham and Claassen (1925), Frison (1942) and Hitchcock (1974) provided ad- equate descriptions of this species. We are providing additional comparative figures of the genitalia and eggs to facilitate identifi- cation of both species (Figs. 3, 4, 11-15, 17). Specimens were examined from the fol- lowing locations: KENTUCKY: Boyd Co., Ashland; Powell Co., Mull Crk, Natural Bridge State Park. NEW YORK: Tompkins Co., Ithaca (holotype); Ringwood. PENN- SYLVANIA: Centre Co., Penn-Roosevelt Dam; Elk Co., Ridgeway Spring; Watercress Spring; Erie Co., 6-Mile Crk; Fulton Co., Crystal Spring; Westmoreland Co., Laugh- intown, Furnace Run. VIRGINIA: Dick- erson Co., Laurel Branch, Breaks Int. Park; Shenandoah Co., Springs, Little Sluice Mountain; Wythe Co., East Fork; Stoney Fork Reed Crk. WEST VIRGINIA: Brax- ton Co., Laurel Run, Falls Mill; Greenbrier 142 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-4. Co., Coats Run, North Fork Cherry Riv; Logan Co., Frogtown Hollow Copperas Mine Fork; Mingo Co., Laurel Fork Pigeon Crk; Pocahontas Co., Fork of Tea Crk. Peltoperla tarteri Stark and Kondratieff, NEw SPECIES Peltoperla arcuata: Stark and Stewart (198 1), in part. Male.—Forewing length 11-12.5 mm. General color brown patterned with pale brown. Tergum 9 unmodified, without dor- sal lobe. Paraprocts and tergum 10 typical of genus. Cerci essentially straight; first seg- ment elongate but not swollen basally (Figs. 5, 8, and 9). Vesicle on sternum 9 slightly wider than long and rounded apically (Fig. Peltoperla eggs. 1, P. tarteri, 300. 2, P. tarteri, detail of chorion and micropyles (M), 1000 x. 3, P. arcuata, 340. 4, P. arcuata, detail of chorion and micropyles, 1000. 6). Aedeagus membranous and multilobed; apex terminates in two sparsely setose lat- eral lobes and 3 mesal finger-like lobes cov- ered with fine spicules; a pair of spiculate mesolateral lobes are occasionally not everted (Fig. 10). Female.— Forewing length 12-13.5 mm. Subgenital plate truncate apically; vaginal sclerite sides parallel (Fig. 16). Egg.—Spherical to slightly ovoid; collar absent. Chorionic surface finely punctate giving surface a rough appearance. Micro- pyles typical of genus (Figs. 1, 2). Nymph.—Presently indistinguishable from P. arcuata. Etymology.—This species is named in honor of Donald C. Tarter, Marshall Uni- versity, Huntington, West Virginia. Material examined.—Holotype 4, allo- VOLUME 89, NUMBER 1 We x é YY \\ GLOEC eK \ LS ra / \ Uf WN; I a \ \ / TARE PESO Mi aces l Meets Ale NNN | i Ot A\/4 (aries a yy } 143 10 Figs. 5-10. P. tarteri, male genitalia. 5, Terminalia dorsal. 6, Sternum 9. 7, Epiproct, lateral. 8, Paraprocts and epiproct, ventral (Giles Co., Virginia). 9, Paraprocts and epiproct, ventral (Wyoming Co., West Virginia). 10, Aedeagus, ventral. type 2, 23 paratype 6 and 43 paratype 8, West Virgina, Fayette Co., Big Hollow of Paint Creek, 19 May 1979, R. F. Kirchner. Additional paratypes: VIRGINIA: Craig Co., Hollow Hill Farms, 26 June 1977, B. Kondratieff, 1 4, 1 2? (VPI); Floyd Co., trib- utary of Little Riv., 8 June 1978, B. Kon- dratieff, 5 6 (VPI); Giles Co., Little Stoney Crk, 14 July 1971, M. Kosztarab, 2 4 (USNM); same location, 26 June 1977, B. Kondratieff, 3 6, 1 @ (VPI); same location 24 June 1978, B. Kondratieff, 7 4, 1 ¢ (VPI); Mountain Lake, small spring on Co. Rt. 700, B. Kondratieff, 1 6, 1 2? (BPS); Mud Branch, Mountain Lake, 26 June 1977, B. Kondra- tieff, 6 6, 2 2? (VPI); Virgin Timber Area, Mountain Lake, 15/18 July 1978, K. A. and C. R. Parker, 5 3, 1 ¢ (VPI); Spring near Co. Rt. 613, 24 June 1978, B. Kondratieff, 9 4, 5 2(VPI); Greene Co., Pocosin Cabin, Shen- 144 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Veal ; oe Mek ball Figs. 11-15. ventral. 14, Aedeagus, ventral. 15, Epiproct, lateral. andoah National Park, 20 July 1973, O. S. Flint, 1 4, 1 ¢ (USNM); Nelson Co., Stony Crk, Co. Rt. 751, 28 June 1983, 1 & (BPS); Patrick Co., Spring into Rock Castle Crk, 10 May 1983, B. Kondratieff, 1 4, 1 ¢(BCK); Rockingham Co., Shenandoah National Park; 30) May: 1976" ©F S.. Flint, 1.6, 12 (USNM); Rappahannock Co., Shenandoah National Park, Skyline Drive MP 71, 24 P. arcuata, male genitalia. 11, Terminalia dorsal. 12, Sternum 9. 13, Paraprocts and epiproct, June 1961, O. S. Flint, 4 4, 1 ¢ (USNM). WEST VIRGINIA: Wyoming Co., Clear Fork Guyandotte Riv, 19 March 1978 (reared), L. Evans, 3 6 (RFK). The holotype and allotype are deposited in the United States National Museum of Natural History (USNM), the paratypes in the authors’ collections (BPS, BCK), and in those of R. F. Kirchner (RFK), Virginia VOLUME 89, NUMBER 1 n / N 7 ‘ i =e aa Nee g ea rar, Figs. 16,17. Peltoperla female genitalia. 16, P. tar- terl. 17, P. arcuata. Polytechnic Institute and State University (VPI), and the Monte L. Bean Museum, Brigham Young University (MLBM). Diagnosis and discussion.— Males of P. tarteri are easily separated from P. arcuata by the absence of the curved cerci and by the lack of a dorsal lobe on tergum 9. The 145 aedeagal apex is also quite different, being apically multilobed in P. tarteri (Fig. 10). The epiprocts of both species are variable. Figures 8, 9 indicate the range of variation for this structure in P. tarteri. Females of both species are similar; how- ever, the subgenital plate of P. tarteri is longer and typically more truncate apically, whereas that of P. arcuata is broadly round- ed apically. Internally the vaginal sclerite of P. arcuata is broader across the anterior margin than posteriorly, whereas in P. tar- teri the sides of the sclerites are parallel. The eggs are similar for both species, although subtle differences are evident in Figs. 2 and 4. Nymphs of both species are very similar and examination of reared material provid- ed no useful characters for separation. The distribution of these species 1s some- what enigmatic, since their ranges overlap broadly in the Blue Ridge, Ridge and Valley and Appalachian Plateau physiographic provinces of Virginia and West Virginia. Both species of Peltoperla typically occur only in crenon habitats (spring sources and springbrooks) of the higher, central Appa- lachians. The two species have not been tak- en together, however, they are often col- lected with species of Tallaperla such as T. anna (Needham and Claassen) and 7. ma- ria (Needham and Claassen). ACKNOWLEDGMENTS We thank the individuals listed above for arranging the loans of specimens. Sarah Fai- son, Univ. of Mississippi School of Den- tistry, assisted in preparing SEM micro- graphs. This study was supported by NSF grant #BSR-8407455. LITERATURE CITED Claassen, P. W. 131. Plecoptera nymphs of North America north of Mexico. Thomas Say Found. Entomol. Soc. Am. 3: 1-199. Frison, T. H. 1942. Studies of North American Ple- coptera with special reference to the fauna of Il- linois. Bull. Ill. Nat. Hist. Surv. 22: 235-355. Hitchcock, S. W. 1974. Guide to the insects of Con- 146 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON necticut. Part VII. The Plecoptera or stoneflies of graph of the Plecoptera or stoneflies of America Connecticut. State Geol. Nat. Hist. Surv. Conn. north of Mexico. Thomas Say Found. Entomol. 107: 1-262. Soc. Am. 2: 1-397. Needham, J. G. 1905. New genera and species of Stark, B. P. and K. W. Stewart. 1981. The nearctic Perlidae. Proc. Biol. Soc. Wash. 18: 107-110. genera of Peltoperlidae (Plecoptera). J. Kans. Needham, J. G. and P. W. Claassen. 1925. A mono- Entomol. Soc. 54: 285-311. PROC. ENTOMOL. SOC. WASH. 89(1), 1987, pp. 147-156 SAWFLIES (HYMENOPTERA: SYMPHYTA) IN URBAN ENVIRONMENTS IN THE WASHINGTON, D.C. AREA DAvID R. SMITH AND EDWARD M. BARROWS (DRS) Systematic Entomology Laboratory, BBII, Agricultural Research Service, U.S. Department of Agriculture, c/o National Museum of Natural History NHB 168, Wash- ington, D.C. 20560; (EMB) Department of Biology, Georgetown University, Washington, DC. 20057¢ Abstract.— Malaise traps were used to determine the species, abundances, and seasonal occurrences of adult sawflies in the Washington, D.C., metropolitan area. The traps, placed in two urban environments for six years and three more natural habitats for two years from March to November, obtained 948 sawflies representing 117 species. Peaks in num- bers of species and specimens were in May. The more abundant species were Acordulecera pellucida (Konow), Schizocerella pilicornis (Holmgren), Allantus nigritibialis (Rohwer), and Ametastegia pallipes (Spinola). Known hosts for all captured species include repre- sentatives of 53 plant genera, with Rosaceae being the most utilized plant family in urban environments. Notes on selected species are given. Urban environments are becoming more common and are of increasing biological in- terest (Frankie and Koehler, 1983), but no quantitative study has yet been done on the urban sawfly fauna. As larvae, all sawfly species are plant feeders, and many are, or have the potential of becoming, important plant pests in urban environments. There are about 1100 species in North America (Smith, 1979). The purpose of this study is to determine what sawfly species are present in urban environments, their abundances, seasonal occurrences, and host-plant utili- zations. The urban environment in our study consisted of relatively well manicured and ornamentally landscaped yards and semi- natural areas within a metropolitan region. MATERIALS AND METHODS Five sites were studied in the Washing- ton, D.C., metropolitan area, two urban en- vironments for six years and three more natural environments for two years. The natural environments were less disturbed, non-landscaped areas. The urban environ- ments were at residences in Glen Echo, Montgomery Co., Maryland (yard-garden trap), and near Annandale, Fairfax Co., Vir- ginia (yard trap). The natural environments were woodland (wooded-stream-habitat trap), woods edge (ecotone trap), and field (field trap), at the David W. Taylor Naval Ship Research Center (TNSRC) in Mont- gomery Co., Maryland, 5 km NNW of Glen Echo. These sites are described in more de- tail by Barrows (1986). Because only three specimens were collected in the field trap, they are combined with the ecotone trap in Table 1. Bioequip® (Santa Monica, Calif.) Mal- alse traps were used at the Glen Echo and TNSRC sites. They are 2 m tall, pyramidal, with four 0.8 m? rectangular openings. In Fairfax Co., a Townes-style Malaise trap was used. It is rectangular with a 1.8 m?’ opening on each side and a killing jar at one 148 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Families, species, number captured, flight times (earliest and latest dates), and recorded hosts of sawflies. YT = yard trap, Fairfax Co., Va.; YGT = yard-garden trap, Glen Echo, Md.; WSHT = woodland- stream habitat trap, TNSRC; ET = ecotone trap, TNSRC. Recorded hosts are from Smith (1979) and Gibson (1980). Families, Species AL VGiseeWwSHE ET Total Dates Recorded Hosts XYELIAE Xyela bakeri Konow 2 0 0 0 2 ~~ ‘III-20; IV-7 Pinus Xyela obscura (Strobl) 13 l 0 0 14 III-20-IV-17 Pinus PAMPHILITDAE Pamphilius middlekauffi Shi- l 0 0 0 1 IV-28 Corylus nohara and Smith Pamphilius ochreipes (Cres- 2 0 1 0 3. ~IV-30-V-9 Viburnum son) Pamphilius pullatus (Cresson) 0 0 1 4 5 V-1-V-8 Viburnum Onycholyda luteicornis (Nor- 8 6 0 0 14. IV-29-V-26 Rubus ton) Onycholyda rufofasciatus 0 l 0 0 1 Ville12 (Norton) PERGIDAE Acordulecera dorsalis Say 27 0 13 8 48 IV-28-VI- Quercus, Corylus, 30; VIII-4 Juglans, Casta- nea Acordulecera maculata Mac- 1 0 0 0 1 V-29 Gillivray Acordulecera mellina Mac- 3 4 0 0 7 VI-12-VIII-28 Gillivray Acordulecera pellucida (Ko- 49 14 0 0 630) IVE25=xel now) ARGIDAE Sphacophilus cellularis (Say) 1 1 0 0 2 ~~ VII-19; VIII-7 Ipomaea, Convol- vulus Arge humeralis (Beauvois) l 2 0) 0) 3. V-10-VI-19 Rhus Arge clavicornis complex 0 0) 0 l 1 V-15 Schizocerella pilicornis 6 29 0 3 138 #£VI-2-X-24 Portulaca (Holmgren) Sterictiphora serotina Smith 1 2 0 0) 3. ~IV-21-V-1 Prunus Atomacera decepta Rohwer 0 l 0 0 1 V-19 Hibiscus Atomacera debilis Say 3 1 0 0 4 V-19-IX-2 Desmodium CIMBICIDAE Zaraea lonicerae (L.) 6 a 4 2 19 IV-14-V-30 Lonicera DIPRIONIDAE Monoctenus sp. 0 ] 0 0 V-6 Juniperus Diprion similis (Hartig) l 0 0 0 IV-27 Pinus XIPHYDRIIDAE Xiphydria maculata Say 20 0 0 0 20 V-14—-VII-24 Acer Xiphydria tibialis Say 2 0 0 0 Dy Viel22 VE? Ulmus, Betula, Fraxinus, Quercus, Rhus, Tilia, Prunus, Crataegus VOLUME 89, NUMBER 1 Table 1. Continued. 149 Families, Species YT YG WSHT “EL Total Dates Recorded Hosts SIRICIDAE Tremex columba (L.) l 0) 0) 0 1 VIII-4 deciduous trees CEPHIDAE Janus integer (Norton) 3 0 0 0 3. V-26-VI-2 Ribes TENTHREDINIDAE Selandriinae Hemitaxonus albidopictus l 0 0 0 1 V-16 Onoclea (Norton) Hemitaxonus dubitatus (Nor- 3 0 0 0 3. =~VI-23-VII-16 Onoclea ton) Aneugmenus flavipes (Norton) 1 0 0 0 1 VII-15 Pteridium Heptamelus ochroleucus (Ste- 0 0 0 3. =V-26; IX-16 ferns phens) Dolerinae Dolerus nitens Zaddach 0 7 1 0 38 =IV-5-IV-16 grasses Dolerus unicolor (Beauvois) 0 2 0 0 2 IV-7: V-8 Phelum Loderus vestigialis apricus 1 0 2 0 3. ~V-22-V-29 Equisetum (Norton) Nematinae Cladius difformis (Panzer) 6 9 0 0 15 IV-22-VIII-15 Rosa Priophorus pallipes (Lepele- 0 1 0 0 1 V-12 Prunus, Cratae- tier) gus, Alnus Hoplocampa marlatti Rohwer 0 3) 0 0 3. ~IV-30-V-8 Prunus ? Craterocercus fraternalis (Nor- 0 2 0) 3. IV-17-IV-24 Quercus ton) Euura sp. 1 0 0 0 1 IV-28 Salix Nematus lipovskyi Smith 2 2 16 0 20 IV-16-V-9 Rhododendron Nematus erythrogaster (Nor- 0 0 1 0 1 IxX-4 Alnus ton) Nematus hudsoniimagnus 0 0 0 1 1 V-22 Populus Dyar Nematus oligospilus Foerster 0 0 0 1 1 SVels Salix Nematus abbotii Kirby l 0 0 3 4 V-8: V-9 Robinia Nematus (ribesii gp.) l 0 0 0 le PaVe23 Nematus tibialis Newman 4 0 0 0 4 VII-26-1X-9 Robinia Neopareophora litura (Klug) l 0 2 0 3. =IV-17-IV-24 Vaccinium Pachynematus corniger (Nor- 12 0) 5 1 18 IV-28-XI-4 grasses ton) Pristiphora rufipes Lepeletier l 1 0 0 2 V-20; VII-14 Ribes Pristiphora bivittata (Norton) DZ) jp) 0 0 4 IV-18-VII-16 Spiraea Pristiphora abbreviata (Hartig) 2 0 0 3. =IV-17; VIII-31 Pyrus Pristiphora banksi Marlatt 10 0 0 0 10 V-1-IX-1 Vaccinium Pristiphora acidovalva Wong 1 0 0 0 1 IV-28 Salix and Ross Pristiphora cincta Newman l 0 0 0 1 We2i Betula, Salix, Vaccinium Pristiphora zella Rohwer Tf 0 0 0 7 Nie5=EX-28 Rubus, Geum, Potentilla Pristiphora chlorea (Norton) 1 0 1 0 2 IV-27; V-14 Quercus 150 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Continued. Families, Species Wale YG “WSHihy VED Total Dates Recorded Hosts Pristiphora sp. (bivittata gp.) 0 1 0) 0) 1 IV-17 Pristiphora micronematica 0 0 0 yD 2 Vell Salix Malaise Amauronematus sp. 0 6 0 0) 6 IV-8-IV-21 Heterarthrinae Caliroa lunata MacGillivray 4 0) 0 0 4 V-26; IX-2-16 Caliroa quercuscoccineae 0 1 l 0) 2 V-29; VIII-3 Quercus (Dyar) Caliroa fasciata (Norton) 0 2 0 1 3. ~=VI-18-VIII-23 Quercus Caliroa obsoleta (Norton) 0 0) 1 0 I) VIT=2 Quercus Endelomyia aethiops (F.) 0 4 0 0 4 JIV-20-V-1 Rosa Metallus rohweri Mac- 15 0 0 0 15 -VII-8-IX-16 Rubus Gillivray Nefusa ambigua (Norton) Z 0 0 0 2 V-14 Viola Fenusa pusilla (Lepeletier) 0 0 0 1 1 V-1 Betula Blennocampinae Eutomostethus ephippium 13 Di] 0 0) 40 IV-18-V-29 Poa, grass (Panzer) Eutomostethus luteiventris 1 0 0 0 1 V-22 Juncus (Klug) Eupareophora parca (Cresson) 0 1 l 0 2 IV-1; V-1 Fraxinus Phymatocera fumipennis 2 0 5) 0 7 V-22-VII-12 Smilacina (Norton) Phymatocera racemosae 1 0 0 0 1 VII-15 Smilacina, Poly- Smith gonatum Paracharactus rudis (Norton) 5 0) 0) 1 6 IV-23-V-29 grasses Monophadnoides geniculatus 3 0 0 0 3. IV-27-VI-10 Rubus (Hartig) Periclista marginicollis (Nor- 0 1 l 0 2 IV-19; IV-23 Carya ton) Periclista albicollis (Norton) 1 1 2 0 4 IV-14-IV-30 Quercus Periclista inaequidens (Nor- 0 0 1 0 1 IV-10 Quercus ton) Halidamia affinis (Fallen) 0 i] 1 1 9 IV-17-V-31 Galium Allantinae Pseudosiobla excavata (Nor- 0 0 0 1 I 3 V-22 Cephalanthus ton) Allantus nigritibialis (Rohwer) 19 32 0 0 S51 IV-15-V-29;TX- = Rosa 9-X-9 Monostegia abdominalis (F.) 1 0) 0 0 1 VIII-26 Lysimachia Ametastegia aperta (Norton) 2 ” 6 0) 10 I[V-21-VIII-28 Ametastegia articulata (Klug) 3 1 0 0 4 VII-8-X-11 Rumex, Polygo- num Ametastegia equiseti (Fallén) 0 l 0 0) 1 VI-27 Rumex Ametastegia becra Smith 0 1 0 0 1 VIII-5 Ametastegia pallipes (Spinola) 56 50 0 0 106 IV-13-X-18 Viola Ametastegia pulchella (Roh- 3 4 0 0 7 IV-20-VI-30 wer) Ametastegia tener (Fallén) 0 l 0 0) I EXe2 1 Rumex Empria maculata (Norton) 6 4 0 1 11 IV-23-VI-5 Fragaria, Poten- tilla Empria multicolor (Norton) 5) 0 0 0 5 V-9-VI-12 Alnus, Betula SS VOLUME 89, NUMBER 1 151 Table 1. Continued. Families, Species Yar YGT WSHIT ET Total Dates Recorded Hosts Macremphytus testaceus (Nor- 4 1 1 1 7 VI-17-VII-14 Cornus ton) Taxonus epicera (Say) 6 10 0 0 16 IV-28-V-15 Taxonus pallidicornis (Nor- 7 0 0 0 2 VI-12 Rubus ton) Taxonus pallipes (Say) 14 0 3 0 17 V-17-IX-4 Taxonus terminalis (Say) 1 0 0 l 2 ~~ VII-10; VIII-21 Rubus Tenthredininae Lagium atroviolaceum (Nor- 0 1 0 1 2 ~ VI-5; VI-19 Sambucus, Vi- ton) burnum Aglaostigma semiluteum l 0 0 0 1 V-22 Impatiens (Norton) Tenthredo rufopecta (Norton) 2 2 6 0 10 IV-20-VII-17 Tenthredo sp. 1 0 0 0 1 VI-3 Macrophya alba MacGillivray l 0 0 0 1 9 Vi=17 Macrophya albomaculata 0 0 1 0 1 VIII-8 Sambucus (Norton) Macrophya cinctula (Norton) 0 | 0 0 1 VI-9 Macrophya flavicoxae (Nor- l 1 0 0 2. VI-10; VI-17 ton) Macrophya flavolineata (Nor- 0 ] 1 0 2 V-8; V-29 ton) Macrophya formosa (Klug) 11 0 2 1 14 ~VI-3-VII-24 Macrophya goniphora (Say) 0 1 Bi 0 3 + VI-14-VII-2 Macrophya lineatana Rohwer ] | 0 0 2 VI-9; VI-29 Macrophya macgillivrayi Gib- 2 0 0 0 2 WWe22 Vilma son Macrophya mensa Gibson 2 0 1 6 V-14—-VII-9 Macrophya mixta Mac- 0 l 0 0 1 V-8 Viburnum Gillivray Macrophya pannosa (Say) 1 0) p 0 3. V-7: V-14 Sambucus Macrophya pulchella (Klug) 3 0 0 0 3. ~=VI-10-VII-1 Macrophya senecca Gibson 0 0 0 1 A EVES Macrophya simillima Rohwer 0 l 0 0 Le Vi-22 Rudbeckia Macrophya succincta Cresson 0 j) 0 0 2 IV-21; V-1 Macrophya tibiator Norton 0 0 2 0 2 V-29; VI-12 Macrophya trisyllaba (Norton) 0 0 0 1 VI-4 Sambucus Macrophya varia (Norton) 6 1 1 1 9 VI-22-VII-17 Macrophya zoe Kirby Pelt LOLIK0" OUT RD PVe2I IV 25 Totals 418 402 89 39 =—948 end about 2 m above the ground. The traps were left up continually from the last week in March to the first week in November from 1980-1985 for the yard traps and 1983-1984 for the TNSRC traps. Collec- tions were made about once a week. Spec- imens were identified by DRS, except for most Pristiphora which were identified by H. R. Wong. Voucher specimens are de- posited in the National Museum of Natural History, Washington, D.C. RESULTS AND DISCUSSION Species and abundance.—Species and numbers of specimens collected from each site are listed in Table 1. At all sites, 948 specimens representing 10 families, 53 gen- era, and 117 species were caught. This rep- 40 30 Number of Species 91011 IZ, Se4-5) 6; 7 14 15 16171819 20 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 38 40 48 51 63 106138 Number of Individuals Fig. 1). resents more than half of the approximately 220 species of Symphyta that are expected to occur in the Washington, D.C., metro- politan area (DRS, personal notes). In the two urban environments, 820 specimens of 104 species were collected. For all sites, 35 species were found only in Fairfax Co., 16 in Glen Echo, and 13 at TNSRC. Fairfax Co. and Glen Echo had 20 species in com- mon; Fairfax Co. and TNSRC, 14 species; and Glen Echo and TNSRC, 9 species. Ten species were found in all three study sites. Differences in species composition at each site (Table 1) are likely due to the different hosts present near the traps, length of sam- pling time (two years at TNSRC), and/or the type of trap used. For example in Glen Echo, 402 specimens were caught in two traps, whereas in Fairfax Co., 418 speci- mens were caught in one trap. Frequencies of specimens per species (Fig. 1) indicated that only a few species are pres- ent in large numbers. For 83 species (81%) only four, or fewer, individuals were caught. The collection of only one to several spec- imens of a species may indicate species rar- ity, accidentally wind-blown individuals from other habitats, or representatives of Frequencies of specimens per species in the total sample. groups, e.g. Macrophya, that are strong fliers and may fly far in search of hosts in other areas. Several species, not listed in Table 1, were hand collected in the Fairfax Co. yard but never collected in the traps. These are Ato- macera decepta Rohwer defoliating Hibis- cus sp., Allantus viennensis (Schrank) on Rosa sp., Urocerus cressoni Norton collect- ed at large, and a Neodiprion sp. on Pinus sp. However, most species collected in the traps were never observed, or hand collect- ed, in the yard. This indicates a limitation of the use of a single sampling method. A combination of trapping and observation gives a more complete picture of the fauna. Seasonal distribution. — The earliest saw- flies collected were Xyvela spp. on March 20 and the latest was Pachynematus corniger on November 4. Sawflies were present throughout the intervening period, but the peak in numbers of species and numbers of specimens was from the second half of April through the second half of May (Figs. 2, 3). The flight time for each species is given in Table 1, and examples of different types of flight periods are shown in Fig. 4. Adults appear for a short time once a year for most VOLUME 89, NUMBER 1 species, indicating that most are univoltine, and most of these appear in the spring: 64 species appeared during April to June, sim- ilar to the flight period of Eutomostethus ephippium (Fig. 4); 16 species appeared mostly during June to July; 11 species ap- peared during July to August; and 4 species were late fliers, appearing only during Au- gust and September, similar to the flight pe- riod of Metallus rohweri (Fig. 4). One species, Allantus nigritibialis, had two flight periods, one in spring and one in late sum- mer (Fig. 4). Seventeen species were col- lected through a large part of the season, and peaks of occurrence were apparent dur- ing several periods, similar to the flight pe- riods of Acordulecera pellucida and Schizo- cerella pilicornis in Fig. 4; these apparently have several generations a year. Host-plants.—Sawflies collected in all sites feed on 53 plant genera in 30 families based on recorded hosts (Tables 1, 2). Host- plant records are from Smith (1979) and Gibson (1980). If more than one host is recorded for a species (e.g. Acordulecera dorsalis), all are listed, and the total speci- mens for the species is given for each host in Table 2 since it is not known which host(s) is utilized in the study sites. For sawflies collected in the urban environments, the most common hosts are those used as or- namentals or those present as weeds, and Rosaceae was the most utilized plant family based on numbers of sawfly species and specimens. At TNSRC, however, only two species for which the host is known are as- sociated with members of Rosaceae. Since sawfly species and abundance are depen- dent on available hosts, the trend in urban environments should be toward dominance of those species associated with cultivated plants. Urban species. — Most species collected in the urban environments also occur in nat- ural environments, though some (Table 1) were much more abundant in the yards sampled. Only two species, Al/antus nigri- tibialis and A. viennensis (both associated 153 40 30 Number of Species ie) (eo) De eee ewe Oe Me ha dy eat MA Mine A SS CON 150 a oS 5 aS) Aa Z 100 = 10 — Sm ie) el vo 2 E 50 Z 0 Ze ovalei2ee Lee le el eae wee eel M A J J AYS "ON Figs. 2, 3. Numbers of species and individuals caught from March (M) to November (N), based on half month intervals. | = first half of the month; 2 = second half of the month. with cultivated Rosa spp.) have yet to be found outside urban environments (DRS, personal notes). NOTES ON SPECIES Xyela spp.— The two species were the ear- liest flying sawflies, caught from March 20 to April 7. Larvae feed in staminate pine cones, and adults fly to flowers of other plants to feed on pollen; their flight time usually coincides with the availability of adult food. When numerous, larvae are sometimes a nuisance when dropping from the pine trees to the ground where they enter the soil and form pupal cells. Onycholyda luteicornis.—This was the 154 Table 2. Host plants and number of species and number of individuals collected associated with those plants. Number Number of of Sawfly — Speci- Plant Hosts Species mens Pterydophyta Equisetaceae Equisetum 1 33 Polypodiaceae Onoclea yD, 4 Pteridium l 1 Softening 1 3 Spermatophyta Gymnospermnae Pinaceae Pinus 3 17 Cupressaceae Juniperus l 1 Dicotyledonae Angiospermae Violaceae Viola 2 108 Tiliaceae Tilia l 2 Malvaceae Hibiscus 1 1] Portulacaceae Portulaca l 138 Polygonaceae Rumex 3 6 Polygonum 1 4 Balsaminaceae Impatiens l l Anacardiaceae Rhus 2 5 Aceraceae Acer 1 20 Rosaceae Spiraea 4 Rubus 6 43 Rosa 3 70 Potentilla 2 17 Pyrus 1 3 Crataegus 2 3 Prunus 4 9 Fragaria ] 10 Geum 1 7 Fabaceae Robinia 2 8 Desmanthus 1 4 Fagaceae Castanea l 48 Quercus 9 66 Fagus l D Betulaceae Betula 4 9 Alnus 3 7 Corylus 2 50 Juglandaceae Carya 2 49 Juglans 1 48 Salicaceae Populus 1 1 Salix 5 6 Ulmaceae Ulmus 1 2 Cornaceae Cornus 1 di Ericaceae Rhododendron l 20 Vaccinium 3 14 Primulaceae Lysimachia 1 1 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 2. Continued. Number Number of of Sawfly — Speci- Plant Hosts Species mens Oleaceae Fraxinus 2 4 Convolvulaceae Convolvulus 1 2 Ipomoea 1 2 Rubiaceae Cephalanthus 1 l Galium 1 7 Caprifoliaceae Sambucus 4 1 Viburnum 4 11 Lonicera 1 18 Compositae Rudbeckia i l Monocotyledonae Liliaceae Polygonatum 1 1 Smilacina D, 8 Juncaceae Juncus 1 l Poaceae ““prasses” 5 105 most common pamphiliid and was cap- tured only in the yard traps. Larvae live and feed in rolled leaves and webs of their own making on Rubus spp. Acordulecera dorsalis.—The peak flight was in May and the species was most com- mon in Fairfax Co. and TNSRC. In Fairfax Co., larvae may feed on Quercus sp. or Car- ya sp. nearby. This group has not been stud- ied, and there may be more than one species under this name. Acordulecera pellucida. —The host of this species is not known. It was collected in both yard traps from April to October (Fig. 4) with peak flight periods twice during the season. Schizocerella pilicornis. —This species represented 14% of the specimens collected (138 specimens), most of which were from the yard-garden trap. It is common in urban environments where its larvae feed on purs- lane (Portulaca spp.), a commonly culti- vated, or weed, plant. Flight time is shown in Fig. 4. Atomacera decepta. — This species is a de- foliator of ornamental Hibiscus on which it was found in Fairfax Co. It was not collected in the trap. It may be present from May to VOLUME 89, NUMBER 1 September, passing through several gener- ations. Zaraea lonicerae.—This adventive Eu- ropean species has become common in the D.C. area during the past 20 years. Larvae feed on Lonicera spp., and adults fly during April and May. Neodiprion sp.— No adults were collected in traps, but larvae were found feeding on Pinus spp. in Fairfax Co. Several species are known to be destructive to ornamental pines. Diprion similis.—One specimen of this adventive European species was collected in Fairfax Co., and it is the first record of this species for northern Virginia. This pine- feeding species was known from northeast- ern United States south to Pennsylvania and from an isolated area in extreme south- western Virginia into North Carolina. Xiphydria maculata. —The larvae of this species bore 1n wood in branches of Acer spp. Adults were found only at the Fairfax Co. site where maple is a dominant tree. Heptamelus ochroleucus.—This adven- tive European species feeds on ferns and was known only from British Columbia and New York. More recently it was found in Maryland, and now in Fairfax Co. Adults were collected in spring and fall. Dolerus nitens.—This adventive Euro- pean species feeds on grasses and has spread rapidly in North America. It is sometimes abundant during its very short flight period. Cladius difformis. —Commonly called the bristly rose slug, it is one of the common defoliators of cultivated roses and was col- lected at both yard trap sites and found feed- ing on roses in Fairfax Co. This species has several generations a year. Amauronematus sp.—This species can- not be identified until taxonomic difficulties are resolved. Known hosts for members of the genus are A/nus spp. and Salix spp. Metallus rohweri.—This species was col- lected only in late summer and early fall (Fig. 4). The larva is a leafminer of Rubus spp. 155 Number of Individuals Zexl MerAca Mi dedi id Ay SS Fig. 4. Adult flight periods for selected species. Eu- tomostethus ephippium (Ee), early-spring; Allantus ni- gritibialis (An), spring and fall; Acordulecera pellucida (Ap), entire season; Schizocerella pilicornis (Sp), pri- marily mid-summer; and Metallus rohweri (Mr), late summer and early fall. Fenusa pusilla.—Known as the birch leafminer, this species is a common pest farther north, though it is sometimes abun- dant on ornamental birch in the D.C. area. A single specimen was collected in the field trap at TNSRC. Eutomostethus ephippium.—This is ad- ventive from Europe and has spread rapidly in North America. It can be abundant dur- ing its short flight period in early spring (Fig. 4). Allantus nigritibialis.—This was a rarely collected species before we initiated trap- ping. It was found on and reared from larvae feeding on cultivated roses in Fairfax Co., and adults were collected in both yard trap 156 sites. Collection records indicate a spring and a late summer to fall generation (Fig. 4). Allantus viennensis. — This species was not collected in the traps but was reared from larvae feeding on cultivated roses in Fairfax Co. It isan adventive European species, first recorded in North America from near Ith- aca, New York. The Virginia specimens represent the second North American rec- ord. Monostegia abdominalis. —Specimens collected in the Fairfax Co. trap represent a new record for Virginia and the southern- most record for the species. Ametastegia pallipes.—This species was collected in large numbers and only in both yard trap sites. Its host, Viola, isa common weed species. Adults were collected throughout the season from April to Oc- tober with peak numbers during the end of May and first of April, the end of July and first of August, and the end of September. Macrophya spp.—Of the approximately 40 species in eastern North America (Gib- son, 1880), we collected 20 during this study. Most collections are represented by very few specimens. Hosts are not known for many of the species. They are strong fliers and many may have been accidental catches and may not be associated with plants in the trap vicinities. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ACKNOWLEDGMENTS H. R. Wong, Canadian Forestry Service, Northern Forestry Centre, Edmonton, Al- berta, identified most of the Pristiphora. J. R. Gauthey permitted us to trap sawflies at the Taylor Naval Ship Research Center. The Washington Field Biologists’ Club and Georgetown University provided part of the financial support for this work. We appre- ciate the comments of the following review- ers: H. R. Wong; H. Goulet, Biosystematics Research Institute, Agriculture Canada, Ot- tawa; Robert D. Gordon and E. E. Grissell, Systematic Entomology Laboratory, USDA, Washington, D.C. LITERATURE CITED Barrows, E. M. 1986. A hornet, paper wasps, and yellowjackets (Hymenoptera: Vespidae) in sub- urban habitats of the Washington, D.C., area. Proc. Entomol. Soc. Wash. 88: 237-243. Frankie, G. W. and C. S. Koehler, eds. 1983. Urban Entomology: An Interdisciplinary Approach. Praeger Press, New York. 493 pp. Gibson, G. A. P. 1980. A revision of the genus Mac- rophya Dahlbom (Hymenoptera: Symphyta, Ten- thredinidae) of North America. Mem. Entomol. Soc. Can. No. 114, 167 pp. Smith, D. R. 1979. Symphyta, pp. 3-137. Jn Krom- bein, K. V. et al., eds., Catalog of Hymenoptera of America north of Mexico. Vol. 1. Smithsonian Institution Press, Washington, D.C. PROC. ENTOMOL. SOC. WASH. 89(1), 1987, pp. 157-166 CROSSING AND CHROMOSOMAL EVIDENCE FOR TWO ADDITIONAL SIBLING SPECIES WITHIN THE TAXON ANOPHELES DIRUS PEYTON AND HARRISON (DIPTERA: CULICIDAE) IN THAILAND V. BAIMAI, R. G. ANDRE, B. A. HARRISON U. KIJCHALAO, AND L. PANTHUSIRI (VB, UK) Department of Biology, Faculty of Science, Mahidol University, Rama VI Road, Bangkok 10400, Thailand; (RGA, BAH) Department of Entomology, Walter Reed Army Institute of Research, Washington, D.C. 20307-5100, USA; (LP) U.S. Army Medical Component, AFRIMS, Rajvithi Road, Bangkok 10400, Thailand. Abstract. —Crossing and chromosomal evidence 1s presented for two additional sibling species, dirus C and dirus D, within the taxon Anopheles dirus Peyton and Harrison, in Thailand. The affinities of the four currently recognized species in this complex in Thailand and the limitations of certain techniques used to identify the species are discussed. Cytogenetics is one of the most useful tools for elucidating cryptic species of insects (Dobzhansky, 1970; White, 1973). The use of this method, together with biochemical, behavioral, ecological and morphological techniques has led to the recognition of a significant number of sibling species com- plexes of anopheline mosquitoes in different parts of the world (Bryan and Coluzzi, 1971; Kitzmiller et.al. 1973: White et al.; 1975; Kitzmiller, 1976; Coluzzi et al., 1979; Steg- nul and Kabanova, 1978; Peyton and Har- rison, 1979, 1980; Green and Miles, 1980; Subbarao et al., 1983; Green and Baimai, 1984; Green et al., 1985). The discovery of these cryptic species is a highly significant step in the development of rational and ef- ficient control programs against the vectors of various mosquito-borne diseases. One of the most renowned vectors of hu- man malaria parasites in Southeast Asia is Anopheles balabacensis Baisas, a member of the widely distributed Leucosphyrus Group. Recently, it was demonstrated that An. balabacensis is a species complex (Pey- ton and Harrison, 1979, 1980; Baimai et al., 1981; Hii, 1982, 1984, 1985). Anopheles dirus Peyton and Harrison, was described as a species distinct from An. balabacensis in 1979, and is considered widespread in peninsular Malaysia and Thailand, while An. balabacensis sensu stricto is confined to the type-locality on Balabac Island and to neighboring areas of Palawan Island, Sabah and northeast Kalimantan (Peyton, unpub- lished data; Peyton and Harrison, 1979; Hii, 1982). Baimai et al. (1981) recently demonstrat- ed that the laboratory colony strain of An. balabacensis Perlis form from The Institute of Medical Research (Kuala Lumpur) showed different sex chromosome charac- ters as seen in mitotic karyotype as well as on salivary gland polytene chromosomes. Genetic incompatibility between dirus and the balabacensis Perlis form also was ob- served (Baimai and Harrison, 1980). The recognition of the balabacensis Perlis form as a distinct genetic species from dirus was confirmed later by the detailed studies of 158 Table 1. indicated) used in this study. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Laboratory family stocks of Anopheles dirus complex from different localities in Thailand (otherwise eeeaals dirus Species Locality (Date) Examined No. of Families Maintained (Code) A Chonburi (1964) and Nakhon Rat- _ Bangkok colony (BK)* chasima (1971) Petchaburi (1983) 17 2 CL) B Perlis State, Malaysia (1965) _ IMR colony (PR)* Trengganu State, Malaysia (1982) 5 1 (MH) Phangnga (1982) 2 — (S Kanchanaburi (1980, 1982) 9 1 (KN) D Ranong (1983, 1984) 59 2 (RN) Phangnga (1982) 5 1 (PG) * The mixed colony. ** The isofemale line now used as the reference stock for dirus A. Hii (1982) who designated the Bangkok col- ony strain as dirus A and the Perlis form as dirus B. In 1979-80, E. L. Peyton, at the Smith- sonian Institution (personal communica- tion) advised the authors that two addition- al members of the Dirus Complex existed in Thailand which could be separated from dirus A and dirus B by morphological char- acters. Subsequently, these members were collected and colonies of both were estab- lished in 1980-81 at the Armed Forces Re- search Institute of Medical Sciences (AF- RIMS), Bangkok. Recently, Wibowo et al. (1984) discov- ered marked differences in the amount and distribution of constitutive heterochroma- tin in the sex chromosomes, as revealed by the Hoechst 33258 staining technique, of specimens from a Kanchanaburi colony de- rived from a single isofemale of one of the two new members. Based on comparative cytological data, these investigators con- firmed Peyton’s morphological findings that the Kanchanaburi isoline represents a dis- tinct genetic species designated as dirus C, which occurred sympatrically with dirus A. Additional cytogenetic studies also have re- vealed that the second undescribed member noted by Peyton is distinct and represents a 4th member, dirus D, of the complex in Thailand. This paper presents crossing and chro- mosomal evidence supporting the existence of two additional sibling species, dirus C and dirus D, within the Dirus Complex in Thailand. MATERIALS AND METHODS Individual wild caught Anopheles females were identified to species by morphology, isolated, and allowed to oviposit. Subse- quent F, larvae were reared in the labora- tory and used for cytogenetic confirmation of the species. Mitotic and salivary gland polytene chromosomes were prepared from 4th instar larvae using the method of Bai- mai et al. (1981). Isofemale lines of each cytotype were set up with respect to the X and Y chromosome configurations and maintained in the laboratory for further crossing experiments (Table 1). The mosquitoes used in this study came from isofemale lines that were determined cytogenetically and maintained at the De- partment of Medical Entomology, AF- RIMS, Bangkok, and at the Department of Biology, Mahidol University, Bangkok. Anopheles dirus A (Bangkok colony strain = VOLUME 89, NUMBER 1 159 Table 2. Crossing combinations among the isolines of “Anopheles dirus” from different geographic origins. “dirus” Crosses Peuaes No. of Mean No. of F, Adults Ovipositing Ovipositions Eggs per Group Q 3 (Total) Hatching Oviposition % Eggs Hatching Q 3 1 B(MH) B(PR) 5 (10) 5 142.2 54.0 168 is (384/711) B(PR) B(MH) 8 (10) 8 85.9 aD 132 Hit. (395/687) 2 (@ A (I) 6 (10) 4 59.5 47.3 69 13 (169/357) A C (ID) 5 (10) 3 96.0 50.0 12 $3 (240/480) 3 B(MH) A 9 (20) ql 103.2 2N9 8 1 (203/929) A B(MH) 9 (20) 8 108.3 4.1 2 ay (40/975) 4 B(PR) Cc 8 (20) 3 54.1 10.2 0 0 (44/433) (e B(PR) 7 (20) 0 88.3 0 = = (0/618) 5 D(PG) B(MH) 5 (10) 1 58.0 0.3 0 0 (1/290) B(MH) D(PG) 0 (10) 0 0 0 _ (0/0) 6 D(RN) @ 2 (10) 1 209.0 127 0 0 (53/418) (e D(RN) 2 (10) 0 85.5 0 0 0 (0/171) 7 A D(RN) 7 (10) 0 75.0 0 0 0 (0/525) D(RN) A 2 (10) 2 30.0 15.0 8 0 (9/60) * Sterile male F, hybrids. BK) was used as the standard stock, whereas the IMR colony strain (Perlis form = PR) and the Trengganu colony strain (= MH) represented An. dirus B in this study. The isoline strain from Kanchanaburi (= KN) represents dirus C. Two other isoline strains of “dirus” were collected from Ranong (= RN) and Phangnga (= PG) in southern Thailand, and represent dirus D (Table 1). Combinations of reciprocal pair-matings (Table 2) among the different cytotype strains were performed by the artificial mat- ing technique of Ow Yang et al. (1963). In each cross pair-mating 10-20 individual fe- males were mated with mature males. After successful copulation, each female was iso- lated in an oviposition vial. The number of females ovipositing, number of eggs ovi- posited and hatching and number of emerged F, adults were scored daily in the same manner as described by Klein et al. (1985). Fertility of F, hybrids was deter- mined later by self-crossing among them- selves as well as backcrossing to the respec- tive parental strains (Table 3). Genetic incompatibility could be inferred on the de- gree of synapsis in salivary gland polytene chromosomes of F, larval hybrids of anoph- 160 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 3. Backcrossing and selfcrossing experiments of F, hybrids from the crosses between isolines of Anoph- eles dirus A and An. dirus C. Crosses No. of Females Q 3 Ovipositing (Total) F, (D* A 3 (5) F, (1) € 4 (5) A F, (I) 2 (5) c F, (I) 25) F, (1) F, (1) 5 (5) F, (D** A 8 (12) F, (11) G 8 (12) A F, (II) 5 (12) G F, (II) 7 (10) F, (II) F, (II) 8 (12) * JT = cross between female dirus C x male dirus A. ** TT = cross between female dirus A x male dirus C. elines, and the degree of viability was de- termined by egg hatch rates. RESULTS Hybridization tests.—The results of cross- mating experiments among the different sibling members of the complex are sum- marized in Table 2. Crosses between the 2 dirus B isoline strains (MH and PR), which showed the same mitotic karyotype, yielded a large number of eggs per female (85.9 and 142.2) and a high percentage of eggs hatch- ing (54.0% and 57.5%). Adult F, hybrids of both sexes from both directions were fully fertile and continued to produce progeny for several generations. There was no evidence of genetic incompatibility between these two strains. The crossing evidence clearly indi- cates that the MH and PR strains belong to the same genetic species, An. dirus B. Crosses between the dirus A and dirus C isoline strains also were successful in both No. of Ovipositions Mean No. of Eggs Hatching per Oviposition % Eggs Hatching 3 102.0 88.6 (271/306) 4 es 69.3 (336/485) 2 78.0 57.1 (89/156) D 134.5 46.8 (126/269) 2) 189.0 76.4 (722/945) 7 130.5 49.9 (521/1044) 7 149.5 60.1 (719/1196) 0 12 0 (0/611) 0 12257 0 (0/859) 0 103.8 0 (0/830) directions, producing a large number of eggs (averaging 59.5 and 96.0 per female, re- spectively) and a high percentage of eggs hatching (47.3% and 50.0%, respectively). Many F, adults of both sexes were obtained in each direction. Crosses between female dirus C x male dirus A (designated as cross I) yielded fully fertile male and female F, hybrids. In contrast, the reciprocal cross be- tween female dirus A and male dirus C (des- ignated as cross II) produced fertile F, fe- males, but completely sterile F, males (Table 2). These results indicate that genetic in- compatibility exists between these two 1so- lines, at least in one direction of hybridiza- tion. These crossing data are supported by cytological evidence described below from the F, hybrids. All combinations of crosses involving the dirus B isoline strains (PR and MH) with the other dirus isolines were less successful than those matings mentioned earlier. VOLUME 89, NUMBER 1 Crosses between dirus B (MH) and dirus A in both directions produced a large number of eggs (average of 103.2 and 108.3 per fe- male). However, very low percentages of the eggs hatched in these crosses. Furthermore, very few adult F, offspring emerged, and of these, the F, males were sterile. Crosses be- tween female dirus B (PR) x male dirus C were less successful because only 10.2% of the eggs hatched, and no adults emerged. The reciprocal cross between female dirus C x male dirus B (PR) gave more eggs (av- erage of 88.3 per female), but none hatched. Furthermore, a remarkable example of ge- netic incompatibility was obtained from the crosses between dirus D (PG) and dirus B (MH). A very small percentage of the eggs hatched (0.3%), and only in one direction. In the reciprocal mating of this cross, none of the 10 artificially inseminated females produced eggs. The dirus D isoline strains (RN and PG) exhibit similar karyotype and polytene band sequences. Unfortunately, the PG strain was lost before the RN strain was obtained, thus cross mating tests between them were not made. However, cytological evidence in- dicates that they are conspecific strains. All combinations of cross matings among the dirus D isoline strains (RN or PG) with oth- er isolines yielded either very small num- bers of F, female hybrids or no F, hybrids at all (see groups 5, 6 and 7 in Table 1). These results clearly indicate that the RN and PG isoline strains of dirus D were ge- netically distinct from the other dirus iso- lines employed in this study. Cytological evidence.— The examination of F, hybrid larval salivary gland polytene chromosomes revealed some differences in banding sequences. Based on the standard salivary gland polytene chromosomes of di- rus A (Baimai et al., 1980), the F, female larvae from the cross between female dirus A x male dirus C exhibited approximately 5-10% asynapsis of the chromosome com- plement (Fig. 1). In addition, marked dif- ferences in polytene banding sequences were 161 observed at zone 6 (Fig. 1, arrow) and at the tip of the X chromosome (Fig. 2, arrow), as well as at the tips of chromosome arm 2L and arm 2R (Figs. 3, 4, respectively). These differences are good chromosome markers for the dirus C karyotype. Larval salivary gland polytene chromo- somes of F, hybrid females from the cross between female dirus A x male dirus B (PR) showed approximately 80% asynapsis of the chromosome elements (Fig. 5). This sug- gests that genetic differentiation at a sub- microscopic level between these two species is more extensive than in the case of dirus A and dirus C. The chromosome complement of the di- rus D (RN and PG) strains is remarkably different from the standard dirus A colony strain. The F, female larval chromosomes from the cross between female dirus A x male dirus D (RN) showed over 90% asyn- apsis along the 5 chromosome arms (Fig. 6). The X chromosome of the dirus D (RN) strain exhibited a fixed inversion covering zones | and 3 of the X chromosome com- pared with the standard sequence of the di- rus A strain (Fig. 7). Zone 6 of the X chro- mosome of the F, female hybrids was asynapsed completely. Moreover, asynapsis in chromosome arm 2R of F, hybrids in this case was more pronounced (Fig. 8) than in the case of dirus C x dirus A F, hybrids. Overall, asynapsis was a persistent feature of the hybrid polytene chromosomes of the Dirus Complex. In addition, an analysis of the mitotic karyotype showed that the X and Y chromosomes of the dirus D (RN) strain are shorter than those of dirus A as can be observed in F, hybrid larval chro- mosomes (Figs. 9, 10). Thus, the cytological observations clearly support the sterility and viability evidence from the hybridization experiments described above. DISCUSSION Recent morphological, genetic and cyto- genetic studies of the taxon dirus have re- vealed that it consists of at least 3 genetic 162 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 4 . ee oar " 4 oe + 4, . Ep . Se tn < ‘ 4 oy : 2 ae ai 7a bad s > > oat < ~ Po 7 od ‘ _— -” 7 s * 5 ' >. Figs. 1-5. Figs. 1-4. Larval salivary gland polytene chromosomes of F, hybrid females from the cross matings between female dirus A x male dirus C. 1, Condition of synapsis along the 5 chromosome elements, zone 6 of the X chromosome (small arrow) is almost totally asynapsed. 2, Complete synapsis of zones 1-5 of the X chromosome with a distinct banding difference at the tip (arrow). 3, 4, Tips of chromosome arm 2L and arm VOLUME 89, NUMBER 1 species namely dirus A, B and C (Peyton, unpublished data; Baimai et al., 1981; Hii, 1982; Wibowo et al., 1984). The present investigation confirms those findings. Fur- ther, the present cytogenetic evidence has confirmed the fourth species recognized morphologically by Peyton (unpublished) within this taxon. Provisionally this species is designated dirus D, and it is represented by the RN and PG isoline strains from southern Thailand populations. Our results, however, seem to be in disagreement with the interpretation of Kanda et al. (1981). Based on their hybridization data, Kanda and co-workers are of the opinion that their colony strains from Chantaburi, Kanchan- abur1 and IMR only represent geographical populations of An. balabacensis. The results of our cross mating experi- ments clearly indicate that An. dirus 1s ac- tually a cluster of closely related species. The divergence of these siblings could have occurred comparatively recently. The pres- ent data suggest that dirus A and dirus C are very closely related and they occur sym- patrically, at least in Kanchanaburi prov- ince. Artificial mating between them is pos- sible under laboratory conditions. Whether gene flow between these two genetic species occurs in nature is not known. The species isolating mechanism for these two siblings probably involves premating isolation, as well as the unidirectional genetic incom- patibility detected in this study. Both dirus A and dirus C exhibit similar banding sequences of salivary gland poly- tene chromosomes and general mitotic karyotypes, although the former shows het- erochromatin variation in the sex chro- mosomes (Baimai et al., 1984). However, striking differences in the polytene chro- mosomes of F, hybrids were observed at the tips of chromosome X and chromosome 163 arms 2L and 2R. Furthermore, differences in the amount of heterochromatin are no- ticeable in the sex chromosomes (Wibowo et al., 1984). In general, the mechanism in the process of species differentiation for sib- lings in the Dirus Complex resembles that for some species groups of the picture- winged Hawaiian Drosophila (Ohta, 1980; Carson and Yoon, 1982). Anopheles dirus B (= Perlis form) is ge- netically distinct from dirus A and dirus C. Hybridizations between dirus B and dirus C were cross-sterile, producing no adult F, hybrids in either direction. Cross mating be- tween dirus B and dirus A, however, yielded very few adult F, hybrids, of which the males were completely sterile. Anopheles dirus B showed cytological differences from dirus A both in salivary gland polytene chromo- some (Fig. 5) and mitotic karyotype (Baimai et al., 1981). So far, distribution records in- dicate that dirus B is confined to southern Thailand. Thus, dirus B may be isolated geographically from dirus A. On the other hand, dirus A seems to be widespread in central and northern Thailand. We now are investigating the boundary limits of dirus A and dirus B in southern peninsular Thai- land. The distribution of dirus C is limited to collection areas in Kanchanaburi Prov- ince, in western Thailand where it coexists with dirus A and dirus D, and an isolated questionable area in southern Thailand where it may be sympatric with dirus B. The highest degree of genetic incompat- ibility was found in all combinations of hy- bridization tests involving dirus D. Most cross matings involving the RN or PG strains of dirus D completely failed to pro- duce F, hybrid adults. However, cross mat- ings between female dirus D (RN) x male dirus A produced a few F, hybrid females which were very weak. Asynapsis in F, lar- — 2R, respectively (chromosomes of dirus C are indicated by arrows). Fig. 5. Larval salivary gland chromosome elements of a F, hybrid female from a cross between female dirus A x male dirus B (PR) showing extensive regions of asynapsis. Figs. 6-10. Salivary gland polytene chromosomes and mitotic karyotypes of F, hybrid larvae from crosses between female dirus A x male dirus D (RN). 6, Asynaptic condition of the whole polytene chromosome complement. 7, A fixed inversion difference on the distal half and the complete asynaptic region of zone 6 of the X chromosome (arrows). 8, Asynapsis in chromosome arm 2R. 9, Comparison of heterochromatin differences of the X chromosomes in a F, female larval neuroblast cell. 10, A short Y chromosome of dirus D compared with the X chromosome of dirus A in a F, hybrid male. VOLUME 89, NUMBER 1 val polytene chromosomes was extensive, covering more than 90% of the chromo- some elements. Our data indicate that dirus D is genetically remote from the other sib- lings in the Dirus Complex. Anopheles dirus D apparently is distributed widely in central and southern Thailand, and northern Ma- laysia, and has been found in sympatry with dirus A, dirus B and dirus C. The use of heterochromatic variation in sex chromosomes as a means for routine identification of our material from the field has distinct limitations. On the other hand, analysis of salivary gland polytene chro- mosomes now provides a better means of species identification of this sibling species complex than mitotic karyotypes, and this method is used routinely in our laboratory. First, X chromosome heterochromatin variation cannot be used routinely because it is difficult to score and is seen only in rare, superb preparations. The Y chromosome variation, on the other hand, is scored much more easily and is available for routine identification of families from wild-caught material. There is a quantitative difference in data from these two sources of variation. The X chromosome data can provide direct evidence for gene flow characteristics in na- ture and thus, evidence for mixtures of cryp- tic species in samples. The Y chromosome data cannot provide such evidence because of the combination of the obvious hemi- zygous condition of the Y in males and the knowledge that most female anophelines are mated successfully only once. Consequent- ly, different Y chromosomes are not ex- pected to occur together in single broods, and we cannot tell from their distribution in broods whether these Y chromosomes represent intra- or interspecific variation. The best we can do is to correlate Y chro- mosome variation with primary evidence for the different species within the Dirus Complex. The most widely used criterion is interspecific sterility as seen in laboratory crossing experiments. Our procedure has been to score Y chro- 165 mosome variation in samples from nature and where a sample or sub-sample of fam- ilies shows the same Y chromosome, cross one of these families to laboratory reference stocks. At first, we were forced to use the non-isoline colonies of the Bangkok strain (species A) and the Perlis form (species B). As identified isofemale lines became avail- able, we replaced the non-isoline colonies as reference stocks and also established iso- female lines as reference stocks for species C and D. The recognition of the existence of cryptic species within the taxon An. dirus has led to a better understanding of the process of species differentiation of the Leucosphyrus Group of Anopheles. Further information on species distributions, behavior and pop- ulation dynamics of these siblings undoubt- edly will lead to a better understanding of malaria transmission and strategies for ef- fective vector control in this region. Differ- ences in biological properties and behavior with respect to the vectorial capacity and the epidemiological significance of the four member species of the Dirus Complex are under investigation. Presently, the identi- fication of these genetic species from natural samples is a problem. A practical taxonom- ic key is now being developed at the Walter Reed Biosystematics Unit (Peyton, personal communication). Another technique which may be valuable in identifying these species is recombinant DNA for species specific DNA probes. ACKNOWLEDGMENTS This investigation was supported partial- ly by the UNDP/World Bank/WHO Special Program for Research and Training in Tropical Diseases and Mahidol University Fund. The reviews of the finished manu- script by R. A. Ward and E. L. Peyton are greatly appreciated. We thank C. A. Green for comments on the manuscript and the entomological field staff and the Medical Audiovisual Section of the AFRIMS for technical assistance. 166 LITERATURE CITED Baimai, V. and B. A. Harrison. 1980. Evidence of sibling speciation in the balabacensis complex of Southeast Asia (Diptera: Culicidae). Abst. 10th Intern. Cong. Trop. Med. Malaria, 9-15 Nov., Manila, pp. 83-84. Baimai, V., R. G. Andre, and B. A. Harrison. 1984. Heterochromatin variation in the sex chromo- somes in Thailand populations of Anopheles dirus A (Diptera: Culicidae). Can. J. Genet. Cytol. 26: 633-636. Baimai, V., B. A. Harrison, and V. Nakavachara. 1980. The salivary gland chromosomes of Anopheles (Cellia) dirus (Diptera: Culicidae) of the Southeast Asian Leucosphyrus Group. Proc. Entomol. Soc. Wash. 82: 319-328. Baimai, V., B. A. Harrison, and L. Somchit. 1981. Karyotype differentiation of 3 anopheline taxa in the Balabacensis complex of Southeast Asia (Dip- tera: Culicidae). Genetica 57: 81-86. Bryan, J. H. and M. Coluzzi. 1971. Cytogenetic ob- servations on Anopheles farauti Laveran. Bull. W.H.O. 45: 266-267. Carson, H. L. and J. S. Yoon. 1982. Genetics and evolution of Hawaiian Drosophila, pp. 297-344. In Ashburner, M., H. L. Carson and J. N. Thomp- son, eds., The genetics and biology of Drosophila, Vol. 3b. Academic Press. New York, NY. Coluzzi, M., A. Sabatini, V. Petrarca, and M. A. Di Deco. 1979. Chromosomal differentiation and adaptation to human environments in the Anoph- eles gambiae complex. Trans R. Soc. Trop. Med. Hyg. 73: 483-497. Dobzhansky, T. 1970. Genetics of the Evolutionary Process. Columbia Univ. Press. New York, 505 pp. Green, C. A. and V. Baimai. 1984. Polytene chro- mosomes and their use in species studies of ma- laria vectors as exemplified by the Anopheles mac- ulatus complex. In B. C. Joshi, R. P. Sharma, H. C. Bansal, and V. L. Chopra, eds., Genetics: new frontiers. Proc. X Vth Int. Cong. Genet. Vol. 3: 89-97. Oxford and IBH Publ. Co., New Delhi. Green, C. A. and S. J. Miles. 1980. Chromosomal evidence for sibling species of the malaria vector Anopheles (Cellia) culicifacies Giles. J. Trop. Med. Hyg. 83: 75-78. Green, C. A., V. Baimai, B. A. Harrison, and R. G. Andre. 1985. Cytogenetic evidence for a com- plex of species within the taxon Anopheles mac- ulatus (Diptera: Culicidae). Biol. J. 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WASH. 89(1), 1987, pp. 167-174 DISTRIBUTIONAL DATA, ILLUSTRATIONS, AND HABITAT OF THE SOUTH AMERICAN WATER-STRIDER MICROVELIA AYACUCHANA (HEMIPTERA: VELIIDAE) PAUL J. SPANGLER AND RICHARD C. FROESCHNER Department of Entomology, Smithsonian Institution, Washington, D.C. 20560. Abstract. —The microveliine water-bug Microvelia ayacuchana Drake and Maldonado Capriles [sic] (1952) is illustrated for the first time. A habitus view, the antennae, and the male and female genitalia are illustrated by pen and ink drawings and the surface sculpture of selected parts of the body are illustrated by scanning electron micrographs. Known distribution records for this species from Brazil, Guyana, Surinam, and Venezuela are reported. Habitats are discussed and a photograph of one of the biotopes is included. During a fieldtrip to collect aquatic Co- leoptera, Hemiptera, and other insects in the Takutu Mountains of Guyana in No- vember and December, 1983, numerous aquatic Hemiptera were collected by mem- bers of an Earthwatch Expedition. Among the water bugs were 10 distinctive winged specimens of a large species of Microvelia. An examination of those specimens and comparison with the type specimens of Mi- crovelia in the Drake collection and the gen- eral collection of aquatic Hemiptera in the U.S. National Museum of Natural History, Smithsonian Institution confirmed that the specimens from Guyana are conspecific with Microvelia ayacuchana Drake and Maldo- nado Capriles [sic] (1952). More recently, 19 and 24 February 1986, Spangler collect- ed 22 more specimens of this species near the type locality, Puerto Ayacucho, T.F.A., Venezuela. Because M. ayacuchana is rare in collec- tions; belongs to a large genus with 81 de- scribed species in the Western Hemisphere that are similar in external morphology, sculpture, and color; has not been previ- ously illustrated; and has not been included in any keys; we have prepared this article with the hope that it will allow the reader to identify this species more easily. The type specimens of M. ayacuchana were collected by J. Maldonado at Puerto Ayacucho, Venezuela, in May and June 1950; and Drake and Hussey (1955), in their checklist of the species of Microvelia, re- ported M. ayacuchana from British Guiana [now Guyana]. There are no specimens of the species from “British Guiana” in the NMNH but the record may have been based on borrowed specimens; unfortunately, the source of the specimens was not given. There has been nothing reported about M. aya- cuchana since Drake and Roze (1958) listed it as one of the seven species of Microvelia reported from Venezuela. The description of Microvelia ayacu- chana by Drake and Maldonado Capriles [sic] (1952) is adequate for the external mor- phological characters as seen under a ste- reoscopic microscope. Additional external characters, some illustrated by scanning electron micrographs, and genitalic char- acters are discussed below. A habitus view (Fig. 1) and the dissected and cleared male 168 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. genitalia (Figs. 13, 14) and female genitalia (Figs. 15, 16) of MM. ayacuchana are illus- trated for the first time. Microvelia ayacuchana Drake and Maldonado Capriles [sic] Additional descriptive characters. — Mac- ropterous adult males longer (2.75 to 3.16 mm) than most members of genus. Head Microvelia ayacuchana, male, habitus view. (Fig. 2) with longitudinal groove on midline distinct, 3 pairs of trichobothria, ocelli ab- sent. Antennae (Fig. 3) long; 4 segmented; internodial piece between segments 2 and 3 and another between segments 3 and 4; an- tennal segment ratios 17:13:17:23. Rostrum 3 segmented, extending to mesocoxae; man- dibular stylets toothed (Fig. 4). Mesoscutellum covered by posterior ex- 851633 15KY¥ ¥600°° "5 Samm 851632 10Ky 5 Figs. 2-6. Microvelia ayacuchana. 2, Head. 3, Antenna. 4, Mandibular stylets. evaporatorium tuft. 6, Methathoracic scent gland channels. , Head, pronotum, and 170 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 18KY¥ VOLUME 89, NUMBER 1 171 Figs. 13, 14. Microvelia ayacuchana. 13, Male genitalia, extended in alcohol, dorsal view. 14, Male genitalia, cleared in KOH, lateral view. — Figs. 7-12. Figs. 7-10. Microvelia ayacuchana. 7, Campaniform sensillum, base of protarsal segment. 8, Campaniform sensillum enlarged. 9, Protibial grooming comb (arrow) and grasping combs, male. 10, Protibial grasping comb, male. Figs. 11, 12. Microvelia ayacuchana. 11, Protibial grasping comb, male, enlarged. 12, Protarsal claws. 172 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Y fom ————— (0). 440ni¢a) Figs. 15, 16. Microvelia ayacuchana. 15, Female genitalia, extended in alcohol, dorsal view. 16, Female genitalia, lateral view. VOLUME 89, NUMBER 1 173 “er 1". Fig. 17; Venezuela. tension of pronotal lobe (Fig. 5). Metatho- racic scent gland channels (Fig. 6) terminate in an evaporatorium bearing a prominent tuft of long setae (Fig. 5, arrow) on pleuron in front of metacoxae. Protarsus with | campaniform sensillum dorsally at base of ultimate segment (Figs. 7, 8). Front tibia of male with a short, trans- verse, apical, grooming comb of stout setae on dorsal margin (Fig. 9, arrow) and a long grasping comb (Figs. 9-11) of short stout spines on inner (ventral) surface of distal third. Claws (Fig. 12) distinctly inserted be- fore apex of last tarsal segment. Male genitalia (Figs. 13, 14) with aedea- gus with membranous vesica; parameres minute. Female genitalia (Figs. 15, 16) with proctiger broad, decurved, covering gono- coxal and genital opening. Specimens examined.—Specimens from the John T. Polhemus collection are indi- cated by (JTP) and those from the National Museum of Natural History, Smithsonian Institution are indicated by (NMNH). Rep- resentative specimens will be deposited in the Instituto de Zoologia Agricola, Univer- = tent 23 Microvelia ayacuchana, pothole biotope at Tobogan area, 40 km south of Puerto Ayacucho, T.F.A., sidad Central de Venezuela, Maracay, Ven- ezuela. BRAZIL: AMAZONAS: Lago Salgado, Jg. duebra d., 29.4.48, H. Sioli, 2 males (JTP). GUYANA: POTARO-MAZAR- UNI DISTRICT: Takutu Mountains, 6°15'N 59°5’W, 3-10 Dec. 1983, P. J. Span- gler, R. A. Faitoute, P. D. Perkins, 3 males, 7 females (1 on SEM stub) (NMNH). SURI- NAM: Sipaliwini River, 13-VI-63, 6 males, 6 females (NMNH). VENEZUELA: TER- RITORIO FEDERAL AMAZONAS: Puer- to Ayacucho, 15 June 1950, J. Maldonado Capriles, holotype male, allotype (NMNH); same data, | male, 2 females (paratypes) (JTP); Puerto Ayacucho (40 Km S) at To- bogan, 19 Feb. 1986, P. J. Spangler, 1 male, 1 female (NMNH); same data except 24 Feb. 1986, 16 males, 4 females (NMNH). One additional paratype with the same data as the holotype except 15 May 1950 has not been found. Habitat.—The specimens from Guyana were collected from the shaded margin of a slowly flowing brook in the rainforest. The series from the Tobogan area south of Puer- 174 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON to Ayacucho, Venezuela, were found on the water of a small pothole (Fig. 17) in bedrock beside a small stream; the pothole was ex- posed to full sunlight for a relatively short time during the day. ACKNOWLEDGMENTS We thank the following for their assis- tance: The Center for Field Research, for sponsoring the fieldwork in Guyana; the ad- ministrators of the Smithsonian Institu- tion’s Research Opportunities Fund for supporting the fieldwork in Venezuela; Young T. Sohn, biological illustrator, for the pen and ink drawings; John T. Polhe- mus, for locality data from his specimens; and Phyllis Spangler for typing the manu- script into the word processor. LITERATURE CITED Drake, C. J. and J. Maldonado Capriles. 1952. Water- striders from Territorio Amazonas of Venezuela (Hemiptera: Hydrometridae, Veliidae). The Great Basin Naturalist 12: 47-54. Drake, C. J. and R. F. Hussey. 1955. Concerning the Genus Microvelia Westwood, with Descriptions of Two New Species and a Check-list of the Amer- ican Forms (Hemiptera: Veliidae). The Florida Entomologist 38: 95-115. Drake, C. J. and J. A. Roze. 1958. Anew Microvelia from Venezuela (Hemiptera: Veliidae). Bulletin of the Southern California Academy of Sciences 57: 47-48. PROC. ENTOMOL. SOC. WASH. 89(1), 1987, pp. 175-176 THE CORRECT IDENTITY OF ACLERIS INANA (ROBINSON) (LEPIDOPTERA: TORTRICIDAE) J. F. GATES CLARKE Department of Entomology, Smithsonian Institution, Washington, D.C. 20560. Abstract.—The true identity of Acleris inana (Robinson) is revealed and the genitalia are figured. While curating the species of Ac/eris in the collection of the United States National Museum of Natural History it soon became abundantly clear that the identity of several species was confused, and that many series were mixed. Contributing to this confusion is the fact that the phenotypes are frequently repeated in several taxa and the study of the genitalia is the only means of correctly iden- tifying the species. Among the similarly marked species is Acleris inana (Robinson) (1869: 281) de- scribed as Teras inana. The number of spec- imens in his series was not indicated al- though he had at least two because he stated ““Habitat.—Mass., N.Y.” This species be- longs to a group of closely similar species consisting of 4A. inana (Robinson), A. fla- vivittana (Clemens), A. robinsoniana (Forbes), and perhaps others. Of inana, McDunnough (1934) stated that the “... determination of this species is based on a specimen compared by the writer in 1925 with a series in the United States National Museum under this name, and by a recent examination of a specimen in the American Museum Collection labelled ““Homotype” by Kearfott.”” Unfortunately, McDunnough did not see or examine the Lectotype, which is in the Academy of Nat- ural Sciences, Philadelphia, but based his identification on specimens that appear to have been incorrectly identified. Mc- Dunnough’s figure of the female genitalia (1934, p. 331, fig. 1) which he supposed to be inana, does not agree with the type, and I have not yet been able to associate defi- nitely any species with his figure. Subse- quently, Razowski (1966) figured as inana (fig. 526) what appears to be exactly the same as that figured by McDunnough (see above). In neither case have I been able to ascertain what species they illustrated. So far, I have been able to associate def- initely with inana, two females from Mis- souri, Jasper County, Sarcoxie, 14 June 1975, in the J. R. Heitzman collection and four specimens in the United States Na- tional Museum as follows: One male and two females are labeled “NY. The male bears a further label inscribed “‘agrees per- fectly with type’”’ and one female bears a label ““Mass. Agrees with type.’ The lecto- type, designated by Klots (1942) and now in the Academy of Natural Sciences, Phila- delphia, bears the following label data: Small, square white label “36.” Small white label ““NY.”’ Red label: Teras inana C. T. Robinson, 7439.” Green label, genitalia slide by JFGC, female, USNM 25553. The gen- italia of the lectotype are illustrated in Fig. 1. I am uncertain whether Acleris flavivit- tana (Clemens) is a distinct species. The type of flavivittana, apparently, is lost. The fe- male genitalia of a series of specimens in the United States National Museum from 176 da PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Acleris inana. a, Ventral view of female genitalia. b, Enlarged view of sterigma, ostium and posterior part of ductus bursae. Pennsylvania, Virginia, and Washington state identified as flavivittana are practically identical to those of inana. Until such time as the status of flavivittana can be estab- lished beyond a doubt I am retaining the two names. I wish to thank the authorities of the Philadelphia Academy of Sciences for the loan of the type of Ac/eris inana (Robinson). Also I wish to thank Victor Krantz for the photographs and Silver West for typing the manuscript, both of the Smithsonian staff. LITERATURE CITED Klots, A. B. 1942. The Material of North American Microlepidoptera Other Than Aegeriidae in the American Museum of Natural History. Bulletin of the American Museum of Natural History 79: 391- 424. McDunnough, J. 1934. The Canadian Species of the Tortricid Genus Peronea. Canadian Journal of Re- search 11: 290-332. Powell, J. A. 1964. Biological and Taxonomic Studies on Tortricine Moths, with Reference to the Species in California. I-IV, 1-317, 15 maps, 108 figures, 8 plates. Razowski, J. 1966. World Fauna of the Tortricini (Lepidoptera, Tortricidae). Zaklad Zoologi Sys- tematycznej} W. Krakowie. Polskie} Akademii Nauk. 1-575, 832 figures, 41 plates. Robinson, C. T. 1869. Notes on American Tortri- cidae. Transactions of the American Entomolog- ical Society 2: 261-288, pls. 1, 4-8. PROC. ENTOMOL. SOC. WASH. 89(1), 1987, pp. 177-184 GENERIC REVISION OF CLOEODES AND DESCRIPTION OF TWO NEW GENERA (EPHEMEROPTERA: BAETIDAE) R. D. WALTZ AND W. P. MCCAFFERTY Department of Entomology, Purdue University, West Lafayette, Indiana 47907. Abstract. —The genus Cloeodes is redefined to include species with hindwings and species without hindwings. Notobaetis Morihara and Edmunds (1980) and certain species pre- viously assigned to Centroptella Braasch and Soldan (1980), including the type species, are congeneric with Cloeodes, necessitating the following recombinations: Cloeodes penai (Morihara and Edmunds) n. comb., Cloeodes longisetosus (Braasch and Soldan) n. comb., and Cloeodes soldani (Miiller-Liebenau) n. comb. A new Neotropical genus, Bernerius n. gen., and its type species, B. incus n. sp., from Peru are described. Bernerius is a sister genus of Cloeodes. Species previously assigned to the ceylonensis-similis-pusilla species group of Centroptella are a monophyletic cluster unrelated to Cloeodes (= Centroptella Braasch and Soldan, in part) and are therefore placed in Chopralla n. gen. that includes: Chopralla ceylonensis (Miller-Liebenau) n. comb. (= type species), Chopralla similis (Miiller-Liebenau) n. comb., and Chopralla pusilla (Miiller-Liebenau) n. comb. Cloeodes was erected by Traver (1938) to include several Puerto Rican species of Bae- tidae with adults very similar to Nearctic species assigned to Pseudocloeon Klapalek (both having paired marginal intercalaries of the forewings and no hindwings) and larvae distinct from any known Nearctic species assigned to Pseudocloeon (Cloeodes having a well-developed median terminal filament). Our revisionary study of Baetidae has shown Cloeodes to be a distinct genus possessing several apomorphic character states. We redefine and describe it herein along with describing a new sister genus of Cloeodes. The revision of Cloeodes has ne- cessitated the revision, and hence dissolu- tion, of the genus Centroptella Braasch and Soldan because certain of those species, in- cluding the type species of Centroptella, are actually Cloeodes. Other unrelated Cen- troptella are placed in an additional new and distinctive genus that is also described herein. Genus Cloeodes Traver Cloeodes Traver, 1938: 32. Type species Cloeodes maculipes Traver, by original designation. Notobaetis Morihara and Edmunds, 1980: 606. Type species N. penai Morihara and Edmunds, by original designation. n. syn. Centroptella Braasch and Soldan, 1980: 123. (in part). Type species C. /ongisetosa Braasch and Soldan, by monotypy. n. syn. Larva.—Labrum slightly broader than long, clearly emarginate and with marginal shelf anteriorly. Left mandible (Fig. 3) with incisors fused apically; prostheca stout, dig- itate; no tuft of setae between incisors and molar area; thumb of molar area triangulate and elevated above plane of incisor base. Right mandible (Fig. 4) with incisors sep- arated apically; prostheca reduced, slender, and variably furcate. Labium with 3-seg- mented palps; terminal segment rounded to oblique; segment 2 with weakly developed 178 3 4, Figs. 1-5. setal tuft. inner apical lobe; glossae subequal to para- glossae; paraglossae with subparallel mar- gins. Femora parallel sided, without ventral se- tal patch and with short, dorsal bristles and similarly shaped distal bristles. Tibiae with subproximal arc of long, fine setae; no fine setae adjoining tibial seam. Foreleg with subtending bristle and tibial seam and with subproximal arc of long, fine setae. Claws (Fig. 17) ca. 0.33 tarsal length, with or without microspines. Abdominal terga with broadly pointed, rectangulate-based scales and fine setae; scales with median length subequal to basal width; posterior marginal spines present on all terga; ventrally abdomen with scales and fine setae, with prominent tufts (i.e. with contiguous setal bases) of long, fine setae on PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Cloeodes spp. 1, Forewing. 2, Metascutellar hump. 3, Left mandible. 4, Right mandible. 5, Ventral segments 2-6 (Figs. 5, 18). Gills (similar to Fig. 13) 1-7, asymmetric, broadly pointed, without marginal spination or ciliation; each gill often 2 or more length of associated tergum. Median terminal filament subequal to cercl. Adult male.—Forewings (Fig. 1) with paired marginal intercalaries in most cells; detached base of vein MA, extending well beyond 0.5 x distance between distal cross- vein and proximal (= MA, to MP,) cross- vein. Hindwings present or absent; when present with acute or distinctly hooked me- dian costal process. Posterior margin of metanotum deeply emarginate; metascutel- lar hump (Fig. 2) not flattened before apex but projecting dorsoposteriorly. Forceps four-segmented; segments 2 and 3 nearly fused; segment 3 ca. 2x length segment 2, VOLUME 89, NUMBER 1 segment 2 subequal to segment | and with basal bulge bearing fine bristle-like setae; no median spine or protuberance between for- ceps bases. Adult female.— Marginal intercalaries of forewing paired or less often single. Poste- rior margin of metanotum not as deeply emarginate as in male; metascutellar pro- cess as in male. Subanal plate not devel- oped. Material examined.—Cloeodes macu- lipes Traver: Holotype, ¢ adult, Puerto Rico, Ludillo Mtns., camp lab 46 (107), VI-14- 1935. J. Garcia-Diaz. in alcohol, genitalia slide mounted in balsam (solvent: xylene); larval exuviae of holotype ¢ slide mounted in euparal (solvent: abs. alc.), C.U. Type No. 1402.1, Cornell University. Paratypes, 6 adult (genitalia missing), subimago exu- viae, larval exuviae (missing), same data and deposition as above; paratype (allotype), 2 adult, in alcohol; wing slide mounted, Puer- to Rico, Trout’s pool, El Yunque Trail, VI- 12-1935. J. Garcia-Diaz. C.U. Type No. 1402.2, same deposition as holotype. Cloeodes consignatus Traver: Holotype, 2 adult, Puerto Rico, Yunez River, VI-21- 1935, J. Garcia-Diaz. Cornell University Type No. 1403.1, in alcohol; one wing slide mounted. Notobaetis penai Morihara and Edmunds: Paratypes, ¢ larva, Argentina, Tucuman N.W. of San Miguel de Tucuman, 1-25-1969, W. L. and J. G. Peters, slide mounted in balsam (solvent: xylene), Pur- due Entomological Research Collection (PERC); 2 larva, Argentina, Cordoba Prov., Copina (ca. 25 km WNW Alta Garcia), elev. 1650 m, IV-11/14-1969, L. Pena, slide mounted in balsam (solvent: xylene) (PERC); 2 adult, legs slide mounted in bal- sam (solvent: xylene), same data as above (PERC); ¢ adult, wings slide mounted, same data as above (PERC); three whole larvae (in alcohol), same data as above, Florida A&M University Collection; two adult ¢ and one subimago ¢ and two associated larval exuviae (in alcohol), same data as above, University of Utah Collection. Centroptella 179 longisetosa Braasch and Soldan: paratype larva (in alcohol), Peoples Republic of China, Liu Chui, Kuj Fon Shan River, 1 1- 12-1959, I. Hrdy, deposited Purdue Ento- mological Research Collection, originally from paratypes in the collection of T. Sol- dan. Centroptella soldani Miiller-Liebenau: paratype larva (in alcohol), Ceylon, FC/ 11/c Rasanawa-Fall, Ratnapura, XI-19- 1970. F. StarmiihIner deposited Florida A&M University. Remarks.— Although previously known only from Puerto Rico, a species revision of this genus by Waltz and McCafferty (in press) indicates that Cloeodes is a wide- spread, primarily tropical taxon of Neo- tropical origin, ranging from central Argen- tina to southwestern United States and southeastern Asia. Notobaetis Morihara and Edmunds and certain species of Centrop- tella Braasch and Soldan are clearly con- generic with Cloeodes Traver as indicated by the synapomorphic possession of ventral setal tufts on abdominal segments 2-6. Nominal species of the genus at this writing include Cloeodes consignatus Traver, C. longisetosus (Braasch and Soldan) n. comb., C. maculipes Traver, C. penai (Morihara and Edmunds) n. comb., and C. soldani (Miller-Liebenau) n. comb. We are describ- ing additional species in our revision of Cloeodes. Taxonomic and phyletic relation- ships of Cloeodes with the additional genera described herein is treated under those gen- era. Bernerius Waltz and McCafferty, New GENUS Larva.—Labrum (Fig. 6) slightly broader than long, clearly emarginate and with mar- ginal shelf anteriorly. Left mandible (Fig. 7) with incisors fused apically; prostheca stout, digitate; no tuft of setae between incisors and molar area; thumb of molar area trian- gulate and not elevated above plane of in- cisor base. Right mandible (Fig. 8) with in- cisors separated apically; prostheca reduced, slender and furcate; no tuft of setae between 180 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 6-13. Bernerius incus larva. 6, Labrum (left, ventral; right, dorsal). 7, Left mandible. 8, Right mandible. 9, Maxilla. 10, Labium (left, ventral; right, dorsal). 11, Foreleg (db—distal bristles, pa—proximal arc, ts—tibial seam, sb—subtending bristle). 12, Claw (ms—microspines). 13, Gill 4. VOLUME 89, NUMBER 1 incisors and molar area. Labium (Fig. 10) with three-segmented palps; terminal seg- ment obliquely truncate; segment 2 with weakly developed inner apical lobe; glossae subequal to paraglossae; paraglossae with weakly convex subparallel margins. Femora (Fig. 11) parallel sided, without ventral setal patch, and with short dorsal bristles and similarly shaped distal bristles. Tibiae with subproximal arc of long, fine setae; no fine setae adjoining tibial seam. Foreleg with subproximal arc of setae, sub- tending bristle, and tibial seam. Claws (Fig. 12) ca. 0.33 tarsal length, excavate and with microspines basally. Abdominal terga with rectangulate-based scales and fine setae; scales with median length ca. 2 basal width; posterior mar- ginal spines present on all terga; ventrally abdomen with scales and fine setae, no setal tufts. Gills (Fig. 13) 1-7 asymmetric, elon- gate, and broadly pointed apically; without marginal spination or ciliation. Median ter- minal filament subequal to cerci. Type species. — Bernerius incus Waltz and McCafferty n. sp. Etymology.— Bernerius is of masculine gender and is based on the sur-name of the American ephemeropterist Lewis Berner. Remarks.—This genus is erected for a Pe- ruvian species previously treated as Baetis sp. B by Berner (in Roback et al., 1980) and redescribed as Bernerius incus n. sp. below. Adults are presently unknown. Bernerius is a sister genus to Cloeodes as demonstrated by the following synapomor- phies: the presence of a left mandible (Figs. 3, 7) with incisors fused apically and a stout prostheca; a right mandible (Fig. 4, 8) with incisors partially fused but separated api- cally and a slender, furcate prostheca; the presence of an arc of long, fine setae sub- proximally on the tibia (Fig. 1 1-pa); simple, asymmetrically lamellate, and broadly pointed gills (Fig. 13). Bernerius retains the pleisomorphous condition of abundant long, fine setae on the venter of most abdominal segments, which in Cloeodes species form 181 distinct tufts (Figs. 5, 18) on abdominal seg- ments 2-6. The elevation of the left mandible thumb above the plane of the incisor bases, the presence of clearly defined setal tufts on sterna 2-6, reductionist tendencies in spi- nation of the legs, increased stability in the length of posterior marginal row spines and differences in the degree of convexity of the exterior margin of the paraglossae (toward a more parallel and less convex condition) are phenoclines clearly differentiating lar- vae of the relatively more derived Cloeodes species from Bernerius. The similarity of Bernerius especially to plesiomorphic Cloeodes species suggests a Neotropical origin for Cloeodes. Species of Cloeodes we have studied that possess hindwing pads are apparently restricted to South America. Apomorphic species, in- cluding those to be described from North America and those transferred herein, C. longisetosus (China) and C. soldani (Sri Lanka), have lost the hindwings. Bernerius incus Waltz and McCafferty, NEw SPECIES Baetis(?) sp. B. Berner, 1980: 190. Larva.— Body 10-11 mm long. Head cap- sule lightly pigmented, pale near oral mar- gin and frontal area below antennal bases. Antennae pale, tapered, ca. 3 x head capsule in length; scape subequal to pedicel in length; flagellar segments darkened apically. La- brum (Fig. 6) with 1 + 2-3 subapical setae. Maxillary palp (Fig. 9) exceeding galea-la- cinea, with evidence of third segment; palp with numerous fine setae. Segment 2 of la- bial palp (Fig. 10) ca. 1.25 x length of seg- ment 3 and with 6-8 dorsal setae; apex of segment 3 obliquely truncate with inner marginal stout bristles; paraglossa with 3 apical rows of setae, 4-5 ventral intero-mar- ginal fine setae, and 5-6 dorsal intero-mar- ginal setae medioapically; glossa with ca. 20 stout inner marginal setae. Prothorax lightly pigmented with medial 182 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON dark hourglass marking; meso- and meta- thorax lightly pigmented, without pattern- ing; hindwing pads present. Legs (Fig. 11) lightly pigmented, tibiae and tarsi darker than femora; trochanters with 20 or more laterally serrated blade-like bristles and nu- merous fine setae, without scales; femora with many pointed, laterally serrate ventral bristles, with fine setae and scales, with short, blunt dorsal bristles, and with 6-7 stout, pointed, distal setae; tibiae with bristle-like setae dorsally and ventrally, (subtending bristle blade-like as other tibial setae); tarsi with scales, fine setae, and many bristle-like stout setae dorsally and ventrally, without dorsal row of fine setae. Claws (Fig. 12) ca. 0.33 x tarsal length, with basal microspines. Dorsal abdomen lightly patterned with brown medially; segment | uniformly col- ored; segment 2 with pale brown trapezoidal medial patch; segments 3, 4, and 5 with pale broken coloration; segments 6 and 7 with dark trapezoidal medial patch; segments 8- 10 pale to cream. Posterior marginal spines present on all tergites, basally with clear area as in Centroptilum sp. No. 4 Demoulin (1970), with base of each ca. 0.5 x length, subequal, evenly tapered; smaller spines sit- uated randomly throughout series. Abdo- men pale ventrally, with scales and long, fine setae randomly scattered over surface, without tufts of setae. Paraproct with spi- nous posteromedial margin (large and small spines alternating); surface with scales and fine setae. Type material.— Holotype, 2 larva, Peru, Llave River at Llave, VII-2-1977. 4000 m. Coll. #21. S. S. Roback, Academy of Nat- ural Sciences of Philadelphia. Slide 1, la- bium in balsam (solvent: xylene), other mouthparts mounted in euparal (solvent: abs. alc.); slide 2, abdominal segments 4— 10 mounted in euparal (solvent: abs. alc.); slide 3, gills 3, 4, 7 mounted in balsam (sol- vent: xylene); slide 4, foreleg in balsam (sol- vent: xylene), abdominal segment 3 in eu- paral (solvent: abs. alc.); slide 5, thorax, abdominal segments | and 2 mounted in euparal (solvent: abs. alc.); head capsule (70% ETOH). Remarks. — Roback (Roback et al., 1980) described the Llave River where B. incus (= Baetis sp. B. Berner) was collected as a large river ca. 200 meters wide and organ- ically enriched, presumably with the offal and raw waste from a local slaughter house. Photographs of the habitat indicate the river had slow to moderate current. Chopralla Waltz and McCafferty, NEw GENUS Centroptella Braasch and Soldan, 1980: 123 (in part). Centroptella, Miiller-Liebenau, 1984b: 96 (ceylonensis-similis-pusilla species group). Genus No. 2 sp. | Miiller-Liebenau, 1984a: DN. Larva.—Labrum slightly broader than long, clearly emarginate and with marginal shelf anteriorly. Left mandible (similar to Fig. 3) with incisors fused apically; pros- theca stout and digitate; no tuft of setae be- tween incisors and molar area; thumb of molar area triangulate and elevated above plane of incisor base. Right mandible (sim- ilar to Fig. 4) with incisors separated api- cally; prostheca stout or reduced and fur- cate. Labium with 3-segmented palps; terminal segment rounded to broadly rounded apically; segment 2 with weakly developed inner apical lobe; glossae sub- equal to paraglossae; paraglossae with sub- parallel margins, not convex. Femora (Fig. 14) parallel sided, without ventral setal patch, and with long, dorsal bristles and similarly shaped distal bristles. Tibiae with subproximal arc (Fig. 14-pa) of long, fine setae and with long, fine setae ad- joining meso- and metatibial seams (Fig. 14-tss). Foreleg with subproximal arc of long, fine setae, subtending bristle, and without tibial seam or with weakly developed seam only and with fine setae adjoining tract of tibial seam. Claws (Figs. 15, 16) ca. 0.33 x tarsal length, with 3-5 paired medial and apical denticles. VOLUME 89, NUMBER 1 15 14 Figs. 14-17. 183 16 17 Chopralla ceylonensis larva. 14, Foreleg (db—distal bristles, pa— proximal arc, tss—tibial seam setae). 15, Claw (ventral) after Miiller-Liebenau (1983). 16, Claw (oblique) after Miller-Liebenau (1983). 17, Cloeodes longisetosus larval claw (lateral). Abdominal terga with rectangulate-based and elongate, broadly pointed scales and with fine setae; scales with median length subequal to basal width or with median length ca. 2.0-2.5 basal width; posterior marginal spines present at least on terga 9 and 10; ventrally abdomen with broadly pointed, rectangulate-based scales and fine setae, no setal tufts. Gills 1-7 asymmetric, simple, and rounded apically; without mar- ginal spination, or ciliation. Median ter- minal filament subequal to cerci. Type species.—Centroptella ceylonensis Miller-Liebenau, 1983: 486. Etymology. — Chopralla is of femine gen- der and is named in recognition of the In- dian ephemeropterist B. Chopra. Material examined.—C. ceylonensis (Miller-Liebenau): eight larvae, Sri Lanka, Kitugala, Hal-oya, FC 35/a, XII-27-1970, det. I. Miiller-Liebenau, 1983. Chopralla sp.: four larvae, Viet Nam, Vinh Phu’ Prov., stream nr. Tam-Dao, X-10-1984, T. Soldan deposited in the Purdue Entomological Re- search Collection. Remarks.—Species herein assigned to the genus Chopralla are clearly separated from Cloeodes (= Centroptella in part) species by the absence of ventral tufts of setae on ab- dominal segments 2-6, the apically rounded gills (versus broadly pointed in Cloeodes species), the peculiar claw structure (unlike edentate claws of Cloeodes), and the pos- session of long, fine tibial seam setae (not present in Cloeodes species). Nominal species of Chopralla include the Sri Lankan species Chopralla ceylonensis (Miiller-Liebenau) n. comb. and C. similis (Miiller-Liebenau) n. comb. and the East Malaysian species C. pusilla (Miller-Lie- benau) n. comb. The West Malaysian species, Genus No. 2 sp. 1 Miiller-Liebenau (1984a) could be placed in Chopralla on the basis of the following apparent synapomor- phies: possession of Chopralla-like claws with paired apical denticles, possession of 184 Fig. 18. Cloeodes sp. larva. SEM of ventral setal tuft on left side of abdominal segment 4. long, fine tibial seam setae, asymmetric ovate gills, and fimbriate paraproct scales. This species differs from other known Cho- pralla species, however, by possessing the following inferred plesiomorphic charac- ters: 1) stout prostheca of the right mandi- ble; 2) posterior marginal spines present on segments 1-10; and 3) tergal scales with me- dian length subequal to basal width. Al- though this latter species appears to clearly belong to Chopralla, we do not name the species herein since we have not yet studied the single specimen described. ACKNOWLEDGMENTS We thank G. F. Edmunds, Jr., (Salt Lake City, Utah), W. Flowers, (Tallahassee, Flor- ida), J. K. Liebherr (Ithaca, New York), I. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Miiller-Liebenau (Pl6n, West Germany), S. S. Roback (Philadelphia, Pennsylvania), T. Soldan (Ceske Budejovice, Czechoslova- kia), and Q. D. Wheeler (Ithaca, New York) for the loan of material; and J. Martin and A. V. Provonsha for Figs. 13 and 14 re- spectively. We especially with to thank T. Soldan for the gift of one paratype larva of Centroptella longisetosa now deposited in the Purdue Entomological Research Collec- tion (PERC), West Lafayette, Indiana. This paper has been assigned Purdue University Experiment Station Journal No. 10,436. LITERATURE CITED Braasch, D.and T. Soldan. 1980. Centroptellan. gen., eine neue Gattung der Eintagsfliegen aus China (Baetidae: Ephemeroptera). Reichenb. 18: 123- 27 Demoulin, G. 1970. Ephemeroptera des faunes Ethiopienne et Malgache. Jn South African Ani- mal Life, Results of the Lund University Expe- dition in 1950-1951. 14: 24-170. Morihara, D. K. and G. F. Edmunds, Jr. 1980. No- tobaetis: a new genus of Baetidae (Ephemerop- tera) from South America. Int. Rev. Gesampten Hydrob. 65: 605-610. Miiller- Liebenau, I. 1983. Three new species of the genus Centroptella Braasch and Soldan, 1980, from Sri Lanka (Insecta: Ephemeroptera). Arch. Hy- drob. 97: 486-500. . 1984a. New genera and species of the family Baetidae from West-Malaysia (River Gombak) (Insecta: Ephemeroptera). Spixiana 7: 253-284. . 1984b. Baetidae from Sabah (East Malaysia) (Ephemeroptera). Proc. Fourth Int. Conf. Ephem- eroptera 1983: 85-99. Roback, S. S., L. Berner, O. S. Flint, Jr., N. Nieser, and P. J. Spangler. 1980. Results of the Cath- erwood Bolivian-Peruvian Expedition Part I. Aquatic insects except Diptera. Proc. Acad. Nat. Sci. Phila. 132: 176-217. Traver, J.R. 1938. Mayflies of Puerto Rico. J. Agric. Univ. P.R. 22: 5-42. Waltz, R. D. and W. P. McCafferty. 1987. Revision of the genus Cloeodes Traver (Ephemeroptera: Baetidae). Ann. Entomol. Soc. Am. (In press.) PROC. ENTOMOL. SOC. WASH. 89(1), 1987, pp. 185-199 THE PHYTOPHAGOUS INSECT FAUNA ASSOCIATED WITH BACCHARIS HALIMIFOLIA L. AND B. NEGLECTA BRITTON IN TEXAS, LOUISIANA, AND NORTHERN MEXICO W. A. PALMER North American Field Station, Queensland Department of Lands, 2714 Pecan Drive, Temple, Texas 76502. Abstract.—A survey of the phytophagous insect fauna on Baccharis halimifolia and B. neglecta was undertaken between 1982 and 1986 as part of a program to find biological control agents for B. halimifolia in Australia. One hundred and thirty three phytophagous insect species were collected and these included 11 species that were considered monoph- agous. The monophagous group of species contained a high proportion of Lepidoptera and endophages, and all monophagous species were collected from B. halimifolia. Over 50% of the monophagous group and 21% of the total number of species were found on both species of Baccharis. The relevance of the survey to the biological control program is discussed. The woody shrub Baccharis halimifolia L. (Family Asteraceae: Tribe Astereae: Sub- Tribe Baccharinae), which was introduced from North America in the latter part of the 19th century (Bailey, 1900), has become one of the most serious weeds in Queensland, Australia. The shrub invades cattle pas- tures, reforested areas, and disturbed sites and is a declared noxious weed under the Stock Routes and Rural Lands Protection Acts (1944-1967) of Queensland (Stanley and Ross, 1986). As part of its efforts to control this weed the Queensland Depart- ment of Lands has supported a long ranging research program by the Alan Fletcher Re- search Station to find suitable biological control agents from the New World where the Baccharinae are native. Although the genus Baccharis is best rep- resented in South America with over 300 species, some 21 species including B. ha- limifolia are native to North America. The Alan Fletcher Research Station set up field stations in Lake Placid, Florida (1968) and Curitiba, Brazil (1974) to survey the phy- tophagous insect fauna on Baccharis and to determine which species were sufficiently stenophagous (i.e. having a limited host range) for introduction to Australia. A num- ber of insects were subsequently introduced (McFadyen, 1981). In 1982 the North American Field Station was established in Temple, Texas to survey B. halimifolia at the western margin of its range and the closely related species B. neglecta Britton which is found in central and western Texas. Various surveys of insects on Baccharis sp. have been reported. Bennett (unpub- lished) surveyed the fauna on B. halimifolia in southeastern United States and on var- ious species of Baccharis in Brazil. Tilden (1951), after a very comprehensive survey, listed the insects associated with the vege- tative parts of B. pilularis in the area to the south of San Francisco, California. Kraft and Denno (1982) listed the major herbi- 186 vores attacking B. halimifolia in Maryland. All the studies, with the exception of that of Kraft and Denno (1982) indicated the Baccharis is associated with a considerable number of insect species and that a number of species were most probably monopha- gous. THE PHENOLOGY AND RANGE OF B. HALIMIFOLIA AND B. NEGLECTA Both B. halimifolia and B. neglecta are perennial, dioecious woody shrubs growing to a height of about 15 feet. Both species produce massive amounts of seed which are dispersed by air and so they are often found colonizing disturbed or denuded areas. They are typically found along watercourses, in neglected pastures, along roadsides and drainage ditches, and in vacant lots in towns. The plants usually maintain their foliage during winter but hard freezes can cause defoliation and stem dieback. New growth begins in late winter. Kraft and Denno (1982) reported that the leaf bio- mass of B. halimifolia increased steadily throughout spring and summer and then dropped slowly during autumn in response to an increase in inflorescence biomass. However, possibly more important, they re- ported that the leaves became significantly tougher and thicker during the growing sea- son while the moisture content and nitrogen content declined. The maturing leaves also increased in the concentration of an acetone soluble secondary chemical that acted as a deterrent to herbivory (Kraft and Denno, 1982). These parameters suggested to these authors that there was a general decrease in the quality and availability of the foliage to herbivorous insects, and they noted that no major herbivore was found to feed on B. halimifolia in Maryland after early summer. Both species flower in autumn. The sta- minate inflorescences, which are a rich creamy color, are first to bloom followed a couple of weeks later by the white pistillate inflorescences. By late autumn the very small achenes, each attached to a feathery pappus, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON are dispersed by air. The reproductive out- put of a stand of B. halimifolia has been estimated as high as 376,000 achenes per m?, a figure exceeding that of any other plant species reported in the literature (Panetta, 1979). B. halimifolia and B. neglecta are mor- phologically very similar. Mahler and Wa- terfall (1964) separate the species on the characters of leaf shape and involucre length: B. halimifolia has elliptic to rhomboid leaves and involucres 4-6 mm in length and B. neglecta, narrowly elliptic, linear or oblan- ceolate leaves and involucres 4-8 mm in length. In areas where the two species over- lap it is often difficult to separate them be- cause of the tremendous variation in leaf shape and because intermediate types exist. These intermediate types are a good indi- cation that the two species are very closely related indeed. The habitat range of B. halimifolia ex- tends along coastal areas from Massachu- setts to Texas (Correll and Johnston, 1979). In Texas it is found east of a line that could be drawn between Victoria, Bryan and Dal- las i.e. in higher rainfall areas with acid soil types. It is abundant in coastal areas, low lying and poorly drained areas, and in dis- turbed habitats in townships and oil drilling areas. It is not found in forested areas be- cause the large trees soon completely dis- place it. B. neglecta on the other hand, is found throughout almost all of Texas to the west and south of that line (Correll and Johnston, 1979); in areas with moderate to low rainfall and alkaline soil types. It also is found along roadsides, creeks, vacant lots in townships and other disturbed areas. In some areas it is known as ‘““New Deal weed” or “Roosevelt weed” (Correll and Johnston, 1979) because it became weedy in the 1930s when farmers, for financial reasons, were unable to properly tend to their pastures. Specimens of B. neglecta have been col- lected from Arizona through to North Car- olina and also into Mexico (Correll and Johnston, 1979). VOLUME 89, NUMBER 1 104° W MEXICO ah B-halimifolia B. neglecta “Monterrey Fig: 1, THE AREA AND METHODS OF SURVEY The area covered by this survey is given in Fig. 1. It might be described as consisting of southern Louisiana, the Gulf Prairies and Marshes of Texas, the Blackland Prairies of Central Texas, the southern Edwards Pla- teau of Texas, the South Texas Plains and northern Mexico (cf. Correll and Johnston (1979) for descriptions of the vegetational areas of Texas). The survey was conducted over a four year period between 1982 and 1986. In 1983 and 1984 regular inspections at about two weekly intervals were conducted at a num- ber of sites in close proximity to Temple, Texas with particular emphasis being placed on a site on Lake Stillhouse Hollow about 15 miles west of Temple. The areas farther afield from Temple were visited on an ir- regular basis by 3-4 day trips to such cities 187 y."sConroe }" aa) The area surveyed in Texas, Louisiana and northern Mexico. Cities that were often visited are shown. as Lafayette, Beaumont, Conroe, Galves- ton, Brownsville, Del Rio and Monterrey, Mexico. Stands of Baccharis were inspected along the roadside on these trips particularly where the plants looked to be unhealthy be- cause of possible insect attack. In addition, certain sites were established near all of the above cities and these were inspected at every visit to that city. Insects were collected by both visually in- specting the plant and by sweeping the fo- liage. When evidence of internal insect in- festation was present, plants were either removed from the ground and dissected or the appropriate limb sawn offand split. Any evidence of feeding by the insect was noted. When immatures were found without adults being present, the immatures were collected and reared through to maturity to obtain adults for identification. This applied par- 188 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Phytophagous insect species collected on either B. halimifolia or B. neglecta in Texas, Louisiana or northern Mexico. Relative Fre- quency Eco- ol- Insect-Host Speci- nomic Species lected! Stages Collected Relationship Baccharis Hosts ficity? Pests? ORTHOPTERA Acrididae Hesperotettix viridis viridis O adult foliage feeder B. neg A Thomas Melanoplus differentialis (Tho- O adult foliage feeder B. neg * * mas) Melanoplus sp. R adult foliage feeder B. hal Schistocerca alutacea albolineata R adult foliage feeder B. neg (Thomas) Schistocerca obscura (F.) O adult foliage feeder B. neg b? HEMIPTERA Alydidae Hyalymenus tarsatus (F.) R nymph, adult ectophagous B. neg; B. hal = Coreidae Acanthocephala declivis (Say) R adult ectophagous B. neg - Acanthocephala terminalis (Dal- R adult ectophagous B. neg las) Acanthocephala thomasi (Uhler) R adult ectophagous B. neg = Leptoglossus phyllopus (L.) Cc adult ectophagous B. neg; B. hal = a Merocoris typhaeus (F.) O adult ectophagous B. neg af Mozena lurida (Dallas) R adult ectophagous B. neg U Corimelaenidae Corimelaena pulicaria (Germar) O adult ectophagous B. neg a &2 Cyndnidae Pangaeus bilineatus (Say) R adult ectophagous B. neg a Largidae Largus cinctus (Herrich-Schaef- R nymph, adult ectophagous B. neg * fer) Lygaeidae Lygaeus kalmii Stal O adult ectophagous B. neg - Melanopleuris belfragei (Stal) R adult ectophagous B. neg U Neocoryphus bicrucis (Say) R adult ectophagous B. neg g Nysius niger Baker R adult foliage feeder B. neg f - Nysius raphanus Howard O adult foliage feeder B. neg; B. hal - Ochrimnus mimulus (Stal) C nymph, adult flower and seed _B. neg; B. hal = feeder Oncopeltus fasciatus (Dallas) O adult flower and seed B. neg * feeder Oncopeltus sexmaculatus Stal O adult ectophagous B. neg he Miridae Lygus lineolaris (Palisot de O adult flower feeder B. neg, B. hal = * Beauvois) Polymerus basalis (Reuter) R adult ectophagous B. neg * Pseudatomoscelis seriatus (Reu- O adult flower feeder B. neg; B. hal « + ter) Talorilygus pallidulus (Blan- O adult flower feeder B. neg; B. hal chard) EEE VOLUME 89, NUMBER 1 189 Table 1. Continued. eS Relative Te- quency : Eco- Col- Insect-Host Speci- nomic Species lected Stages Collected Relationship Baccharis Hosts ficity? Pests? Pentatomidae Euschistus servus (Say) Mecidea major Sailer Nezara viridula (L.) Thyanta accerra McAtee Thyreocoridae Galgupha sp. HOMOPTERA Acanaloniidae Acanalonia bivittata (Say) Acanalonia conica (Say) Acanalonia laticosta Doering Acanalonia parva Doering Aphididae Aphis pr. baccharicola HRL Aphis coreopsidis (Thomas) Cercopidae Clastoptera xanthocephala Ger- mar Lepyronia quadrangularis (Say) Cicadellidae Aceratagallia calcaris Oman Balclutha sp. Chlorotettix viridius Van Duzee Empoasca fabae (Harris) Homalodisca coagulata (Say) Menosoma cinctum (Osborn & Ball) Cixiidae Oecleus productus Metcalf Oliaris aridus Ball Delphacidae Stobaera pallida Osborn Dictyopharidae Rhynchomitra recurva (Metcalf) Flatidae Anormenis septentrionalis (Spino- la) Metcalfa pruinosa (Say) Ormenis saucia Van Duzee Ormenis sp. Ormenoides venustus (Melichar) Fulgoridae Poblicia fuliginosa (Olivier) Issidae Hysteropterum auroreum (Uhler) Gre AROR o) ANCA AR AW OAC Oy «Oo R R adult adult adult adult adult adult nymph, adult adult adult nymph, adult nymph, adult nymph, adult adult adult adult adult nymph, adult adult adult adult adult nymph, adult nymph, adult adult adult adult adult adult adult adult ectophagous ectophagous ectophagous ectophagous ectophagous ectophagous ectophagous ectophagous ectophagous ectophagous ectophagous ectophagous ectophagous ectophagous ectophagous ectophagous ectophagous ectophagous ectophagous ectophagous ectophagous ectophagous ectophagous ectophagous ectophagous ectophagous ectophagous ectophagous ectophagous ectophagous . neg; B. hal . neg . hal . neg Dow jee) . neg neg hal neg hal Dow wm ee) . neg; B. hal B. hal B. neg; B. hal B. neg neg neg neg . neg; B. hal . neg; B. hal neg B. neg B. neg B. neg; B. hal B. hal B. hal . neg; B. hal neg hal hal wo jee) . neg B. neg * *& + * & & & “GCG Cc ix 190 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Continued. EE EEE EEE Relative Fre- quency Eco- Col- Insect-Host Speci- nomic Species lected! Stages Collected Relationship Baccharis Hosts ficity? Pests? Se EE ee EE EE EE EEE EE ES EEE EE Membracidae Tortistilus abnormus (Caldwell) R adult ectophagous B. neg U Vanduzeea segmentata (Fowler) O nymph, adult ectophagous B. neg st LEPIDOPTERA Gelechiidae Aristotelia ivae Busck O larva foliage feeder B. neg; B. hal ete Geometridae Anacamptodes defectaria (Gue- O larva foliage feeder B. neg = née) Anavitrinelia pampinaria (Gue- R larva foliage feeder B. neg a 2 née) Eupithecia miserulata Grote R larva foliage feeder B. neg = Itame varadaria (Walker) O larva foliage feeder B. hal 2% Pero meskearia Packard R larva foliage feeder B. neg U Pleuroprucha insulsaria (Gue- O larva flower feeder B. neg; B. hal a née) Lyonetiidae Bucculatrix ivella Busck C larva, pupa leaf miner B. neg; B. hal sit Noctuidae Platysenta videns (Guenée) O larva foliage feeder B. neg; B. hal =? Spodoptera frugiperda (Smith) O adult foliage feeder B. neg Spodoptera ornithogalli (Guenée) O larva foliage feeder B. neg; B. hal a7 Pterophoridae Oidaematophorus balanotes € larva, pupa stem borer B. neg; B. hal piel (Meyrick) Oidaematophorus kellicotti R larva stem borer B. neg se (Fish) Pyralidae Homoeosoma electellum (Hulst) R larva flower feeder B. neg af t? Tortricidae Epiblema discretivana (Heinrich) c larva stem gall B. hal stat Platynota sp. R pupa B. neg U Sonia paraplesiana Blanchard © larva, pupa root feeder B. hal U COLEOPTERA Cerambycidae Amniscus perplexus (Haldeman) C larva, pupa stem borer B. hal sats Ancylocera bicolor (Olivier) R adult B. neg = Anelaphus sp. R adult B. neg Dendrobias mandibularis Ser- R adult B. hal * ville Dorcasta cinerea (Horn) R adult B. neg U Eliphidion linsleyi Knull O larva stem borer B. neg Eliphidionoides incertus New- O larva stem borer B. neg man Euderces pini (Olivier) R adult B. neg U Stenosphenus dolosus Horn R adult B. neg U Tragidion coquus L. R adult B. neg U eee eee VOLUME 89, NUMBER 1 191 Table 1. Continued. Relative Fre- quency Eco- Col- Insect-Host Speci- nomic Species lected! Stages Collected Relationship Baccharis Hosts ficity? ‘Pests? Chrysomelidae Altica sp. R adult ectophagous B. hal U Calomicrus prob. blakeae Wilcox R adult B. neg U Colaspis planicostata Blake R adult B. hal U Colaspoides opacicollis Horn R adult B. neg U Cryptocephalus cribripennis Le R adult B. neg ss Conte Cryptocephalus nr. pumilis Gc adult ectophagous B. neg; B. hal . Haldeman Diabrotica balteata Le Conte O adult ectophagous B. hal ig - Diabrotica connexa Le Conte R adult ectophagous B. hal a Diabrotica undecimpunctata ho- ‘e adult ectophagous B. neg; B. hal wardi Barber Diachus auratus (Fab.) O adult flower feeder B. neg; B. hal As Exema elliptica Karren (C larva, adult foliage feeder B. hal axx Microtheca ochroloma Stal R adult ectophagous B. hal U Monoxia sp. R adult ectophagous B. hal U Nodonota rotundicollis Schaeffer S adult foliage feeder B. neg * Nodonota texana Schaeffer R adult ectophagous B. neg Nodonota tristis (Olivier) O adult ectophagous B. neg a Systena blanda Melsheimer O adult foliage feeder B. neg aS « Ophraella sexvittata Le Conte R adult foliage feeder B. hal aay Paria thoracica (Melsheimer) R adult ectophagous B. hal a) Trirhabda bacharidis (Weber) EC larva, adult foliage feeder B. neg; B. hal abt Curculionidae Baris sp. R adult B. neg U Compsus auricephalus (Say) O adult B. neg - Cophes texanus Sleeper R adult endophagous? B. neg U Eudiagogus pulcher Fahraeus R adult ectophagous B. hal = Tsodacrys burkei Howden O adult B. neg; B. hal * Lixus scrobicollis Boheman R adult endophagous? B. neg Mitostylus setosus (Sharp) R adult ectophagous B. neg * Prosaldius blanditus (Casey) R adult B. hal U Prosaldius deplanatus (Casey) O adult B. neg; B. hal U Dermestidae Cryptorhopalum uteanum Casey R adult B. neg U Elateridae Melanotes indistinctus Quate R adult B. hal U Lampyridae Lucidota sp. R adult B. neg U Pyropyga sp. R adult B. hal U Scarabaeidae Cotinus mutabilis Gory and Per- O adult ectophagous B. hal * cheron Euphoria sepulchralis (F.) O adult ectophagous B. hal Tenebrionidae Bothrotes canaliculatus acutus R adult B. neg +) (Le Conte) 192 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Continued. Relative Fre- quency Eco- Col- Insect-Host Speci- nomic Species lected! Stages Collected Relationship Baccharis Hosts ficity? Pests? DIPTERA Agromyzidae Phytobia sp. O pupa leaf miner B. neg U Cecidomyiidae Neolasioptera lathami Gagné C larva stem galler B. neg; B. hal Ad Tephritidae Acinia picturata (Snow) R adult seed feeder B. hal = Dioxyna picciola (Bigot) R adult B. hal U Euarestoides acutangulus R adult B. neg U (Thomson) Neaspilota dolosa Benjamin R adult B. neg U Neotephritis finalis (Loew) R adult B. neg U Tephritis new sp. € larva stem galler B. neg; B. hal oa% Tephritis subpura (Johnson) Cc larva stem galler B. hal pt Trupanea nr. actinobola (Loew) R adult B. neg U Tomoplagia obliqua (Say) R adult B. hal U 'R = rare, O = occasional, C = common. 2 *** — monophagous (host plants apparently restricted to the genus Baccharis); ** = oligaphagous (host-plants apparently restricted to the tribe Astereae; * polyphagous (having a wider host range than above two categories); *? = specificity unknown but very likely polyphagous; U = specificity unknown. 3 * = pest species. ticularly to caterpillars, leafminers, and gall formers. Samples of inflorescences were also collected in the autumn and placed in emer- gence cages and any resulting insects col- lected. All insect specimens were first submitted to the Biosystematic and Beneficial Insects Institute, Agricultural Research Service, USDA, Beltsville, Maryland for expert identification by specialists of that Insti- tute’s Systematic Entomology Laboratory. When species could not be fully identified by this laboratory the specimens were later forwarded elsewhere to other taxonomists expert with the particular group in question. After the insects had been properly iden- tified, entomologists knowledgeable about the particular species or group and the lit- erature were consulted to determine the de- gree of stenophagy exhibited by the species. Species that appeared to be sufficiently ste- nophagous were then selected for formal host specificity testing in order to obtain per- mission to introduce the insect into Aus- tralia (e.g. Palmer, 1986). RESULTS The phytophagous species (excluding pol- len and nectar gatherers) found on either species of Baccharis are shown in Table 1. One hundred and thirty three species were collected representing six orders. The Or- thoptera, Hemiptera, Homoptera, Lepidop- tera, Coleoptera and Diptera were repre- sented by 5 (or 4% of the species), 27 (20%), 27 (20%), 17 (13%), 46 (35%) and 11 (8%) species respectively. The insects were classified as monopha- gous if restricted to Baccharis, oligophagous if the host range was restricted to the Tribe Astereae and polyphagous if having a wider host range. Evidence of host range was ob- tained from formal host testing, observa- tions during the course of the survey, con- sultations with acknowledged experts on specific groups of insects, examination of VOLUME 89, NUMBER 1 major insect collections, and texts such as Arnett (1985), Slater and Baranowski (1978), Arnett et al. (1980) and Borrer et al. (1981). Eleven insects were considered monopha- gous, and interestingly, all were found on B. halimifolia. Six of these eleven were also found on B. neglecta but only one monoph- agous species, Tephritis new sp., waS more commonly found on B. neglecta than on B. halimifolia. The monophagous fauna thus represented about 8% of the total phytoph- agous fauna. Six of the 11 monophagous insects (or approximately 55%) were endophagous for at least part of their lifecycle. Of ten species that were definitely endophagous on Bac- charis, six were monophagous, one was oli- gophagous, two were polyphagous, and the hosts of one were not known although it also was quite probably monophagous. A very high proportion (80%) of the endophages therefore had a limited host range. Only two insects were classified as oli- gophagous. The remaining | 18 species were considered either polyphagous, host un- known or hosts unknown but probably po- lyphagous. The proportion of oligophagous to polyphagous species depends, of course, on what arbitrary criteria are set for oligoph- agy. Five of 11 monophagous species were also Lepidoptera. This proportion is consider- ably higher than the proportion of Lepi- doptera found in the total number of species. While six of 11 (or 55%) monophagous species were common to both B. halimifolia and B. neglecta, only 28 of the total 133 species (21%) were common to both species. This perhaps indicates that many of the po- lyphagous insects did not have any sub- stantial relationship with these hosts but rather their occurance (or absence) was de- pendent primarily on other factors. A number of well known crop pests were collected on Baccharis. These included the differential grasshopper, Melanoplus differ- entialis Thomas; the lygus bug Lygus lineo- laris (Palisot de Beauvois); the cotton flea- 193 hopper, Pseudatomoscelis seriatus (Reuter); the brown stinkbug, Euschistus servus (Say); the southern green stinkbug, Nezara viri- dula (L.); the southern corn rootworm, Dia- brotica undecimpunctata howardi Barber and the fall armyworm, Spodoptera frugi- perda (Smith). During the course of the study many non- phytophagous insects were collected. These included known predator and flower feeding species as well as many insects that were probably only casually associated with the plant. These species are listed in Table 2. The collection of these insects was only a very secondary aspect of the project and quite likely there were many more such species present on Baccharis than are listed. NOTES ON THE More IMPORTANT SPECIES By far the most important phytophage was the chrysomelid Trirhabda bacharidis (We- ber) which was found throughout the survey area except for the lower Rio Grande Valley and northern Mexico. It is univoltine in the study area with larvae occurring in late win- ter and adults being found from late April to early August. However in one year, 1984, following an unusually wet autumn, early instar larvae were found in mid-December but these were killed during winter. It there- fore appears that at least some individuals in the population do not have a diapause mechanism and that the regularity of emer- gence at the end of winter may be more a function of extreme mortality of early emerging individuals rather than a dia- pause. Both larvae and adults can occur in tremendous numbers and are capable of completely defoliating a bush. The effect is particularly destructive if late winter freezes occur while the bushes are regenerating their foliage following larval attack. In this situ- ation the stems are frequently killed. The case bearing chrysomelid Exema el- liptica Karren was common throughout the B. halimifolia area in spring and summer. Larvae were found in April and May and adults thereafter. Both stages fed on foliage 194 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 2. Non-phytophagous insect species collect- ed on either B. halimifolia or B. neglecta in Texas, Louisiana or Northern Mexico. Species Habit HEMIPTERA Anthocoridae Orius insidiosus (Say) predator Miridae Deraeocoris nebulosus (Uhler) predator Pentatomidae Euthyrhynchus floridanus (L.) predator Podisus maculiventris (Say) predator Phymatidae Phymata americana Melin predator Phymata sp. predator Reduviidae Zelus bilobus (Say) predator Thyreocoridae Galgupha sp. incidental COLEOPTERA Bruchidae Merobruchis major (Fall) Cantharidae Cantharis sp. Cantharis sp. Chauliognathus basalis Le Conte Chauliognathus marginatus (F.) Chauliognathus scutellaris Le Conte Chauliognathus sp. Podabrus sp. Carabidae Colliuris pennsylvanica (Linne) Coccinellidae Coleomegilla maculata fuscila- bris (Mulsant) Cycloneda sanguinea (L.) Hippodamia convergens Guerin Olla v nigrum Mulsant Scymnus loewii Mulsant Meloidae Epicauta pennsylvanica (De- Geer) Melyridae Collops balteatus LeConte Collops quadrimaculatus F. Nitidulidae Carpophilus nr. transitans Sharp pollen feeder pollen feeder pollen feeder pollen feeder pollen feeder pollen feeder pollen feeder pollen feeder predator predator predator predator predator predator predator predator predator incidental Table 2. Continued. Species Habit Phalacridae Phalacrus sp. incidental DIPTERA Bibionidae Plecia nearctica Harcy incidental Bombylidae Villa sp. incidental Calliphoridae Chrysomya rufifacies (Macquart) incidental Cochliomyia macellaria (Fab.) incidental Chironomidae Procladius bellus (Loew) incidental Chloropidae Apallates particeps (Becker) incidental Conioscinella grisescens (Sa- brosky) incidental Conioscinella nuda (Adams) incidental Liohippelates pusio (Loew) incidental Thaumatomyia glabra (Mg.) incidental Ephydridae Ditrichophora argyrostoma (Cresson) incidental Ochthera lauta Wheeler incidental Philygria debilis Loew incidental Muscidae Musca domestica (L.) incidental Sepsidae Palaeosepsis pusio (Schiner) incidental Syrphidae Allograpta obliqua (Say) Palpada agrorum (Fab.) Palpada pusilla (Macquart) Palpada vinetorum (Fab.) Syrphus rectus O. S. Toxomerus politus (Say) Tachinidae Angiorhina sp. Pseudomyothria nr. ancilla (Walker) Ptilodexia sp. HYMENOPTERA Apoidae Apis mellifera L. Vespidae Dasymutilla sp. Polistes apachus Saussure Polistes sp. pollen feeder pollen feeder pollen feeder pollen feeder pollen feeder pollen feeder incidental incidental incidental pollen feeder incidental incidental incidental VOLUME 89, NUMBER 1 and heavily infested small plants may ex- hibit damage to their terminals. Although the type series for E. elliptica was reported from Iva frutescens L. (Karren, 1966), it was not found on J. frutescens in this survey even though this plant was growing in close proximity to infested B. halimifolia on many occasions. It is therefore considered that a misidentification of the morphologically similar plant species may have occurred and that FE. elliptica may be specific to Bac- charis. Two other genera of chrysomelids are quite commonly found on Baccharis. Adults of Nodonota spp. were found on B. neglecta in the spring. Infestations of N. ro- tundicollis Schaeffer were often seen along the Rio Grande Valley. Damage was in- variably noticeable but of little significance. Adults of the three polyphagous Diabrotica species were taken quite commonly in the autumn but never in damaging numbers. The lepidopterous foliage feeders caused at most only minor damage to the plant. Bucculatrix ivella Busck was the most abun- dant of these; and in April populations of several hundred per plant were sometimes seen, particularly on B. halimifolia. The first three instars feed inside a serpentine mine while the last two instars are external feed- ers. The very characteristic ribbed pupal co- coons were also found on the plant. During the rest of the year only very occasional specimens were seen. Greater detail on the biology and host specificity is given by Palmer and Diatloff (in press). The leaf web- bing caterpillar, Aristotelia ivae Busck, was also quite commonly found in spring but there were rarely more than one or two per plant. These small greenish larvae feed un- der a web on the leaf and become explo- sively active when touched. The geometrid, Itame varadaria (Walker) was collected from B. halimifolia at a number of sites by sweep- ing the foliage. It had three generations per year with larvae being present in April, July and October. It was never very abundant: a collection of half a dozen larvae after an hour’s sweeping was a typical result. 195 The most abundant stem borer was the plume moth Oidaematophorus balanotes (Meyrick), which was found throughout the survey area. The phenology of this univol- tine species was clearly defined. Moths were active in late summer and early autumn. Early instar larvae were often seen in inflo- rescences placed in emergence cages. They were also found in damaged vegetative ter- minals. Later instar larvae bored into the woody tissue of the stem and created a char- acteristic gallery which were up to a meter in length. The exit hole was covered with woody frass which had been removed from the gallery. Pupation and eclosion of the moth occured in the gallery. Occasionally bushes were heavily infested with this insect and on one occasion 15 larvae were found in the one stem. However, it was much more common to find plants infested with just one or two larvae. The related species, O. kellicotti (Fish) was found in B. neglecta in northern Mexico. This is a new host rec- ord for this species that has previously been reported only from Solidago spp. (Cashatt, 1972). The cerambycid, Amniscus perplexus (Haldeman), was found to infest a large pro- portion of B. halimifolia plants at just a few sites. It was also univoltine with adult ac- tivity in late spring and early summer. Eggs were Oviposited under the bark, usually near the crown of the plant and within 30 cm of ground level. Larval feeding continued from summer to the following spring when both pupae and teneral adults were found in the larval galleries. A characteristic finely pow- dered frass was found at the base of infested plants. Both large and small plants were at- tacked and it was quite common to find 2- 3 larvae in quite small plants. The larvae significantly weakened the stems and pre- disposed the plants to attack by disease or- ganisms. Two cerambycids Eliphidion linsleyi Knull and Eliphidionoides incertus were found in B. neglecta stems along the Rio Grande Valley. Both species were associ- 196 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ated with weakened or dying branches, but it was not ascertained whether they had caused this damage or whether they had at- tacked already dying material. Four gall forming species were commonly found. The cecidomyiid Neolasioptera la- thami Gagné was found on both Baccharis species throughout the survey area. It forms a soft globular gall on the stems and ter- minals. A large gall (3 cm in diameter) might contain as many as 15 larvae, each in an individual chamber. Isolated patches of Baccharis were infested with up to 10 galls per plant in spring. For the rest of the year only very occasional galls were found. Sig- nificant damage to the plant was not ob- served within this survey area. A more de- tailed account of this insect is given by Diatloff and Palmer (1987). The two tephritids, Tephritis subpura (Johnson) and 7. new sp., appeared to oc- cupy very similar ecological niches on B. halimifolia and B. neglecta respectively. Both fed in the terminal stems in spring and caused characteristic swelling of the final few centimeters of the stem and ultimately terminal die-off. Flies of both species emerged in autumn from inflorescences placed in emergence cages. The autumn adults of both species were smaller and darker than the individuals emerging in the spring. The tortricid Epiblema discretivana (Teinrich) occurred in elongate woody stem galls, approximately 3 cm in length, on B. halimifolia. This insect was also univoltine, with adults emerging in early spring and the insect overwintering as larvae. E. discreti- vana is generally distributed throughout the habitat of B. halimifolia. As many as ten galls have been found on plants, but damage attributable to them was not discerned. The delphacid Stobaera pallida Osborn was found to be quite abundant on B. ha- limifolia and much less abundant on B. ne- glecta. There appears to be three generations a year with population peaks occurring in May, July and September. It was possible to collect over 100 individuals by sweeping one large bush. The life cycle is similar to that of other species of Stobaera (McClay, 1983; Reimer and Goeden, 1982). Eggs are Oviposited into the pith of stems and both nymphs and adults remain on the plant. With the onset of flowering the lygaeid Ochrimnus mimulus (Stal) adults were pres- ent in tremendous numbers on both male and female inflorescences. Later in autumn nymphs were found by dissecting the female inflorescences. The insect overwintered as late instar nymphs or adults, which were quite commonly found throughout the spring and summer. A more detailed ac- count of this insect is given by Palmer (1986). The coreid Leptoglossus phyllopus (Say) was also very commonly associated with these plant species while they were flowering. PROSPECTS FOR BIOLOGICAL CONTROL Four of the insects, Trirhabda bacharidis, Aristotelia ivae, Oidaematophorus bala- notes and Neolasioptera lathami had pre- viously been found elsewhere in the United States and had been proved host specific by various officers of the Queensland Depart- ment of Lands. T. bacharidis was released in Queensland where it now occurs in dam- aging populations in some localized areas. A. ivae became generally distributed but has only been found at low, non-damaging pop- ulation levels. O. balanotes is at present being released in the field and N. lathami has not yet been reared in the laboratory in Australia. A further five insects have been tested at the North American Field Station, Temple and permission to introduce these species into quarantine in Australia has been ap- proved or is anticipated. The species are Tephritis new sp., Stobaera pallida, Buccu- latrix ivella, Itame varadaria and Amniscus perplexus. The remaining monophagous in- sects will be tested in the near future. These 11 insects were rated in two ways in an attempt to predict their eventual ef- fectiveness as biocontrol agents in Austra- VOLUME 89, NUMBER 1 lia. Ideally it would be highly desirable if a reliable quantitative formula were available for use by biocontrol researchers. Harris (1973) devised a formula that was later modified by Goeden (1983), and this latter formula is possibly the best available at this stage. All the insects were therefore scored by Goeden’s formula (Table 3). The scores ranged from 34 to 53. In this system N. lathami, O. balanotes and B. ivella, by scor- ing more than 50 points, would be consid- ered superior prospects and the rest were predicted to be partially effective agents. The insects were also assessed subjec- tively by the author based on observations in the United States only and rated from | (poor prospect) to 5 (superior prospect) after considering such aspects as damage ob- served, ecoclimatic similarity to Australia, potential reproductive rate and degree of parasitism observed. 7. bacharidis, B. ivella and A. perplexus were considered to be the best prospects. It should be pointed out, however, that a general and reliable method for predicting eventual effectiveness of potential biocon- trol agents has not yet been devised and its reliability proved. Aspects of Goeden’s for- mula have been criticized by both Palmer (unpublished) and Wapshere (1985) who questioned whether it was indeed possible to quantify potential effectiveness in a new habitat. Perhaps the point to be made is that, while it is highly desirable to attempt to predict the best possibilities, preferably by quantitative methods, all sufficiently stenophagous agents should ultimately be utilized when at all possible. DISCUSSION Faunal richness of species inhabiting a plant species is determined by many factors but Strong et al. (1984) considered the two most important factors to be the size of the geographic range and the plant “‘architec- ture”’ (i.e. 1ts size and growth form). Both Baccharis species are rather large, woody, perennial shrubs that occupy an extensive 197 Table 3. The potential effectiveness of the mono- phagous species as biocontrol agents as predicted by the formula of Goeden (1983) and by the author’s sub- jective assessment (with a poor candidate scoring | and a superior prospect scoring 5). Author’s Goeden’s Assess- Species Formula ment Amniscus perplexus 47 5) Trirhabda bacharidis 45 5 Exema elliptica 34 3 Aristotelia ivae 49 3 Oidaematophorus balanotes 53 4 Bucculatrix ivella 51 5 Itame varadaria 44 ? Epiblema discretivana 36 1 Stobaera pallida 41 2 Tephritis subpura 40 3 Tephritis palmeri n. sp. 35 2 geographic habitat and these factors should indicate a rich insect fauna such as was found in the survey. Perhaps it could also be ar- gued that as the geographic area occupied by B. halimifolia is much greater than that of B. neglecta a greater number of monoph- agous insects might be associated with B. halimifolia, as was found in this study. Another factor influencing the number of species found is, of course, the length of time devoted to the survey. New species were still being found in the last year of this project and undoubtedly the faunal list would have been longer had the survey been continued for a longer period. Nevertheless, four years represents a very adequate time frame for such a survey. The number of insect species common to both species of Baccharis clearly indicates that these plant species are very similar chemically as well as morphologically. In fact, the association is even closer than the data indicate. In the laboratory A. perple- xus, I. varadaria, and E. elliptica, found only on B. halimifolia in the field, fed readily on B. neglecta. Furthermore, a number of ste- nophagous insects collected from B. pilu- laris D.C. have fed equally well on both local species of Baccharis and also B. sar- 198 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON athoides Gray, which is found in Arizona. It is therefore very probable that the differ- ences in insect fauna found between the two Baccharis species in this survey are due to different climatic factors or factors other than intrinsic differences between the species themselves. A very close association between steno- phagy and endophagy was evident. Endo- phages by their very nature are specialized with adaptions for internal living and must develop a close relationship with their host. It 1s therefore not surprising that a signifi- cant proportion of them are highly stenoph- agous. The high proportion of endophages that were also monophagous in this survey highlights the need for those involved in biological control programs such as this to place great importance on searching for en- dophages. ACKNOWLEDGMENTS I particularly thank the expert taxono- mists who identified the insects. Most tax- onomists were associated with the System- atic Entomology Laboratory (SEL), Agricultural Research Service, USDA or the associated Department of Entomology, Na- tional Museum of Natural History (NMNH), Smithsonian Institution. I also thank the Biosystematics and Beneficial Insects Insti- tute for providing such a fine service for collectors in North America through these two institutions. Identifications were performed by the fol- lowing taxonomists many of whom also of- fered valuable advice on the biology and habits of the insects they had identified: R. L. Brown, Mississippi Entomological Mu- seum (Lepidoptera: Tortricidae); E. V. Cashatt, Illinois State Museum (Lepidop- tera: Pterophoridae); J. A. Chemsak, UC Berkeley (Coleoptera: Cerambycidae); D. R. Davis, NMNH (Lepidoptera: Lyonetiidae); D. C. Ferguson, SEL (Lepidoptera: Geo- metridae, Pterophoridae, Pyralidae); R. H. Foote, SEL (Diptera: Tephritidae); R. J. Gagné, SEL (Diptera: Cecidomyiidae, Cal- liphoridae); R. D. Gordon, SEL (Coleop- tera: Scarabaeoidae, Meloidae, Lampyri- dae, Coccinellidac):) Ts J. Henry sb (Hemiptera); R. W. Hodges, SEL (Lepidop- tera: Gelechoidae); F. Hovore, Placerita Canyon Nature Center (Coleoptera: Cer- ambycidae); J. Jenkins, Washington State University (Diptera: Tephritidae); J. M. Kingsolver, SEL (Coleoptera: Dermestidae, Bruchidae); L. Knutson, SEL (Diptera: Bombyliidae); J. P. Kramer, SEL (Homop- tera: Fulgoridae, Cicadellidae, Membraci- dae, Cercopidae); D. R. Miller, SEL (Ho- moptera: Coccoidae); D. A. Nickle, SEL (Orthoptera: Acrididae); R. W. Poole, SEL (Lepidoptera: Noctuidae); C. W. Sabrosky, SEL (Diptera: Chloropidae); F. C. Thomp- son, SEL (Diptera: Syrphidae); T. J. Spil- man, SEL (Coleoptera: Cerambycidae, Ela- teridae); M. B. Stoetzel, SEL (Homoptera: Aphididae); W. W. Wirth, SEL (Diptera: Chironomidae); R. E. White, SEL (Coleop- tera: Chrysomelidae, Cerambycidae); and D. R. Whitehead, SEL (Coleoptera: Cur- culionidae). LITERATURE CITED Arnett, R. H. 1985. American insects. A handbook of the insects of America north of Mexico. Van Norstrand Reinhold Co. New York. 850 pp. Arnett, R. H., N. M. Downie, and H.E. Jacques. 1980. How to know the beetles. W. C. Brown Co. Du- buque, IA. 417 pp. Bailey, F. 1900. The Queensland flora, Part 3. A. J. Diddams & Co. Brisbane, Qld. Borrer, D. J., D. M. De Long, and C. A. Triplehorn. 1981. An introduction to the study of insects. 5th Edition. Saunders College Publishing. Philadel- phia, PA. 827 pp. Cashatt, E. D. 1972. Notes on the balanotes (Meyr- ick) group of Oidaematophorus Wallengren with description of a new species (Pterophoridae). J. Lepid. Soc. 26: 1-13.11 Correll, D. S. and M. C. Johnston. 1979. Manual of the vascular plants of Texas. University of Texas at Dallas, Dallas, TX. 1881 pp. Diatloff, G. and W. A. Palmer. 1987. The host spec- ificity of Neolasioptera lathami Gagné (Diptera: Cecidomyiidae) with notes on its biology. Proc. Entomol. Soc. Wash. 89: 185-199. VOLUME 89, NUMBER 1 Goeden, R. D. 1983. Critique and revision of Harris’ scoring system for selection of insect agents in bi- ological control of weeds. Prot. Ecol. 5: 287-301. Harris, P. 1973. The selection of effective agents for the control of weeds. Can. Entomol. 105: 1495- 1503. Karren, J. B. 1966. A revision of the genus Exema of America, North of Mexico. Univ. Kansas Sci- ence Bull. 46: 672. Kraft, S. K. and R. F. Denno. 1982. Feeding re- sponses of adapted and non-adapted insects to the defensive properties of Baccharis halimifolia L. (Compositae). Oecol. 52: 156-163. Mahler, W. F. and U. T. Waterfall. 1964. Baccharis (Compositae) in Oklahoma, Texas and New Mex- ico. Southwest. Nat. 9: 189-202. McClay, A. S. 1983. Biology and host specificity of Stobaera concinna (Stal) (Homoptera: Delphaci- dae), A potential biocontrol agent for Parthenium hysterophorus L. (Compositae). Fol. Entomol. Mex. 56: 21-30. McFadyen, P.J. 1981. Current status of the biological control programme against groundsel bush (Bac- charis halimifolia). Proc. 6th. Aust. Weeds Conf. Vol. 1. 151-154. Palmer, W. A. 1986. Host specificity of Ochrimnus mimulus (Stal) (Hemiptera: Lygaeidae) with notes on its phenology. Proc. Entomol. Soc. Wash. 88: 451-454. Palmer, W. A. In press. Host specificity of Buccu- 199 latrix wella Busck (Lyonetiidae): a potential bio- control agent for Baccharis halimifolia L. in Aus- tralia. J. Lepid. Soc. Panetta, F.D. 1979. The effects of vegetation devel- opment upon achene production in the woody weed, groundsel bush (Baccharis halimifolia L.). Aust. J. Agric. Res. 30: 1053-1065. Reimer, N. J. and R. D. Goeden. 1982. Life history of the delphacid planthopper Stobaera tricarinata (Say) on western ragweed, Ambrosia psilostachya DC, in southern California (Hemiptera-Homop- tera: Delphacidae). Pan-Pac. Entomol. 58: 105- 108. Slater, J. A. and R. M. Baranowski. 1978. How to know the true bugs. W. C. Brown Co. Iowa. 256 pp. Stanley, T. D. and E. M. Ross. 1986. Flora of South- Eastern Queensland Vol. 2. Queensland Depart- ment of Primary Industries. Brisbane. Misc. Pub. QM84007. 623 pp. Strong, D. R., J. H. Lawton, and R. Southwood. 1984. Insects on plants. Community patterns and mech- anisms. Harvard University Press. Cambridge, MA. 313 pp. Tilden, J. W. 1951. The insect associates of Baccharis pilularis De Candolle. Microentomology 16: 149- 188. Wapshere, A. J. 1985. Effectiveness of biological con- trol agents for weeds: present quandaries. Agric. Ecosystems Environ. 13: 261-280. PROC. ENTOMOL. SOC. WASH. 89(1), 1987, p. 200 NOTE Hedge Bindweed, Calystegia sepium (Convolvulaceae), an Adventitious Host of the Chrysanthemum Lace Bug, Corythucha marmorata (Heteroptera: Tingidae) The chrysanthemum lace bug, Corythu- cha marmorata (Uhler), ranges throughout most of the United States and southern Canada (Slater and Baranowski, 1978. How to know the true bugs (Hemiptera-Heter- optera). Wm. C. Brown, Dubuque, Iowa, 256 pp.). As the common name suggests, this tingid feeds on Compositae (= Aster- aceae), particularly species of Ambrosia, As- ter, Chrysanthemum, Helianthus, and Sol- idago (Bailey, 1951. Entomol. Am. 31: I- 140). Drake and Ruhoff (1965. U.S. Nat. Mus. Bull. 243: 1-634) listed Echinops, Rudbeckia, and Tanacetum spp. as addi- tional composite hosts, and Horn et al. (1979. No. Car. Agric. Exp. Stn. Tech. Bull. 257: 1-22) added Silphium to the known hosts. In Pennsylvania, C. marmorata 1s common on spotted knapweed, Centaurea maculosa Lam., and mugwort, Artemisia vulgaris L.; in herb gardens on the Cornell University campus, Ithaca, N.Y., it causes extensive foliar chlorosis on wormwood, Artemisia absinthium L., and southern- wood, 4. abrotanum L. (unpubl. observa- tions). The record of this tingid from oak, the only noncomposite host given in the literature, probably should be referred to the oak lace bug, C. arcuata (Say) (see Bailey, ibid.: 88). On 7 August 1986, I observed a large pop- ulation of the chrysanthemum lace bug damaging hedge bindweed, Calystegia se- pium (L.) R. Br. (formerly in Convolvulus) at Milan (Bradford Co.), Pennsylvania. All life stages of the tingid were present; eggs were inserted near the midrib and lateral veins on abaxial and adaxial surfaces. The hedge bindweed, growing on a fence above a mugwort plant infested with the lace bug, showed severe chlorosis and accumulation of cast skins and black excrement. Rather than a host shift, or the adding of a convolvulaceous plant to the diet of C. marmorata, my observations probably rep- resent adaptive behavior allowing the bug to cope with the apparent deterioration of an isolated mugwort plant. The hawthorn lace bug, C. cydoniae (Fitch), has been re- ported to temporarily colonize and injure roses growing near heavily damaged plants of a preferred rosaceous host, Cotoneaster sp. (Wheeler, 1981. Great Lakes Entomol. 14: 37-43). In the present case the adven- titious host, hedge bindweed, belongs to a different family (Convolvulaceae) and order (Polemoniales) from the typical hosts of C. marmorata; the Convolvulaceae and Com- positae, however, are members of the same subclass (Asteridae) (Cronquist, 1968. The evolution and classification of flowering plants. Houghton Mifflin, Boston, 396 pp.). For C. morrilli Osborn and Drake, another species of the genus that feeds on herba- ceous composites (Silverman and Goeden, 1979. Pan-Pac. Entomol. 55: 305-308), the only known noncomposite host also belongs to the Convolvulaceae: sweet potato, [po- moea batatas (L.) Lam. (Drake and Ruhoff, Ibids? 155): Adults and nymphs have been deposited in the insect collection of the Pennsylvania Department of Agriculture (PDA). A. G. Wheeler, Jr., Bureau of Plant In- dustry, Pennsylvania Department of Agri- culture, Harrisburg, Pennsylvania 17110. PROC. ENTOMOL. SOC. WASH. 89(1), 1987, pp. 201-203 Book REVIEW Coevolution of Parasitic Arthropods and Mammals, edited by Ke Chung Kim. Wi- ley-Interscience, a division of John Wiley & Sons, Inc., 605 Third Avenue, New York, NY 10158. xvi + 800 pp. 1985. $69.95/cloth. ISBN: 0 471-08546-4. This collection of essays marks the latest effort to summarize and analyze the increas- ingly daunting literature on host associa- tions of parasitic arthropods. As such, it continues a genre begun with the First Sym- posium on Host Specificity among Parasites of Vertebrates (Neuchatel, Switzerland, 1957) and recently elaborated by Adrian Marshall in his masterly survey, The Ecol- ogy of Ectoparasitic Insects (1981). The style of Kim’s book is immediately revealed in his list of contributors, among whom are several renowned arthropod systematists, each the doyen of his field: Emerson (Mal- lophaga), Fain (Astigmata), Hoogstraal (Ixodoidea), Radovsky (Mesostigmata), Traub (Siphonaptera). Himself an expert on Anoplura, Kim is the sole author of four chapters (1, 5, 7, 13) and co-author of two more (4, 10). The text is divided into four parts. Part One (chapters 1-3) is an introduction to the broad spectrum of evolutionary relation- ships between arthropods and mammals. Topics covered include parasite and host anatomy and morphology, reproductive cycles, population dynamics, dispersal, and geographic radiation—particularly as af- fected by continental drift. Part Two (chap- ters 4-8) examines the host associations, evolution, and zoogeography of ectopara- sitic Insecta, chiefly Phthiraptera and Si- phonaptera. Part Three (chapters 9-12) covers the Acari, with emphasis on pro-, meso-, and astigmatid mites as well as the Ixodoidea (ticks). Part Four (chapter 13) is an overview of the evolutionary pathways detailed in Parts Two and Three. Each chap- ter comes with its own list of references, which together run to 79 pages. Following the text are two remarkable appendices: A, an alphabetical list by family and genus of the world’s parasitic arthropods and their mammal hosts; and B, the reverse of the preceding, an alphabetical list of the orders and families of mammals and their arthro- pod parasites. The work concludes with a 56-page index to arthropods, mammals, and all subject headings. Most chapters contain numerous illustrations, the best being those of flea morphology (chapter 8) and the his- tory of continental movements (chapter 3). However, several figures have not repro- duced well, either because the originals were crudely executed (chapter 4) or excessively reduced (chapter 10). Students of a partic- ular group will also quickly note a number of minor typographical errors, as on page 664 where the amblyommine tick genus Aponomma is misspelled twice (as Apo- nemma and Aponema) in the same sen- tence. Such blemishes seldom impede un- derstanding. My criticism of this work stems from its title, which unduly stresses coevolution. As Dan Janzen makes clear in his short but engaging essay “Coevolution as a Process: What Parasites of Animals and Plants Do Not Have in Common,” this is not a col- lection of coevolutionary studies in the strict reciprocal sense advocated by many who contributed to Futuyma and Slatkin’s sem- inal synthesis Coevolution (1983). Rather, it is a review of parallelisms, especially those illustrative of Fahrenholz’s Rule and re- source tracking. In fact, Fain and Hyland refrain from even mentioning coevolution in their chapter, “Evolution of Astigmatid Mites on Mammals”; they prefer “‘parallel evolution.”” My concern over definitions may seem trivial, but it arises from a basic difference between Kim’s text and that of 202 Futuyma and Slatkin. The latter gathered authorities from a wide range of fields with the object of formulating coevolutionary concepts. Kim, on the other hand, has chief- ly collaborated with systematic specialists — taxonomists—each of whom has focused on a particular ectoparasite group in order to describe (as he sees it) evolution in that group. One result is that much of this book is unnecessarily Darlingtonian in tone: hundreds of pages are given over to lists of taxa, their hosts and distribution, often with minimal evolutionary follow-up. Another problem in working with spe- cialists is “expert opinion”: anything said is automatically ex cathedra. A striking ex- ample is Hoogstraal’s depiction of evolu- tion in the Ixodoidea (pp. 508-516). Bearing in mind that no pre-Eocene fossil ticks have ever been found (a few forms resembling extant species are known from amber), we read that ancestral ticks were eyeless para- sites of large, “glabrous” reptiles living communally during the late Paleozoic or early Mesozoic eras. The argasid line was represented by Argas and Ornithodoros “partially as we know them today,” but oth- er argasids “probably did not evolve until the Tertiary.” “‘Modern”’ ixodids (the Hae- maphysalinae, Ixodinae, and Rhipicepha- linae) evolved from spiderlike amblyom- mines and were “probably as large as the largest extant Amblyomma,” while xeroph- ilous Hyalomma “may have appeared later, close to the Cretaceous period of Mesozoic environmental stresses.” This entirely con- jectural scenario, capped by a regrettable dendrogram reminiscent of something by Ernst Haeckel, is based on the author’s ex- pert knowledge of tick morphology and host associations, but it does not address con- flicting evidence from other fields. For ex- ample, Hoogstraal believes that prostriate ixodids (i.e. the genus /xodes) are advanced because of their smaller size, shorter palps, streamlined morphology, and presumably recent radiation with the Rodentia. How- ever, he overlooks the rich chaetotaxy of PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Ixodes larvae, occasional mating off the host, and the absence in this genus of a cement feeding cone, all characters suggestive of the supposedly primitive argasids. Available karyotypes for Ixodes also are similar to those in the Argasidae, as are the systems of sex determination (XX-XY) and sper- matogenesis (males in both groups may re- main aphagous). How might these authors have injected greater objectivity into their arguments? The obvious answer 1s through cladistic (or phy- logenetic) analysis, the only repeatable method of biotic classification that enables its users to generate testable hypotheses of phylogeny. This is not to say that references to cladistics are missing from Kim’s book. The now familiar terms (apomorphy, ple- siomorphy, etc.) are there, and we are even treated to a small cladogram (p. 272) for the five genera of Echinophthiriidae (Ano- plura). For the most part, however, these are evolutionary “‘just-so” stories, products of traditional synthetic taxonomic proce- dures that are neither repeatable nor test- able and depend entirely on the opinions of experts. It follows that all the zoogeographic arguments would also have greatly benefited from application of modern vicariance methodology—the union of cladistics and Leon Croizat’s track analysis—which is blissfully free of any a priori assumptions concerning dispersal or centers of origin. Though thin on evolutionary analysis, this book is a veritable encyclopedia of ecto- parasitology, worthy of a prominent posi- tion in any entomologist’s library. Kim’s compilation is not likely to be superseded in our time, which should be comforting to those who must purchase it. But the flip side of literary immortality is the sad realization that contemporary society no longer seems interested in a sequel. As one painfully aware of this problem, Frank Radovsky deserves the last word (p. 496): “A pattern of ... evolution has emerged, and I have attempt- ed to interpret it here. However, forms that are significant in understanding this pattern VOLUME 89, NUMBER | continue to be discovered. Our inventory of these is at best sketchy; the gaps in knowl- edge of bionomics and basic host-parasite relationships are especially glaring. Follow- ing a period of considerable interest in the 1950s and early 1960s, there has been a decline in biological studies of vertebrate- associated [arthropods]. This important area of research should be revitalized.” 203 Richard G. Robbins, Department of Health and Human Services, Public Health Service, National Institutes of Health, Na- tional Institute of Allergy and Infectious Diseases, % Department of Entomology, Museum Support Center, Smithsonian In- stitution, Washington, D.C. 20560. PROC. ENTOMOL. SOC. WASH. 89(1), 1987, pp. 203-204 Book REVIEW The Pleasures of Entomology. Portraits of Insects and the People Who Study Them. By Howard E. Evans. Smithsonian Institution Press, Washington DC. 1985. 238 pp., illus. Price: $14.95, paperbound only. The title says it all—studying insects is a pleasure. Not only studying them, reading about the studies of others is a pleasure. That’s how Howard Evans, high expert on the Psammocharidae and other wasps at Colorado State University, feels about it. And he makes others feel the same, what with his low-key enthusiasm for the subject and his easy style of writing. He isn’t a ran- ter, nor does he want us to act as if we were entering a cathedral of science. Put simply, he simply likes our animals. He devotes 12 chapters to 12 different insects, things such as gypsy moth, love bug, killer bee, boll weevil, and some less no- torious beasts. Each life history portrait is “painted” in a most engaging way, as is the story of the scientist who worked out that life history. There are chapters on garden insects, past entomologists, and himself. These are not dry accounts of insects; they are accounts of the positive or negative im- pacts that insects have made on the public and on the people who studied them. Evans knew or knows many scientists who worked out life histories or complicated control measures. Even when he discusses those he didn’t know, usually long before his time, he does it with appreciation and freshness. He stresses the personal involvement of the entomologist with the insect, putting a hu- man touch in the science of entomology; he loves the lover (amateur), sometimes even more than the professional, and rues his passing. Because many entomologists came accidentally to their calling and suffered and enjoyed much along the way, the evolution of each is almost as interesting as the life history of the insect. You don’t have to be a good bug to be interesting to Howard Evans, for he some- times welcomes pests into his home garden for the sheer pleasure of watching them. He watches for what he has read and looks for something new. A few extra plants, so the crop yield is still adequate, is a small price to pay for a pest’s presence. He even wishes he could attract a few more that sound in- teresting, like the harlequin cabbage bug or imbricated snout beetle. And as for control, he admits to the pleasure of popping and oozing tobacco hornworms between his toes, just as he did as a boy. Evans often implies or states emphati- cally that our society is too materialistic, 204 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON that we are too much concerned with the GNP (gross national product) when we should be concerned with the PGN (pro- founder grasp of nature). Surely this book and others like it will help toward a better balance of our priorities. I have recently become interested in the history of entomology and the Americans who made it, so I was happy to read his chapters devoted to Abbot, Say, Peck, Har- ris, Cockerell, Williams, and Dietrich. As in all history, there is no set formula for success. Different personalities, such as Cockerell and Williams, with very different methods of attack can make tremendous contributions. Evans tells us that the only requirements for success in our field are great amounts of inquisitiveness, patience, and love of insects. Not only are insects interesting to ento- mologists, they provide a source of liveli- hood. Therefore, for all the information that Drosophila has supplied and for the careers that it has made possible, Evans awarded those fruitflies the Evans Prize: a bunch of bananas. I'll turn that line back on him, so to speak; for giving pleasure with this book, I award Howard E. Evans the Spilman Prize: a sharp nib, a full jar of ink, and an ento- mological author’s rating of 9.0 (in a scale of 1-10). (He’s in good company; I once awarded Shakespeare a 9.9 and the author of the Bible a 10). It is my duty as reviewer to find an error. On page 164, the Colorado Potato Beetle reached Illinois by 1864, not 1964. Thus have I justified receiving this book free. Even so, I feel guilty; now that I have read it, I almost want to pay for it. It’s that good. T. J. Spilman, Systematic Entomology Laboratory, BBII, Agricultural Research Service, USDA, % National Museum of Natural History, NHB-168, Washington DiE220560) PUBLICATIONS FOR SALE BY THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MISCELLANEOUS PUBLICATIONS Cynipid Galls of the Eastern United States, by Lewis H. Weld nieces ce tececeeneceeeenentneneneltnnen Geena Galls of the Southwest)iby; Lewis: Weld.0 oo) BOLEEDADEES/ ON CYMIDIC: Calis! Eine Ae eC Dope BONES RU MU Pe PRC Identification of Alaskan Black Fly Larvae, by Kathryn M. Somme rman oiie..oocc.csccscsssssessneessesveeeseeeesesssecseeecesese Unusual Scalp Dermatitis in Humans Caused by the Mite Dermatophagoides, by Jay R. DEANE Ts) be Pe SeLEMLOS ADORED RATE USSR EAT ES ELV ed Bl Pee EU PLO Ee DE EAE EEE RY A Short History of the Entomological Society of Washington, by Ashley B. Gurey -0.2.0 Pictorial Key to Species of the Genus Anastrepha (Diptera: Tephritidae), by George C. SEDATE EB Sy TLE MN A PN TL Taxonomic Studies on Fruit Flies of the Genus Urophora (Diptera: Tephritidae), by George C. SERPS RUNS OR TLE i ed PAA AR A LR WN A LS AMS MOMENTA RO NE 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. Pap hoy ong 0) Uy: RMA AS DRT LE Dat SMS 0 A ON ANNA ee SN NP SL AY RN IEA YA AE SR No. 3. The Nearctic Leafhoppers, a Generic Classification and Check List, by Paul Wilson Oman. DISS) 0) ey COL ONIN N A RY STAD A 8 OO SR ON ee OE PR NO A A A A Ree EN Si 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. OR er Urea Bowes Lata nie Moe te ECR Deyo LO TOPE a ek OU ds RV ea ay eb Ee BL A ee No. 6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi akandsi eo SOL ps (LOGO tue wa WER BC be eas UP ee EGR Se REO NGO ae MOR De Ree URE A ge 1 No. 7. Ant Larvae: Review and Synthesis, by George C. Wheeler and Jeanette Wheeler. 108 pp. MOSS MMe SONS CaN LA Td a SEO Ay SCAM R SNA SSUR ORT OF LI Oak CY Id DAL UN LL Se 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. 1979.0 No. 9. The Flower Flies of the West Indies (Diptera: Syrphidae), by F. Christian Thompson. 200 TDPsg UO Bal eee ANY ERE ROC PRUNE Py Be TUR PY CO I BPC PAU Ie PCN, Sk WP De BR CNR AN 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. 1982.00. No. 11. A Systematic Study of the Japanese Chloropidae (Diptera), by Kenkichi Kanmiya. 370 pp. SDS hee We eer LER ESP SUI ae me DY BENE! EA OURS Were ee RN RecN WCB BB No. 12. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff. PSP TOR Ee yet ee OU Note MPG ENA RDN Ey AU es BD WG PE WV BED UL BN Re MAIN 2.00 15.00 11.00 12.00 10.00 11.00 18.00 5.00 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) PETERS, T. M.—A new Nearctic Dixa (Diptera: Dixidae) from Pennsylvania .............. 137i; Wt SCHAUFF, M. E.—Taxonomy and identification of the egg parasites (Hymenoptera: Platygas- tridae, Trichogrammatidae, Mymaridae, and Eulophidae) of citrus weevils (Coleoptera: Curculionidae)! s/h. LAREN Ue AT OE aD A ST Sh AO eS eR a 31 | i! SCHWARTZ, M. D. and G. M. STONEDAHL—Oaxacacoris, a new plant bug genus and three _ 4K new species of Orthotylini from Mexico (Heteroptera: Miridae) ....................-.. ; 5 SHARKEY, M. J.—Agathis thompsoni n. sp., a Nearctic species of Agathidinae (Hymenoptera: Braconidae) parasitic on Greya subalba (Braun) (Lepidoptera: Incurvariidae) ........... 47 SMITH, D. R. and E. M. BARROWS-—Sawflies (Hymenoptera: Symphyta) in urban environ- ments iin the’ Washington :@) aned hiya Dae POU eA ea i ee SOP a a 147 SPANGLER, P. J. and R. C. FROESCHNER — Distributional data, illustrations, and habitat of , the South American water-strider Microvelia ayacuchana (Hemiptera: Veliidae) ......... 167 STARK, B. P. and B. C. KONDRATIEFF—A new species of Peltoperla from eastern North American (Blecoptera: Peltoperlidae) 2:4, 45. 4a a A eR eee 141 TURNER, C. E., R. W. PEMBERTON, and S. S. ROSENTHAL—Host range and new host records for the plume moth Platyptilia carduidactyla (Lepidoptera: Pterophoridae) from California thisties (Asteraceae) | 8p 4 Ys WO oe UY ed a 132 WALTZ, R. D. and W. P. MCCAFFERTY —New genera of Baetidae (Ephemeroptera) from ATRIOS D QL W SA Ree MOLI RMN ak A eh A RCE Odd 95 WALTZ, R. D. and W. P. MCCAFFERTY —Generic revision of Cloeodes and description of two! new genera (Ephemeroptera: Baetidae)ii, 4.1) ki) LS IA eo ee Th Maas ay 177 WHARTON, R. A.—Changes in nomenclature and classification of some opiine Braconidae (CElymenoptera) eee SV eae EE Te NEON a Re or RAE det aed ok Tae le 61 WOODLEY, N. E.—The Afrotropical pachygastrine genera Ashantina Kertész and Meristo- meringina James, with two new generic synonyms (Diptera: Stratiomyidae) ............ 103 WOOLLEY, J. B. and H. W. BROWNING—Morphometric analysis of uniparental Aphytis reared from chaff scale, Parlatoria pergandii Comstock, on Texas citrus (Hymenoptera: Aphelinidae; Homoptera: (Diaspididae) iy) eS ee OG Ne A RN PE Ti NOTES HALSTEAD, J. A.—Brachymeria discretoidea, a new junior synonym of Brachymeria discreta (Hymenoptera:/Chaleididae)} re 2 Pas RAS a Pg ag es Eg ae 131 WHEELER, A. G., JR.— Hedge bindweed, Calystegia sepium (Convolvulaceae), an adventitious host of the chrysanthemum lace bug, Corythucha marmorata (Heteroptera: Tingidae) ... 200 BOOK REVIEWS ROBBINS, R. G.— Coevolution of Parasitic Arthropods and Mammals ..................... 201 SPILMAN, T. J.— The Pleasures of Entomology. Portraits of Insects and the People Who Study PREM MAO A SON TE A RE SERN AL PTS Se Sn Teh Me VL 203 Iw fs -* VOL. 89 APRIL 1987 NO. 2 (ISSN 0013-8797) | PROCEEDINGS. of the ENTOMOLOGICAL SOCIETY ot WASHINGTON PUBLISHED QUARTERLY CONTENTS if ADAMSKI, D. and R. L. BROWN —A new Nearctic G/yphidocera with descriptions of all stages } fi (epidopteras Blastobasidae: Syminocinae) yk ae eo Te TOO Ne 329 H BURROWS, W. L.—A new species of Ame/etus (Ephemeroptera: Siphlonuridae) from eastern INOTEORASUELICN) CE PRL MLE UR Oya wee REC ES tC Re Leh ae LR ale Bed eae ty Lav 284 DAVIS, D. R.—Neotropical Tineidae, IV: Three new Acrolophus species from Cuba and the rediscovery of Acrolophus niveipunctatus Walsingham (Lepidoptera) ................... 275 DONNELLY, T. W.—Structural variation of Ophiogomphus mainensis: description of a new : subspecies and relationship to sibling species (Odonata: Gomphidae) .................. 205 | : FENNAH, R. G.—A new subfamily of Nogodinidae (Homoptera: Fulgoroidea) with the de- | SCHIPUODNGE a HE WPSDCCIES OL GASEFINIIG ! Nie PU bares all hid Uk a Leto ed ie SLO ene) EE 363 ian GOEDEN, R. D.—Host-plant relations of native Urophora spp. (Diptera: Tephritidae) in south- My STV CCA LONNIE URUK PRCT C EN eee ALR Me ideey EY LR TELE PARE eC ds Dnt sie APE fal Nee ey 269 - GRISSELL, E. E. and L. DESANTIS—A new species of Erixestus (Hymenoptera: Pteromalidae), V, an egg parasitoid of Calligrapha polyspila (Coleoptera: Chrysomelidae) in Argentina .... 264 : re | HOEBEKE, E. R., Q. D. WHEELER, and R. L. GILBERTSON—Second Eucinetidae-Conio- phoraceae association (Coleoptera: Basidiomycetes), with notes on the biology of Eucinetus ty oviformis LeConte (Eucinetidae) and on two species of Endomychidae ................. 215 | i _ HURYN, A. D.—A new species of Notiphila (Notiphila) (Diptera: Ephyridae) from Ohio .... 322 iN KROMBEIN, K. V.—Synonymic notes on the Bethylidae described by V. de Motschulsky : t Cig mlenootara Acetate Ply We ate PU ULM Pa tee ehh Sm mean Ae) DR DA Ue an th 356 | __ MASON, W. R. M.—Discovery of female Apozyx (Hymenoptera: Apozygidae) and comments ig Olds LAR ONOMMC POSIMON (IU MH GAL GLB Nici OL NUN Fe a Darah More Uh UT 226 aN i MASON, W. R. M.— Vadum, a new genus of Nearctic Braconidae (Hymenoptera) .......... 325 _ MILLER, J. S.—A revision of the genus Phryganidia Packard, with description of a new species h Pe ApIR tae VAN SL HME RE EY Mi Bh Ey Ber Me 303 | (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ORGANIZED MARCH 12, 1884 OFFICERS FOR 1987 THOMAS E. WALLENMAIER, President MICHAEL J. RAupp, Program Chairman F. EUGENE Woop, President-Elect GEOFFREY B. WHITE, Membership Chairman PAUL M. MArsH, Recording Secretary VicTOR L. BLACKBURN, Custodian RICHARD G. RosBINs, Corresponding Secretary MANYA B. STOETZEL, Delegate, Wash. Acad. Sci. NORMAN E. WoobDLey, 7reasurer ( _ RAYMOND J. GAGNE, Editor M Publications Committee DAVID R. SMITH THEODORE J. SPILMAN GEORGE C. STEYSKAL Associate Editor ROBERT W. CARLSON Honorary President C. F. W. MUESEBECK Honorary Members FREDERICK W. Poos ASHLEY B. GURNEY THEODORE L. BISSELL All correspondence concerning Society business should be mailed to the appropriate officer at the following a‘ address: Entomological Society of Washington, c/o Department of Entomology, NHB 168, Smithsonian Insti- Y tution, Washington, D.C. 20560. if regularly in the Proceedings. oh MEMBERSHIP.— Members shall be persons who have demonstrated interest in the science of entomoligla Annual dues for members are $20.00 (U.S. currency) of which $18.00 is for a subscription to the Proceedings 1 of the Entomological Society of Washington for one year. PROCEEDINGS.-— Published quarterly beginning with January by the Society at Washington, D.C. Members 7 | in good standing receive the Proceedings of the Entomological Society of Washington. Nonmember subscriptions — are $35.00 per year, domestic, and $40.00 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. 730 of the July 1984 issue for information regarding preparation of manuscripts. STATEMENT OF OWNERSHIP iy, Title of Publication: Proceedings of the Entomological Society of Washington. ; Frequency of Issue: Quarterly (January, April, July, October). i) Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of Washeit ington, c/o Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash- ington, D.C. 20560. Editor: Raymond J. Gagné, Systematic Entomology Laboratory, c/o U.S. National Museum NHB 168, Wash- ington, D.C. 20560. Managing Editor and Known Bondholders or other Security Holders: none. ; ( \G This issue was mailed 16 April 1987 \ iy Second Class Postage Paid at Washington, D.C. and additional mailing office. y + rae PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA Se ee PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 205-214 STRUCTURAL VARIATION OF OPHIOGOMPHUS MAINENSIS: DESCRIPTION OF A NEW SUBSPECIES AND RELATIONSHIP TO SIBLING SPECIES (ODONATA: GOMPHIDAE) THOMAS W. DONNELLY 2091 Partridge Lane, Binghamton, New York 13903. Abstract. —Ophiogomphus mainensis mainensis Packard, new status, consists of a large, stable population (New England and adjacent parts of New Jersey, Pennsylvania, New York, Quebec, and New Brunswick) and a disjunct population in the high mountains of North and South Carolina. An allopatric population in central Pennsylvania and West Virginia is named as a new subspecies fastigiatus. Structurally intermediate specimens from the vicinity of the boundary of the two ranges have been found in Pennsylvania and New York. Ophiogomphus incurvatus Carle may have been derived from O. m. fastigiatus, with the subspecies O. i. alleghaniensis Carle the bridging taxon. Ophiogomphus acumi- natus Carle occurs southwest of these two species; its relation to the other two species is uncertain. The geographic pattern of variation of O. mainensis sspp. is unusual for odo- nates of the eastern United States, and is reminiscent of “leapfrog” variation of Andean birds. The gomphid Ophiogomphus mainensis was described by Packard (1863) from a fe- male collected in Maine. A male was de- termined as mainensis by Hagen (Selys, 1878; Howe, 1918); however, Howe’s (1918) figure shows that it is not this species, and Garman (1927) identified it as carolus Needham. Needham (1897) reared a male of mainensis but described it as O. johan- nus. The two species johannus and mainen- sis were considered to be distinct until Gar- man (1927) established the synonymy. The name Ophiogomphus incurvatus was proposed by Carle (1982) to replace the in- correctly named QO. carolinus Hagen, a species of the piedmont of Georgia to Mary- land. Carle also described the subspecies O. i. alleghaniensis, which occurs to the west of the nominate subspecies. Ophiogomphus acuminatus was de- scribed by Carle (1981) from specimens from Tennessee. O. bouchardi, which was de- scribed by Louton (1982), is a synonym. Although placed by Carle with O. edmundo Needham, it seems to be closer to O. mai- nensis s. lat. and incurvatus Ss. lat. The present study began with the recog- nition of a distinct taxon occurring in cen- tral and western Pennsylvania south to high elevations in West Virginia (Fig. 1). Al- though this taxon has some resemblance to O. i. alleghaniensis, it is closely related to mainensis. Preliminary studies suggested that it might merit specific status, but a few specimens of this taxon and typical mai- nensis close to their geographic boundary in northeastern Pennsylvania and southern New York show intermediate characters. Thus, the new taxon is considered a sub- species of mainensis. Abbreviations for collectors’ names in the descriptions are: TD = T. Donnelly; CS = C. Shiffer; HW = H. B. White III; FC = F. Carle; SD = S. Dunkle; CU = Cornell Univ.; 206 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON a ’ mM@® M. MAINENSIS O M. FASTIGIATUS 4 |. INCURVATUS = A i. ALLEGHANIENSIS A © ACUMINATUS Fig. 1. Map showing distribution of Ophiogomphus m. mainensis (U.S. specimens examined for this study only; ““m” for Canadian localities reported in Walker, 1958); O. m. fastigiatus; O. i. incurvatus (localities from Carle, 1982 and from my collection); O. i. alleghaniensis (localities from Carle, 1982, with additional localities from specimens in J. Louton Collection); and O. acuminatus. Arrows show localities with structurally inter- mediate specimens. VOLUME 89, NUMBER 2 OF = O.S. Flint, Jr., U.S. Nat. Mus.; SB = S. W. Bromley, U.S. Nat. Mus.; ED = E. M. Davis, Mus. Comp. Zool.; EW = E. B. Williamson, Univ. Mich. Mus. Zool. Ophiogomphus mainensis fastigiatus Donnelly, NEw SUBSPECIES Figs. 2-5 Ophiogomphus mainensis, Ahrens et al., 1968: 107. Butler Co. specimen included here. Ophiogomphus mainensis, Beatty et al., 1969: 131. In part this subspecies. Holotype male.— Head: pale yellow green, very dark brown as follows: thin marginal line on labrum, vertex except for a pale sub- basal medial spot. Thorax: proepisternum very dark brown with pale lateral spots, proepimeron yellow brown. Pterothorax yellow green, with a very dark brown mid-dorsal line, which does not meet the anterior margin. Mesepisternum with a dark curved line extending to % the distance to rear and not touching meso- pleural suture. Mesopleural and metapleu- ral sutures and margins of mesepimeron and metepisternum dark brown. Legs black ex- cept for obscure pale of coxae and interior surface of profemora. Abdomen: Each segment mainly black, green as follows: sides of 1 and 2; basal- lateral and medial-lateral spots on 3 to 7, decreasing in size rearwards; fused lateral spots on 8 and 9; medial-basal lines on | to 8, thin and short on 3 to 8; tiny middorsal spot on 9; 10 with postero-lateral line and small posterior middorsal spot. Appendages brown, superiors indistinguishable from m. mainensis, in lateral view each appendage straight, tapering, divergent in dorsal view (Fig. 5, no. 11), sub-parallel sided in lateral view with prominent, small posteriorly di- rected apical point. Inferior appendage in lateral view with lateral spine low and less prominent than apical spine, located about Y) the distance from base (Fig. 2, no. 9); in 207 ventral view (Fig. 3, no. 9) the appendage is distinctly wedge-shaped, in sharp contrast to typical m. mainensis, in which the ap- pendage is obtusely truncated (Fig. 3, nos. 1 to 4). Allotype female.— Coloration as in male, with pale colors slightly more extensive. The occiput with paired, anteriorly directed spines as in m. mainensis. The vulvar lam- ina resembles that of m. mainensis; in lat- eral view the tips are elevated and then de- flected caudally, forming a sigmoid bend (Fig. 5, no. 10). A reliable distinction be- tween females of m. fastigiatus and m. mai- nensis has not been found. Types.— Holotype 6, PENNSYLVANIA: Sullivan Co., Loyalsock Cr. 3 mi. N. of LaPorte, 4 July 1983 (TD). Allotype 2, same loc., 19 June 1983 (TD). Paratypes: PENN- SYLVANIA: same loc. as holotype: 19 June and 4 July, 5 6 2 2 1983 (TD and CS); Clin- ton Co., Kettle Cr., 2 mi. E. of Hammersley Fork, 21 June 1969, 21 June 1975, 22 June 1977, 25 June 1983, 5 8 4 2 (CS); Centre Co., Black Moshannon Creek nr. Black Moshannon Dam, 4 Aug. 1983, 1 é (CS); Somerset Co., Rockwood, 29 June 1900, 1 6 (EW); WEST VIRGINIA: Nicholas Co., Cranberry R. 5 mi. N. of Richwood (2100’), 2) June 1969;,.1 ¢ (1D); Randolph Ga., Shaver Fork of Cheat R. at highway 250 (3600’), 23 June 1973, 1 6 (HW). The ho- lotype and allotype are deposited in the Florida State Collection of Arthropods. Dimensions. — The holotype male has ab- domen and appendages 34 mm long, and hind wing 26 mm long. The allotype female has these dimensions 32 and 27 mm. The total group examined (14 males, 7 females) has these dimensions 32.7 mm (.85) and 25.65 (.5) mm for the males and 31.8 (.4) and 27.1 (.7) for the females. The figure in parentheses is standard deviation. Variations in the type series.— The color patterns show little variation. In about half the specimens the posterior end of the mes- episternal dark stripe is free, and in half it 208 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Profiles of appendages of male Ophiogomphus m. mainensis, 1-5, and O. m. fastigiatus, 6-12. Specimens 5 (mainensis) and 6-8 (fastigiatus) show intermediate characteristics. Localities are 1 = Essex Co., Vt.; 2 = Perry Co., Pa.; 3 = Pike Co., Pa.; 4 = Haywood Co., N.C.; 5 = Sullivan Co., N.Y.; 6 and 7 = Sullivan Co., Pa.; 8 = Centre Co., Pa.; 9 = Sullivan Co., Pa. (holotype); 10 = Somerset Co., Pa.; 11 = Nicholas Co., W. Va.; 12 = Randolph Co., W. Va. is fused with the humeral stripe. The three pale spots on abdominal segment 10 are fused in many specimens, forming a pos- terior band. The appendages show some variation, as seen in Figs. 2, 3, and 4. In three specimens (two from Sullivan Co., Pennsylvania, and one from Centre Co.., Pennsylvania; Figs. 2 and 3, nos. 6 to 8) the lateral spine on the inferior appendage is set posterior to the midpoint, somewhat as in m. mainensis. Although these specimens are more like m. fastigiatus than m. mainensis in the ventral view of the appendages (Fig. 3, nos. 6 to 8) and in the relatively small medial spine of the inferior appendage, their intermediate character is significant. Remarks.—The subspecies m. fastigiatus is distinguishable from the nominate sub- species by the form of the inferior append- age, which has a low, blunt lateral spine set at about the midpoint of the appendage (Fig. 2, nos. 9 to 12). In m. mainensis this ap- pendage is variable, but always has a very prominent, pointed lateral spine located distinctly caudal to the mid point of the inferior appendage (Fig. 2, nos. | to 4). In VOLUME 89, NUMBER 2 10 Fig. 3. Ventral views of male appendages of same specimens shown in Fig. 2. ventral view (Fig. 3, nos. 9 to 12) the wedge of the inferior appendage is distinctive (the name fastigiatus refers to a gable). Ophiogomphus mainensis mainensis Packard, NEw STaTus Figs. 2-4 Ophiogomphus mainensis, Packard, 1863. Holotype @. Ophiogomphus johannus, Needham, 1897. Holotype ¢. Ophiogomphus johannus, Howe, 1918. Ophiogomphus mainensis, Garman, 1927. Established synonymy of mainensis and Johannus. Descr. é and 2, figs. Ophiogomphus mainensis, Walker, 1958. Descr. 6 and 9, figs. Ophiogomphus mainensis, Carle 1982. De- scriptive notes, figs. The holotype is a female, which, as has been the case with many other gomphids, has resulted in continuing confusion. The male of the nominate subspecies has the thorax marked as follows: yellow green, with a dark brown mid-dorsal stripe ending pos- terior to anterior margin; dark brown stripes on mesopleural (humeral) and metapleural sutures, the latter covering the anterior half of suture; dark curved stripe on mesepi- sternum adjacent to mesopleural suture and 210 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Inclined views of male appendages of O. m. mainensis, 1-2, and O. m. fastigiatus, 3-4. Specimen 3 shows intermediate characteristics. Localities are 1 = Perry Co., Pa.; 2 = Pike Co., Pa.; 3 = Sullivan Co., Pa.; 4 = Sullivan Co., Pa. (holotype). extending 73 distance towards wing bases. The distinctive inferior appendage is shown in Figs. 2, 3, and 4. Needham’s holotype of O. johannus was examined and found to be m. mainensis. However, the holotype is a reared male pre- served in alcohol, and the specimen has sub- sequently turned to a jellied mass, of which the sclerotized abdominal appendages are among the few portions recognizable as parts of an insect. Material examined.— MAINE: Aroos- took Co., 18 mi. W. Ashland, 19 July 1982, 1 6(TD); Washington Co., E. Machias R. 3 mi. N. of Wesley, 11 July 1982, 2 6 (TD); Penobscot Co., Seboeis R. west of Shin Pond, 28 July 1959, 2 6 1 2 (TD). MAS- SACHUSETTS: Franklin Co., West Haw- ley, 23 June 1971, 1 ¢(HW); Middlesex Co., Ashby, 9 July 1939, 1 (ED). NEW YORK: Hamilton Co., nr. L. Durant, 3 July 1980, 1 6 1 2@(HW); Herkimer Co., Wilmurt, n.d., 1 6 (holotype of O. johannus) (CU); Dela- ware Co., Beaver Kill, nr. Roscoe, 31 July 1967, 1 (TD); Sullivan Co., Beaver Kill at Lewbeach, 31 July 1967, 2 6 (somewhat intermediate to fastigiatus) (TD). VER- MONT: Essex Co., nr. Island Pond, 14—15 July 1982, 5 6 3 2 (SD); Brunswick, 7 July 1985, 1 6 1 2 (HW); Outlet Dennis Pond, 24 June 1983, 2 6 (FC); Caledonia Co., Pea- cham; 27 June 1983; 2.6 (O3NEW HAMPSHIRE: Hillsborough Co., Wilton, 22 June 1969, 1 (HW). CONNECTICUT: Fairfield Co., Stamford, 16 June 1940, 1 3 (SB). NEW JERSEY: Morris Co., Hackle- barney St. Pk., 21 June 1959, 1 6 (TD). PENNSYLVANIA: Pike Co., and Wayne VOLUME 89, NUMBER 2 Figs 5: 12, and O. acuminatus, 5-8. 1 and 5 are lateral, 2 and 6 ventral, and 3, 7, and 11 are ventral views of male appendages. 9 and 10 are inclined views of female vulvar laminae. 4, 8, and 12 are the vesicle of the penis. Co., nr. Greentown, 4-5 July 1985, 2 (TD); Perry Co., nr. New Bloomfield, 23 June 1965, 3-10 July 1966, 3 June 1975,441¢2 (CS). NORTH CAROLINA: Haywood Co., nr. Sunburst, 15 June 1984, 2 6 (TD and CS); Yancey Co., nr. Busick, 11 June 1958, 6 6 (TD); Avery Co., nr. Minneapolis, 13 June 1958, 23(TD). SOUTH CAROLINA: Oconee Co., Burrell’s Ford, Chattooga R.., 19 May 1970, 1 ¢ (OF). Variations in Ophiogomphus m. mainen- sis. —Specimens from three disjunct popu- lations were examined: a northern, extend- ing from Maine to New Jersey and northeastern Pennsylvania (26 males, 6 fe- males), a small group from Perry Co., Penn- sylvania (4 males, 1 female), and a group from western North Carolina and South Carolina (11 males). Sizes of the three pop- ulations are virtually the same, and I de- Structural details of Ophiogomphus incurvatus alleghaniensis, 1-4, 9, O. mainensis fastigiatus, 10- tected no distinction in color pattern nor structure between the northern and south- ern populations. The Perry Co. specimens were slightly larger but identical in color- ation. The mean dimensions for males of the three groups of m. mainensis are as fol- lows: northern group abdomens 32.4 (.6) mm and hind wings 25.7 (.6) mm; for the southern group 33.3 (.9) and 25.9 (.45) mm; for Perry Co., Pennsylvania 34.25 (.3) and 26:255(@3) mm, Remarks. — The two subspecies are indis- tinguishable in color pattern, and both show parallel variations in the mesepisternal dark stripe and pale color of segment 10. The sizes are very similar, except for the slightly larger Perry Co. population of m. mainen- SIS. The inferior appendages show some vari- ation in the size of the lateral spine, but in 212 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON all specimens this spine is prominent, point- ed, and located posterior to the mid point of the appendage. In ventral view the ap- pendage varies from a nearly oblique ter- mination to a more acute wedge shape. Two specimens from Sullivan Co., New York (Figs. 2, 3, no. 5) are extreme in this regard and appear to vary in the direction of m. fastigiatus, while still retaining a predomi- nantly m. mainensis appendage. In summary, the two subspecies m. mai- nensis and m. fastigiatus are allopatric with a tendency for structurally intermediate specimens to occur near the boundary be- tween their ranges. The disjunct southern population is separated from West Virginia m. fastigiatus by a gap occupied by Ophio- gomphus incurvatus alleghaniensis. DISTINCTION AMONG TAXA OF THE OPHIOGOMPHUS MAINENSIS GROUP The Ophiogomphus mainensis group contains three species (mainensis, incurva- tus, and acuminatus), two of which have subspecies. Males of the three species are most readily identified by the form of the superior appendage (cerci), as shown in Fig. 5. Ophiogomphus mainensis and incurvatus (Fig. 5, nos. 1-4) are distinguished by the pale femora, and rounded, blunt-tipped superior appendages of incurvatus. The inferior appendage viewed ventrally is wed- ge-shaped as in m. fastigiatus but has a prominent lateral spine as in m. mainensis. The abdomen is paler in incurvatus than in mainensis. The vesicle of the penis is black and broader in mainensis and narrower and centrally pale in incurvatus. The two sub- species of imcurvatus have been character- ized by Carle (1982). The similarity in the inferior appendages of i. alleghaniensis and m. fastigiatus is noteworthy and might have a bearing on their relationship. Ophiogom- phus acuminatus is a lesser known species found in Tennessee. The male is readily dis- tinguished by its broad superior appendage with a distinctively pointed tip (Fig. 5, nos. 5-7). The vesicle of the penis has narrower paired cylindrical processes than either in- curvatus Or mainensis. Females of mainensis and incurvatus are best distinguished by the paler colors of the latter, as discussed by Carle (1982). The vul- var lamina of i. alleghaniensis (Fig. 5, no. 10) is similar to that of m. fastigiatus (Fig. 5, no. 9) but appears on the basis of one specimen to be distinguishable by its tip which is deflected ventrally rather than sig- moidally ventrally and caudally. However, this distinction may be a poor one; this sclerite is subject to both in vivo and post mortem changes in shape. The female of acuminatus has not been described. Ophiogomphus m. mainensis and m. fas- tigiatus are allopatric and are distinguished from each other only by the form of the male inferior appendages (epiproct). Although this appendage shows some variability through- out both populations, all specimens exam- ined are referrable to one or the other sub- species by size of the lateral spine of the inferior appendage and by shape of this ap- pendage in ventral view, as discussed above and shown in Figs. 2, 3 and 4. The existence in the boundary area between the subspecies of some specimens with appendages of in- termediate shape shows that genetic isola- tion has not been achieved and reinforces the conclusion that the two taxa are of sub- specific rank. THE SYSTEMATIC RELATIONSHIPS AND PUTATIVE DERIVATION OF THE TAXA IN THE OPHIOGOMPHUS MAINENSIS GROUP Ophiogomphus mainensis occurs from Quebec and New Brunswick southward in the high Appalachian Mountains to South Carolina, a distribution shared with many other Odonata. However, the subspecies mainensis occurs as two disjunct popula- tions: the northern is widespread in New England, extending southward to Harris- burg at high elevations. The southern oc- cupies the high mountains of western North and South Carolina. The subspecies fasti- VOLUME 89, NUMBER 2 giatus occupies much of the intervening area, occurring from Pennsylvania south to the New River valley, which cuts through the Appalachians in southern West Virginia and southwestern Virginia. I suggest that m. mainensis 1s the ancestral form and origi- nally had a continuous distribution throughout the entire range. This type of distribution is well known among Andean birds and has been called “leapfrog varia- tion” by Remsen (1984), who suggested that the very young tectonics of the Andes might provide opportunities for restriction in gene flows, with the consequent appearance of derivative forms in the center of a once- continuous range. The tentative explanation offered here be- gins with late Cenozoic uplift of the Ap- palachian mountains which was concen- trated in the Carolinas along the Cape Fear arch. The uplift and accompanying west- ward tilting caused major tributaries of the Mississippi River system (Tennessee, French Broad, Holston, and New rivers) to penetrate the mountains and capture former eastward drainages. Continued develop- ment of these river systems left the Appa- lachian chain divided into segments. The New River created the broadest low-ele- vation division of the mountain chain. Sub- sequent to Pleistocene glaciation, odonate species in a putative southern Appalachian refugium spread northward along the mountain chain. Continued amelioration of the climate drove many species into higher areas, and the gaps created by cross-cutting river systems left some of these populations genetically isolated. The originally contin- uous population of Ophiogomphus mainen- sis was broadly divided by the New River gap. One possible result was the appearance of a relatively low-elevation form (O. mai- nensis fastigiatus) in this gap. Further cli- matic amelioration drove the boundary be- tween mainensis and fastigiatus further north to its present position, where the two subspecies face each other across a relatively short boundary region. 213 At a still later time, a new form, O. in- curvatus, supplanted fastigiatus at lower el- evations in this gap. It subsequently divided into two forms, i. incurvatus and i. alle- ghaniensis, across the drainage divide be- tween the New and Roanoke River systems in southwestern Virgina. At present 7. in- curvatus 1s confined to Atlantic drainages, and 7. alleghaniensis occupies Gulfand Mis- sissippi drainages. The origin of acuminatus remains elusive, but a derivation from in- curvatus 1s plausible. The similarity be- tween the inferior appendages of m. fasti- giatus and i. alleghaniensis is consistent with these taxa being annectant forms between the two species but is not compelling. Other schemes might be proposed; a final reso- lution of this question would best be ap- proached through an assessment of protein similarities. ACKNOWLEDGMENTS I am most grateful for the loan or gift of specimens by O. S. Flint, Jr., J. Louton, H. B. White III, S. Dunkle, F. Carle, C. Shiffer, and L. K. Gloyd. I am further indebted to Frank Carle for extended discussions on speciation in Ophiogomphus and related genera and to J. Louton, S. Dunkle, R. Gar- rison, F. Carle, and C. Shiffer for stimulating criticism of this paper. LITERATURE CITED Ahrens, C., G. H. Beatty, and A. F. Beatty. 1968. A survey of the Odonata of western Pennsylvania. Proc. Pa. Acad. Sci. 42: 103-109. Beatty, G. H., A. F. Beatty, and C. Shiffer. 1969. A survey of the Odonata of central Pennsylvania. Proc. Pa. Acad. Sci. 43: 127-136. Carle, F. C. 1981. A new species of Ophiogomphus from eastern North America, with a key to the regional species (Anisoptera: Gomphidae). Odon- atologica 10: 271-278. Carle, F. L. 1982. Ophiogomphus incurvatus: A new name for Ophiogomphus carolinus Hagen (Odo- nata: Gomphidae). Ann. Entomol. Soc. Am. 75: 335-339. Garman, P. 1927. The Odonata or Dragonflies of Connecticut. Conn. State Geol. Nat. Hist. Surv. Bull. 39. 331 pp. Howe, R. H. Jr. 1918. Manual of the Odonata of 214 New England, part III. Mem. Thoreau Mus. Nat. Hist: II, pp. 25-40. Louton, J. A. 1982. A new species of Ophiogomphus (Insecta: Odonata: Gomphidae) from the western highland rim of Tennessee. Proc. Biol. Soc. Wash. 95: 198-202. Needham, J. G. 1897. Preliminary studies on North American Gomphinae. Can. Entomol. 29: 181- 186. Packard, A. S. 1863. Jn Walsh, B. D., ed., Obser- vations on Certain N.A. Neuroptera. Proc. Ento- mol Soc. Phila. 2: 167-272. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Remsen, J. V. Jr. 1984. High incidence of “leapfrog” pattern of geographic variation in Andean birds: Implications for the speciation process. Science 224: 171-173. Selys Longchamps, E. de. 1878. Quatriémes addition au synopsis de gomphines. Bull. Acad. R. Belg. (2)46: 408-698. Walker, E. M. 1958. The Odonata of Canada and Alaska, volume 2. Univ. of Toronto Press. 318 Pp. PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 215-218 SECOND EUCINETIDAE-CONIOPHORACEAE ASSOCIATION (COLEOPTERA; BASIDIOMYCETES), WITH NOTES ON THE BIOLOGY OF EUCINETUS OVIFORMIS LECONTE (EUCINETIDAE) AND ON TWO SPECIES OF ENDOMYCHIDAE E. RICHARD HOEBEKE, QUENTIN D. WHEELER, AND ROBERT L. GILBERTSON (ERH, QDW) Department of Entomology, Cornell University, Ithaca, New York 14853: (RLG) Department of Plant Pathology, University of Arizona, Tucson, Arizona 85721. Abstract. —A breeding population (adults and larvae) of the eucinetid beetle, Eucinetus oviformis LeConte, was found in association with basidiocarps of the wood-rotting fungus Coniophora arida (Fr.) Karst. var. arida (Basidiomycetes, Coniophoraceae) in the Finger Lakes region of New York in late summer 1986. Larvae of E. oviformis, maintained in the laboratory, aggregated upon the substrate and pupated in one closely-packed group. Pupae hung inverted from the last larval exuvia which were attached to the substrate by the caudal end. In addition, adults and larvae of the endomychids, Mycetina perpulchra (Newman) and Aphorista vittata (F.), also were found in association with the same wood- rotting fungus. The Coniophoraceae-F. oviformis association and the two breeding species of Endomychidae on a host in the Coniophoraceae provide additional evidence for the diversity of Coleoptera feeding on minute Basidiomycetes. Wheeler and Hoebeke (1984) recently re- ported the discovery of adults, larvae and pupae of Eucinetus oviformis LeConte in association with basidiocarps of the wood- rotting fungus Coniophora olivacea (Pers.) Karst. near Highlands, North Carolina (Ma- con County). This represented only the sec- ond breeding record for a eucinetid beetle on a Basidiomycete, although fungus-feed- ing habits have been suggested for more than a century (Perris, 1851). The first substan- tiated record was for E. punctulatus Le- Conte, which breeds in boletes (Bruns, 1984), although fungus associations were also reported by Klausnitzer (1971, 1975). In this paper we report the discovery of a breeding population of FE. oviformis near Trumansburg, New York (Tompkins Coun- ty) also on a host in the Coniophoraceae, and report the association of the endomy- chids Mycetina perpulchra (Newman) and Aphorista vittata (F.) with the same fungus. EUCINETIDAE-CONIOPHORACEAE ASSOCIATION Until our previous report on E. oviformis (Wheeler and Hoebeke, 1984), no Eucine- tidae were known to feed on wood-rotting Basidiomycetes (Gilbertson, 1984). The collection of another breeding population (including adults and larvae) of this euci- netid beetle on a related species of the same genus of fungus, Coniophora, provides fur- ther confirmation of this association. On August 26, 1986, one of us (ERH) found a large fallen trunk of hemlock (7suga canadensis (L.) Carr.) with a very extensive development of basidiocarps of Coniophora arida (Fr.) Karst. var. arida. Found on the surface of the fungal fruiting bodies were 216 adults and larvae of Eucinetus oviformis. Mature larvae and some adults were still present as late as October 18, 1986. The collection site was in a large virgin woodlot (known as the Henry A. Smith Woods, founded in 1909), located on the southern village limit of Trumansburg, New York. This woodlot is a small (ca. 30 acres) but impressive stand of old growth forest. The principal soil type is an Arkport fine sandy loam (deep, well-drained and acidic throughout the profile). The forest is dom- inated by beech (Fagus grandifolia Ehr.), sugar maple (Acer saccharum Marsh.), tulip tree (Liriodendron tulipifera L.), and hem- lock (Tsuga canadensis (L.) Carr.). The woodland floor is shadowed by an extensive canopy and moisture content is relatively high. These conditions support an under- story dominated by elder (Sambucus), spikenard (Aralia racemosa L.) and numer- ous herbaceous plants. In addition to the various life stages of the eucinetid, the ba- sidiocarps of C. arida were supporting large breeding populations of two species of En- domychidae that are discussed below. Until its association with Coniophora oli- vacea, the hosts of Eucinetus oviformis were unknown. Although other members of the Eucinetidae feed on either fungi or slime molds, there is no clear evidence to suggest that any eucinetids have particularly broad feeding habits. Our two records of Conioph- ora hosts may suggest that E. oviformis has a restricted host range—within the Conio- phoraceae or related taxa. This hypothesis, of course, remains to be tested with future field work. Both species of Coniophora reported as hosts for eucinetid beetles have a wide range in North America. Coniophora arida var. arida is circumglobal in the North and South Temperate zones, and C. olivacea is circum- global in the North Temperate Zone (Ginns, 1982). They occur primarily on dead co- nifers but may also decay dead hardwoods, particularly those in coniferous forest eco- systems. Their basidiocarps are annual, and PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON in most parts of North America would de- velop primarily in the summer and early fall and would deteriorate rapidly with the advent of low temperatures. They overwin- ter as mycelia in the wood. Coniophora species cause a brown rot, selectively re- moving the cellulose and hemicelluloses from wood and leaving a stable residue of slightly modified lignin. ORIENTATION OF EUCINETID PUPAE We maintained live Eucinetus oviformis larvae in the laboratory, taken from the Tru- mansburg population. These larvae suc- cessfully pupated, and we noticed an inter- esting phenomenon that has not been previously reported in the Eucinetidae. Lar- vae pupated in the rearing container gre- gariously, and pupae hung inverted by an attachment to the last larval exuvium (Fig. 1). The same attachment was apparent, upon reexamination, in the material collected in North Carolina. We are uncertain how widespread this pupation behavior is in oth- er Coleoptera, but have noted a similar case (involving gregarious behavior and inverted hanging in the larval exuvium) in a species of Neotropical Ni/io (Nilionidae or Tenebri- onidae) (Wheeler, unpublished data). ENDOMYCHIDAE ASSOCIATED WITH CONIOPHORACEAE Relatively few specific fungal associa- tions are recorded for species of Endomy- chidae; those that are recorded include various Basidiomycetes (Scheerpeltz and HOfler, 1948; Benick, 1952). The lack of much specific host data suggested to Crow- son (1984) that many endomychids might be associated with Ascomycetes. Our ob- servations on two genera (below), however, add another possible explanation for the lack of published host records. At the Trumans- burg collecting site and intermingled with the eucinetids, breeding populations of two species of Endomychidae were also found on the same host, Coniophora arida. On August 26, 1986, numerous larvae of My- VOLUME 89, NUMBER 2 217, Figs. 1,2. 1, Aggregation of pupae of Eucinetus oviformis LeConte; pupae hang inverted from the last larval exuvia which are attached by the caudal ends to the substrate. Scale line = 4 mm. 2, Mature larvae of Aphorista vittata (F.) (4 large, dark larvae) and Mycetina perpulchra (Newman) (2 small, pale larvae) grazing on basidiocarps of the wood-rotting fungus Coniophora arida var. arida. Scale line = | cm. 218 cetina perpulchra (Newman) and Aphorista vittata (F.) were found moving over basi- diocarps of the wood-rotting fungus (Fig. 2). Adult pairs of both species were found in copula upon the felled hemlock trunk. Lar- val identification was made by association with adults. Considerably more larvae and adults of M. perpulchra than A. vittata were present on this first collecting date. On later visits to the same site (September 27 and October 18, 1986), ERH found larvae of A. vittata in much greater abundance. Also on October 18, adults of M. perpulchra were again extremely numerous. So far as we know, this is the first published record of Endomychidae on a host in the Conio- phoraceae. Voucher specimens of larvae and adults of both endomychids are in the Cor- nell University Insect Collection along with samples of the wood-rotting fungus. CONCLUSIONS Many wood-rotting Basidiomycetes nev- er produce large fruiting bodies (Gilbertson, 1984) and yet occur with such abundance and frequency that they must pose a sizable resource for mycophagous insects. This confirmation of the Coniophoraceae-Euci- netidae association and the new breeding reports of two species of Endomychidae on a host in the Coniophoraceae provide added evidence for the diversity of Coleoptera feeding on minute Basidiomycetes, a fauna that also includes Dasyceridae (Wheeler, 1984). Similar field studies are needed to further elucidate relationships between bee- tles and these wood-rotting fungi. ACKNOWLEDGMENTS This research was supported, in part, by NSF Grant No. BSR-8315457 and Hatch Project no. NY(C)139426 to Q. D. Whee- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ler. We thank Peter Marks (Cornell Univer- sity) for providing us with information about the abiotic and biotic communities of the Henry Smith Woods. LITERATURE CITED Benick, L. 1952. Pilzkafer und Kaferpilze. Acta Zool. Fennica, no. 70, 250 pp. Bruns, T.D. 1984. Insect mycophagy in the Boletales: Fungivore diversity and the mushroom habitat, pp. 91-129. In Wheeler, Q. and M. Blackwell, eds., Fungus-Insect Relationships. Columbia Univer- sity Press, New York. Crowson, R. A. 1984. The associations of Coleoptera with Ascomycetes, pp. 256-285. Jn Wheeler, Q. and M. Blackwell, eds. Fungus-Insect Relation- ships. Columbia University Press, New York. Gilbertson, R. L. 1984. Relationships between in- sects and wood-rotting Basidiomycetes, pp. 130- 165. In Wheeler, Q. and M. Blackwell, eds., Fun- gus-Insect Relationships. Columbia University Press, New York. Ginns, J. 1982. A monograph of the genus Coniopho- ra (Aphyllophorales, Basidiomycetes). Opera Bot. 61: 1-61. Klausnitzer, B. 1971. Zur Biologie einheimischer Ka- ferfamilien, 7. Eucinetidae. Entomol. Ber. 15: 73- 74. 1975. Beitrage zur Insektenfauna der DDR: Coleoptera-Eucinetidae. Beitr. Entomol., Berl. 25(2): 325-327. Perris, E. 1851. Quelques mots de les metamorphoses de Coleopteres mycetophages, le Triphyllus punc- tatus, Fab.; le Diphyllus lunatus, Fab.; Agathi- dium seminulum, Linn., et ?Eucinetus (Nycteus Latr.) meridionalis de Castelnau. Ann. Soc. Ento- mol. France (2)9: 39-53. Scheerpeltz, O. and K. Hofler. 1948. Kafer und Pilze. Verlag fur Jugend und Volk, Wien. 351 pp. + IX plates. Wheeler, Q. D. 1984. Notes on host associations and habitats of Dasyceridae (Coleoptera) in the south- ern Appalachian Mountains. Coleopt. Bull. 38: 227-231. Wheeler, Q. D. and E. R. Hoebeke. 1984. A review of mycophagy in the Eucinetoidea (Coleoptera), with notes on an association of the eucinetid bee- tle, Eucinetus oviformis, with a Coniophoraceae fungus (Basidiomycetes: Aphyllophorales). Proc. Entomol. Soc. Wash. 86: 274-277. PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 219-225 A NEW SPECIES OF WATER PENNY BEETLE, PHENEPS CURSITATUS, FROM CERRO DE LA NEBLINA, VENEZUELA (COLEOPTERA: DRYOPOIDEA: PSEPHENIDAE) PAUL J. SPANGLER Department of Entomology, National Museum of Natural History, Washington, D.C. 20560. Abstract. —A new species of psephenid beetle, Pheneps cursitatus Spangler, from Cerro de la Neblina, Venezuela, is described, illustrated with line drawings and scanning electron micrographs, and compared to its closest relative, Pheneps antennalis Spangler & Steiner (1983), from Surinam. A photograph and notes on the habitat are provided. Insect collections made during a biolog- ical survey of the plants and animals of the tepui Cerro de la Neblina, in southern Ven- ezuela, yielded numerous new taxa, distri- bution records, and biological data. The new aquatic beetle described in this paper is another example of the rich and relatively unstudied fauna of the Amazon basin. The genus Pheneps and two species, P. gracilis from Haiti and P. cubanus from Cuba, were described by Darlington (1936); a third species, P. antennalis from Surinam, was described more recently by Spangler and Steiner (1983). Although the male of P. cu- banus is unknown, the males of P. gracilis, P. antennalis, and P. cursitatus, new species, are known and all have exceptionally long antennae (Figs. 1, 2) as described for the genus by Darlington. Pheneps cursitatus Spangler, NEw SPECIES Figs. 1-12, 15 Holotype male.—Form and size: Body flattened; thorax narrowed anteriorly; elytra diverging slightly posteriorly, widest at about posterior three-fourths, apices rounded. Length, 2.47 mm; width, 1.31 mm. Coloration: Covered with fine, dense, short, recurved, golden pubescence. Head, antennal segments 3-11, pronotum, and elytra dark brown; basal two antennal seg- ments, labium, and labial palpi yellowish brown. Ventral surface light yellowish brown except maxillary palpi, prosternum, sides of metasternum, apices of femora, and tibia gray brown. Abdomen dark gray brown ex- cept slightly yellowish medially. Tarsi yel- lowish brown. Head: Almost flat behind eyes, decurved between eyes; finely microreticulate and finely punctate, more densely so anteriorly; labroclypeal suture distinctly depressed. Clypeus with anterior margin subtruncate. Labrum narrow, broadly subrectangular, and indistinctly arcuately emarginate on an- terior margin. Eyes prominent, hemispher- ical. Antenna (Fig. 5) densely pubescent; fi- liform; exceptionally long, as long as length of body. Maxillary palpus (Fig. 2) 4 seg- mented, filiform; about three-fourths the length of antennal segments 1-3 combined; first segment shortest, about a fifth as long as second segment; second segment longest, only slightly longer than fourth segment; third segment about three-fourths as long as second segment; fourth segment slightly longer than third segment, slightly com- 220 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-4. Pheneps cursitatus, new species, male Maxillary palpus, apical segment. 4, Protarsus. pressed laterally (Fig. 3). Labial palpus very small, 3 segmented; first segment about a third as long as second segment; second seg- ment swollen and slightly longer than third segment; third segment partially com- pressed apically. Thorax: Pronotum widest at base; discal . 1, Habitus, dorsal view. 2, Habitus, ventral view. 3, area densely, finely punctate; discal punc- tures separated by 12 to 1 times their di- ameter; disc moderately convex, moderate- ly depressed adjacent to posterolateral angles; finely and indistinctly margined ventrolaterally; anterolateral angles strong- ly rounded; posterolateral angles forming VOLUME 89, NUMBER 2 right angles but slightly rounded; with apical and basal margins moderately bisinuate. Scutellum slightly wider than long, flat; fine- ly and densely punctate; rounded poste- riorly. Elytra about 3.5 times longer than pronotum and about a fourth wider at wid- est point than pronotum; humeri slightly gibbous; apices broadly rounded; each ely- tron with 5 indistinct striae on discal area paralleling elytral suture; punctures fine and very dense. Prosternum (Fig. 2) moderately short in front of procoxae; covered by plas- tron setae. Prosternal process narrow, keel- like; apex extending into mesosternum. Mesosternum narrow between mesocoxae, about an eighth as wide as mesocoxal width (Fig. 8); with a very narrow, median, lon- gitudinal cleft. Metasternum behind me- socoxae abruptly and strongly raised above plane of mesosternum; with a very fine, me- dian, longitudinal line extending along length of raised portion; surface microretic- ulate and finely, densely punctate. Femora robust and swollen. All tibiae very slender and each with a low but distinct carina pos- terolaterally (Figs. 2, 3, arrows). Mesotibiae and metatibiae each with a bifurcate carina. Protibiae and mesotibiae grooved laterally on apical three-fifths. Protarsal segments (Fig. 4) and mesotarsal segments 1 and 2 much broader than segments 3, 4, and 5 and densely pubescent ventrally; metatarsi un- modified. Tarsal claws small, slender, and sharp; each with a subbasal tooth. Abdomen: All sterna with surface sculp- ture as on metasternum and covered with plastron setae. Second visible sternum strongly emarginate in medial third. Fifth visible sternum broadly emarginate along posterior margin. Sixth visible sternum broadly triangularly incised medially, re- sulting in 2 subtriangular lateral lobes. Sev- enth visible sternum rounded at apex. Male genitalia: Bulbous basally and tr- lobate apically as illustrated (Figs. 11, 12). Female.—The female differs from the male as follows. Larger size, 3.90 mm long, 1.51 mm wide. Antennae (Fig. 6) only 221 slightly longer than length of pronotum and segments submoniliform. Maxillary palpi short; about as long as combined length of basal 2 antennal segments and about half as long as palpi of male. Mesosternum wide between mesocoxae, about half as wide as mesocoxal width (Fig. 7). Visible abdomi- nal sterna 1-5 subequal in length (Fig. 9); only fifth sternum shallowly emarginate ap- icomedially; sixth sternum broadly round- ed. Tarsal segments all equally very slender. Variation.— The specimens in the type se- ries are very similar except for the slightly variable extent of the yellowish color of the abdominal sterna and variations in length from 2.47 to 2.90 mm and width from 1.20 to 1.51 mm. Comparative notes.—The new species is very similar to Pheneps antennalis Spangler & Steiner (1983). All specimens in the type series of P. antennalis have a reddish-brown head and pronotum which contrasts with the dark brown elytra; P. cursitatus 1s com- pletely dark brown dorsally. Also, P. cur- sitatus 1s a smaller species (male length, 2.47 to 2.97 vs 3.1 to 3.8 mm). The shapes of the median lobe and parameres of the male genitalia are distinctive (Figs. 11-14). Fe- males of P. antennalis are unknown. Type-data.— Holotype ¢ and allotype: VENEZUELA: TERRITORIO FEDERAL AMAZONAS: Departamento Rio Negro: Cerro de la Neblina, Base camp, Rio Baria, 00°50'N 66°10’W, 140 m, 11 February 1985, P. J. and P. M. Spangler, R. A. Faitoute, W. E. Steiner; deposited in the National Mu- seum of Natural History, Smithsonian In- stitution. Paratypes: Same data as holotype, 90 4, 3 2; same data except as follows: 27 January 1985, 2 9; 10 February 1985, 5 4; 12 February 1985, 15 4; 14 February 1985, 1 6; 20 February 1985, 1 3; 5 February 1985, W. E. Steiner, 1 9; 6 February 1985, W. E. Steiner, 1 3. Paratypes will be deposited in the collec- tions of the Instituto de Zoologia Agricola, Facultad de Agronomia, Maracay, Vene- zuela; American Museum of Natural His- 222 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON — Ljnlai—|~inid uh ee = I \ | Poy ! / | : DD \ Figs. 5-10. Pheneps cursitatus, new species. 5, Antenna, male. 6, Antenna, female. 7, Mesosternum, female. 8, Mesosternum, male. 9, Abdomen, female. 10, Abdomen, male. VOLUME 89, NUMBER 2 i) tN Ww Figs. 11-14. 11, 12, Pheneps cursitatus, new species. 11, Male genitalia, dorsal view. 12, Male genitalia, lateral view. 13, 14. Pheneps antennalis Spangler and Steiner. 13, Male genitalia, dorsal view. 14, Male genitalia, lateral view. 224 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 15. tory, New York; California Academy of Sci- ences, San Francisco; Canadian National Collection, Ottawa; Institut Royal de His- toire Naturelle de Belgique, Bruxelles; Mu- seum National de Histoire Naturelle, Paris; Museo Argentina de Ciencias Naturales, Buenos Aires; Zoologische Sammlung Bay- erischen Staates, Munchen; and the collec- tion of Harley P. Brown, Norman, Okla- homa. Etymology.—The trivial name, cursita- tus, 1s derived from Latin meaning “run hither and thither” in reference to the rapid, erratic movements of these beetles over the rocks in riffles in their habitat. Habitat.—Specimens of Pheneps cursi- tatus were collected with aerial nets or oc- casionally in a seine from bare emergent rocks in the riffles; others were attracted to black lights operated on the bank of the Rio Baria or in a clearing in the rainforest near the river. In bright sunlight on February 11 and 12, we found specimens of P. cursitatus in rath- er large aggregations running over rocks in an area of shallow riffles in the Rio Baria (Fig. 15). The water level of the river was dropping rapidly during that time and tem- Biotope; riffle area in Rio Baria, T. F. Amazonas, Venezuela. perature differences were distinctly evident as we waded through the warm, shallow water into the cool, deeper water and faster current. The beetles were difficult to capture because they ran rapidly and took flight quickly when attempts were made to cap- ture them in an aerial net. The decreasing water level and increasing temperature of the exposed rocks seems to have triggered eclosion from the pupal stage because 66% of the specimens obtained were collected during those two days. An additional 15 specimens were collected at our blacklight operated on shore beside the riffle on the evening of February 11. Colorimetric water chemistry analyses provided the following data: pH, 4; oxygen, 12-15 ppm; hardness, 0. When these anal- yses were made, the day was sunny, the air temperature was 28.5°C, the water temper- ature was 24.5°C, and the water around the rocks where the beetles congregated was about 15 cm deep but kept getting shallower throughout the day. As the water level dropped and the rocks became exposed, the psephenid beetles kept moving to other rocks still surrounded by water at their bases. The elevation at the stream was about 120 m. VOLUME 89, NUMBER 2 ACKNOWLEDGMENTS The specimens of this new species were collected during a biological survey of Cerro de la Neblina. The expedition to Cerro de la Neblina Park was organized and directed by the Foundation for the Development of Physics, Mathematics, and Natural Sci- ences of Venezuela, with the patronage of the following Venezuelan organizations: The Ministry of Education, The Ministry of the Environment, the National Council of Sci- entific and Technological Research, the Venezuelan Air Force, and the National In- stitute of Parks. The entire project was co- ordinated by Dr. Charles Brewer-Carias and was conducted in collaboration with the Na- tional Science Foundation of the United States, the American Museum of Natural History, the Field Museum of Natural His- tory, the Missouri Botanical Garden, the New York Botanical Garden, and the 225 Smithsonian Institution. Biologists from several universities and other institutions also participated. I thank all of the above organizations and their administrators for the extensive contributions to the survey. I also thank Young T. Sohn for the line drawings, Robin A. Faitoute and Heidi Wolf for the scanning electron micrographs, Roy W. McDiarmid for reviewing the manu- script; and Phyllis M. Spangler for typing the manuscript. LITERATURE CITED Darlington, P. J. 1936. A list of the West Indian Dryopidae (Coleoptera) with a new genus and eight new species, including one from Colombia. Psyche 43: 64-83. Spangler, P. J. and W. E. Steiner. 1983. New species of water beetles of the genera E/moparnus and Pheneps from Suriname (Coleoptera: Dryopidae: Psephenidae). Proceedings of the Entomological Society of Washington 85: 826-839. PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 226-229 DISCOVERY OF FEMALE APOZYX (HYMENOPTERA: APOZYGIDAE) AND COMMENTS ON ITS TAXONOMIC POSITION W. R. M. MAson Biosystematics Research Centre, Agriculture Canada, Ottawa, Ontario K1A 0C6, Can- ada. Abstract. —Previously unknown, the female of Apozyx penyai Mason (Apozygidae) is found to be very similar to the male. Several more males from south-central Chile are reported. A further examination of characters of Apozyx shows it to be undoubtedly a member of Ichneumonoidea but not of Microhymenoptera nor Aculeata. Differences between Apozygidae and Braconidae are tabulated. In the eight years since I described Apozyx penyai Mason (Mason, 1978) several more males and a female have been collected in Chile. Additional males have been taken in Nuble (Las Trancas, 5 km E.S.E. of Recinto at 1250 m) in a Malaise trap operated by A. Newton and M. Thayer in Dec. 1982 in Nothofagus forest. Later a male and a female were taken in Nahuelbuta National Park, 50 km W. of Angol (12-1500 m, flight in- tercept trap operated from Dec. 1984 to 17 Feb. 1985 by S. and J. Peck), the site being in Nothofagus-Araucaria forest. Apozyx penyai Mason, SUPPLEMENTARY DESCRIPTION Description of male, see Mason (1978). The sexes are very similar except for the large ovipositor (Fig. 4), the stouter flagellar articles, and smaller eyes of the female (Figs. 1D): Abdomen of 2 (Fig. 4) resembling that of 6 but hypopygium large and triangular in profile. Ovipositor sheaths hairy, slightly deeper apically, about as long as metasoma; Ovipositor weakly and evenly curved up- ward, uniformly deep except subapically where it is slightly deeper, the apex sharply pointed and bearing a few small teeth ven- trally. Flagellum of 2 with 17 articles, about 20% shorter than that of 4; antipenultimate ar- ticle 1.4 times as long as wide (in é 2.3-2.8); second article of female 3 times as long as wide (in 4 2.5—3.5). Compound eye in 2 con- siderably smaller and less protuberant than in 6 (Figs. 1, 2); thus in dorsal or anterior aspect outer part of the eyes slightly closer than temples (in ¢ eyes bulging distinctly farther apart than temples) and the vertex elevated above top of eyes for a distance almost equal to eye height (in 6 vertex ele- vated for a distance about half eye height). In both sexes second abdominal (first metasomal) sternum including 2 parts: 1, an anterior heavily sclerotized section fused with the tergite into a tubular structure with component parts not easily distinguishable; 2, a small lenticular posterior section formed of 2 subtriangular plates separated medially by narrow membranous area (Figs. 5, 6). During flexion of the part of the abdomen behind tergum 2 this pair of plates overlaps externally the tubular fused tergite and ster- nite 2. Metasomal terga | and 2 articulate laterally by pair of submarginal condyles VOLUME 89, NUMBER 2 Figs. 1-4. Apozyx penyai. 1, 2, Head of female, frontal and lateral view. 3, Paramere and volsella, inner side. 4, Female abdomen. (Fig. 5). Sterna not involved in articulation. Because sterna of holotype could not be counted, original description fails to men- tion that the metasomal sternites 2 and 3 are fused exactly as their respective tergites. On p. 609, line 9, the terms (2r-m) and (3r- m) should be corrected to (1r-m) and (2r- m) respectively. Male genitalia (Fig. 3) with weakly ex- panded and rounded parameres. Cuspis broad, truncate and very short, digitus ta- pered, extending far beyond cuspis. DISCUSSION In 1978, I was uncertain about the place- ment of this family but studies during in- tervening years and recent knowledge of the female anatomy have confirmed my first opinion. Apozyx shares the following char- acter states with other Ichneumonoidea: 1, flagellar articles with large longitudinal placodes; 2, mandibles with two apical teeth; 3, pronotum much shorter medially than laterally, the posterodorsal corners touching the tegulae; 4, pronotum (plus prepectus) rigidly attached to the mesoscutum and mesopleuron; 5, posterodorsal corner of pronotum developed into a lobe behind which lies the mesothoracic spiracle; 6, pre- pectus fused to the posterior vertical pro- notal margin, the pit that indicates exter- nally the site of the apodeme bearing the origin of the spiracular occlusor muscle lying very near the posterior margin of the fused 228 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 5,6. Apozyx penyai, junction of abdominal segments 2 and 3, showing abdominal tergites 2 and 3 (T2, T3), sternites 2 anterior (S2a) and posterior (S2p), sternite 3 (S3), and condyle (C). 5, Anterolateral view. 6, Ventrolateral view. pronotum plus prepectus (for definition of prepectus and discussion of characters 4-6, see Gibson, 1985); 7, costa and radius of fore wing fused; 8, sternum of metasoma one divided into a heavily sclerotized an- terior section and comparatively weakly sclerotized posterior section (Mason, 1981, 1983); 9, metasomal segments | and 2 ar- ticulated by a pair of sublateral condyles on the hind margin of tergum | and anterior margin of tergum 2, the sterna playing no part in the articulation. The presence in the female of complete metasomal terga 7 and 8, the latter bearing large cerci, eliminates Apozyx from the Acu- leata (Koenigsmann, 1978). The presence of spiracles on each of abdominal tergites | to 8 as well as the step-like posterior deflection of fore wing Cu at its junction with m-cu eliminates Apozyx from Microhymenop- tera because all Microhymenoptera have a comparatively straightened Cu (if it is not reduced to invisibility) and their abdomen has only 2 pairs of spiracles on terga 1 and 8, or only one pair on tergum | (propo- deum), the other spiracles being absent or vestigial. The general aspect of Apozyx is similar to that ofa braconid but there are important differences: 1, in Apozyx abdominal sterna 3 and following are rounded and sclerotized as strongly as the terga, a basic condition shared with most other Hymenoptera, whereas the same sterna of Braconidae are extensively soft, membranous and usually divided into several weakly sclerotized plates, a derived condition found only in Braconidae, Ichneumonidae and Paxylom- matidae; 2, vein 2r-m (2nd recurrent) is present as a strong tubular vein in the fore wing, a basic condition (Mason, 1981), whereas in Braconidae this vein is missing, a derived condition. ACKNOWLEDGMENTS My thanks go to the technicians of the Electron Microscope Center (Canada Ag- riculture) and to H. E. Bisdee, B.R.C., for VOLUME 89, NUMBER 2 229 assistance in making the illustrations. I also der Hymenoptera, 4. Dtsch. Entomol. Zeit. 25: thank my colleagues J. R. Barron and M. J. 365-435. f ; d : fth Mason, W. R. M. 1978. A new genus, species and Sharkey or suggestions and review of the family of Hymenoptera (Ichneumonidae) from Manuscript. Chile. Proc. Entomol. Soc. Wash. 80: 606-610. 1981. Paxylommatidae: The correct family- LITERATURE CITED group name for Hybrizon Fallén (Hymenoptera: Ichneumonoidea), with figures of unusual anten- nal sensilla. Can. Entomol. 113: 433-439. . 1983. Anew South African subfamily related to Cardiochilinae (Hymenoptera: Braconidae). Contrib. Am. Entomol. Inst. 20: 49-62. Gibson, G. A. P. 1985. Some pro- and mesothoracic structures important for phylogenetic analysis of Hymenoptera, with a review of terms used for the structures. Can. Entomol. 117: 1395-1443. Koenigsmann, E. 1978. Das phylogenetische system PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 230-241 A REVIEW OF THE SPECIES OF SYNERGUS FROM GUATEMALA, WITH NOTES ON CYNIPS GUATEMALENSIS CAMERON (HYMENOPTERA: CYNIPIDAE) A. J. RITCHIE AND J. D. SHORTHOUSE (AJR) Department of Biology, Carleton University, Ottawa, Ontario (Current address: 47 Deepwood Drive, Hamden, Connecticut 06417); (JDS) Department of Biology, Lau- rentian University, Sudbury, Ontario P3E 2C6, Canada. Abstract. —Three new species of Synergus Hartig (Hymenoptera: Cynipidae) from Gua- temala are described and Synergus filicornis Cameron is redescribed. A key to S. cultratus new species, S. kinseyi new species, S. mesoamericanus new species, and S. filicornis Cameron is presented. Biological notes, when available, are given for each species. Cynips guatemalensis Cameron is transferred to Andricus and is a senior synonym of Andricus mexicana Bassett, A. mexicanus Kinsey new synonym, and Synergus dorsalis Cameron new synonym. The nomenclature for Synergus dorsalis Cameron and S. dorsalis (Pro- vancher) is discussed. During a visit to the American Museum of Natural History, one of us (AJR) discov- ered a series of undescribed inquiline gall wasps belonging to the genus Synergus Har- tig (Hymenoptera: Cynipidae) from Gua- temala. These species are of interest for sev- eral reasons. Guatemala is the southernmost point of the range of inquiline cynipids in the New World and is isolated from other cynipid habitats by the lowland Isthmus of Tehuantepec. Secondly, some of the most primitive oak gall wasps come from Gua- temala and southern Mexico. Finally, two of the three new species are extremely large for inquiline Cynipidae. Very little is known about the gall wasps of Guatemala. Cameron (1883) described the woody oak-stem galls of Cynips cham- pioni and C. imitator and the oak-leaf galls of C. guatemalensis. Cameron described also two species of Synergus from Guatemala: S. filicornis from an unknown gall, and S. dorsalis from galls of C. guatemalensis. Kinsey (1920) cut insects from galls that he considered to be made by C. guatemalensis. Kinsey (1936) later described Acraspis fu- giens, Atrusca lucaris, and Atrusca lumi- naris, all of which form leaf galls on oaks. Weld (1913) described Synergus furnessana from Mexico but Weld (1930) later decided that this was synonymous with S. filicornis Cameron. The only other inquiline known from Mexico is S. dugesi Ashmead (holo- type in USNM; examined), which has been reared from galls of Andricus championi (Cameron). Kinsey made important contributions to our knowledge of the taxonomy and biology of gall wasps. Although his taxonomic con- cepts and nomenclature were radical and have not been followed since, his collection of Cynipidae, now in the American Mu- seum of Natural History, is one of the most important in the world. He was an avid col- lector, and some estimates of the size of his collection reach 5 million specimens (Weld, VOLUME 89, NUMBER 2 1952). Kinsey never described or reported any of the inquilines or parasites that he reared from galls. He exhaustively labelled them with locality, collection and emer- gence dates, host plant and host gall, but curated them under the name of the host gall inducer. Unfortunately, he assigned manuscript names to many species of gall inducer which he never described. This is apparently the case with the Guatemalan gall wasps, as two of the four galls from which he reared inquilines cannot be posi- tively identified; one (brelandi) is a Kinsey manuscript name (Weld, 1952), and the other (ruginos.) is abbreviated. Tentative host identifications are discussed in the remarks for S. cultratus. Collection dates were given in the following form, “‘month.day.year,” and the symbol ® was used to indicate the collection date of the gall and “ing.” for the date of emergence for an inquiline. Kinsey usually put a num- ber and letter after the locality; these indi- cate the number of miles and direction from the nearest town (i.e. Sacapulus 9S means 9 miles south of Sacapulus). This has been followed in the present paper. Morphological terms are those of Ritchie and Peters (1981) and Shorthouse and Rit- chie (1984), with the following exceptions: supracoxal carina = carina on propodeum running from metacoxal socket dorsally to anterior of the flap over propodeal spiracle; metanotal median fovea = the small me- dian fovea posterior and between the lateral foveae, and above the area between the pro- podeal carinae. The majority of specimens examined in this study, including the holotypes of the new species, are deposited in the American Museum of Natural History, New York, New York (AMNH) except as noted. The holotypes of Cynips guatemalensis Cam- eron and Synergus dorsalis Cameron are de- posited in the British Museum (Natural History) (BMNH), the holotype of Cer- optres dorsalis Provancher is in the Public 231 Museum of Quebec, Quebec City, Quebec (PMQ), and the holotype of Synergus fur- nessana Weld is in the Philadelphia Acad- emy of Natural Sciences, Philadelphia, Pennsylvania (PANS). Kinsey collected all specimens and made all host-gall determin- ations except as noted. All Kinsey material was collected from galls on Quercus pili- caulis. Representative specimens of the four Guatemalan species of Synergus have been retained by both authors. KEY TO THE SPECIES OF SYNERGUS FROM GUATEMALA 1. Female metasomal tergite 2+ 3 (T2+3) deeply excavated along posterodorsal margin, poster- ior third sparsely and weakly punctate (Figs. 9, 10); meso- and metasoma black and white, black and yellow, or entirely yellow ........ 2 — Female T2+3 at most weakly excavated along posterodorsal margin, posterior third entirely smooth (Figs. 11, 12); mesosoma entirely black, metasoma black and rufous 2. Crossvein r-m in hind wing recurved (Fig. 17); radial sector (Rs) 1 and Rs2 nearly parallel (Fig. 19); third male antennomere (A3) elongated beyond notch (Fig. 13); mesopleuron entirely vellowsabovercoxallibaSeSiew ence eae BRE, eae cultratus Ritchie and Shorthouse, n. sp. — Crossvein r-m in hind wing normal (as in Fig. 18); Rsl and Rs2 divergent (Fig. 20); male A3 not elongated beyond notch (Fig. 14); meso- pleuron with some black above coxal bases . Sapien ek eRe gS eee ORR A filicornis Cameron 3. Female with 15 antennomeres; radial cell length 3.0 times width; Rs2 straight (at most weakly bent distally); areolet absent or elongate (Fig. 2) susually over:d:Oimmelongis.. essence kinseyi Ritchie and Shorthouse, n. sp. — Female with 14 antennomeres; radial cell length usually 2.0 times width; Rs2 distinctly bent; areolet equilateral (Fig. 22); under 4.1 mm .. .. Mesoamericanus Ritchie and Shorthouse, n. sp. Synergus cultratus Ritchie and Shorthouse, NEw SPECIES igss ll) 9. Pe yd Female (holotype).—Length 7.0 mm. Yellow; tip of mandible, frons above anten- nal socket and between frontal ridges, me- dian third of vertex and occiput, pronotal PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 4 Car ry agi pi ve me — UN fd i tae T ae) Sy Wows!” iS oe * . ~ a 44 VOLUME 89, NUMBER 2 plate, scutum between anterior lines, meso- pleuron between coxal sockets, anterodorsal margin of T2+3, posterodorsal margin of T7 black; distal half of mandible except tip, A1-A7, tarsal claw, T1, posterodorsal mar- gin of T2 +3, exposed dorsal margins of T4- T6 piceous brown. Head slightly narrower than thorax (slightly wider in all other species); malar space 0.56 of eye height; frontal ridge weak, reaching ectal margin of posterior ocellus; LOL = 0.8D, POL = 1.4D, OOL = 1.4D; antenna with 14 antenno- meres, 5.29 mm long; Al-A14 (mm): 0.33, 0.17, 0.56, 0.57, 0.58, 0.56, 0.46, 0.40, 0.35, 0229570:25.0225;,-0:2:1; 0:31. Mesoscutum (Fig. 1) transversely rugulose, punctate be- tween anterior lines; notauli percurrent, ir- regularly interrupted by transverse rugulae; median groove represented by weak crease extending about one third of mesoscutum; anterior lines slightly divergent, extending about one fourth of mesoscutum. Meso- scutellum (Fig. 1) rugose, rugosity stronger posteriorly, median depression absent; up- turned posterior carina strong, complete; fovea large, shallow, rugulose. Mesopleuron (Fig. 5) entirely striate. Metanotal median fovea with longitudinal carina. Propodeum with an extra weak carina parallel to and between supracoxal carina and propodeal carina; propodeal carinae very strong, thick- ened, strongly coriaceous. Metasoma (Fig. 9) longer than head plus mesosoma. T2+3 covering about two thirds of metasoma (lat- eral view), strongly excavated along pos- terodorsal margin, posterior third sparsely, weakly punctate. Forewing (Fig. 19) 6.0 mm long; radial cell length 3.0 times width; Rs1 nearly parallel to Rs2 (divergent in all other species); Rs2 very weakly bent near wing margin, almost straight; areolet normal; hind wing 4.0 mm long, r-m strongly recurved — Figs. 1-8. 233 (Fig. 17) (weakly recurved in all other species; Fig. 18). Tarsal claw with distinct tooth. Male (allotype).— Differs from holotype as follows: length 4.75 mm; antenna 4.9 mm long, with 15 antennomeres, Al—A15 (mm): 0:38; 05155035050550;,0:50,:0550;10.42 0:33; O31; O:2680:2350 2107021 0118022743 weakly bent and notched (Fig. 13), elon- gated beyond notch; T2+3 covering entire metasoma (dorsal view), T7 (posterior view) with setigerous punctures; forewings 5.6 mm long, hind wings 3.4 mm long. Variation. — Female: Length 5.0-7.25 mm, average of 50 specimens = 6.3 mm; mesoscutum from yellow with small black area between anterior lines to black with small yellow areas at posterior ends of no- tauli; scutellum sometimes amber or light brown; propodeal carinae yellow or black, black area sometimes extending ectal of ca- rinae; mediodorsal black area of metasoma sometimes expanded to occupy entire dor- sal margin. Male: Length 3.3-5.1 mm, average of 50 specimens = 4.4 mm; coloration as in fe- male except propodeum, mesoscutum, mesoscutellum, and metapleuron some- times entirely black. Type material.—Holotype, 2? (AMNH): Huehuetenango 3S, Guate[mala], 7300’, ® 12.22.35, ing. 8.18.37, ex gall of brelandi. Allotype, ¢: with same data as holotype. Paratypes (142 2, 145 6; AMNH): 71 9, 74 é with same data as holoypte; 13 2, 16 6 with same data as holoytpe but emerged 7.1- 9.10.36; 20 2, 26 6 with same data as ho- lotype but emerged 7.1—-9.1.38; 13 2, 3 4 with same data as holotype but emerged 8.22-8.27.37, ex gall of ruginos.; 5 2, 10 4, Sacapulus 9S, Guate., 6000’, ® 12.28.35, ing. 7.3-10.22.37, ex gall of brelandi; 1 8, 1-4, Mesonota of Synergus spp., dorsal view. 1, Synergus cultratus. 2, Synergus filicornis. 3, Synergus kinseyi. 4, Synergus mesoamericanus. 5-8, Mesopleura of Synergus spp., lateral view. 5, Synergus cultratus. 6, Synergus filicornis. 7, Synergus kinseyi. 8, Synergus mesoamericanus. 234 1 6 with same data as preceding series but emerged 9.1.38; 1 2 with same data as pre- ceding series but emerged 7.1.39; 2 2, 2 6, Quiche 2N, Guate., 7500’, ® 12.27.35, inq. 8.10-9.10.37, ex gall of brelandi; 4 2, 2 6 with same data as preceding series but emerged 7.1—7.15.38; 1 2, Guatemala City 3W, Guate., 63007; @ 12.30:35, ing. 8.3.37, ex gall of brelandi; 11 2, 1 & (Label lost), Guatemala, 1935, ing. 9.20.37, ex gall of brelandi. Excluded from type series: 89 specimens from the above localities and emergence dates, in various but poor condition; ten- tatively identified as S. cultratus. Etymology.—The species name is based on the Greek word for knife in reference to the strongly compressed metasoma. Diagnosis.—Synergus cultratus can be distinguished easily from other Guatemalan species by the following characters: r-m in the hind wing strongly recurved; Rs1 nearly parallel to Rs2; mesopleuron entirely yel- low; male A3 elongate beyond notch; and the extra carina between the supracoxal and propodeal carinae. Some specimens of this species are the largest known inquiline cyni- pids, being over 7.0 mm in length. Remarks. — Host relationships for S. cul- tratus are unclear, as neither of the reported hosts on the labels can be definitely iden- tified. One labelled host, bre/andi, is a Kin- sey manuscript name for a species of 4An- dricus (Weld, 1952). The other labelled host, ruginos., may be an abbreviation for An- dricus ruginosus Bassett, which produces a hard polythalmous stem gall (Weld, 1957) or more likely, a variety of ruginosus (see Kinsey, 1930, 1936 for his taxonomic con- cepts and use of varietal names). The emer- gence dates for this species are also unusual, as most species of Synergus emerge after one or two winters in the gall (Eady and Quin- lan, 1963). Synergus cultratus appears to have a one to four year variable life cycle, although it is possible that first year emer- gents were from older galls and that the life cycle is from two to four years in length. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Most individuals (65% of types) emerged after two winters in the gall. Synergus filicornis Cameron Figs. 25:6, 10; 145 20 Synergus filicornis Cameron, 1883: 71 (8, 4). TYPE: Holotype, 2 (BMNH). Exam- ined. Synergus furnessana Weld, 1913: 134 (8, 8); synonymized by Weld, 1930: 143. TYPE: Holotype, + (PANS). Examined. Female. —2.0-—3.6 mm. Black and yellow; Head yellow; frons above antennal socket and between frontal ridges, ocellar area, me- dian third of vertex, median third of occiput usually black (rarely entirely yellow); anter- ior of pronotum, anterior of pronotal plate piceous brown to black; posterior of pro- notal plate yellow to black, remainder of pronotum yellow to amber; mesoscutum usually yellow or amber, with black or pi- ceous brown between notauli (rarely entire- ly yellow or mostly piceous brown); meso- scutum usually yellow to amber, rarely dark piceous brown; mesopleuron black, fre- quently with median yellow area, rarely en- tirely yellow; metanotum and metapleuron usually black, sometimes piceous brown, rarely yellow; propodeum piceous brown to black, rarely yellow; spiracular flap almost white, always lighter than surrounding col- oration; metasoma yellowish, with broad dorsal band piceous brown to black, band sometimes reduced to anterior half of T2+ 3; legs yellow, metatarsus amber to brown; wings hyaline, veins yellow to light brown. Head as wide or slightly wider than thorax; frons below antennal socket and beneath eye with strong radiating striae (relatively stronger than in other species), striae weaker between anterior tentorial pits; malar space about 0.54 of eye height; posteroventral margin of gena with weak short carina; fron- tal ridge present, moderately strong, not quite reaching posterior ocellus; LOL = 0.91D, POL = 2.0D, OOL = 1.36D; anten- na with 14 antennomeres; A3—A6 relatively VOLUME 89, NUMBER 2 thin, subsequent antennomeres stouter; rel- ative length of antennomeres (Al-A14): OZ 50010" O8s230:52,0:32- 0:29" 025,0.2 1, ONO. tS 0532 Os 12.012. (0.20: Mesoscu- tum (Fig. 2) transversely rugulose; notauli percurrent, grooves sometimes interrupted by transverse rugulae, bottom of grooves weakly rugulose; median groove, lateral lines absent; anterior lines parallel, extending about one fourth of mesoscutum. Mesoscu- tellum (Fig. 2) rugose, median depression absent, weak posterior carina not upturned; fovea large, shallow, sometimes weakly ru- gulose, not well defined. Mesopleuron (Fig. 6) entirely striate, striations stronger ven- trally. Metanotal median fovea broadened, broader than in other species. Propodeum with moderately weak transverse carinae betwen supracoxal and propodeal carinae, no extra longitudinal carina; propodeal ca- rinae very strong, thick, strongly coriaceous. Metasoma (Fig. 10) slightly longer than head plus mesosoma; T2+3 covering about two thirds of metasoma, posterior third punc- tate, posterodorsal margin deeply excavat- ed; T4-T7 exposed along posterodorsal margin, more densely punctate than T2+3. Forewing (Fig. 20) about 1.0-1.1 length of body; radial cell length 3.0 times width; Rs2 very weakly bent near wing margin, almost straight; areolet of normal shape, posterior margin very weak to missing. Tarsal claw with a distinct tooth. Male.—Differs from female as follows: length 2.0-—2.5 mm; frontal ridge sometimes complete, running to anterior margin of posterior ocellus; antenna with 15 anten- nomeres, about equal to body length, A3 prominently notched and bent (Fig. 14), rel- ative lengths of antennomeres (Al-—A15): O2'60:08022 5.0.23; 0.21,.0.200515,0.12, Oa 20. Fie O51 O10) “0510; 0:09: 20:13: mesoscutum sometimes entirely black, with stronger transverse rugosity; mesopleuron sometimes entirely black, with stronger striae, propodeal carinae not as strong or as thickened, not as strongly coriaceous; meta- soma sometimes entirely black; T2+3 cov- 235 ering entire metasoma, not posterodorsally exavated; posterior third of T2+3 usually with band of punctures interrupted by pos- terodorsal smooth area, punctures some- times stronger and band not interrupted. Type material examined.—Synergus fili- cornis Cameron. Holotype, ? (BMNH, Type No. 7.110), labelled: ““San Geronimo, Gua- temala, Champion,” “Synergus filicornis.” Synergus furnessana Weld. Holotype, 2 (PANS), “Michoacan, Mex.,” “Synergus furnessana,” “‘Holotype”’ (red label). Allo- type, ¢ (PANS), with same data as holotype. Other material examined (AMNH).—18 2, 1 6, Huehuetenango 14S, 7500’, Guate., @® 12:23:35, mq. 1.7=—1.14.36, ‘ex gall or mexicanus; 1 2, 1 6, aS previous series but emerged 7.1.36; 23 2, 1 6, Sacapulus 9S, 6000’, Guate., ® 12.28.35, ing. 1.7—1.14.36, ex gall of mexicanus; 10 2, 1 6, Quiche 2N, 7500", Guate., © 12-27-35, nq. 1.7—1-22:36, ex gall of mexicanus; | 6, Guatemala City 5W, 6500’, Guate., ® 12.30.35 (no emer- gence date), ex gall of mexicanus. Diagnosis. —Synergus filicornis Cameron is most similar to S. cu/tratus and can be distinguished from it and other species by the following characters: under 4.0 mm; posterior third of T2+3 punctate, postero- dorsal margin of female T2+3 deeply ex- cavated; A3-A5 elongate and thin. The type of Synergus filicornis is in poor condition; the following structures are missing: left A2- Al4, right A8-A14, and the metasoma. Remarks.—This species is the most vari- able of the Guatemalan species of Synergus, especially in coloration. Very small females (~2.0 mm) are almost entirely yellow and have much finer sculpture of the mesopleu- ron and punctures on T2+3, whereas some of the larger females have much more brown or black and have stronger sculpturing. Sim- ilarly, two of the larger males have the meso- and metasoma entirely black, very strong mesopleural striations, and have a complete band of punctations in the posterior third of T2+3. Unlike the other species, S. fili- cornis has been reared from both a leaf gall, 236 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VOLUME 89, NUMBER 2 Andricus guatemalensis (Cameron), and a stem gall, 4. furnessana Weld. The life cycle of this species appears to be limited to one year and the rapid emergence of some spe- cimens may indicate that S. filicornis has more than one generation per year. Synergus kinseyi Ritchie and Shorthouse, NEw SPECIES Rigs 9 dl lose oe | Female (holotype).—Length 6.25 mm. Head yellow; frons above antennal sockets median third vertex, and median third of occiput black; antenna and mandible brown. Mesosoma black, legs except metacoxal bases yellow, wings hyaline, veins brown. Metasoma rufus except for dorsal margin of T2+3 and exposed dorsal margins of T4— T7 black. Head slightly wider than meso- soma; malar space 0.73 of eye height; pos- teroventral margin of gena with short, weak carina; frontal ridge absent; LOL = 0.8D, POL = 1.7D, OOL = 2.0D; antenna with 15 antennomeres, 3.88 mm long; Al-A15 Gnim)3+0542, .0.21..0.49; 0:31.°0.33...0.39, 032750225, 0:20.0:20.0.20,.0.17,,0:17 0:17, 0.20. Mesoscutum (Fig. 3) with weak trans- verse rugulae; notauli percurrent, grooves shining smooth, not interrupted by trans- verse rugulae, distinctly widened posterior- ly; median groove, lateral lines absent; an- terior lines slightly convex, extending about one third of mesoscutum. Mesoscutellum (Fig. 3) weakly rugose, median longitudinal depression and posterior upturned carina well developed; fovea well developed, dis- tinctly closed posteriorly. Mesopleuron (Fig. 7) aciculate above, weakly striate below. Metanotal medial fovea without longitu- dinal carina. Propodeum ruglose; supracox- — Figs. 9-18. 2351, al carina strong; propodeal carinae strong, not thickened, finely coriaceous. Metasoma (Fig. 11) longer than head plus mesosoma; T2+3 covering about three fourths of meta- soma, weakly excavated along posterodor- sal margin, posterior third glabrous. Fore- wing (Fig. 21) 5.25 mm long; radial cell length 3.0 times width; Rs2 weakly but dis- tinctly bent near wing margin; areolet ab- sent; hind wing (Fig. 18) 3.6 mm long. Tar- sal claw with a distinct tooth. Male (allotype).— Differs from holotype as follows: length 4.25 mm; malar space 0.53 of eye height; antenna 3.41 mm long; Al- AVS.(mm):0:3270591750:42.,0.2550.25; 025. 023; 0.22 02180210519) 0:17.02 16.0M15. 0.21; A3 (Fig. 15) slightly notched and bent, elongate beyond notch; anterior lines more distinct; median groove reduced to notch between raised ental margins of notauli; mesopleuron evenly, finely striate; supra- coxal carina weaker; propodeal carinae not as strong; T2+3 covering entire metasoma, dorsal black area larger, yellowish ventrally; radial cell length about 2.7 times width. Variation.— Female: Length 4.75-6.7 mm, average of 50 specimens = 5.8 mm; head sometimes amber; areolet absent or elongate, posterior margin very weak (both sexes); metasoma from rufus to dark rufus, dorsal black area sometimes expanded ven- trally. Male: Length 3.5—4.5 mm, average of 34 specimens = 4.2 mm; coloration similar to female except metasoma ventrally dark ru- fus to dark amber, dorsal black area some- times expanded. Type material.—Holotype, °, Huehue- tenango 3S, Guate., 7300’, ® 12.22.35, Q. pilicaulis, Kinsey coll., ex gall of ruginos., 9-12, Metasoma of Synergus spp., lateral view. Scale bar = 1.0 mm. 9, Synergus cultratus. 10, Synergus filicornis. 11, Synergus kinseyi. 12, Synergus mesoamericanus. 13-16, Male antennomere 3 (A3) of Synergus spp., lateral view. Scale bar = 0.1 mm. 13, Synergus cultratus. 14, Synergus filicornis. 15, Synergus kinseyi. 16, Synergus mesoamericanus. 17, 18, Hind wings of Synergus spp. Scale bar = 1.0 mm. 17, Synergus cultratus. 18, Synergus kinseyi. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 22 Figs. 19-22. Forewings (radial cell) of Synergus spp. 19, Synergus cultratus. 20, Synergus filicornis. 21, Synergus kinseyi (inset shows variant without areolet). 22, Synergus mesoamericanus. Kinsey det. Allotype, 3, with same data as holoytpe. Paratypes (86 2, 34 6: AMNH): 36 2, 12 6 with same data as holotype; 45 2, 22 6, Sacapulus 9S, Guate., 6000’, ® 12.28.35, ing. spr. 37, ex gall of brelandi; 5 2, (Label lost), Guatemala, 12.36, fall °36, ex gall of peredurus. Excluded from type se- ries: 133 specimens from above localities, ® 12.22—28.35, inq. fall °36—spr. ’37; in var- ious but poor condition; tentatively iden- tified as S. kinseyi. Etymology.—This species is named for Dr. A. C. Kinsey to commemorate his con- tribution to our knowledge of the Cynipi- dae. Diagnosis.—Synergus kinseyi is easily distinguished from other Guatemalan species by the following characters: areolet absent or elongate with the posterior margin very weak; mesopleuron entirely black; fe- male antenna with 15 antennomeres. This species is also very large, exceeded only by S. cultratus in size. Remarks.—Synergus kinseyi has been reared from three hosts and appears to be restricted to woody stem galls, two of which (brelandi and ruginos.) are the same as those for S. cultratus (see remarks for S. cultratus for problems in identifying the hosts). The third recorded host, Andricus peredurus Kinsey, produces a hard polythalmous stem gall. The presence of A. peredurus in Gua- temala is a major range extension, as this species previously known only from San Louis Potosi, Mexico (Kinsey, 1920). There is a contradiction in the labelling of the five specimens reared from the galls of A. peredurus, in that the label shows them being collected in December of 1936 and emerging in the fall of 1936. It is known that Kinsey collected in Guatemala during 1935 (Kinsey, 1936). Therefore, it is prob- able that these insects were collected in 1935 and emerged in the fall of 1936. This agrees with the dates of some of the specimens excluded from the type series. Less is known VOLUME 89, NUMBER 2 about the life cycle of this species than for other Guatemalan Synergus spp., as only one series has emergence dates. These data indicate that S. kinseyi has a one to two year life cycle. Synergus mesoamericanus Ritchie and Shorthouse, NEW SPECIES Figs. 4.8, 12,.16,.22 Female (holotype).—Length 3.4 mm. Coloration similar to S. kinseyi. Head slightly wider than mesosoma; malar space 0.57 of eye height; posteroventral margin of gena with moderately long carina; frontal ridge absent; LOL = 0.9D, POL = 1.7D, OOL = 2.3D; antenna with 14 antenno- meres, 2.75 mm long; Al-A14 (mm): 0.29, Ou22022920) 19803215 02150:215.0. 19,081.73 Oni7anOnli7= O215:. O52 .0:23: Mesoscutum (Fig. 4) with strong transverse rugulae (stronger than in other species); notauli strong, percurrent, bottom of grooves smooth, not interrupted by transverse ru- gae; median groove, lateral lines absent; an- terior lines convergent, extending about one third of mesoscutum. Mesoscutellum (Fig. 4) strongly rugulose, posterior of disc de- pressed, posterior upturned carina moder- ately developed; fovea large, deep anterior- ly, open posteriorly. Mesopleuron (Fig. 8) entirely striate, striations stronger ventrally. Metanotal median fovea without longitu- dinal carina. Propodeum with supracoxal carina weak ventrally, stronger dorsally; propodeal carinae moderately strong, not thick, finely coriaceous. Metasoma (Fig. 12) longer than head plus mesosoma; T2+3 covering about three fourths of metasoma, weakly excavated along posterodorsal mar- gin, posterior third smooth. Forewing (Fig. 22), 3.8 mm long; radial cell length 2.0 times width, Rsl very weak along anterior margin of wing (distinct in other species); Rs2 strongly bent near wing margin; areolet nor- mal; hind wing 2.4 mm long. Tarsal claw with a distinct tooth. Male (allotype).— Differs from holotype 239 as follows: length 3.7 mm; antenna 3.05 mm long, with 15 antennomeres, Al—A15 (mm): 0:35,015.0:352021002170:2170 2150 21¢ O21 Ou Oo Ons. -0 505152017243 (Fig. 16) notched and bent, not elongate af- ter notch; upturned carina on posterior mar- gin of scutellum not as strong; T2+3 cov- ering entire metasoma, black dorsally, dark rufus ventrally; forewing 3.8 mm long, hind wing 2.5 mm long; radial cell length 2.4 times width. Variation. — Female length 3.1-4.0 mm, average of 5 specimens = 3.7 mm; male length 2.75—-3.7 mm, average of 2 specimens 3.2 mm; head from yellow to amber; meso- scutum and mesoscutellum from black to dark rufus; mesoscutellum sometimes with fovea partly closed posteriorly, a weak me- dian depression, and posterior upturned ca- rina stronger; radial cell sometimes more elongate with Rs2 less strongly bent, ante- rior margin sometimes indistinctly closed. Type Material.—Holotype, 2°, Sacapulus 9S, Guate., 6000’, @ 12.28.35, ing. spr. °37, Q. pilicaulis, Kinsey coll., ex gall of brelan- di, Kinsey det. Allotype, ¢, with same data as holotype. Paratypes: 14 °, 4 6 with same data as holotype (AMNH). Excluded from type series: 3 specimens from type locality in various but poor condition; tentatively identified as S. mesoamericanus. Etymology.—The specific name means ‘‘middle America” and refers to this species distribution. Diagnosis. — Synergus mesoamericanus can be distinguished from all other Guate- malan species by the following characters: female antenna with 14 antennomeres, A3 1.5 times length of A4; radial cell length 2.0 times width, Rs2 distinctly bent; female metasoma rufus below, male metasoma en- tirely black. The radial cell appears to be open in many specimens as Rs| is often very weak along the anterior margin of the wing (Fig. 22). Remarks.—This species is one of the two smaller species of Synergus occurring in Guatemala. All the specimens examined 240 were reared from galls of ‘“‘brelandi” and emerged during the second spring after col- lection. Andricus guatemalensis (Cameron), NEw COMBINATION Cynips guatemalensis Cameron, 1883: 71, gall. TYPE: Holotype, gall (BMNH). Ex- amined. Andricus? mexicana Bassett, 1890: 78 (gall). Andricus mexicanus Bassett, Dalla Torre, 1893: 91. Andricus mexicanus Kinsey, 1920: 308, figs. 25-27 (2, 4, gall). New Synonymy. Synergus dorsalis Cameron, 1883: 72 (not Provancher, 1888: 398). (2, 3). New Syn- onymy. TYPE: Holotype, @ (BMNH). Examined. Remarks. — Cameron (1883) described the galls of C. guatemalensis which were sent to him from Guatemala. Bassett (1890) de- scribed the galls of Andricus? mexicana from galls collected in Mexico. Neither Cameron nor Bassett described what they thought was the gall inducer, although Cameron (1883) described Synergus dorsalis, which he thought was an inquiline in galls of C. gua- temalensis (see below). Kinsey (1920) was the first to describe the adult gall former, which he obtained by cutting from galls that had been sent to the American Museum of Natural History. Kinsey recognized that the galls he had were the same as those of C. guatemalensis Cameron and also those of Andricus mexicanus Bassett. However, Kinsey did not believe that names based on only the galls were valid as he described Andricus mexicanus as a new species while listing C. guatemalensis Cameron 1883 and Andricus mexicanus Bassett 1890 as syn- onyms. Thus, A. mexicanus Kinsey is a ju- nior synonym of both C. guatemalensis Cameron and A. mexicanus Bassett, and a junior homonym of A. mexicanus Bassett. Anew name for Andricus mexicanus Kinsey 1920 would needlessly burden the already confused nomenclature. Both Bassett and PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Kinsey were correct to assign the species to Andricus and C. guatemalensis is hereby transferred to this genus. The holotype of S. dorsalis (in the British Museum (Natural History)) belongs to the genus Andricus and is a synonym of Cynips guatemalensis Cameron. Although the type of guatemalensis is a gall while the type of dorsalis Cameron is an insect and it is de- sirable to base species of Cynipidae on the insect rather than the gall, we retain gua- temalensis as the valid name in the interest of stability. There are two reasons for doing this. First, the name guatemalensis has been associated with a gall-inducing species while dorsalis Cameron has been associated with an inquiline species. Second, retention of guatemalensis simplifies a problem of sec- ondary homonymy between S. dorsalis Cameron 1883 and S. dorsalis (Provancher) 1888. When Weld (1951) transferred Ceroptres dorsalis Provancher to Synergus, this species became a secondary homonym of S. dor- salis Cameron. However, Weld did not re- name S. dorsalis (Provancher) either be- cause he did not know that it was a homonym, or because he knew that S. dor- salis Cameron should not have been placed in Synergus. Weld visited the British Mu- seum (Natural History), after which he syn- onymized S. filicornis Cameron and S. fur- nessana Weld (Weld, 1930). It is quite likely that he also examined the type of S. dorsalis Cameron as well. A new name for S. dorsalis (Provancher) is unnecessary as S. dorsalis Cameron is now considered a junior syn- onym of Andricus guatemalensis Cameron and therefore the two species are no longer congeneric. ACKNOWLEDGMENTS We thank the following curators for mak- ing types and other material available for study: D. Azuma, Philadelphia Academy of Natural Sciences, Philadelphia, Pennsy]l- vania; A. Menke, Systematic Entomology Laboratory, Agricultural Research Service, VOLUME 89, NUMBER 2 USDA, United States National Museum, Washington, D.C.; J. Perron, Laval Uni- versity, Quebec, Quebec; J. Quinlan, British Museum of Natural History, London, United Kingdom; and R. T. Schuh, Amer- ican Museum of Natural History, New York, New York. The scanning electron micro- graphs were taken by Lewis Ling (Carleton University). This research was supported in part by an operating grant from the Natural Sciences and Engineering Research Council of Canada (No. A0230) awarded to J. D. Shorthouse. LITERATURE CITED Bassett, H. 1890. New species of North American Cynipidae. Trans. Entomol. Soc. Am. 17: 59-72. Cameron, P. 1883. Hymenoptera. Biologi Centrali- Americana, vol. 1, 497 pp., 120 Plates. Dalla Torre, K. W. 1893. Cynipidae. Catalogus Hy- menoptorum II. 140 pp. Dalla Torre, K. W. and J. J. Kieffer. 1910. Cynipidae. Das Tierreich 24: 1-891. Eady, R. D. and J. Quinlan. 1963. Hymenoptera: Cynipoidea. Handbooks for the Identification of British Insects, VIII (1a). Royal Entomological So- ciety, London. 86 pp. Kinsey, A. C. 1920. New species and synonymy of 241 American Cynipidae. Bull. Am. Mus. Nat. Hist. 42: 293-317. . 1930. The gall wasp genus Cynips. Ind. Univ. Studies 84-86: 1-577. 1936. The origin of the higher categories in Cynips. Indiana Univ. Pubs., Sci. Ser. No. 4: 1- 334. Provancher, L. 1888. Additions et Corrections au Volume II de la Faune de la Province de Québec. Hymenoptera. Family VI. Cynipidae, pp. 397-398. Ritchie, A. J. and T. M. Peters. 1981. The external morphology of Diplolepis rosae (L.). Ann. Ento- mol. Soc. Am. 74: 191-199. Shorthouse, J.D.and A.J. Ritchie. 1984. Description and biology ofa new species of Dip/olepis Fourcroy (Hymenoptera: Cynipidae) inducing galls on the stems of Rosa acicularis. Can. Entomol. 116: 1623- 1636. Weld, L. H. 1913. A new oak gall from Mexico. In- secutor Inscitiae Menstruus 1: 132-134. . 1930. Notes on types (Hymenoptera: Cynipi- dae). Proc. Entomol. Soc. Wash. 32: 137-141. 1951. Cynipoidea. Jn Muesebeck, C. F. W., K. V. Krombein, and H. K. Townes, eds., Hy- menoptera of America north of Mexico, United States Department of Agriculture, Monograph 2. 1420 pp. . 1952. Cynipoidea. Ann Arbor, Michigan. 351 pp. . 1957. Cynipid Galls of the Pacific Slope. Ann Arbor, Michigan. 80 pp. PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 242-243 A NEW SPECIES OF LEPTOMETOPA (DIPTERA, MILICHIIDAE) FROM MADAGASCAR POLLINATING CEROPEGIA (ASCLEPIADACEAE) CurTIS W. SABROSKY Systematic Entomology Laboratory, BBII, Agricultural Research Service, USDA, % U.S. National Museum NHB 168, Washington, D.C. 20560. Abstract. —Leptometopa nilssoni Sabrosky, new species, is described from Madagascar. Apparently this is the first published record of the genus from that island. A new species of the milichiid genus Lep- tometopa is described to make the name available for L. Anders Nilsson of the In- stitute of Systematic Botany, Uppsala Uni- versity, Sweden, who investigated the pol- lination of Ceropegia albisepta Jum. & H. Perr. (Asclepiadaceae) in Madagascar. Pol- linaria from this plant were found attached to the proboscis of flies entrapped in the trap flowers. This paper appears to be the first published record of Leptometopa from Madagascar. Leptometopa nilssoni Sabrosky, New SPECIES Polished black species, with black hal- teres and a pteropleural (anepimeral) bris- tle: Female.— Almost entirely shining black, except for white epistomal triangle on lower part of face, palpus, proboscis, and stalk of halter yellowish to brownish, halter knob dull black, mid and hind tarsi yellowish, and wing pale with yellowish to whitish-yellow veins. Head with frons highly polished but mi- croscopically striate; 2 interfrontal rows with weak interfrontal hairs in tiny punctures, otherwise scarcely evident; cheek polished, its height 4 that of an eye and less than breadth of third antennal segment, lower margin with a row of 5 long setae that in- crease slightly in length toward a slightly longer and stronger vibrissa; chaetotaxy as usual for genus: inner and outer vertical, postocellar, and ocellar pairs of bristles, and on each side of frons 2 lateroclinate orbital (upper) and 2 mesoclinate frontal (lower) bristles, all approximately same length and strength. Mesoscutum only moderately covered with hairs, in about 12 rows; scutellum bare; mesopleuron (anepisternum) with fine hairs on posterior portion; chaetotaxy: | humer- al, 1+1 notopleural, | supra-alar, | post- alar, 1 dorsocentral and | pteropleural (ane- pimeral) pairs of bristles, and | basal and 1 apical pairs of scutellar bristles. Hind tibia broadening from slender base to apex, at its widest about three times as broad as basal portion, distal half postero- dorsally with broad yellowish tibial organ. Wing as usual for genus, with two costal breaks, costa extending to 4th vein(M) slightly beyond apex of wing, crossveins well separated, and anal vein not extended be- yond anal cell; vein 4(M) converging slightly toward preceding vein (R 4+5), narrowing cell toward apical margin of wing and mak- ing the 4th section of costa slightly shorter than the 3rd. Length.—1.25-1.5 mm. VOLUME 89, NUMBER 2 Holotype and two paratypes.—All fe- males, Madagascar Centre; Mandraka, April 6, 1985 (L. A. Nilsson), in flowers of Cer- opegia albisepta. Type series deposited in the U.S. National Museum of Natural His- tory, by courtesy of the collector. The polished black head and body of L. nilssoni, so unlike the densely gray micro- tomentose type species, Leptometopa la- tipes (Meigen), at first glance suggest a species of the genus Madiza. However, there are polished black species 1n the genus, notably the Nearctic Leptometopa halteralis (Co- quillett). Moreover, some intermediate species neatly bridge the gap between pol- ished and microtomentose species. Lepto- metopa beardsleyi Hardy and Delfinado from the Hawaiian Islands has a polished black head but gray microtomentose thorax, although the latter is not as densely gray as in /Jatipes and therefore appears darker. Also, Leptometopa albipennis (Lamb) from the Seychelles has the head and most of the tho- rax polished black but the scutellum is 243 brownish microtomentose. The latter com- bination of characters is also found in an undescribed species from New South Wales, Australia, but its relationship to a/bipennis has not been studied. Incidentally, L. albi- pennis shows other differences from nils- soni, such as having the frons reddish an- teriorly, knob of halter whitish yellow, and wing very milky white. Males are not available, unfortunately, because the peculiarly broadened hind tibia in that sex is peculiar to the genus Lepto- metopa. However, the head of the female is characteristic of Leptometopa, with the lu- nule extending ventrad as a wedge-shaped divider between deeply-set antennae, its apex nearly touching the apex of the trian- gular epistomal area of the lower face. Fur- thermore, a pteropleural (anepimeral) bris- tle is present, a common, although not uni- versal, feature of Leptometopa. The specific name, a noun in the genitive case, 1s dedicated to L. Anders Nilsson, the botanist who collected the specimens. PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 244-249 CHILACIS TYPHAE (HETEROPTERA: LYGAEIDAE) AND THE SUBFAMILY ARTHENEINAE NEW TO NORTH AMERICA A. G. WHEELER, JR. AND JONATHAN E. FETTER Bureau of Plant Industry, Pennsylvania Department of Agriculture, Harrisburg, Penn- sylvania 17110. Abstract. —Chilacis typhae (Perris), an Old World lygaeid restricted to cattails (Typha spp.), is reported new to the Western Hemisphere based on collections from Delaware (1 county), Maryland (3), New York (3), and Pennsylvania (25). The known Nearctic dis- tribution is mapped, notes on seasonal history and habits in Pennsylvania are given, and morphological characters facilitating recognition of this immigrant heteropteran are pro- vided. Chilacis typhae (Perris) is an Old World artheneine lygaeid occurring throughout England but uncommon in Wales and Ire- land (Southwood and Leston, 1959). On the Continent it ranges from France east to Bul- garia, Rumania, and the European U.S.S.R., including Caucasus to the south, and north to Scandinavia; in Asia it has been recorded from Jordan (Slater, 1964). As the specific name implies, C. typhae is a specialist on plants of the genus Typha (Typhaceae), par- ticularly common cattail, 7. J/atifolia L. (called reedmace or bulrush in Britain). Jor- dan (1935) and Cobben (1953) discussed the habits of this seed-feeding bug; see Slater (1964) for additional references to papers containing biological information on C. ty- phae. The discovery of this lygaeid in Pennsyl- vania and its subsequent collection in Del- aware, Maryland, and New York represent the first Nearctic records for the species. The only other artheneine known from the New World is Polychisme ferruginosus (Stal). This Neotropical bug, recorded from Colombia and Venezuela, was recently removed from the Ischnorhynchinae and transferred to the Artheneinae, thus providing the first West- ern Hemisphere record and making the subfamily known from all major zoogeo- graphic areas except the Nearctic (Slater and Brailovsky, 1986). Here, we give collection data and map the known North American distribution of C. typhae; give notes on its seasonal history, relative abundance, and habits on cattail in Pennsylvania; and list adult and nymphal characters that allow recognition of this ad- ventive lygaeid in the Nearctic fauna. North American distribution.—The first known collection of C. typhae in the New World was made on 4 June 1986 (by JEF) near Lickdale, Pennsylvania. Further col- lecting showed that this species is well es- tablished in eastern Pennsylvania (but ap- parently scarce in the western counties) and present in southern and central New York and northern Delaware and Maryland. We did not encounter C. typhae during limited collecting in Ohio, western New York, and in more southern areas of Maryland (Fig. 1), or in Indiana or Virginia (negative sites not shown in Fig. 1). The following records are available for C. typhae in eastern United States. All collections were made in 1986 from Typha latifolia; voucher specimens VOLUME 89, NUMBER 2 Fig. 1. in North America. Closed circles represent 1986 collection sites; open circles, sites where cattail heads did not yleld C. typhae. have been deposited in the insect collections of Cornell University, Pennsylvania De- partment of Agriculture, U.S. National Mu- seum of Natural History, and University of Connecticut. DELAWARE: New Castle Co., e. of New- ark, 5 Nov., F. G. Stearns. MARYLAND: Allegany Co., nr. Flintstone, 30 July, A. G. Wheeler, Jr.; Frederick Co., Emmitsburg, 18 June, AGW; Washington Co., nr. Han- cock, 30 July, AGW. NEW YORK: Broome Co., s. of Kirkwood and nr. Castle Creek, 26 June, AGW; Tioga Co., e. of Caroline Center, 26 June, AGW; Tompkins Co., Be- semer, 26 June and s. of Ithaca, 29 June, AGW; Town of Ulysses, 9 mi. n. of Ithaca, 7 July, E. R. Hoebeke. PENNSYLVANIA: Adams Co., s. of East Berlin, 16 June, AGW Jet THRERS Map of the Middle Atlantic and New England States showing known distribution of Chilacis typhae and nr. Gettysburg and Hunterstown, 18 June, AGW; Berks Co., nr. Frystown and Rehrersburg, 17 June, AGW and nr. Ham- burg, 24 June, J. F. Stimmel; Bradford Co., Towanda, 7 Aug. AGW and n. of Stevens- ville, 7 Aug., R. J. Gallagher; Bucks Co., s. of Quakertown, 24 June, J. F. Stimmel; Car- bon Co., Nesquehoning, 3 July, AGW; Centre Co., 1-80 e. of Clearfield Co. line nr. Lanse, 10 June, AGW and nr. Martha Fur- nace, 20 July, JEF; Chester Co., Longwood Gardens, 5 Nov., F. G. Stearns; Columbia Co., e. of Bloomsburg, 5 Aug. and 29 Oct., AGW; Cumberland Co., nr. Wertzville, 16 June, AGW; Dauphin Co., nr. Harrisburg Area Community College, 9 June, AGW and Dauphin, 12 June, JEF; Delaware Co., Me- dia, 5 Nov., F. G. Stearns; Juniata Co., nr. 246 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Port Royal, 12 June, JEF; Lackawanna Co., nr. Clarks Summit, 24 June, R. J. Gallagher ande. of Factoryville, 26 June, AGW; Leba- non Co., I-81 at Exit 30, w. of Lickdale, 4 June and Rts. 22 and 934, nr. Harper Tav- ern, 11 June, JEF; e. of Fredericksburg, 17 June, AGW; Lehigh Co., e. of Fogelsville, 3 July, AGW; Luzerne Co., Frances Slocum St. Pk., 24 June, R. J. Gallagher and I-80 at junc. Rt. 309, nr. Freeland, 26 June, AGW: Monroe Co., Sciota, 3 July, AGW; Montour Co., n. of Danville, 5 Aug., AGW; Schuylkill Co., I-81 at junc. Rt. 125, nr. Ravine, 6 June and Barnesville, 3 July, AGW; Susquehanna Co., Great Bend, 26 June, AGW; Washington Co., nr. Bulger, 17 June, L. L. Garrett; Wayne Co., Beach Lake and s. of Hoadleys, 19 June, K. Valley; Westmoreland Co., Delmont, 2 Dec., L. L. Garrett; Wyoming Co., Overfield Twp. n.e. of L. Winola, 24 June, R. J. Gallagher and e. of Dixon, 26 June, AGW; York Co., s.w. of Kralltown, 16 June and s. of Dillsburg, 18 June, AGW. Chilacis typhae appears to be a relatively recent introduction in North America, even though its principal host, Typha latifolia, 1s naturally Holarctic (Gleason and Cronquist, 1963; Everett, 1982). The tribe to which C. typhae belongs, Artheneini, is considered a Palearctic element (Slater, 1964; Slater and Brailovsky, 1986). Claassen (1921) collect- ed the lygaeid Kleidocerys resedae (Panzer) during his work on cattail insects, and it 1s unlikely he would have overlooked C. ty- phae in his thorough survey of Typha, which was conducted mainly in the Ithaca, New York area where this bug is now established. Biological notes.—We observed C. fty- phae in the field and brought back cattail heads to the laboratory so that nymphs and adults could be extracted with a Berlese fun- nel. In early June we found adults and nymphs ofall stages in heads of the previous season. The bugs were restricted to old heads fluffed out by the feeding of the cosmopte- rygid Limnaecia phragmitella Stainton (Fig. 2) (see Claassen, 1921). Four or five heads collected in Schuylkill Co. on 6 June yielded 29 first, 27 second, 35 third, 67 fourth, and 28 fifth instars. Some old heads harbored several hundred adults and nymphs; cast skins often were conspicuous on heavily in- fested heads. The ischnorhynchine K. resedae also was present in old heads taken in several cattail colonies and occasionally occurred in the same head with C. typhae. Three such heads contained 200 individuals of C. typhae and 107 of K. resedae. The latter species feeds on seeds of numerous plants, particularly birch, rhododendron and other ericaceous shrubs, and cattail (see Wheeler, 1976). The interaction and possible competition be- tween these lygaeids of similar feeding hab- its (Sweet, 1960), although difficult to as- sess, seem worthy of study. By mid-July to early August fewer old heads contained large numbers of adults; nymphs, nearly all late instars, became scarce. During this time adults were ob- served in newly formed staminate spikes where they usually were concealed beneath the stamens. The staminate or upper por- tion of the fruiting head is not persistent, the stamens soon being shed, and adults are more likely to be detected on the new pis- tillate spikes. Mating occurred on the sur- face, the pairs typically clustered near white fluffy patches that mark tunnels of L. phrag- mitella (Fig. 3). Adults push their stylets through the dense flowers; during feeding the body is elevated and the abdomen held nearly perpendicular to the surface, as noted by Cobben (1953). The bugs also use lepi- dopteran tunnels to burrow into the new fruiting spikes, which are composed of nu- merous, dense minute flowers. Thus, larvae of the Holarctic moth L. phragmitella are important to C. typhae for permitting access to the interior of hard newly formed heads, and also in spinning silk to hold the downy material together so that seeds are retained; uninfested heads lose their seeds during winter and spring (Claassen, 1921). Other Lepidoptera are known to develop in heads VOLUME 89, NUMBER 2 247 Figs. 2, 3. Pistillate spikes of Typha latifolia. 2, Head of previous season fluffed out by the tunneling and feeding of larvae of the cosmopterygid Limnaecia phragmitella. 3, Current-season head with white, fluffly patches indicating new larval tunnels of L. phragmitella. of T. /atifolia in North America, but L. phragmitella “is the most common and the most abundant of the insects infesting the cat-tail” (Claassen, 1921). One even won- ders whether C. typhae could have become established in North America in the absence of the cosmopterygid. Chilacis typhae oviposits on the seed or pappus (Southwood and Leston, 1959). In early August we collected large numbers of nymphs in pistillate heads, mainly early in- stars of a presumed second generation. The heads contained adults, some of them tener- al, or adults and fourth and fifth instars in late October and early November; adults only were present at one site sampled in early December. Because we did not sample a cattail col- ony at regular intervals throughout a season, our phenological profile of C. typhae in east- ern North America is tentative. In England adults are known to overwinter, oviposition begins in late May or early June, and nymphs become adult by mid-July. There is a single annual generation, although in Central Eu- rope a partial second generation may be produced in autumn, the nymphs overwin- tering (Southwood and Leston, 1959). The adults collected during early June in Penn- sylvania may have been a combination of overwintered individuals and those of the season’s first brood since fifth instars were present. Collection of all nymphal stages, especially early instars, from new pistillate spikes in late July-early August suggests the beginning of a second generation; and the 248 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Adult habitus of Chilacis typhae, dorsal aspect. Scale line = 1.0 mm. presence of teneral adults and late instars in late October, the completion of a second brood. This presumed bivoltinism, with adults overwintering and occurring throughout the summer, contrasts with the life cycle Jordan (1935) outlined for C. typhae in Germany. He emphasized that because this lygaeid de- pends on fruit-bearing stands of cattail, its development takes place during the cold season. Copulation and some development occur in winter, and all nymphs become adult the following May. Only adults are found during summer when they are said to be inactive beneath cattails or rest on other marsh plants. In the Netherlands, however, Cobben (1953) reported that adults are abundant and mate during July and August. VOLUME 89, NUMBER 2 Additional study is needed to determine whether North American populations are bivoltine and whether only adults survive the winter. Recognition features.—The adult of C. typhae (Fig. 4) is 3.0 to 4.8 mm long, gla- brous, shining yellowish brown or ochreous and strongly punctured above, with the head and thorax fuscous beneath. It is easily rec- ognized by the juga that nearly attain the apex of the tylus, the latter extending be- yond the apex of antennal segment I; the prominent fuscous grooves separating tylus from juga; the nearly trapezoidal pronotum with sublaminately produced edges; and the dark scutellum with a pale V-shaped mark. Nymphs are a pale yellowish brown with orangeish or reddish markings (or some- times transverse reddish bands) on the ab- domen and have dorsal abdominal scent glands that open between tergites II-IV, IV-V, and V-VI; the anterior gland is re- duced (Usinger, 1938). Péneau (1909) and Collett (1927) illustrated the fifth instar of C. typhae, and Jordan (1935) described four of the five nymphal stages; his (and Col- lett’s) instar IV is actually V (Puchkov, 1958). For references to additional descrip- tions and illustrations of the immature stages of C. typhae the reader is referred to Slater’s (1964) catalog. The nymph of K. resedae can be distinguished from that of C. typhae by the brown or almost fuscous head and pronotum and dark red or reddish-brown abdomen, and by the well-developed an- terior scent gland. ACKNOWLEDGMENTS We are grateful to J. A. Slater (University of Connecticut) for confirming the deter- mination of C. typhae; R. J. Gallagher, L. ie, Garrett; Ro J; Henry, F. G. Stearns, J. F-. Stimmel, and K. Valley (PDA, BPI) and E. R. Hoebeke (Cornell University) for helping 249 collect cattail heads; and Slater, Valley, and Hoebeke for commenting on the manu- script. LITERATURE CITED Claassen, P.W. 1921. Typha insects: Their ecological relationships. Cornell Univ. Agric. Exp. Stn. Mem. 47. pp. 459-531. Cobben, R. H. 1953. Bemerkungen zur Lebensweise einiger hollandischen Wanzen (Hemiptera-Het- eroptera). Tijdschr. Entomol. 96: 169-198. Collett, H.R. P. 1927. The earlier stages of Chilacis typhae Perr. Entomol. Mon. Mag. 63: 155-157. Everett, T. H. 1982. The New York Botanical Garden illustrated encyclopedia of horticulture. Vol. 10, Ste-Zy. Garland Publishing, New York. pp. 3225- 3601. Gleason, H. A. and A. Cronquist. 1963. Manual of Vascular Plants of Northeastern United States and Adjacent Canada. D. Van Nostrand, Princeton, N.J. 810 pp. Jordan, K. H.C. 1935. Beitrag zur Lebensweise der Wanzen auf feuchten Béden. (Heteropt.). Stett. Entomol. Ztg. 96: 1-26. Péneau, J. 1909. Notules hémiptérologiques (3). I.— Sur la capture de Chilacis Typhae (Perris). Bull. Soc. Sci. Nat. Quest France (2) 9: 511-514. Puchkov. V. G. 1958. Larvae of Hemiptera-Heter- optera. I. Lygaeidae. Entomol. Rev. 37(2): 332- B58 Slater, J. A. 1964. A Catalogue of the Lygaeidae of the World. 2 vols. Univ. of Connecticut. Storrs. 1668 pp. Slater, J. A. and H. Brailovsky. 1986. The first oc- currence of the subfamily Artheneinae in the Western Hemisphere with the description of a new tribe (Hemiptera: Lygaeidae). J. N.Y. Entomol. Soc. 94: 409-415. Southwood, T. R. E. and D. Leston. 1959. Land and Water Bugs of the British Isles. Frederick Warne & Co., London. 436 pp. Sweet, M. H. 1960. The seed bugs: A contribution to the feeding habits of the Lygaeidae (Hemiptera: Heteroptera). Ann. Entomol. Soc. Am. 53: 317- 321. Usinger, R. L. 1938. Dorsal abdominal scent glands in nymphs of Lygaeidae. Pan-Pac. Entomol. 14: 83. Wheeler, A. G., Jr. 1976. Life history of Kleidocerys resedae on European white birch and ericaceous shrubs. Ann. Entomol. Soc. Am. 69: 459-463. PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 250-263 A REVISION OF THE NEARCTIC SPECIES OF ARTHROCERAS (DIPTERA: RHAGIONIDAE) DONALD W. WEBB Section of Faunistic Surveys and Insect Identification, Illinois Natural History Survey, Champaign, Illinois 61820. Abstract. —The genus Arthroceras Williston contains seven species and is Holarctic in distribution. This paper revises the four species recorded by Nagatomi (1966) for the Nearctic Region. Of these four species, one species and two subspecies are relegated to synonymy. The immature stages of this genus are unknown and little is known of its biology. Keys to species, with descriptions and distributions of each Nearctic species, are provided. The genus Arthroceras was erected by Williston (1886) and currently contains sev- en species (Nagatomi, 1966). It is Holarctic in its distribution, although it does not oc- cur in Europe. This paper revises the four species recorded by Nagatomi (1966) for the Nearctic Region. Of these four species, one species and two subspecies are relegated to synonymy. The immature stages of the ge- nus are unknown and little is known of its biology. The terminology used here follows McAlpine (1981) and for the male termi- nalia, Stuckenberg (1973). The range foreach measurement or ratio is followed by the av- erage. Arthroceras Williston Arthroceras Williston (1886: 107): Coquil- lett (1910: 510); James (1965: 298); James and Turner (1981: 486); Leonard (1930: 52); Nagatomi (1966: 44, 1970: 293, 1982: 142, 1984: 142); Nagatomi and Iwata (1976: 20). Type-species: A. pollinosum Williston by original description. Pseudocoenomyia Ouchi (1943: 493): Na- gatomi (1955: 57, 1966: 44). Type-species: P. sinensis Ouchi. Ussuriella Paramonov (1929: 181): Naga- tomi (1970: 293). Type-species: U. gadi Paramonov. Head in lateral view hemispherical. Ver- tex truncate, not emarginate lateral to ocel- lar tubercle. Ocelli dark red to black; ocellar tubercle subtriangular, distinctly raised above vertex in male, not raised above ver- tex in female. Eyes holoptic in male, dich- optic in female; facets in male smaller on ventral third, equal in female; setae absent or very short, scattered; median margin sin- uate, ventral half divergent; in lateral view eyes hemispherical to subtriangular. Frons reduced in male, broad in female, divergent dorsally; setae generally absent in male, scattered in female. Antenna (Figs. 1-6) shorter than length of head; scape in both sexes separated by distance greater than width of median ocellus, short, length less than width, shorter than pedicel, setae ab- sent; pedicel short, globose, shorter than wide, setae shorter than length of segment; flagellum subulate, longer than combined length of scape and pedicel, annulate, setae scattered. Gena broad; parafacial setae ab- sent. Clypeus broad, anterior surface dis- VOLUME 89, NUMBER 2 tinctly convex, forming deep lateral grooves; setae scattered. Maxillary palpus 2 seg- mented; basal segment cylindrical, longer than wide; apical segment cylindrical, curved ventrally, apex rounded, longer than wide, longer than basal segment. Thorax with dorsum rounded; vitta gen- erally indistinct; setae elongate, scattered, in no distinctive pattern. Postpronotal lobe concolorous with thorax. Postmetaspiracu- lar scale and suprametacoxal pit lacking. Scutellum with caudal margin broadly rounded. Laterotergite of postnotum with elongate setae. Wing (Fig. 14) longer than wide; mem- brane opaque; pterostigma variable; micro- setae minute, over entire membrane; setu- late dorsal on length of R,; thyridium absent. Costa circumambient, broader along ante- rior margin, setae fuscous, appressed. Hu- meral crossvein distinct. Subcosta ends dis- tad to middle of wing. Subcostal and marginal cells elongate, open. Radial sector originates from basal fourth of first basal cell, r-m situated above basal third of discal cell. R, ends distad to fork of R,,; and apex of discal cell. Fork of R,,; originates basad to apex of R,, distad to apex of discal cell, angle variable. Cell r, elongate, enclosing apex of wing. R, ends anterior to apex of wing. R,; ends posterior to apex of wing. M, and M, petiolate, contiguous or separate from discal cell. Cell m; open. M, and CuA, parallel. Posterior cells 5. Discal cell elon- gate, apical margin truncate. Posterior cubi- tal cell open. First basal cell elongate, apex truncate, ends distad to apex of second basal cell. Second basal cell emits 4 veins from apex. Anal lobe broadly rounded, right-an- gled. Alula rounded. Squama large; margin- al setae elongate, entire. Legs with tibial spurs 0-2-1. Hind coxa with distinct anterior tubercle. Empodium pulvilliform. Apical claws on tarsomere 5 fuscous, paired, simple. Hind legs not rap- torial. Abdomen with tergite | subrectangular, anterior margin broadly emarginate. Male 251 terminalia with tergite 8 broad, rectangular, length 1.5 times width, caudal margin trun- cate. Epandrium (Figs. 7, 15) with lateral and caudal margins rounded, anterior mar- gin broadly emarginate, length 1.5 times width. Tergite 10 absent. Cerci simple, flat- tened dorsoventrally. Ventral plate of proc- tiger subtriangular. Gonocoxite in ventral view (Figs. 8, 16) broad, lateral margins rounded, fused anteriorly with oval scler- otized median plate; in dorsal view (Figs. 9, 17) gonocoxite with narrow caudal arch joining inner margins, aedeagal apodeme elongate, extending anteriorly to anterior margin of gonocoxite. Gonostylus narrow, tapered apically, reflexed. Aedeagus com- posed of aedeagal sheath, endophallus, and endophallic hilts. Aedeagal sheath broad basally, fused to inner margin of gonocoxite, tapered caudally to form narrow endophal- lic guide. Endophallus elongate anteriorly forming endophallic apodeme, caudal third oval. Endophallic hilts separated medially, thick, heavily sclerotized, tapered anterior- ly. Penis valves and endophallic tines lack- ing. Female terminalia with tergite 9 (Fig. 19) quadrate, as long as wide, caudal margin truncate. Tergite 10 reduced, caudal margin truncate to broadly rounded. Cerci 2 seg- mented; basal segment broad, equal in length to apical segment; apical segment clavate with apical depression. Sternite 8 (Fig. 20) broad, widened caudally, 1.3 times longer than wide, caudal margin rounded with deep, median emargination. Sternite 9 greatly modified, invaginated beneath ster- nite 8 to form internal furca. Sternite 10 broad, membranous, caudal margin sinuate with median groove. Internal reproductive organs (Figs. 11, 21) with furca “Y” shaped, anterior apodeme short, not attached lat- erally to tergite 9. Common spermathecal duct short, trifurcating anteriorly to form 3 spermathecal ducts. Each spermathecal duct membranous, elongate, ending anteriorly in dark brown, spherical spermatheca (Figs. 11, 2A): Immature stages unknown. 252 KEY TO THE NEARCTIC SPECIES OF ARTHROCERAS 1. Length of apical flagellomere greater than 1.5 times width of flagellum (Fig. 6); setae on hind femur pale yellow, elongate, suberect, length greater than 0.5 times greatest width of femur Sgt eee Ree Ce ey pollinosum Williston — Length of apical flagellomere less than 1.5 times width of flagellum (Figs. 1-5); setae on hind femur predominately fuscous, short, ap- pressed, length much less than 0.5 times great- est width of femur . In males, maxillary palpus pale to dark yellow, setae predominately stramineous, apical seg- ment at least 1.5 times length of basal segment. In females, head, antenna, maxillary palpus, thorax, femora and abdomen yellow pollinose toxdarkavellowse se eeeeeey. fulvicorne Nagatomi — Inmales, maxillary palpus brown, setae black, apical segment about 1.2 times length of basal segment. In females, head, antenna, maxillary palpus, thorax, femora, and abdomen brown tovdarkibrownys--. 545.44: leptis (Osten Sacken) i) ARTHROCERAS FULVICORNE NAGATOMI Arthroceras fulvicorne Nagatomi (1966: 46). Arthroceras fulvicorne nigricapite Nagatom1 (1966: 48). NEw SyYNoNymMy. Arthroceras fulvicorne subsolanum Naga- tomi (1966: 49). NEw SYNONyYMy. Arthroceras subaquilum Nagatomi (1966: 59). NEw SYNONYMY. Arthroceras fulvicorne is separated from A. pollinosum by having the length of the apical flagellomere less than 1.5 times as wide as the flagellum and by having the se- tae on the hind femur predominately fus- cous, short, and appressed. Arthroceras ful- vicorne and A. leptis can be separated by the characters given in the key. Nagatomi (1966) separated the females of A. fulvicorne and A. subaquilum on the basis of the coloration of the antennal flagellum and the abdomen. At that time he examined four specimens of 4. subaquilum and 31 females of A. fulvicorne. In the 57 females of A. fulvicorne from northern Utah that I examined, the antennal flagellum varied from brown to black; the abdominal tergites were generally dark yellow and concolorous, but variation ranged from tergites 2—4 being PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON dark brown with the caudal margins dark yellowish brown to the entire abdomen being dark brown. On the basis of this variation, I have synonymized A. subaquilum with A. fulvicorne. Nagatomi (1966) separated A. fulvicorne fulvicorne and A. fulvicorne ni- gricapite on the coloration of the female head. In female specimens from California and Utah, the ground coloration of the head varied from fuscous to black. Because of this variation, I have synonymized A. fulvicorne nigricapite with A. fulvicorne. Nagatomi (1966) separated A. fulvicorne subsolanum and A. fulvicorne fulvicorne on the basis of the antennal flagellum being 7 or 8 seg- mented in A. fulvicorne fulvicorne and 5 or 6 segmented in A. fulvicorne subsolanum. Table 1 shows the variation in the number of completely separated flagellomeres (fla- gellum cleared in 10% KOH). Because of the variation in flagellum segmentation in specimens examined from Nova Scotia to British Columbia and California, I have synonymized A. fulvicorne subsolanum with A. fulvicorne. Male.—Length 6.2-8.2, 7.2 mm. Head with ocellar tubercle fuscous to black, prui- nosity light yellow grey; setae fuscous to Table 1. Segmentation and fusion in the female fla- gellum of Arthroceras fulvicorne. Flagella were cleared in 10% KOH. Distinct Flagel- lomeres Fusion Pattern of Location Flagellomeres 12S Ose 1-52, 354555 O52 [23 4 Seon Nova Scotia Vermont Utah Dds Bip Ilsa? bes 1+2,3+4, 5,6,7 1 2B NARS Ones 1+2,3+4, 5, 6,7 1253).4) 56-8 1+2,3+4, 5,6,7 British Columbia Washington California ~~ + oo MrAANANYANAYATNAAMNAAD & VOLUME 89, NUMBER 2 black, elongate, abundant. Eyes dark brown to black. Frons fuscous in ground color with dense silver pile. Antenna length 0.6-0.8, 0.7 times length of head; scape fuscous to dark brown, pruinosity light yellow, length 0.4—0.6, 0.5 times width, 0.5—1.0, 0.8 times length of pedicel; pedicel fuscous to dark brown, pruinosity light yellow, length 0.4- 0.8, 0.6 times width, setae fuscous, scat- tered; flagellum fuscous to brown, pruinos- ity light yellow, basal flagellomere (1, 2, or 1 +2) yellowish orange to dark yellow, length 3.0-4.1, 3.5 times width, 2.7-3.9, 3.3 times combined length of scape and pedicel, setae silver and fuscous, variable; annuli 7, apical flagellomere length 0.9-1.3, 1.1 times width of flagellum. Gena pruinosity dense silver grey; facial setae white to pale yellow, elon- gate, abundant. Clypeus fuscous in ground color, pruinosity dense silver grey; setae white to pale yellow, elongate, scattered on dorsal third. Maxillary palpus with setae pale yellow, elongate, abundant, occasionally with scattered black setae; basal segment generally pale to dark yellow, occasionally brown, length 1.8—2.2, 2.0 times width; api- cal segment pale to dark yellow, length 2.7- 3.3, 3.0 times width, 1.3-1.8, 1.5 times length of basal segment. Labellum fuscous, pruinosity light yellow; setae fuscous, mod- erately long, scattered. Postocular setae white to pale yellow, elongate, abundant, becoming shorter dorsally. Thorax fuscous to black in ground color, pruinosity light grey to yellowish grey; setae yellow, elongate, scattered; vitta indistinct, occasionally pale fuscous. Postpronotal lobe with setae yellow, elongate, abundant. Pleu- ron fuscous in ground color, pruinosity vari- able in density; setae pale yellow, elongate, abundant on propleuron, scattered on dor- sal two-thirds to half of anepisternum, scat- tered ventrally and dorsally on katepister- num, absent on anepimeron, scattered on ventral third and in caudolateral area of meron, scattered along caudal margin of metapleuron. Halter stalk dark yellow to dark yellowish brown, capitulum fuscous to 253 dark brown, pruinosity light grey; setae fus- cous, short, scattered. Scutellum fuscous, pruinosity yellowish grey; setae pale yellow, elongate, abundant. Postnotum fuscous, pruinosity grey; laterotergite fuscous to dark brown, pruinosity light yellowish grey; setae pale yellow, elongate, abundant. Wingas in A. pollinosum (Fig. 14). Length 6.5-7.5, 7.0 mm, width 2.3-2.8, 2.6 mm, length 2.7—3.0, 2.8 times width. Membrane pale brown; veins yellowish brown; ptero- stigma slightly darker than membrane. Fork of R,,; almost right angled. M,, M,, M; orig- inate separately from discal cell. Coxae fuscous, pruinosity grey, setae on forecoxa pale yellow, elongate, abundant; femora pale to dark yellow, glossy; tibiae variable from fuscous to dark yellowish brown, becoming fuscous apically; tarsi fus- cous. Empodia and pulvilli fuscous to dark brown. Abdomen fuscous in ground color, prui- nosity yellowish grey to brownish grey; setae fuscous, moderately long, scattered medi- ally, with white to pale yellow setae laterally on tergites | and 2. Terminalia (Figs. 7-10). Gonocoxite in ventral view with mediolat- eral enlargement on caudolateral projec- tion. Female.— Length 7.5-11.4, 9.4 mm. Head with ocellar tubercle dark yellow to black in ground color, pruinosity grey to yellowish grey; setae pale to dark yellow, elongate, scattered. Eyes dark red to fuscous. Frons variable, pruinosity greenish grey pollinose; setae white to dark yellow, short to elongate, scattered. Antenna (Figs. 1-4), length 0.7- 0.9, 0.8 times length of head; scape dark yellow, pruinosity light yellow grey, length 0.7-0.9, 0.8 times length of head; scape dark yellow, pruinosity light yellow grey, length 0.5—-0.7, 0.6 times width, length 0.7—1.0, 0.9 times length of pedicel; pedicel dark yellow, pruinosity light yellow grey, length 0.5-0.8, 0.6 times width, setae pale to fuscous, mod- erately long; flagellum brown to black, basal half of flagellomere 1 or 1+2 dark yellow, length 3.2—4.0, 3.7 times width, 2.5-3.8, 3.1 254 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-6. Female antenna, lateral view. 1-4, Arthroceras fulvicorne. 1, New Hampshire. 2, Utah. 3, Utah. 4, California. 5, Arthroceras leptis. 6, Arthroceras pollinosum. Abbreviations: (Flag) Flagellum, flagellomeres 1- 8. (Ped) Pedicel. (Sca) Scape. VOLUME 89, NUMBER 2 259 11 Figs. 7-11. Arthroceras fulvicorne. 7, Male epandrium, cerci, ventral plate of proctiger. 8, Male gonocoxite, ventral view. 9, Male gonocoxite, dorsal view. 10, Male endophallus, lateral view. 11, Female furca and sper- matheca. Abbreviations: (AA) Aedeagal apodeme. (C) Cerci. (E) Endophallus. (EH) Endophallic hilt. (Epa) Epandrium. (Gs) Gonostylus. (Gx) Gonocoxite. (VPP) Ventral plate of proctiger. 256 Fig. 12. Arthroceras fulvicorne, distribution. times combined length of scape and pedicel, setae on flagellomere | or 1+2 fuscous, short, appressed, on apical flagellomeres se- tae silver to pale yellow, very short, ap- pressed; apical flagellomere length 1.0-1.2, 1.1 times width of flagellum. Gena polli- nose, occasionally pruinosity dense grey; fa- cial setae white to pale yellow, elongate, abundant. Clypeus pollinose, occasionally pruinosity dense grey; setae white to pale yellow, elongate, scattered. Maxillary pal- pus dark yellow, pollinose, setae pale yel- low; basal segment length 1.1—2.2, 1.8 times width; apical segment length 3.1—4.2, 3.6 times width, 1.7-2.8, 2.1 times length of basal segment. Postocular setae pale yellow to golden, elongate, abundant, becoming shorter dorsally but reaching vertex. Thorax variable, pollinose, yellowish grey, or greenish yellow-grey; vitta variable, in- distinct, to faintly dark brown, to grey brown. Pleuron pale to dark yellow, prui- nosity light yellow grey; setae as in male. Halter pale to dark yellow brown. Scutellum dark yellow, pruinosity grey. Postnotum variable, dark yellow, occasionally with ventral fourth fuscous; laterotergite dark yellow, pruinosity grey. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Wing length 8.0-10.5, 9.1 mm, width 2.5— 3.8, 3.1 mm, length 2.8-3.4, 3.0 times width. Membrane pale yellow to pale yellow brown. Cell r, more elongate than in male. Coxae yellow to dark yellow, pruinosity light grey; femora dark yellow to brown; tarsi fuscous to black. Color pattern of abdomen highly vari- able, generally dark yellow, subshiny but in many specimens tergites 2—4 are dark brown, pruinosity grey, caudal margin dark yellow- ish brown, tergite 5 dark yellow, and oc- casionally the entire abdomen is dark brown; setae as in male. Terminalia as in A. polli- nosum (Figs. 19-20). Internal reproductive organs (Fig. 11). Furca length 0.40 mm. Spermatheca length 0.10 mm., 1.0 times width. Type material.—The holotype female of Arthroceras fulvicorne (CNC, no. 9100) was collected at Robson, British Columbia on 21 June, 1948 by H. R. Foxlee. The holo- type female of Arthroceras fulvicorne nigri- capite (USNM, no. 68171) was collected at Yosemite, California on 11 June, 1935 by A. L. Melander. The holotype female of Ar- throceras fulvicorne subsolanum (USNM, no. 68172) was collected at Mt. Washing- VOLUME 89, NUMBER 2 ton, New Hampshire, by A. T. Slosson. The holotype female of Arthroceras subaquilum (USNM, no. 68173) was collected at Banff, Alberta on 27 August, 1935 by A. L. Me- lander. Seasonal activity.— Adults of Arthroceras fulvicorne have been collected with aerial nets, in Malaise traps baited with dry ice (carbon dioxide), and at lights. In the col- lections examined adults were taken from 20 May until 20 August, with the majority of specimens collected in July. Females (129) were collected 2.7 times more often than males (47) with no evidence of protandry. Distribution (Fig. 12).— Arthroceras ful- vicorne is a widespread species found from Maine to western Ontario in the east and from Colorado to California north to Al- berta and British Columbia in the west. Specimens examined (176).—UNITED STATES: CALIFORNIA: Phillips Station (Placer County); Snowline Camp (Eldorado County); Blodgett Forest, 13 mi E George- town; Bumble Bee; Giant Forest Camp- ground, Sequoia National Park; Yosemite; Carson Pass (Alpine County). COLO- RADO: West slope of Loveland Pass, 9850’; Great Sand Dunes National Monument, 7600’; St. Louis Creek Campground, 3 mi SW Fraser, 8800’. IDAHO: Moscow Mt.; Long Valley, Alpha; Moscow; Chatcolet; Cub River Canyon, Thomas Spring; Bear Valley (Valley County). MAINE: Allagast Point, St. Francis. MICHIGAN: Pequam- ing; shore of Lake Superior, Marquette. MONTANA: Glacier National Park, Med- icine Lake; Glacier National Park, Lake MacDonald. NEW HAMPSHIRE: Mt. Washington; Mt. Alpine. NEW YORK: Mt. Slide (Ulster County), 3500’. OREGON: Crater Lake; Chief Joseph Mt., Joseph; Wallowa Lake; Velvet Creek, 28 mi SE Union; Ladd Canyon, 14 mi S La Grande, 4280’; Butte Lookout Station, 25 air mi E Medford; Lake of the Woods; Jordan Creek, 28 mi SSW La Grande, 4840’. UTAH: Lo- gan Canyon; Spring Hollow; Tony Grove Junction; Blacksmith Fork Canyon; Willard 251 Basin, 9300’; Monte Cristo; Mendon Cold Spring; Green Canyon; Smithfield Canyon; Logan; Millcreek Canyon; Kaller Hollow Camp, 22 mi NNW Vernel, 8900’. VER- MONT: Camels Hump, Bolton, 4100’. WASHINGTON: Nooksack River, Mt. Baker; Summerland Trail, Mt. Ranier; Gla- cier Peak Wilderness, 27 mi ESE Darring- ton; Moscow Mts.; Field Spring S. P., 4 mi S Anatone, 3500-4000’; Bald Knob Camp- ground, Mt. Spokane S.P., 4800-5200’; Mt. Ranier National Park. WYOMING: Elk Mountain; Old Faithful area, Yellowstone National Park; Trout Lake, 1 mi SW Round Prairie, Yellowstone National Park, 6900’; Grand Teton National Park; Bottle Creek Camp, 7 mi SW Encampment, 8800’: Low- er Green River Lake, Wind River Range; Old Faithful area, Yellowstone National Park, 7000-7500’; South Brush Creek Campground, Medicine Bow National For- est, 8000-8500’. CANADA: ALBERTA: Island Lake, Coleman, 4500’; Banff; Wa- terton; Island Lake, Coleman, 4500’; Mtn. View. BRITISH COLUMBIA: Robson; Hope Mts.; Terrace Mtn. NOVA SCOTIA: Mile 15, Highland Road. ONTARIO: Sud- bury; Macdiarmid; Burke Falls. Arthroceras leptis (Osten Sacken) Arthropeas leptis Osten Sacken (1878: 223). Arthroceras leptis (Osten Sacken) (1878: 223); Aldrich (1905: 213); James (1965: 298); Leonard (1930: 52). Arthroceras leptis is easily separated from A. pollinosum by the length of the apical flagellomere being less than 1.5 times as wide as the flagellum and the setae on the hind femur being predominately fuscous, short, and appressed. Arthroceras leptis and A. ful- vicorne can be easily separated by the char- acters in the key. Male.—Length 5.8-6.5, 6.2 mm. Head with ocellar tubercle fuscous to black, prui- nosity light grey; setae pale yellow to pale fuscous, elongate, abundant. Eyes brown to fuscous. Frons fuscous, pruinosity grey; se- 258 tae pale yellow to pale fuscous, elongate, scattered dorsally. Antenna length 0.46- 0.92, 0.67 times length of head; scape dark brown, length 0.3-0.6, 0.4 times width, 0.5- 0.8, 0.6 times length of pedicel, setae absent; pedicel length 0.5—0.8, 0.6 times width, se- tae pale yellow to pale brown, moderately long, subapical; flagellum length 1.9-3.9, 3.3 times width, 2.1—4.3, 3.0 times combined length of scape and pedicel, setae pale brown, scattered subapically; 7 annuli, apical flag- ellomere length 0.9-1.4, 1.2 times flagellum width. Gena with dense grey pruinosity; fa- cial setae pale yellow, elongate, abundant. Clypeus with dense grey pruinosity; setae pale yellow to gold, elongate, abundant. Maxillary palpus fuscous, pruinosity grey, setae black, elongate, abundant; basal seg- ment length 2.4—3.0, 2.5 times width; apical segment length 3.2—3.3, 3.2 times width, 1.1- 1.4, 1.2 times length of basal segment. La- bellum fuscous, pruinosity grey; setae fus- cous, elongate, scattered. Postocular setae pale yellow to gold, elongate, abundant ven- trally, becoming shorter dorsally, not reach- ing vertex. Thorax dark brown, pruinosity yellowish brown to grey; setae gold, elongate, abun- dant; vitta generally indistinct, occasionally dark brown, faint. Postpronotal lobe with setae brown, elongate, on anterior half. Pleura fuscous in ground color, pruinosity grey; setae dark yellow to gold, elongate, abundant on propleuron, on dorsal half of anepisternum, scattered dorsally and ven- trally on katepisternum, absent on anepi- meron, in caudodorsal patch on meron, scattered on dorsal half of metapleuron. Halter stalk dark yellowish brown, capitu- lum fuscous, pruinosity light grey. Scutellum fuscous, with grey pruinosity; setae brown, elongate, scattered. Postnotum fuscous, with dense grey pruinosity; setae dark yellow to brown, elongate, abundant. Wingas in A. pollinosum (Fig. 14). Length 5.7-6.3, 6.1 mm, width 1.8-3.0, 2.2 mm, length 2.2—3.2, 2.9 times length. Membrane pale brown; veins brown; pterostigma PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON slightly darker than membrane. M, and M, generally petiolate, occasionally originate contiguously from discal cell. Squama pale brown; setae pale yellow, elongate, entire. Coxae fuscous, pruinosity grey, setae gold, elongate, abundant; femora fuscous to dark brown, glossy or with pruinosity light grey; tibiae and tarsi pale brown to fuscous. Em- podia and pulvilla dark brown. Abdomen dark brown, pruinosity light grey; setae pale yellow, elongate, abundant. Terminalia as in A. pollinosum (Figs. 7-9). Female.— Length 7.7-8.9, 8.2 mm. Head with ocellar tubercle dark brown; setae gold- en, shorter than in male. Eyes brown. Frons fuscous, pruinosity pale brown, pruinosity on lateral and ventral margins silver to grey; setae gold, moderately long, scattered sub- apically. Antenna (Fig. 5), length 0.78—0.86, 0.82 times length of head; scapal length 0.3- 0.6, 0.5 times width, 0.4-1.0, 0.7 times length of pedicel; pedicel length 0.5—0.8, 0.6 times width; flagellum length 3.1-3.9, 3.5 times width, 2.8—4.4, 3.3 times combined length of scape and pedicel; apical flagello- mere length 0.8-1.3, 1.1 times flagellum width. Clypeal setae not as abundant as in male. Basal segment of maxillary palpus length 1.9-3.0, 2.4 times width; apical seg- ment length 2.7-3.1, 2.9 times width, 1.5- 2.7, 1.8 times length of basal segment. Post- ocular setae reach vertex. Thoracic setae shorter than in male. Vitta dark brown, separated by yellowish grey pruinosity. Metapleural setae scattered along dorsal margin. Wing length 7.0-8.0, 7.6 mm, width 2.7- 3.0, 2.9 mm, length 1.2—1.6, 1.3 times width. Veins M,, M,;, M; originate separately from discal cell. Terminalia and internal reproductive or- gans as in A. pollinosum (Figs. 19-21). Type material.—The lectotype female, here designated, of Arthropeas leptis (MCZ) was collected in the White Mountains, New Hampshire, by E. P. Austin. Seasonal activity.—In the collections ex- amined Arthroceras leptis was taken from 7 VOLUME 89, NUMBER 2 Fig. 13. Arthroceras leptis, distribution. July until 14 August. Females (19) were col- lected 1.4 times more often than males (14), with no evidence of protandry. Distribution (Fig. 13).—Arthroceras lep- tis is found from New Hampshire to Mich- igan in the east and from Oregon to Wash- ington in the west. Specimens examined (33).—UNITED STATES: MICHIGAN: St. Ignace. NEW HAMPSHIRE: Mt. Washington; White Mts. NEW YORK: Whiteface Mts., Adiron- dacks; Lake Tear (Essex County). ORE- GON: Mt. Hood. VERMONT: Jay Peak, 3400-3800’. WASHINGTON: Mt. Ranier; Sunrise, Mt. Ranier; Deer Lake near Chew- elah; 42 mi SE Randle; Bald Knob Camp- ground, Mt. Spokane S. P., 4800-5200’. CANADA: ALBERTA: Johnston Canyon, Banff. BRITISH COLUMBIA: Manning Park, Valle View. QUEBEC: Mt. Oxford. Arthroceras pollinosum Williston Arthroceras pollinosum Williston (1886: 108); James (1965: 298); Leonard (1930: 53, 1931: 322); Nagatomi (1966: 54). Leptis pruinosa Bigot (1887: 115); Leonard (1931: 322); Nagatomi (1966: 54). Arthroceras pollinosum is easily separat- ed from both A. fulvicorne and A. leptis by the length of the apical flagellomere being greater than 1.5 times the width of the fla- gellum and the setae on the hind femur being pale yellow, elongate, and suberect. Male.—Length 5.2-5.8, 5.6 mm. Head with ocellar tubercle fuscous to black, pru- inosity grey; setae stramineous to brown, elongate. Eyes fuscous. Frons fuscous with dense silver grey pile. Antenna fuscous, pruinosity light grey, length 0.84—0.95, 0.89 times length of head; scapal length 0.6-0.8, 0.7 times width, 0.8-1.0, 0.9 times length of pedicel, setae pale yellow, short; pedicel length 0.6-0.7, 0.7 times width, setae pale yellow, short; flagellum length 4.7-6.0, 5.3 times width, 3.5—4.3, 4.0 times combined length of scape and pedicel, annuli 7, apical flagellomere length 1.6-2.2, 1.9 times width of flagellum, setae fuscous, short, apical. Gena fuscous, pruinosity dense silver grey; facial setae white to pale yellow, elongate, abundant. Clypeus pruinosity dense silver grey; setae pale yellow, elongate, on lateral margins. Maxillary palpus fuscous, pru- inosity light grey, setae white to pale yellow, elongate, abundant; basal segment length 1.4-2.2, 2.0 times width; apical segment length 2.4—4.3, 3.3 times width, 1.1—2.1, 1.5 times length of basal segment. Labellum dark brown to fuscous, pruinosity grey; se- tae a mixture of short fuscous setae and elongate, pale yellow setae. Postocular setae pale yellow, elongate, abundant ventrally, absent dorsally. Thorax fuscous in ground color, pruinos- ity dense grey yellow; vitta indistinct; setae pale yellow, elongate, scattered. Postprono- 260 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 14-21. Arthroceras pollinosum. 14, Wing. 15, Male epandrium, cerci, ventral plate of proctiger. 16, Male gonocoxite, ventral view. 17, Male gonocoxite, dorsal view. 18, Male endophallus, lateral view. 19, Female terminalia, dorsal view. 20, Female terminalia, ventral view. 21, Female furca, spermatheca. Abbreviations: (C) Cerci. (F) Furca. (Sp) Spermatheca. (S8) Sternite 8. (S10) Sternite 10. (T9) Tergite 9. (T10) Tergite 10. VOLUME 89, NUMBER 2 tal lobe with setae pale yellow, elongate on anterior half. Pleura fuscous in ground col- or, pruinosity dense silver grey; setae pale yellow on propleuron, dorsal and caudal margin of anepisternum, scattered along dorsal margin of katepisternum, along dor- socaudal margin of meron, scattered dor- socaudally on metapleuron, absent on ane- pimeron. Halter stalk yellow to dark brown, capitulum fuscous. Scutellum with pru- inosity dense grey yellow; setae pale yellow, elongate, abundant. Postnotum with pru- inosity grey, laterotergite with pruinosity grey; setae pale yellow, elongate, abundant. Wing (Fig. 14) length 4.8-5.7, 5.2 mm, width 1.7—2.0, 1.9 mm, length 2.7—2.9, 2.8 times width. Membrane pale yellow; veins pale brown; pterostigma absent to pale brown. Fork of R,,; originates above or dis- tad to apex of discal cell. Squama and mar- ginal setae pale yellow. Coxa fuscous, pruinosity dense grey; fem- ora fuscous, pruinosity grey, setae pale yel- low, elongate; tibiae pale brown; tarsi fus- cous. Empodia and pulvilli fuscous. Abdomen fuscous in ground color, pru- inosity dense grey; setae pale yellow, elon- gate, abundant. Epandrium, cerci, and ven- tral plate of proctiger (Fig. 15). Gonocoxite in ventral view (Fig. 16); in dorsal view (Fig. 17). Endophallus (Fig. 17) flattened laterally in dorsal view, length 0.36 mm, elongate; in lateral view endophallus (Fig. 18) with anterior third expanded. Endophallic hilts (Fig. 17) thick, heavily sclerotized, tapered anteriorly, separated medially. Female.— Length 5.5—7.5, 6.7 mm. Head with ocellar tubercle pollinose; setae pale to dark yellow. Frons pollinose; setae pale to dark yellow, elongate. Antenna (Fig. 6) fus- cous, apical third of pedicel and basal half of flagellomere 1 + 2 yellow, length 1.0- 1.1, 1.1 times length of head; scapal length 0.6-0.8, 0.7 times width, 1.3 times length of pedicel; pedicel length 0.4—-0.6, 0.5 times width; flagellum length 5.3-6.0, 5.6 times width, 4.1-6.0, 4.9 times combined length of scape and pedicel, apical flagellomere 261 length 1.9-2.5, 2.2 times width of flagellum. Gena pollinose. Clypeus pollinose, setae pale to dark yellow, elongate, lateral. Maxillary palpus pollinose to fuscous, setae pale to dark yellow, elongate, abundant; basal seg- ment length 1.8—2.4; 2.1 times width; apical segment length 2.3-2.8, 2.5 times width, 1.5- 1.7, 1.6 times length of basal segment. Post- ocular setae pale to golden yellow, elongate, abundant, extending to vertex. Thorax pollinose, setae fuscous, elongate, with small fuscous spot at base of each. Postpronotal lobe with setae entire. Pleura with pruinosity dark yellow to grey. Halter dark yellow, pruinosity light grey. Scutellum dark yellow, glossy or pruinosity light grey. Postnotum dark yellow, pruinosity of me- dian area light grey. Wing length 6.0-7.0, 6.5 mm; width 2.2- 2.5, 2.4 mm; length 2.6-3.0, 2.8 times width. Coxa dark yellow, pruinosity grey; fem- ora dark yellow, glossy; tibiae and tarsi pale brown to fuscous, pruinosity light grey. Abdomen fuscous in ground color, pru- inosity dense yellowish grey. Terminalia in dorsal view (Fig. 19), in ventral view (Fig. 20). Internal reproductive organs (Fig. 21). Furca ““Y”’-shaped, anterior apodeme short. Spermatheca length 0.09 mm, width 0.10 mm. Type material. — The lectotype male, here designated, of Arthroceras pollinosum (SEM, no. 5846) has no locality data on the spec- imen. The lectotype male, here designated, of Leptis pruinosa (BMNH) was collected at Mount Hood, Oregon. Seasonal activity. — Adults of Arthroceras pollinosum have been collected at eleva- tions as high as 9800 feet, from Phacelia, with an aerial net, or in Malaise traps, and at lights. In the collections examined, adults were taken from 6 June until 27 July. Males (45) were collected 1.1 times more often than females (40), with no evidence of pro- tandry. Distribution (Fig. 22).—Arthroceras pol- linosum is a western species found from California and New Mexico north to Wy- 262 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 22. Arthroceras pollinosum, distribution. oming and Washington. A single female was collected in Wisconsin. Specimens examined (85).—UNITED STATES: CALIFORNIA: Smith River, Rowdy Creek; Prairie Creek; Weott; moun- tains near Orick; Dry Lagoon Beach State Park; Blacksburg Road; Arcata. COLO- RADO: St. Louis Creek Camp, 3 mi SW Fraser, 8800’; Aspen; Dolittle Ranch, Mt. Evans, 9800’; Chicago Creek, 8800’; Happy Hollow; Little Beaver; Clear Creek; 7 mi N Ward; Lake City, 9000’; Science Lodge; west end of Grand Mesa. NEW MEXICO: Ther- ma; Tajique; Hyde State Park, 8 mi NE San- ta Fe, 8700’; Santa Fe Campground near Santa Fe. OREGON: Robinson Butte Look Out, 25 air mi E Medford; Oregon Creek; Portland; Corvallis; Forest Grove; 14 miS Ranier; 20 mi E Waldport; Black Rock, 10 mi SW Dallas; Beverly Beach. UTAH: Kal- er Hollow Camp, 22 mi NNW Vernal. WASHINGTON: Canyon Creek; Seattle; VOLUME 89, NUMBER 2 Vancouver; Electron; 7 mi E Randle, Rt. 12. WISCONSIN. WYOMING: Bottle Creek Camp, 7 mi SW Encampment, 8800’; Lower Green River Lake, Wind River Range, 8000’. ACKNOWLEDGMENTS I thank L. M. Page, G. L. Godfrey, and W. E. LaBerge for reviewing this manu- script and E. Steger for her editorial com- ments. I thank the curators of the following institutions and collections for the loan of material relevant to this study: American Museum of Natural History, P. Wygodzin- sky; British Museum of Natural History (BMNH), J. Chainey; California Academy of Sciences, P. H. Arnaud, Jr.; Canadian National Collection (CNC), H. J. Teskey; Cornell University, L. L. Pechuman; Flor- ida State Collection of Arthropods, H. V. Weems, Jr.; Kansas State University, H. D. Blocker; Museum of Comparative Zoology, Harvard University; Ohio State University, C. A. Triplehorn; Oregon State University, J. D. Lattin; Royal Ontario Museum, G. Wiggins; W. J. Turner Collection; United States National Museum (USNM), W. Mathis; University of British Columbia, S. G. Cannings; University of California, Berkeley (California Insect Survey), E. I. Schlinger; University of California, Davis, R. O. Schuster; University of Colorado, U. N. Lanham; University of Georgia, C. L. Smith; University of Idaho, J. B. Johnson; University of Kansas, Snow Entomological Museum (SEM), G. W. Byers; University of Michigan, T. E. Moore; University of New Hampshire, D. S. Chandler; Univer- sity of Vermont, R. T. Bell; Utah State Uni- versity, W. J. Hanson; Washington State University, W. J. Turner. This paper is a contribution of the Illinois Natural History Survey supported in part by a grant from the University of Illinois Research Board and an Ernst Mayr Grant from the Museum of Comparative Zoology, Harvard Univer- sity. 263 LITERATURE CITED Aldrich, J. M. 1905. A Catalogue of North American Diptera (or Two-Winged Flies). Smithson. Misc. Coll. 46(1444): 1-680. Bigot, J. M. F. 1887 Diptéres nouveaux ou peu con- nus. Leptidi, Muscidi. Bull. Soc. Zool. France 12: 97-118. Coquillett, D.W. 1910. The type-species of the North American genera of Diptera. Proc. U.S. Natl. Mus. 27: 499-647. James, M. T. 1965. Family Xylophagidae, pp. 296- 298. In Stone, A. et al., eds., A Catalog of the Diptera of America North of Mexico. U.S. Dep. Agric., Agric. Handb. 276: 1-1696. James, M. T. and W. J. Turner. 1981. Family Rhagi- onidae. 33, pp. 483-488. In McAlpine, J. R. et al., eds., Manual of Nearctic Diptera. Res. Br., Agric. Canada Monogr. 27(1): 1-674. Leonard, M. D. 1930. A revision of the dipterous family Rhagionidae (Leptidae) in the United States and Canada. Mem. Am. Entomol. Soc. 7: 1-181. 1931. Some notes on my revision of the Rhagionidae (Diptera). Trans. Am. Entomol. Soc. 57: 321-323. McAlpine, J. F. 1981. Morphology and terminolo- gy—Adults. 2, pp. 9-63. In McAlpine, J. F. et al., eds., Manual of Nearctic Diptera. Res. Br., Agric. Canada Monogr. 27(1): 1-674. Nagatomi, A. 1955. A new genus and species of the dipterous family Coenomyiidae from Japan. Mu- shi 29: 57-60. . 1966. The Arthroceras of the world (Diptera, Rhagionidae). Pac. Insects 8: 43-60. . 1970. Ussuriella Paramonov, anew synonym of Arthroceras (Diptera, Rhagionidae). Mem. Fac. Agr. Kagoshima Univ. 7: 293. . 1982. Geographical distribution of the lower Brachycera (Diptera). Pac. Insects 24: 139-150. . 1984. Male genitalia of the lower Brachycera (Diptera). Beitr. Entomol. 34(1): 99-157. Nagatomi, A. and K. Iwata. 1976. Female terminalia of lower Brachycera. I. (Diptera). Beitr. Entomol. Berlin 26: 5-46. Osten Sacken, C.R. 1878. Catalogue of the Described Diptera of North America (2nd ed.). Smithson. Misc. Coll. 16(270): 1-276. Ouchi, Y. 1943. Diptera Sinica. Coenomyiidae 1. On a new genus belonging to the family Coenomyiidae from east China. Shanghai Sizenkagaku Kenkyu- sho Tho 13: 493-495. Paramonov, S. J. 1929. Dipterologische Fragmente. Trav. Mus. Zool. Kieff. 7: 181-182. Stuckenberg, B.R. 1973. The Athericidae, a new fam- ily in the lower Brachycera (Diptera). Ann. Natal Mus. 21(3): 649-673. Williston, S. W. 1886. On two interesting new genera of Leptidae. Entomol. Am. 2: 105-108. PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 264-268 A NEW SPECIES OF ERIXESTUS (HYMENOPTERA: PTEROMALIDAE), AN EGG PARASITOID OF CALLIGRAPHA POLYSPILA (COLEOPTERA: CHRYSOMELIDAE) IN ARGENTINA E. E. GRISSELL AND L. DE SANTIS (EEG) Systematic Entomology Laboratory, BBII, Agricultural Research Service, U.S.D.A., % U.S. National Museum NHB 168, Washington, D.C. 20560; (LDS) Director, Museo de La Plata, Paseo del Bosque, 1900 La Plata, Republica Argentina. Abstract. — Erixestus pachyneuron Grissell and De Santis, n. sp., is described from Ar- gentina. This species was reared from eggs of Calligrapha polyspila (Germar) (Chryso- melidae) collected on Sida rhombifolia L. (Malvaceae). The genus Erixestus heretofore has been known only from the type species, E. winnemana Crawford, from the north- eastern Nearctic, reared from eggs of Calligrapha spp. The two species of Erixestus are compared and the geographic range of E. winnemana 1s expanded westward to Idaho and New Mexico. Since its description in 1910, the genus Erixestus has been represented by a single species distributed in the northeastern Nearctic (Burks, 1979). The genus was de- scribed for the species winnemana (Craw- ford, 1910), an internal parasitoid of eggs of the chrysomelid genus Calligrapha. Among New World pteromalids, Erixestus iS unique as an internal egg parasite. Recent discovery of an undescribed species of Erix- estus in Argentina, also an internal egg par- asite of Calligrapha, suggests that the genus is both widespread and perhaps quite spe- cialized in its biology. The large number of Calligrapha species (ca. 100) and their ex- tensive geographic distribution throughout the New World (Blackwelder, 1944; Wilcox, 1972) suggest that Erixestus may be more widespread than present records indicate. The new species of Erixestus described below was collected by H. A. Cordo (U.S. Department of Agriculture, Biological Con- trol of Weeds Laboratory, Hurlingham, Ar- gentina) in connection with studies of Cal- ligrapha polyspila (Germar) on Sida rhombifolia L. in Argentina. The chryso- melid is being studied as a possible biocon- trol agent for Sida spinosa L. (Malvaceae), an introduced weed of southeastern United States. Erixestus pachyneuron Grissell and De Santis, NEw SPECIES Figs. 1-4 Female.— Body length 0.9 to 1.2 mm. Black with faint blue or green reflections on the head. Abdomen brownish black. Scape, antenna, mouth parts, tegula, and legs (in- cluding coxae) pale yellowish white. Wings hyaline with yellowish veins. Body polished except occiput, anterior part of mesoscu- tum, scutellum, and axillae faintly aluta- ceous. Propodeum polished except nucha reticulate, medially with 3 complete longi- tudinal carinae intersected by transverse ca- rina, partial longitudinal carinae present lat- erally. Head and thorax with sparse, but obvious VOLUME 89, NUMBER 2 265 Figs. 1, 2. Erixestus pachyneuron (female). 1, Forewing. 2, Antenna. setae (some 2 x or more as long as diameter of ocellus). Eyes glabrous. Midlobe of meso- scutum with irregularly placed long setae (ca. 8 to 16), sidelobes with 6 or 7, axillae each with 5 or 6, scutellum with 3 pairs. Hindcoxae and lateral regions of propo- deum covered with long white setae. Head as wide as thorax. Frontovertex width greater than eye height. Clypeus with anterior edge bilobed, malar distance about as long as longitudinal diameter of an eye, without genal suture. Ocelli in obtuse tri- angle, posterior ocellus at least 2 x own di- ameter from inner margin of eye. Dorsal edge of occiput rounded. Antennae inserted midway between median ocellus and free edge of clypeus; antennal proportions as shown in Fig. 2. Collar of pronotum with anterior edge vaguely carinate. Spiracle el- liptic, anterior border obscured by meta- notum which overhangs it. Wing with dis- tribution of setae as in Fig. 1. Ratio of 266 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Von. Figs. 3, 4. Erixestus pachyneuron (male). 1, Forewing. 2, Antenna. postmarginal : marginal veins 0.57 + SD = 0.05 (range 0.53-0.64, n = 10), of stigmal : marginal veins 0.48 + 0.03 (range 0.45— 0.50, n = 10). Tibial spur ca. half length of basitarsus. Petiole of abdomen short (scarcely visible), much wider than long. Gaster oval, nearly as long as head and tho- rax together; first three terga subequal in length, remainder together shorter than T1. Male.— Body length 0.7 to 1.1 mm. Sim- ilar to female, except body with intense green to bronze reflections. Sculpture of head and thorax more strongly developed than that of female. Antennal club less thickened (Fig. 4). Forewing marginal vein much thickened (Fig. 3). Ratio of postmarginal : marginal veins 0.60 + 0.05 (range 0.53-0.67, n = 10), for stigmal: marginal veins 0.44 + 0.04 (range 0.38-0.55, n = 10). Type material.—Holotype 2, allotype 4, VOLUME 89, NUMBER 2 32 6 and 23 2 paratypes from ARGEN- TINA, Buenos Aires Province, Dique Lu- jan, 10 February and 24 and 27 March 1976, H. A. Cordo, reared from egg mass of Cal- ligrapha polyspila on Sida rhombifolia; 40 6, 50 2 paratypes from ARGENTINA, Entre Rios Province, 40 km S. Gualenguaychu, 4 January 1976, H. Cordo, reared from eggs of Calligrapha sp. The holotype, allotype, and 36 paratypes are deposited in the Mu- seo de La Plata, La Plata, Argentina. One hundred and nine additional paratypes are deposited in the U.S. National Museum, Washington, D.C., the British Museum (Natural History), and the Canadian Na- tional Collection. Variation.—Specimens of Erixestus pachyneuron vary little in size or coloration except that males are slightly smaller than females and are metallic blue or green rather than black. The abdomens of specimens preserved in alcohol tend to have a brown or yellowish cast. Biology.—Erixestus pachyneuron has been reared from Calligrapha polyspila eggs on Sida rhombifolia. This species of Sida is widespread throughout the tropics (M. H. Sachet, Department of Botany, Smithsoni- an Institution, personal communication) and is associated with disturbed areas. Discussion. — Erixestus pachyneuron may be distinguished from E. winnemana by the following characters: in both sexes of pachy- neuron the postmarginal vein averages less than two-thirds the length of the marginal vein (0.59 + 0.05, n = 20), whereas in win- nemana these veins (Figs. 5, 6) are nearly equal (0.97 + 0.04, n = 20); in both sexes of pachyneuron the stigmal vein averages less than one-half the marginal vein (0.45 + 0.04, n = 20), whereas in winnemana the stigmal vein averages two-thirds the mar- ginal (0.66 + 0.4, n = 20); male pachyneu- ron have the marginal vein noticeably ex- panded throughout (cf. Fig. 3, pachyneuron with Fig. 6, winnemana); in both sexes of pachyneuron the antennae are white (dark brown to black in winnemana); in male 6 Figs. 5,6. Erixestus winnemana, forewing vena- tion. 5, Female. 6, Male. pachyneuron the flagellomeres are quadrate to transverse with short setae (Fig. 4), whereas in winnemana each flagellomere is longer than wide with long setae (Fig. 7). Erixestus winnemana Crawford, 1910 Figs. 5-7 Discussion.—This species was described from 10 specimens reared from eggs of Cal- ligrapha bigsbyana (Kirby) (now C. multi- punctata bigsbyana) and 12 specimens from C. scalaris (LeConte). All material was col- lected on Plummers Island, Maryland. At present there are a total of 16 syntypic spec- imens in the U.S. National Museum. As Crawford did not select a holotype at the time of description we herein designate and label a female as LECTOTYPE. The re- maining 15 specimens are paralectotypes. Distinguishing characters of this species are discussed under E. pachyneuron. Distribution.— This species is currently reported from Quebec, Maryland, and Vir- ginia (Burks, 1979). We have examined specimens from the following states which are all new records: New York, Illinois, Pennsylvania (ex Calligrapha spiraceae (Say)), Michigan, North Dakota (ex Cal- ligrapha scalaris), Kansas (ex Calligrapha scalaris), Wyoming (ex Calligrapha sp.), Idaho, and New Mexico. Erixestus winnemana might be expected to occur wherever its hosts do: Calligrapha scalaris is reported from throughout eastern 268 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 7. Erixestus winnemana (male). Antenna. United States and Canada from Quebec in the north to Georgia and Texas in the south; C. multipunctata bigsbyana occurs from Nova Scotia and British Columbia in the north, south to Oregon and Georgia; C. spi- raceae occurs from Maine and Michigan south to Pennsylvania (Wilcox, 1972). Biology. — This species is associated in the mid and eastern Nearctic with Calligrapha found on plants of northern hardwood for- ests: Calligrapha scalaris on Ulmus; C. m. bigsbyana on Salix, Tilia, and rarely Pop- ulus; and C. spiraceae on Physocarpus (Wil- cox, 1972). Western specimens do not pro- vide enough data from which to draw conclusions about host relations. ACKNOWLEDGMENTS We thank H. A. Cordo for specimens used in this study, Richard White for informa- tion on chrysomelid distribution and hosts, M. H. Sachet for information on the distri- bution of Sida, and Steve Heydon for his critical review of the manuscript. LITERATURE CITED Blackwelder, R. E. 1944. Checklist of the coleopter- ous insects of Mexico, Central America, the West Indies, and South America. Part 4. Bull. U.S. Natl. Mus. 185: 551-763. Burks, B. D. 1979. Pteromalidae, pp. 768-835. In Krombein, K. V. et al., eds., Catalog of Hyme- noptera in America North of Mexico. Vol. I. Sym- phyta and Apocrita (Parasitica). Smithsonian In- stitution Press, Washington, D.C. 1198 pp. Crawford, J.C. 1910. Three new genera and species of parasitic Hymenoptera. Proc. U.S. Natl. Mus. 38: 87-90. Wilcox, J. A. 1972. A review of the North American chrysomeline leaf beetles. Bull. N.Y. State Univ. 421: 1-37. PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 269-274 HOST-PLANT RELATIONS OF NATIVE UROPHORA SPP. (DIPTERA: TEPHRITIDAE) IN SOUTHERN CALIFORNIA RICHARD D. GOEDEN Department of Entomology, University of California, Riverside, California 92521. Abstract.—New (some initial) host-plant records are reported for Urophora caurina (Doane), U. formosa (Coquillett), U. rufipes (Curran), U. stenoparia Steyskal, and U. timberlakei Blanc and Foote. Urophora rufipes is initially reported from California. The host plants of these tephritids include one or more species of Acamptopappus, Chryso- thamnus, Grindelia, Gutierrezia, and Haplopappus, all noted to belong to the Subtribe Solidagininae of the Tribe Astereae of the Asteraceae. Synphagy among Urophora, Neaspi- lota, Procecidochares, Tephritis, and Trupanea species is documented and discussed. This is a Companion paper to one to be written by G. C. Steyskal (retired), System- atic Entomology Laboratory, Agricultural Research Service, USDA, % National Mu- seum of Natural History, Washington, D.C., who will treat the taxonomy of some of the specimens reported herein. I report here new information on the host-plant relations of several native Urophora spp. (Diptera: Te- phritidae) resulting from the past six years of field studies on nonfrugivorous fruit flies in southern California, that section of the state defined and treated botanically by Munz (1974). MATERIALS AND METHODS The materials and methods used in the sampling of mature flower heads of Aster- aceae and the rearing of Tephritidae from samples were described by Goeden (1985). Sweep net collections of adults limitedly supplemented these rearings. Identifica- tions of most flies mentioned in this report were confirmed or made by G. C. Steyskal. All host-plant identifications were con- firmed or made by Andrew C. Saunders, Curator of the Herbarium of the University of California, Riverside. The plant nomen- clature used is that of Munz and Keck (1959) and Munz (1974); the insect nomenclature, that of Steyskal (1979). RESULTS AND DISCUSSION Steyskal (1979) reviewed what little was known about the host plants of native North American species of Urophora. Except for the atypical species, U. acuticornis Steyskal reared from Lycium berlandieri Dunal (So- lanaceae), all known host plants of Neo- tropical Urophora are Asteraceae. Unlike the Palearctic Urophora, no North American Urophora has been reported from astera- ceous thistles. At least four Palearctic species have been introduced from Europe to Can- ada and the United States for the biological control of accidentally introduced, weedy thistles in the genera Carduus, Centaurea, and Cirsium (Steyskal, 1979; Julien, 1982). So far, no European species of Urophora has successfully been established in California for thistle control (Julien, 1982). Moreover, recent surveys of insects infesting the flower heads of native Cirsium thistles in northern California (Pemberton et al., 1985) and the above-ground shoots (including heads) in southern California (Goeden and Ricker, 270 1986b, 1987a, b) yielded no rearing records of native Urophora. It may be as Steyskal (1979, p. 25) has suggested, that “... the American species of Urophora, at least in part, may eventually be referred to other genera inasmuch as none of them seem to be very closely related to any Palearctic species, ...” I offer the following informa- tion on host-plant relations of native Uro- phora in southern California as a contri- bution to a better understanding of the biology and ecology of the genus. The fol- lowing treatment of flies is alphabetical by species. Urophora caurina (Doane).—Steyskal (1979) listed the host of U. caurina as the genus Grindelia. | never reared this species from a total of 13 samples of flower heads of five species of Grindelia that I have sam- pled since 1980. However, 2 6 of U. caurina were reared from a quantity of flower heads of Gutierrezia sarothrae (Pursh) Britton and Rusby collected in the Chihauhua Valley, NE San Diego Co., 11.xi.1982. Also, 8 é and 8 2 of U. caurina were reared from a quantity of flower heads of Haplopappus er- icoides (Lessing) Hooker and Arnott ssp. blakei C. B. Wolf collected near Orcutt, San- ta Barbara Co., 12.xi.1980. Both records represent new host-plant genera for this te- phritid. Urophora formosa (Coquillett).— Was- bauer (1972) listed Chrysothamnus viscidi- florus (Hooker) Nuttall, Grindelia campo- rum Greene, G. nana Nuttall, Grindelia sp., Haplopappus squarrosus Hooker and Ar- nott ssp. grindelioides (deCandolle) Keck, and H. venetus (Humboldt) Blake as hosts of U. formosa. In reviewing these host gen- era, Steyskal (1979) noted that some of these records may properly refer to U. caurina. I have reared the following specimens of U. formosa: 9 6 and 11 2? from flower heads of H. squarrosus spp. grindelioides collected above upper end of Kitchen Creek, Cleve- land Nat. Forest, San Diego Co., 9.1x.1980; 7 6 and 7 2 from flower heads of H. venetus prob. spp. furfuraceus (Greene) Hall col- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON lected in Proctor Valley, SW San Diego Co., 28.x.1981; 15 ¢and 11 2 reared from flower heads of H. venetus spp. vernonioides (Nut- tall) Hall collected NW of Temecula, SW Riverside Co., 15.x.1980; 1 ¢éand 5 2 reared from flower heads of H. venetus prob. spp. vernonioides collected at Cardiff-by-the-Sea, San Diego Co., 15.x.1980; 12 6 and 16 2 reared from a quantity of flower heads of H. venetus ssp. vernonioides collected near U.S. Navy facility on Santa Cruz Island, Santa Barbara Co., 8.x.1985. My rearing records definitely confirm Haplopappus as hosts of U. formosa; however, as with U. caurina, I have not reared U. formosa from any sample of Grindelia to date (including a sample of G. camporum). As the host rec- ords for U. formosa from Grindelia in Was- bauer (1972) are based on two or three sep- arate plant species and three independent sources and F. L. Blanc (in litt. 1986) has reconfirmed the record for G. camporum in Foote and Blanc (1963), this suggests a def- inite relationship under as yet undefined conditions with this fly. This host-plant re- lationship warrants additional study. The published ‘“‘unpublished”’ host record for C. viscidiflorus in Wasbauer (1972) remains unconfirmed. It may properly refer to a sweep record or, perhaps, to U. timberlakei Blanc and Foote (see discussion below). Urophoran. sp. Steyskal.—To complicate matters further, the only tephritid that I have reared from Grindelia to date in southern California apparently is undescribed (Stey- skal, in litt. 1986). The sole host plant found to date for the robust individuals of this species appears to be G. hallii Steyermark, a plant species confined to dry flats in the Cuyamaca Mountains, San Diego Co. (Munz, 1974), and especially common around Lake Cuyamaca, where mature flower heads sampled in quantity on 9.1x.1980 and 17.vi1.1985 yielded only 2 4 and 4 2 and 2 éand | °, respectively. Flower heads sampled in bulk on 2.vii, 31.vii, and 14.vii.1985 from the same area yielded no flies. Factors involved in oviposition site VOLUME 89, NUMBER 2 271 Table 1. Synphagy among Tephritidae reared with Urophora from samples of mature heads of Asteraceae from southern California 1980-1985. Tephritid Genera (No. Species) Total No. (%) Adults Reared Reared Host Plant* Sample Date? Acamptopappus shockleyi 27 V 1982 A. sphaerocephalus 21 V 1982 Chrysothamnus nauseosus 3 IX 1982 C. viscidiflorus 3 IX 1981 Grindelia hallii 9 IX 1981 17 VII 1985 Gutierrezia microcephala 9 IX 1981 Gutierrezia sarothrae 18 IX 1980 11 X 1982 Haplopappus acradenius 8 XII 1983 28 XI 1984 11 XII 1984 H. cuneatus 16 X 1980 H. laricifolius 21 X 1982 H. squarrosus 9 IX 1980 H. venetus 15 X 1980 15 X 1980 28 X 1981 8 X 1985 @ See text for complete rearing record for Urophora. selection will be studied along with other aspects of the life history of this tephritid beginning in 1987. This species is especially suitable for a life history study among Neo- tropical Urophora because apparently no other species of Tephritidae infests the flow- Neaspilota 13 (11) Trupanea (2) 71 (58), 21 (17) Urophora 17 (14) Neaspilota 3 (11) Trupanea 20 (71), 2 (7) Urophora 3 (11) Neaspilota 1 (6) Procecidochares 3 (18) Tephritis 6 (35) Trupanea (2) 4 (24), 2 (12) Urophora 1 (6) Trupanea 7 (9) Urophora 73 (91) Urophora 6 (100) Urophora 6 (100) Urophora 4 (100) Urophora 2 (100) Urophora 2 (100) Neaspilota 2 (67) Urophora 1 (33) Trupanea 8 (62) Urophora 5 (38) Neaspilota 1 (14) Urophora 6 (86) Tephritis 33 (72) Trupanea 4 (9) Urophora (2) Si) se) Neaspilota 25 (63) Urophora 15 (37) Neaspilota 13 (21) Trupanea 30 (48) Urophora 20 (32) Trupanea 3 (10) Urophora 26 (90) Tephritis 1 (6) Trupanea (2) 8 (47), 2 (12) Urophora 6 (35) Trupanea (2) $iG2); 31(1:2) Urophora 14 (56) Trupanea 2 (7) Urophora 28 (93) er heads of G. hallii (Table 1); whereas, as discussed below, the other species of Uro- phora commonly are synphagous with other genera of Tephritidae. Urophora rufipes (Curran).—The single of U. rufipes reared from a small quantity 272 of flower heads of Haplopappus acradenius (Curran) prob. spp. eremophilus (Greene) Hall collected at Mountain Springs, SW Im- perial Co., 8.x1i.1983, represented the first host-plant record for this species (Was- bauer, 1972; Steyskal, 1979). Since then I have reared 4 6 and 2 2 and 4 éand 1 ? from quantities of flower heads of H. acradenius collected along the south shore of Clark Dry Lake, SE San Diego Co., 11.xii.1984, and along the Coachella Canal above the ther- mal springs area, Riverside Co., 28.x1.1984, respectively. These also are the first Cali- fornia records of U. rufipes, known previ- ously only from Arizona (Steyskal, 1979). Urophora stenoparia Steyskal.—Similar- ly, | éand 1 2 of U. stenoparia reared from a quantity of flower heads of Gutierrezia sarothrae collected at Pine Valley, Cleve- land Nat. Forest, San Diego Co., 18.1x.1980, represented the first host-plant record for this species (Steyskal, 1979). I additionally have reared 3 6 and | 2 from flower heads of Gutierrezia microcephala (deCandolle) Gray and 1 2 from the sample of flower heads of H. cuneatus Gray along with the 8 specimens of U. timberlakei Blanc and Foote reported below. I also swept 3 6 and | 2 of U. stenoparia from Hymenoclea_ salsola Torrey and Gray during extensive field studies of the insect fauna of this common desert shrub (Goeden and Ricker, 1986a), which is not a host plant of this tephritid. Nineteen additional species of Tephritidae were swept from H. salsola, only one species of which infests the flower heads or other- wise reproduces on this common desert shrub (Goeden and Ricker, 1986, unpub. data). Urophora timberlakei. —Wasbauer (1972) listed Gutierrezia microcephala (de- Candolle) Gray as a host plant of U. tim- berlakei, and Steyskal (1979) cited this ge- nus and Chrysothamnus as hosts. I have reared 10 6 and 7 2 of U. timberlakei from flower heads of Acamptopappus shockleyi Gray, collected at the SE end of Kingston PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Mountains, NE San Bernardino Co., 27.v.1982; 2 6and 1 2? from flower heads of A. sphaerocephalus (Harvey and Gray) Gray collected at Snow Creek, Riverside Co.., 21.v.1982; 1 2 from flower heads of C. nau- seosus (Pallas) Britton collected in Landers Meadow, Sequoia Nat. Forest, Kern Co., 3.1x.1981; 1 6 and 2 2 from flower heads of C. teretifolius (Durand and Hilgard) Hall collected in Westgard Pass, Inyo Nat. For- est, Inyo Co., 9.1x.1986; 31 6 and 42 2 and 3 6 and 3 2 from flower heads of C. visci- diflorus also collected in Landers Meadow on 3.ix.1981 and at Antelope Spring, NE Inyo Co., 15.i1x.1982, respectively; 4 6 and 4 2 from flower heads of Haplopappus cu- neatus collected in Lark Canyon, San Diego Co., 16.x.1980; and 8 6 and 7 2 from flower heads of H. laricifolius Gray, 1 km NW of Kessler Peak, S end of Ivanpah Mountains, NE San Bernardino Co., 21.x.1982. All my rearing records are for new host species. Two new host genera also are represented. The record for G. microcephala in Wasbauer (1972), originating from a host listing of R. H. Foote, but noted only as a sweep record by Blanc and Foo.e (1961) and Foote and Blanc (1963), still lacks confirmation and is suspect. My field observations and rearing data suggest that all Urophora species native to southern California encountered to date are flower head-infesting, seed-feeding species, with the exception of U. acuticornis, which doubtfully belongs in the genus, judging from its distinctive host-plant affinities and at- tendant mode of development. Steyskal (1979) suggested that U. acuticornis likely will be referred to a distinct genus when more is known about American Myopitin- ae. Allen L. Norrbom (in litt. 1986) states that U. acuticornis not only is not a Uro- phora or myopitine, but rather 1s a trype- tine. I have detected no sign of galls on the excised compound inflorescences that mainly constituted the flower head samples VOLUME 89, NUMBER 2 of Chrysothamnus, Gutierrezia, and Hap- lopappus reported above. Nor have I noted any enlarged flower heads in these and other host-plant genera mentioned in this report that are symptomatic of some other gallico- lous Tephritidae, e.g. Procecidochares in Chrysothamnus (Table 1). Of evolutionary and taxonomic signifi- cance is my observation that all of the con- firmed host plants of the southern Califor- nia Urophora mentioned in this report belong to the Subtribe Solidagininae of the Tribe Astereae (Munz and Keck, 1959). Thus, like so many Eurasian Urophora con- fined to hosts in the Tribe Cynareae, Sub- tribes Carduinae and Centaurinae, these southern California Urophora show a com- mon affinity for a definable group of host plants in the Asteraceae (Zwo6lfer, 1965; Steyskal, 1979). No life history of any Nearctic Urophora has been published. One difficulty involved is illustrated in Table 1. In southern Cali- fornia, at least, flower heads of Urophora host-plant species at most locations were commonly infested with other Tephritidae (synphagy). Genera commonly associated with Urophora are Neaspilota, Tephritis, and Trupanea. Rarely was more than one species of Urophora reared from a single sample. Associated genera differed qualitatively and quantitatively among some host-plant pop- ulations sampled, e.g. Haplopappus acra- denius and H. venetus (Table 1). Urophora commonly were reared in small numbers and usually have been poorly represented in my sweep collections, reflecting, perhaps, their low population densities in nature. In some flower head samples, however, Uro- phora outnumbered at least one associated tephritid species; in Chrysothamnus visci- diflorus, overwhelmingly so (Table 1). Sam- ples from two different species of Acamp- topappus appeared similarly composed. The undescribed Urophora apparently lacks te- phritid competitors for heads of Grindelia hallii; whereas, Gutierrezia heads yield 273 another genus of Tephritidae, 1.e. 7rupanea (Goeden, 1985). Much remains to be learned about the host-plant relations of these and other Nearctic Urophora. ACKNOWLEDGMENTS My thanks to George Steyskal for his taxonomic help noted in the text, to Louis Blanc, Dick Foote and Al Norrbom for their reviews of early drafts of this paper, and to Don Ricker for his patience during stops to allow my many collecting forays. LITERATURE CITED Blanc, F. L. and R. H. Foote. 1961. A new genus and five new species of California Tephritidae. Pan- Pac. Entomol. 37: 73-83. Foote, R. H. and F. L. Blanc. 1963. The fruit flies or Tephritidae of California. Bull. Calif. Insect Surv. T.A1S pp: Goeden, R. D. 1985. Host-plant relations of Tru- panea spp. (Diptera: Tephritidae) in southern Cal- ifornia. Proc. Entomol. Soc. Wash. 87: 564-571. Goeden, R. D. and D. W. Ricker. 1986a. Phytoph- agous insect fauna of the desert shrub Hymenoclea salsola in southern California. Ann. Entomol. Soc. Am. 79: 39-47. 1986b. Phytophagous insect faunas of the two most common, native Cirsium thistles, C. cal- ifornicum and C. proteanum, in southern Califor- nia. Ann. Entomol. Soc. Am. 79. (In press.) 1987a. Phytophagous insect faunas of the native thistles, Cirsium brevistylum, C. congdonii, C. occidentale, and C. tioganum, in southern Cal- ifornia. Ann. Entomol. Soc. Am. 80. (In press.) . 1987b. Phytophagous insect faunas of native Cirsium thistles, C. mohavense, C. neomexican- um, and C. nidulum, in the Mojave Desert of southern California. Ann. Entomol. Soc. Am. 80. (In press). Julien, M. H. (ed.) 1982. Biological control of weeds: A world catalogue of agents and their target weeds. Commonw. Agric. Bur., Commonw. Inst. Biol. Control, Farnham Royal, Slough. 108 pp. Munz, P. A. 1974. A Flora of Southern California. Univ. Calif. Press, Berkeley, Los Angeles, London. 1086 pp. Munz, P. A. and D. D. Keck. 1959. A California Flora. Univ. Calif. Press, Berkeley and Los An- geles. 1681 pp. Pemberton, R. W., C. E. Turner, and S. S. Rosenthal. 1985. New host records for tephritid flies (Dip- tera) from Cirsium and Saussurea thistles (Aster- 274 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON aceae) in California. Proc. Entomol. Soc. Wash. tera: Tephritidae). Calif. Dep. Agric. Bur. Ento- 87: 790-794. mol. Occas. Pap. 19. 172 pp. Steyskal, G.C. 1979. Taxonomic studies on fruit flies Zwolfer, H. 1965. Preliminary list of phytophagous of the genus Urophora (Diptera: Tephritidae). insects attacking wild Cynareae (Compositae) in Entomol. Soc. Wash. Misc. Pub. 61 pp. Europe. Commonw. Inst. Biol. Control Tech. Bull. Wasbauer, M. W. 1972. An annotated host catalog 6: 81-153. of the fruit flies of America north of Mexico (Dip- PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 275-283 NEOTROPICAL TINEIDAE, IV: THREE NEW ACROLOPHUS SPECIES FROM CUBA AND THE REDISCOVERY OF ACROLOPHUS NIVEIPUNCTATUS WALSINGHAM (LEPIDOPTERA) DONALD R. DAvis Department of Entomology, National Museum of Natural History, Smithsonian Insti- tution, Washington, D.C. 20560. Abstract.— Examination of a small collection of Acrolophus in the collections of the Instituto de Zoologia in Havana, Cuba, has revealed the presence of three new species (A. fuscisignatus, A. guttatus, and A. basistriatus) and the first specimens collected of A. niveipunctatus Walsingham since 1891. All species are fully described and illustrated. In 1980, the Academia de Ciencias de Cuba and Smithsonian Institution signed an agreement designed to promote cooperative research in the natural sciences. A vital part of this agreement encourages an exchange of scholars between our two countries. In February 1981, I was privileged to be one of the first entomologists to participate in this program. Although February was certainly not the most opportune season for Lepidoptera, a respectable sample of specimens, including 28 species of Tineidae, was collected at five sites in western Cuba over an | 1 day period. The purpose of this brief account is not to report upon what is hoped to be only the first of several trips to Cuba, but instead upon a small but interesting series of Ac- rolophus found in the collections of the In- stituto de Zoologia in Havana. Most of the specimens were collected by Pastor Alayo and associates; all but one species were pre- viously undescribed; and none was encoun- tered during my brief excursion. Only five species of Acrolophus have been described from Cuba (Davis, 1984). Be- cause the unique holotypes of four of these (A. dimidiella Wlsm., A. niveipunctatus Wlsm., A. noctuina Wlsm., A. vitellus Poey) are now lost, their identities have been un- clear. Fortunately, one of the four species found in the collections of the Instituto de Zoologia has been determined as conspe- cific with A. niveipunctatus. At present, these represent the only examples of this species collected in nearly a century, which merely indicates again how poorly known the Cu- ban microlepidoptera are. It is further in- teresting to note that all of the species treat- ed herein, with the exception of A. fuscisignatus, new species, are members of the North American p/umifrontellus group. This group, which was previously known to contain only A. plumifrontellus Clemens (Hasbrouck, 1964), is characterized pri- marily by the bifid apex of the male valva. Institutional acronyms referred to in this paper are: IZAC for Instituto de Zoologia, Academia de Ciencias de Cuba, Havana; and USNM for National Museum of Nat- ural History (formerly United States Na- tional Museum), Smithsonian Institution, Washington, D.C., USA. Acrolophus fuscisignatus Davis, NEw SPECIES Figs. 1-3, 7-10, 23 Adult (Figs. 1-3).— Length of forewing: 4, 10.0-14.5 mm; @ 15-17 mm. A relatively large species with strongly recurved labial 276 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-6. Adults. 1, Acrolophus fuscisignatus, paratype male, Pico Turquino, length of forewing 17.7 mm. 2, A. fuscisignatus, paratype male, Moa, length of forewing 10 mm. 3, A. fuscisignatus, paratype male, Pico Turquino, length of forewing 15.6 mm. 4, A. guttatus, holotype male, Finca La Ciega, length of forewing 12 mm. 5, A. niveipunctatus, male, Hongolosongo, length of forewing 13.9 mm. 6, A. basistriatus, holotype male, Cuabal de Gulindo, length of forewing 12.5 mm. VOLUME 89, NUMBER 2 palpi in the male. The forewing is white, variously marked with fuscous. Head: Vestiture mostly white with a fringe of fuscous hairs around eyes. Eye smooth with long dark lashes from posterior rim. Antenna approximately 0.4—-0.5 the length of forewing, 61-64 segmented; scape en- tirely white; flagellum subserrate with gray- ish-white scales dorsally and often indis- tinctly banded with light brown; naked ventrally except for dense, very short whit- ish sensory setae. Labial palpus 3 segment- ed, extremely long in male and recurved over thorax to metanotum; relatively short in female, length about 2 x diameter of eye and porrect; vestiture mostly white with lat- eral surfaces of basal segment dark fuscous. Thorax: Pronotum mostly white, some- times with heavy suffusion of fuscous an- teriorly and medially. Venter white. Fore- wing white, variously marked by fuscous as shown in Figs. 1-3; most specimens with a prominent sinuate fuscous band extending longitudinally for most of the length of the wing along the medial vein; cilia white with scattered patches of fuscous. Hindwing uni- formly pale to dark gray; cilia sometimes paler and with whitish apices. Foreleg fus- cous to buff dorsally, white ventrally. Mid- leg slightly paler; femur brown dorsally; tib- ia mostly white usually with two dorsal brown spots; tarsi brown with white band- ing. Hindleg pale brown to nearly white dor- sally, white ventrally. Abdomen: Pale gray to buff dorsally, white ventrally. Male genitalia: As shown in Figs. 7-10. Uncus relatively deeply bifid. Gnathos en- tire, consisting of a broadly rounded lobe. Valva simple, straight with apex evenly rounded. Aedoeagus moderately short, about two-thirds the length of valva, with an elongate cluster of 30 or more minute spines extending from middle to apex. Female genitalia: As shown in Fig. 23. Seventh sternum evenly rounded posterior- ly. Bursa copulatrix short, approximately 277 1.5x the length of posterior apophyses; ductus bursae very short, with heavily fur- rowed walls, corpus bursa simple, without signum or spicules. Immature stages unknown. Holotype.—é, Trinidad Mts., San Blas, Cienfuego Province, Cuba; 5 May 1932, S. C. Bruner and A. Otero, E.E.A. Cuba Ento. No. 9967, USNM Type No. 100676 (USNM). Paratypes.—CUBA: Specific locality un- known: | 4, W. Schaus Coll. (USNM). Ca- maguey Prov. [?]: Las Animas, Sierra Ran- gel, 1500° ft.) [457 mi; 1%) Aug. 1922. Roberto (USNM). Cienfuego Prov.: Same data as holotype; 2 6 (USNM). La Habana Prov.: Arroyo Nararrjo; 1 4, 13 May 1934, L. C. Scaramuzza (USNM). Santiago de las Vegas; 1 2°, 15 May 1934, A. R. Otero (USNM); 1 9, 24 May 1932, A. R. Otero, E.E.A. Cuba Ento. No. 9967 (USNM). Hol- guin Prov.: Moa, El Johnson; | 6, June 1954, Zayas & Alayo (IZAC). Santiago de Cuba Prov.; Pico Turquino; 3 6, 1 2, June 1963; Alayo and Garcia IZAC, USNM); 1 6, June 1964, Garcia (IZAC); 2 4, 10-29 June 1936, J. Aoulla (USNM). Sierra Maestra, 1000 ft. [305 m]; 2 4, 7-21 June 1930, O. Querci (USNM). Host. — Unknown. Flight period. — May to August. Most rec- ords indicate a May-June emergence, al- though this may be only a collecting artifact. Distribution. — This species appears to be widely distributed over Cuba, occurring at rather low elevations to as high as 450 m. It may also occur in the Bahamas, but the single male examined in the USNM from Mangrove Cay, Andros Island, is in too poor condition (rubbed and without genitalia) to confirm. Etymology.—The specific name is de- rived from Latin fuscus (dusky, dark) and signatus (mark, stamp) in reference to the forewing markings. Discussion.— This species, with its white forewings heavily streaked with fuscous, ap- 278 pears distinct from all other West Indian Acrolophus. Consequently, its nearest affin- ities remain uncertain, although the male genitalia closely resemble a few currently unnamed species from the Virgin Islands. The forewing pattern can vary consider- ably from the most common expression (Fig. 1) in which a heavy, sinuate streak of dark fuscous extends from the basal fourth of the costa through the discal cell to the termen. The smallest male examined (Fig. 2) from Holguin Province possesses the most re- duced markings but exhibited genitalia in- separable from the typical form (Fig. 1). Acrolophus guttatus Davis, NEw SPECIES Figs. 4, 11-14 Adult (Fig. 4).—Length of forewing: 4, 12.0-12.5 mm. A moderately large species with strongly recurved labial palpi in the male. The forewing is reddish brown marked by 4—5 large, reticulated whitish spots. Head: Vestiture densely hairy, brown where exposed; frons and vertex largely hid- den by labial palpi. Eye hairy; lashes not evident. Antenna approximately 0.5 x the length of forewing, 56-59 segmented; scape brown dorsally, pale buff to white ventrally; flagellum laminate, with pale buff to white scales dorsally; naked ventrally except for dense, very pale, short sensory setae. Labial palpus 3 segmented, extremely long in male and recurved over thorax to metanotum; vestiture brown with lateral portions of dis- tal half mostly suffused with pale buff to white. Thorax: Pronotum brown with scattered, indistinctly white tipped scales. Tegula dark brown. Venter pale buff to white, very hairy. Forewing predominantly orange brown with 4-5 large variegated whitish spots arranged as in Fig. 4; each spot mostly cream colored with small scattered patches of silvery-white scales and a reticulate pattern of smaller or- ange scales; fringe brown. Hindwing pale gray with fringe only slightly darker. Foreleg and midleg brown dorsally, pale buff to PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON nearly white ventrally. Hindleg paler, most- ly whitish buff with spurs and tarsi brown. Abdomen: Pale whitish gray dorsally, slightly darker, more buff ventrally. Male genitalia: As shown in Figs. 11-14. Uncus shortly bifid. Gnathos divided into a pair of elongate, rounded lobes. Valva di- vided less than a third its length at apex into a pair of similar, nearly straight lobes. Ae- doeagus moderately stout, about two-thirds the length of valva, and without cornuti. Female and immature stages unknown. Holotype.—é. Finca La Ciega, Camaguey Province, Cuba; 18 June 1955, P. Alayo, at light (IZAC). Paratypes.—Cuba: Pinar del Rio Prov- ince: Guanahacabibes Peninsula: Cabo Cor- rientes thickets: | 46, 15 May 1956, P. Alayo, at light (USNM). Host.— Unknown. Flight period.— May to June. Distribution.—Known only from two disjunct localities, at the western tip of Cuba and from east-central Cuba. Etymology.—The specific name is de- rived from the Latin guttatus (spotted, speckled) in reference to the conspicuous spotted pattern on the forewing. Discussion. — The presence of 4 to 5 large, reticulated whitish spots on the forewing of this species readily distinguishes it from all other Acrolophus. The divided valva in the male genitalia amply demonstrates its affin- ities to the plumifrontellus group. The rel- atively straight, unswollen cucullar lobe and the absence of discernible cornuti is diag- nostic of the species. Acrolophus niveipunctatus Walsingham Figs. 5, 15-18 Acrolophus niveipunctata Walsingham, 1892: 513; Davis, 1984: 20, no. 113. Acrolophus niveipunctatus Walsingham, 1897: 174. Adult (Fig. 5).— Length of forewing, 4, 13- 14 mm. A moderately large species with VOLUME 89, NUMBER 2 219 Figs. 7-14. Male genitalia. 7, Acrolophus fuscisignatus, ventral view. 8, Lateral view. 9, Lateral view of valva. 10, Aedoeagus. 11, A. guttatus, ventral view. 12, Lateral view. 13, Lateral view of valva. 14, Aedoeagus. All scales = 0.5 mm. 280 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 21 Figs. 15-22. Male genitalia. 15, Acrolophus niveipunctatus, ventral view. 16, Lateral view. 17, Lateral view of valva. 18, Aedoeagus. 19, 4. basistriatus, ventral view. 20, Lateral view. 21, Lateral view of valva. 22, Aedoeagus. All scales = 0.5 mm. VOLUME 89, NUMBER 2 strongly recurved labial palpi in the male. The forewing is pale reddish brown with a whitish-buff colored anal area possessing a bicrenulate anterior margin bordered by an interrupted line of broad, white scales. Head: Vestiture densely hairy, reddish brown where exposed; frons and vertex of male largely covered by labial palpi. Eye hairy with long lashes. Antenna approxi- mately 0.5 x the length of forewing, 60-62 segmented; scape reddish brown dorsally, buff to white ventrally; flagellum laminate, with pale buff to white scales dorsally, naked ventrally except for dense, very pale, short sensory setae. Labial palpus 3 segmented, extremely long in male and recurved over thorax to metanotum. Vestiture dark red- dish brown over basal two-thirds and apex; subapical region with white-tipped, pale buff scales laterally. Thorax: Pronotum generally pale brown with a mixture of piliform, dark brown- tipped scales and paler, broader, white- tipped scales. Tegula reddish brown. Venter mostly reddish brown, very hairy. Forewing anteriad of CuA dark reddish brown, grad- ually fading to pale buff along subterminal margin; anal area posteriad of CuA con- trastingly pale gray to buff except for dark reddish-brown suffusion across base of anal area and a semicircular protrusion below middle of wing from Cu; margin between the two areas continues obliquely to tornus along CuA2; also situated disjunctly along this margin is a thin line of broad, silvery- white scales along base of CuA and more distally along CuA2 to tornus but inter- rupted medially by the semicircular lobe of dark scales; cilia dark reddish brown. Hindwing uniformly pale grayish brown ex- cept for slightly darker cilia. Foreleg and midleg reddish brown dorsally, buff colored ventrally. Hindleg much paler, pale buff with mixture of indistinctly white-tipped scales; spurs and tarsi more brown. Abdomen: Pale gray dorsally; reddish brown ventrally. Male genitalia: As shown in Figs. 15-18. 281 Uncus shortly bifid. Gnathos divided into a pair of elongate rounded lobes. Valva di- vided less than a third its length at apex into a pair of straight lobes; ventral lobe (cucul- lus) with slender base and slightly enlarged apex. Aedoeagus moderately broad, about 0.75 x the length of valva; a cluster of 10- 12 minute spines present near apex. Female and immature stages unknown. Type.— Holotype, ¢. ““Museum Staudin- ger,” present deposition unknown. Type locality.—Cuba. Host. — Unknown. Flight period.— May to June. Distribution.—Known only from the Sierra Maestra of Santiago de Cuba Prov- ince. Material examined.—3 6. CUBA: Santia- go de Cuba Prov.: Honglosongo, Cobre, Loma de Gato; | 6, 20 June 1952 (IZAC). Gran Piedra, Caney; | 6, June 1954, Zayas and Alayo (IZAC). Gran Piedra Mt., Sierra Maestra; 1 6, 30 May 1959, P. Alayo, at light (USNM). Discussion.— The discovery of the above specimens in the Instituto de Zoologia de Cuba Is significant because it establishes the identity and relationship of a previously named but otherwise unknown species. Most of the uncertainty surrounding 4. nivei- punctatus 1s caused by the absence of the type specimen, present whereabouts of which remains unknown. All specimens bearing this name in the collections that I have examined were found to be misiden- tified. Fortunately, this species possesses a rather distinctive wing pattern in which an interrupted series of broad white scales are most diagnostic. This pattern, as carefully described by Walsingham, agrees complete- ly with the specimens before me. It should be noted that even if the male holotype is found, it would still be impossible to deter- mine the specific group relationship of niv- eipunctatus solely on that specimen, assum- ing Walsingham’s original statement is correct: ““Abdomen missing. (a female ab- domen is stuck on to this specimen).” 282 Fig. 23. Acrolophus fuscisignatus, female genitalia, ventral view. Scale = 0.5 mm. Acrolophus basistriatus Davis, NEw SPECIES Figs. 6, 19-22 Adult (Fig. 6).— Length of forewing: 6, 1 1- 12 mm. A moderately large species with strongly recurved labial palpi in the male. The forewing is reddish brown with a prom- inent white streak extending along the base of the cubital vein. Head: Vestiture densely hairy, reddish brown with white-tipped piliform scales where exposed; frons and vertex of male largely hidden by labial palpi. Eye hairy with long lashes. Antenna approximately 0.4 x the length of forewing, 59-62 segmented; scape reddish brown dorsally, mostly white, irrorated with reddish-brown scales ven- trally; flagellum laminate, with light brown scales dorsally, naked ventrally, except for PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON dense, very pale, short sensory setae. Labial palpus 3 segmented, extremely long in male, recurved over thorax to metanotum. Ves- titure similar to head, consisting of dense piliform scales of reddish brown with mi- nute white tips. Thorax: Pronotum similar to head in col- or and vestiture. Venter paler in color, pale reddish brown to buff. Forewing uniformly reddish brown except for a prominent white streak extending along the base of CuA; dis- tal half of streak tends to diffuse into anal area. Cilia consisting of very short, broad scales with dull white tips. Hindwing uni- formly pale brown. Foreleg reddish brown with paler, more wooly vestiture ventrally. Midleg similar to foreleg in color. Hindleg much paler in color, pale buff with indis- tinctly white-tipped scales dorsally; vesti- ture mostly white ventrally. Abdomen: Densely covered with light brown wooly scales dorsally and ventrally; usually a concentration of darker brown scales along mid-venter. Male genitalia: As shown in Figs. 19-22. Uncus shortly bifid. Gnathos divided into a pair of elongate, rounded lobes. Valva di- vided about one-third its length at apex into a straight, relatively broad, costal lobe and a more slender, strongly curved cucullar lobe. Aedoeagus moderately slender, nearly as long as valva, and with a central apical mass of approximately 12-14 short spines. Female and immature stages unknown. Holotype.—é. Cuabal de Gulindo, Valle del Yumuri, Matanzas, Cuba; June 1970, P. Alayo (IZAC). Paratypes.—Same data as holotype, 5 4 (IZAC, USNM). Host.— Unknown. Flight period.—June. Distribution.— Known only from the type locality in Matanzas Province. Etymology.—The specific name is de- rived from the Labin basis (base, bottom) and stria (furrow, line, stripe) in reference to the single white streak along the base of the forewing cubital vein. VOLUME 89, NUMBER 2 Discussion.—Acrolophus basistriatus is the only Cuban member of the genus with a whitish streak along the cubital vein. The male genitalia are diagnostic in possessing the largest cornuti and the most curved cu- cullar lobe (of the valva) of any member of the plumifrontellus group. ACKNOWLEDGMENTS I am grateful to the Instituto de Zoologia, Academia de Ciencias de Cuba, for spon- soring my trip to Cuba in 1981 and to the Smithsonian Institution for a Fluid Re- search Grant that allowed me to participate in this program. I am especially thankful to Pastor Alayo of the Instituto de Zoologia for allowing me to study his material. The line drawings were executed by my former assistant, Biruta Akerbergs Hansen, and the photographs are by Victor Kranz of the 283 Smithsonian Photographic Laboratory. Sil- ver West of our department typed the final draft of the manuscript, which was reviewed by J. F. Gates Clarke, and an anonymous reviewer. LITERATURE CITED Davis, D.R. 1984. Tineidae, pp. 19-24. Jn Heppner, J. R., ed., Atlas of Neotropical Lepidoptera, vol. 2, Checklist, pt. 1 Micropterigoidea to Immoidea. W. Junk, The Hague. Hasbrouck, F. F. 1964. Moths of the family Acro- lophidae in America north of Mexico (Microlep- idoptera). Proceedings of the United States Na- tional Museum 1 14(3475): 487-706. Walsingham, Lord (Thomas de Grey). 1892. On the Micro-Lepidoptera of the West Indies, pp. 511- 517, 544-545. In Proceedings of the Zoological Society of London. 1897. Revision of the West-Indian Micro- Lepidoptera, with descriptions of new species, pp. 169-175. In Proceedings of the Zoological Society of London. PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 284-287 A NEW SPECIES OF AMELETUS (EPHEMEROPTERA: SIPHLONURIDAE) FROM EASTERN NORTH AMERICA WELDON L. BURROWS Consulting Aquatic Biologist, 902 Pennsylvania Ave., St. Albans, West Virginia 25177. Abstract.— Nymphs and reared imagoes of Ameletus tarteri, new species, are described. The new species is presently known from West Virginia, Virginia, and New York. Mor- phological characters of the male imago and nymph of the new species are illustrated and compared with the nominally described eastern Nearctic species and with three western Nearctic species that possess similar male genitalia. Notes on the ecology and habits of the nymph of the new species are included. The genus Ameletus was established by Eaton (1885). Thirty-two species of Ame- letus have previously been described from North America including six species from eastern North America (Edmunds et al., 1976; Carle, 1978). During an ecological survey of streams in central West Virginia, nymphs of an apparent new species of Ame- letus were discovered. Subsequent rearings revealed characters of the male imago that differed from those of the known North American species of Ameletus. Since little is known of the biogeography of Ameletus, available type specimens of three western species with similar male genitalia, as well as five of the six nominal eastern species, were examined. The male of the eastern species A. walleyi Harper has complex gen- italia (see Harper, 1970, figs. 1 and 2) quite unlike that of the new species and was not examined; the nymph of 4. walleyi is un- described. Ameletus tarteri Burrows, NEw SPECIES Figs. 1-5 Male imago (in alcohol).—Body length 7.5-10.0 mm. Forewing length 8.0-9.5 mm. Head brown. Scapes and pedicels of anten- nae tan; flagella light purple. Compound eyes yellow-green dorsally, tan ventrally. Thorax mostly brown with lighter and darker areas and streaks. Legs tan with forelegs darker. Forewings hyaline; costal and subcostal cells suffused with amber; bullae darkly mar- gined. Hind wings hyaline; costal cells suf- fused with amber. Abdominal tergum 1 brown; terga 2-10 pale, shading to tan at posterior and lateral margins and each with submedian dark marks; sterna pale with dark marks submedian on 2-8 and in ganglionic areas; sternum 9 with gray-brown anterior and lateral margins. Genitalia (Figs. 1 and 2) mostly gray-brown; medial area and an- terior margin of subgenital plate lighter, with median apical emargination, and with apex of mesad apical processes attaining apex of first forceps segment; dorsal sclerotization extending mesad from bases of penis lobes and contiguous mesally; dorsal sclerotized bar at fused base of penes; pair of straight, acuminate, laterad titilators with 1-3 den- ticles along ventral margin of each. Caudal filaments tan. Female imago (in alcohol). — Body length 8.0-10.5 mm. Forewing length 9.0-11.5 VOLUME 89, NUMBER 2 285 Figs. 1-5. Ameletus tarteri. 1, Male genitalia, dorsal, ap (mesad apical process), bs (sclerotized bar at fused base of penes), /s (mesad extension of sclerotization from bases of penis lobes), p/ (penis lobe), ¢ (titilators, hidden). 2, Left penis lobe, lateral, d (denticle), p/ (penis lobe), ¢ (titilator). 3, Abdomen of mature nymph, dorsal. 4, Abdomen of mature nymph, ventral. 5, Left mandible of mature nymph, i/ (inner incisor), /m (lacinia mobilis), mb (molar brush), sg (setal gap), sr (setal row). mm. Head tan with black submedian lines on vertex. Compound eyes olive-brown dorsally, olive ventrally. Abdomen (with eggs) orange except segment | brown; sterna with dark marks submedian on 2-7, 9 and in ganglionic areas; subanal plate variable but often with anterior and lateral margins entire. Remainder similar to male imago. Mature nymph (in alcohol). — Body length 7.0-11.5 mm; caudal filaments about 0.5 x body length. Head gray-brown; lighter areas on frons and mouthparts; female with sub- median dark lines on vertex extending to lateral ocelli. Antennae tan, shading distally to purple. Labrum tan, often with median “Vv” outlined darker. Distal seta of inner margin of mandibles (Fig. 5) about 1.5 x length of proximal seta of row; length of setal row about 2.8 x length of setal gap; gap between proximal seta of row and molar setal brush about 1.7 length of proximal seta. Thorax tan with lighter and darker streaks and areas. Legs pale with gray-tan shading; each leg from foreleg progressively lighter; tarsi with apical black band. Ab- dominal terga (Fig. 3) tan with lighter and darker areas; dark submedian marks on 2- 10; spinules on posterior margins of 3 or 4— 10; lateral margins of | or 2-9 posteriorly produced, spinelike on 2 or 3-9. Gills hya- line with amber tracheae in fresh specimens; sclerotized at ventral margin and near dor- 286 sal margin. Abdominal sterna (Fig. 4) pale with darker anterior margins; dark median, submedian, and sublateral markings vari- ably on 3-9. Caudal filaments alternately banded light-dark-light-dark from base; proximal dark band dark gray with all ar- ticulations similar; distal dark band purple- gray; each light band tan on cerci, white on terminal filament; hairlike setae of proximal half of each filament purple-brown, white in distal half of each. Holotype.—é¢ imago, West Virginia, Greenbrier Co., Hamrick Run at West Vir- ginia Route 39/55 near confluence with North Fork of Cherry River, 900 m el. (38°13'40”N, 80°24'04”W), 15 June 1983, W. L. Burrows. Paratypes.— 4 6 imagoes, 3 2 imagoes, same data as holotype; 7 nymphs, same data as holotype but 13 July 1985; 2 4 imagoes, 3 2imagoes, Greenbrier Co., Carpenter Run at WV Route 39/55 near confluence with North Fork of Cherry River, 750 m up- stream from Hamrick Run, 15 June 1983, W. L. Burrows; 2 ¢ imagoes, same data but 9 July 1982; 16 nymphs, same data but 13 July 1985. All material in alcohol; all adults reared with exuviae included. The holotype will be deposited at the U.S. National Mu- seum of Natural History (USNMNH); para- types will be deposited at USNMNH, Ca- nadian National Collection, West Virginia Benthological Survey (Marshall Universi- ty), Purdue University, Florida A&M Uni- versity, and the Academy of Natural Sci- ences, Philadelphia. Other material examined.— Nymph, New York, Chemung Co., McCorn Creek, 7 April 1976, Lamb, Cornell University collection; 4 nymphs, Virginia, Giles Co., Stony Creek, above White Rock Branch, 10 March 1985, Burrows, author’s collection. Etymology.—I am pleased to name the new species after Donald C. Tarter, Chair- man, Dept. of Biology, Marshall Universi- ty. His work, as both student and professor, has contributed much to the knowledge of the regional aquatic fauna. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Discussion. — The penes (Figs. 1 and 2) of the male imago of A. tarteri resemble those of the western species A. cooki Mc- Dunnough, A. imbellis Day, and A. vernalis McDunnough in that each has a pair of straight, acuminate, laterad titilators. The penes of the eastern species A. cryptostim- ulus Carle are somewhat similar, but ex- amination of the genitalia of the holotype reveals that the titilators are not only nearly contiguous as in Carle’s (1978) figure, but that they are apparently flattened through- out their lengths. The male imago of A. tar- teri can be distinguished from all known Ameletus from North America by the pres- ence of denticles (Fig. 2) on the ventral mar- gins of the acuminate titilators. The nymphal tarsi of all species examined have a dark band at both the base and apex except for A. tarteri and A. cryptostimulus, which have only a dark apical band. Al- though the maculation of the dorsal abdo- men of the nymph of 4. cryptostimulus is similar to A. tarteri (Fig. 3), the following should reliably distinguish the two species: on tergum 7 the dark, submedian, curved marks are followed by a median spot in A. cryptostimulus but by submedian spots or, in many cases, by no such posterad spots in A. tarteri; on tergum 10 the dark submedian marks are expanded anterolaterally in A. tarteri, but not in A. cryptostimulus, to form a dark margin extending to the lateral mar- gin of the tergum. The labra of A. tarteri and A. cryptostimulus are mostly tan, often with a darker median “‘V” in the former and nearly always with a dark macula reminis- cent of the small form of the Greek letter omega in the latter. Spinules occur on the posterior margins of abdominal terga 3 or 4-10 in A. tarteri, on 5 or 6-10 in A. cryp- tostimulus, and on 1-10 or 2-10 in the other seven species of Ameletus examined. Sev- eral characters of the inner mandibular setal row (Fig. 5), including numbers of setae and relative lengths of setae, setal row, and setal gap, are useful for distinguishing A. tarteri from the other species examined. For ex- VOLUME 89, NUMBER 2 ample the setal row of the species examined is less than 1.5 x the setal gap except in A. tarteri (2.8 x), A. ludens Needham (4.5 x), A. lineatus Traver (4.8), and A. tertius McDunnough, whose setal row is continu- ous to the molar setal brush with no gap. Ecology and habits.—Nymphs of A. tar- teri were found in rocky first and second order streams. These streams typically had low pH, low alkalinity, and low specific con- ductivity. The immature nymphs were often found on horizontal substrate; later instars were typically found on nearly vertical faces of boulders; pre-emergent nymphs, how- ever, were often found congregated on boul- der faces angled beyond the perpendicular. The nymphs were most efficiently collected by sweeping those surfaces with a small aquarium net. ACKNOWLEDGMENTS I thank the following people for making specimens available for study: D. Azuma and D. Otte, the Academy of Natural Sci- ences, Philadelphia; N. D. Penny and W. J. Pulawski, California Academy of Sciences; R. Foottit and J. E. H. Martin, Canadian National Collection; J. K. Liebherr, Cornell 287 University; R. D. Davic, personal collec- tion; O. S. Flint and G. F. Hevel, U.S. Na- tional Museum of Natural History; B. C. Kondratieff and J. R. Voshell, Jr., Virginia Polytechnic Inst. and State University. I thank Dean Adkins, Dept. of Biology, Mar- shall University and Jan Hacker, Plant Pest Control Div., W.V. Dept. of Agriculture, for critically reviewing the manuscript. I thank Steve Lawton for illustrating the nymphal abdomen and Vicki Crager for help with typing. I am grateful to several mayfly work- ers, including anonymous reviewers, for the generous sharing of their expertise, insight, and time. LITERATURE CITED Carle, F.C. 1978. Anew species of Ameletus (Ephem- eroptera: Siphlonuridae) from western Virginia. Ann. Entomol. Soc. Am. 71: 581-584. Eaton, A.E. 1885. Arevisional monograph of Recent Ephemeridae or mayflies. Trans. Linn. Soc. Lond. (2) Zool. 3: 210. Edmunds, G. F., Jr.,S. L. Jensen, and L. Berner. 1976. The Mayflies of North and Central America. Univ. Minn. Press, Minneapolis, MN. 330 pp. Harper, F. 1970. A new species of Ameletus (Ephem- eroptera: Siphlonuridae) from southern Ontario. Can. J. Zool. 48: 603-604. PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 288-295 LIFE HISTORY AND LABORATORY REARING OF PELOCORIS FEMORATUS (HEMIPTERA: NAUCORIDAE), WITH DESCRIPTIONS OF IMMATURE STAGES J. E. McPHERSON, R. J. PACKAUSKAS, AND P. P. Korcu, III Department of Zoology, Southern Illinois University, Carbondale, Illinois 62901; RJP, present address: Biological Sciences Group, University of Connecticut, Storrs, Connecticut 06268. Abstract.— The life history of Pelocoris femoratus (Palisot de Beauvois) was studied in southern Illinois, and the immature stages were described. The bug was reared from egg to adult in the laboratory. Adults of this univoltine species overwintered in mud and detritus at the bottom of their aquatic habitat and became active in early March. Eggs were found between late April and mid-May and had been glued singly to leaves of Ceratophyllum demersum L. beneath the surface of the water. First instars appeared in mid-May followed by marked overlapping of the subsequent instars. Active adults were last observed in November. This species was reared on Chaoborus americanus (Johannsen) larvae under a 16L:8D photoperiod at 26.7 + 1.5°C. The incubation period averaged 17.7 days. Durations of the five subsequent stadia averaged 10.5, 9.7, 11.0, 12.6, and 19.5 days, respectively. The naucorid Pelocoris femoratus (Pali- sot de Beauvois) ranges in continental United States from New England south to Florida and west to the Dakotas (Slater and Baranowski, 1978), Kansas, Oklahoma, and Texas (Sanderson, 1982). It occurs through- out Illinois (Lauck, 1959). Scattered notes have been published on this bug’s field life history. It is predaceous, feeding on small mollusks, dragonfly naiads, and other aquatic animals (Lauck, 1959; Uhler, 1884). It inhabits various lentic (e.g. lakes, ponds, pools) (Blatchley, 1926; Bobb, 1974; Ellis, 1952; Froeschner, 1962; Gon- soulin, 1973; Hungerford, 1927 [as P. car- olinensis Torre-Bueno; see La Rivers, 1948]; Lauck, 1959; Polhemus, 1979; Slater and Baranowski, 1978; Torre-Bueno, 1923; Wilson, 1958) and the sluggish parts of lotic habitats (Gonsoulin, 1973; Polhemus, 1979) where it is usually well concealed amidst thick growths of aquatic plants (e.g. Alter- nanthera, Chara, Lemna, Myriophyllum, Nitella) (Bobb, 1974; Ellis, 1952; Gonsou- lin, 1973; Hungerford, 1927; Lauck, 1959; Polhemus, 1979; Slater and Baranowski, 1978; Torre-Bueno, 1903, 1905, 1923; Wil- son, 1958). Adults overwinter at the bottoms of ponds and pools in muck and detritus (Blatchley, 1926; Bobb, 1974; Uhler, 1884) and emerge in spring to feed and reproduce. Oviposition begins in spring (Bobb, 1974; Torre-Bueno, 1903) and continues at least until the middle of the summer (Torre-Bueno, 1903). Nymphs have been found during the sum- mer (Bobb, 1974; Torre-Bueno, 1903). There appears to be only one generation per year (Sanderson, 1982) although nymphs may be collected in several stages at the same time (Sanderson, 1982; Torre-Bueno, 1903;,1923): VOLUME 89, NUMBER 2 Pelocoris femoratus has been reared in the laboratory under uncontrolled condi- tions from egg to adult and the immature stages have been briefly described (Hunger- ford, 1927; Torre-Bueno, 1903). For the past three years (i.e. 1983-1985), we have studied the life history of a popu- lation of P. femoratus occurring in the La Rue-Pine Hills Ecological Area. This area, located ca. 30 km northeast of Cape Gi- rardeau, Missouri, in the northwest corner of Union County, Illinois, is part of the Shawnee National Forest. It includes both heavily forested areas atop limestone bluffs, and moist forests at the base of these bluffs that surround La Rue Swamp and Winters Pond. These aquatic habitats are continu- ous and the naucorid occurs throughout the area. Much of the study area is blanketed with duckweeds (i.e. Lemna, Spirodela, Wolffia, and Wolffiella) along the shoreline. This paper presents information on the life history and laboratory rearing of P. fe- moratus and includes descriptions of the immature stages. MATERIALS AND METHODS Life history.—The study began in March 1983, before the bugs emerged from over- wintering sites that year. Samples of adults and/or nymphs were taken with an aquatic net at ca. weekly intervals at six sites along the edge of the study area into November after all nymphs had disappeared and adult activity had markedly decreased. Sampling during the following two years was con- ducted similarly. All samples were pre- served in 75% ethanol and examined in the laboratory to accurately determine the de- velopmental stages present in each sample. Eggs were collected by hand-picking and re- turned to the laboratory for incubation to confirm their identity when the first instars emerged. Occasional collections were made also during the winter months to determine overwintering stage(s) and sites. Data gath- ered during the three years of this study were 289 combined to gain a better understanding of the annual life cycle. Laboratory rearing. — Eighteen adults were collected during late March 1985, returned to the laboratory, and divided between two aquaria (8, 10 adults; no attempt was made to sex the individuals). Each aquarium (ca. 30 x 20.5 x 15 cm) was covered on the bottom with aquarium gravel and filled with ca. 6 cm of dechlorinated water. Cerato- phyllum demersum L. (Ceratophyllaceae) was provided as an oviposition site. Adults were maintained on larvae of Chaoborus americanus (Johannsen) (Diptera: Cha- oboridae). Aquaria were cleaned as the water became fouled, ca. every 20 days. The C. demersum was replaced daily; any sections with attached eggs were removed and placed in petri dishes. Each dish (ca. 9 cm diam, 4 cm depth) was covered on the bottom with filter paper and filled with ca. 1 cm of distilled water, which was sufficient to keep the eggs submerged. Upon hatching, the first instars were also placed in petri dishes. Each dish was again covered on the bottom with filter paper and filled with ca. 1 cm of distilled water, more than sufficient to cover the bugs. Later instars were also provided sufficient water to just keep them submerged. Five first instars were placed in each dish but further separated as they de- veloped through subsequent instars. One C. americanus larva was provided daily as food for first and second instars, increased to two larvae for third and fourth instars, and to three for fifth instars. Dishes were checked daily for exuviae and prey car- casses removed. Clean dishes, paper, and water were provided at each molt; water was changed more frequently if it became fouled. Three pairs of newly emerged F, adults were placed in finger bowls (one male and one female per bowl) to determine if they would reproduce. Each finger bowl (10.5 cm ID, 3.5 cm depth) was filled with 2.5 cm of dechlorinated water, and C. americanus larvae and a section of Elodea canadensis Michaux (Hydrocharitaceae) were provided 290 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MARCH APRIL MAY JUNE JULY SEPT. OcT. NOV. ADULT of ECS 1 (N=10) 254 50 25 1ST O4INSTAR | #£| | Qari ——---.—-..- — 25 (N=47) 50 25 2ND 0-4 INSTAR 25 (N=103) PERCENT OF INDIVIDUALS 25— 3rd 0-4 INSTAR (N=202) 25—| 4TH 0-4 INSTAR (N=243) 5TH 07 INSTAR (N=339) PERCENT OF INDIVIDUALS 2 Figs. 1, 2. Field life cycle of P. femoratus during 1983-1985 combined seasons in Union Co., Illinois. 1, Percent of individuals in each stage per sample. 2, Percent in each sample of total individuals of same stage collected during season. For both figures, dashed lines between first and last dates for first instar indicate no specimens were found. VOLUME 89, NUMBER 2 as food and an oviposition site, respectively. The finger bowls were cleaned as the water became fouled (ca. every 20 days). The aquaria, petri dishes, and finger bowls were kept in incubators maintained at ca. 26.7 + 1.5°C anda 16L:8D photoperiod (ca. 260 ft-c). Descriptions of immature stages.—First to fifth instars were selected from field sam- ples, eggs from those deposited in the lab- oratory by field-collected adults; all speci- mens had been preserved in 75% ethanol. The description of each stage is based on ten individuals. Drawings were made with the aid of a camera lucida, measurements with an ocular micrometer. Dimensions are expressed in mm as X + SE. RESULTS AND DISCUSSION Life history.—This species overwintered as adults at the bottom of the swamp in mud and detritus and became active in early March (Figs. 1, 2). Eggs (10) were found between late April and mid-May but un- doubtedly were laid well beyond this period, based on the late seasonal appearance of first instars. All had been glued singly (i.e. not in clusters) to leaves of C. demersum beneath the surface of the water. The first instars were found from mid- May to the third week of August, second instars from the third week of May to late August, third instars from late May to early September, fourth instars from early June to early September, and fifth instars from mid-June to late September. No active adults were found after November. This species is apparently univoltine. Our conclusion is based in part on the fluctua- tions in numbers of the adults and nymphs during the season. Although there was marked overlapping of the various stages and, thus, any particular sample could have any combination of individuals (Fig. 1), weekly plotting of data for each stage showed only one peak for the first, fourth, and fifth instars, and adults (i.e. a buildup of new generation adults); the second and third in- 291 Table 1. Duration (in days) of each immature stage of P. femoratus under controlled laboratory conditions. Number pleting tive Stage Stadium Range T= SE Mean Age Egg 32535 A222 E006) a Nymph Ist instar 206 8-20 10.5+0.11 28.2 2nd instar 174 6-14 Oe == OBZ 3729 3rd instar 156 6-16 11.0+0.13 48.9 4th instar 144 9-24 12.6+0.16 61.5 Sth instar 124 16-26 19.5+0.18 81.0 * 369 eggs were laid. stars show more fluctuations in seasonal numbers but this may be a reflection of small sample sizes which, by chance, was not ap- parent with the first instars (Fig. 2). Laboratory rearing.— As in the field, eggs were glued singly to the leaves of C. de- mersum beneath the surface of the water. They were white at oviposition but dark- ened to yellowish with brown markings dur- ing maturation. The incubation period av- eraged 17.7 days (Table 1). The first instar emerged through a slit in the cephalic end of the egg. It was whitish at this time but soon darkened to its normal color (see description). It usually fed on its first C. americanus larva within one day. The first through fifth stadia averaged LOD 293577 TdeO12%65-ands 19.5 xdaysjre- spectively (Table 1). The total develop- mental period averaged 81.0 days. F, adults maintained in the laboratory for six weeks never reproduced. This further supports our conclusion that this species is univoltine in southern Illinois. DESCRIPTIONS OF IMMATURE STAGES Egg (Fig. 3).—Length, 1.37 + 0.01; width, 0.77 + 0.01. Eggs laid singly and glued to aquatic vegetation, each egg white at ovi- position, but turning yellowish with brown markings during maturation; chorion with primarily irregular hexagonal pattern; mi- cropylar plug at cephalic end. 292 Table 2. Measurements (mm)? of P. femoratus instars. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Nymph Ist Instar 2nd Instar 3rd Instar 4th Instar Sth Instar Body length 25) = O05 B25 9 OOM 4.74 + 0.05 6.24 + 0.08 8.17 + 0.09 Body width? le 8a== OLO2 DAS) == 0103 2.97 + 0.02 3.93 + 0.06 5533) == 0105 Width at eyes 0.90 + 0.01 120122 OYO1 1.56 + 0.01 2NO==OO2 21S OIO2 Synthlipsis 0.49 + 0.02 0.58 + 0.01 0.85 + 0.02 0.95 + 0.03 A se (00) Head lengths 0.60 + 0.02 0.66 + 0.02 0.94 + 0.02 FOS ==" 0108 1.14 + 0.04 Pronotal lengths 0255-2 010) 0.39 + 0.01 0.64 + 0.01 0.93 + 0.02 esas (00 Mesonotal lengths OZOEEOO 0.25 + 0.01 0.44 + 0.01 OF70'==10103 1.16 + 0.03 Metanotal lengths 0.31 + 0.00 0.41 + 0.01 0.60 + 0.01 0.80 + 0.02 1.02 + 0.01 Leg lengths: profemur 0.60 + 0.01 OR Sie 020) ValSses20:011 1S 1EeEs OO 1.93 + 0.02 protibia 0.34 + 0.01 0.49 + 0.01 OW 2e== 0:01 LO =Es0L02 1.36 + 0.01 protarsus O21 450:00 0.23 = 0:01 O27 = 0101 0.28 + 0.01 0.33 + 0.01 mesofemur 0.50 + 0.01 0.69 + 0.01 1202721001 1.39 + 0.01 1-8 50102 mesotibia 0.41 + 0.01 0.55 + 0.01 0.83 + 0.01 ellshss (0)(0 1.46 + 0.01 mesotarsus 0:25) == 0.00 O32)7== 02011 0.43 + 0.01 0.59 + 0.01 0.76 + 0.01 metafemur 0.64 + 0.01 0.89 + 0.01 1-32) A001 1.80 + 0.02 23332-2008 metatibia ONAG==- 010i 0.95 + 0.01 1.40 + 0.02 N87 330102 2.49 + 0.03 metatarsus 0.39 + 0.01 0.50 + 0.01 0.68 + 0.01 0.93 + 0.01 1-23) 0/01 a x + SE; SE values rounding off to less than 0.005 listed as 0.00. > Measured across metanotum. © Measured along midline. Nymphal instars.—The first instar is de- scribed in detail, but only major changes that have occurred from previous instars are described for subsequent instars. Length is measured from tip of tylus to tip of abdo- men; width across metanotum for Ist—4th instars, across mesonotal wing pads at level of metanotum for Sth. Additional measure- ments are given in Table 2. First instar (Fig. 4).—Length, 2.55 + 0.05; width, 1.58 + 0.02. Body elongate oval, general appearance dorsoventrally flat- tened, greatest width at metathorax; dor- sally convex; ventrally, flattened to slightly concave laterally, convex medially; brown- ish with yellowish white markings dorsally, yellowish white with brown markings ven- trally. Head broadly triangular, recessed into prothorax to level of eyes; anterior margin convex, continuous with lateral margins of prothorax; posterior margin arcuate, often subtruncate medially. Dorsally, head yel- lowish white with posteriorly directed, broad, brown, anchor-shaped mark medi- ally, and thin yellowish line originating near midpoint of inner margin of each eye, both lines continuing posteriorly and converging medially near back of head to form broad V-shaped pattern and giving rise posteriorly to short line that is continuous with mid- dorsal yellow line of thorax. Eyes red, synth- lipsis (taken near anterior margin of eyes) ca. 2.5x width of one eye. Antennae yel- lowish, 3-segmented, segment 2 ca. 2x length of segment | and 0.7 x length of seg- ment 3. Beak yellowish, broadly triangular, 3-segmented, segment 2 ca. 3x length of segment | and ca. 1.5 x length of segment 3. Thoracic nota generally brownish with yellowish or whitish markings, more whit- ish laterally, these lateral areas usually lack- ing brown spots. Anterior margin of prono- tum concave; posterior margin nearly straight. Mesonotum shortest of thoracic nota, ca. 0.8x length of pronotum along VOLUME 89, NUMBER 2 midline, posterior margin straight medially, arcuate laterally; mesonotal wing pads ev- ident. Metanotum longest of the thoracic nota, ca. 1.6 length of mesonotum along midline; posterior margin nearly straight; ratio of mesonotal wing pad to metanotum along lateral edge ca. 3:4. Metanotum often overlapping abdominal segment | and much of Prothoracic leg raptorial, yellowish, fre- quently with brownish streak on outer mar- gin of femur. Procoxa ca. 2 x length of tro- chanter and 0.5 x length of femur. Profemur flattened laterally, markedly tapering dis- tally, basally ca. 4 x width distally; ventrally with 2 contiguous rows of pegs. Protibia and tarsus much narrower than femur, together equal to length of femur ventrally. Tibia with ventral surface forming shallow groove, this groove extending onto tarsus; pegs of femur approximating edge of groove of tibia and tarsus when these segments are ap- posed; tarsus 1-segmented with single mi- nute claw. Meso- and metathoracic legs yellowish; all leg segments on metathorax longer than respective segments on mesothorax. Me- socoxa elongate, ca. 2 x length of trochanter and 0.8x length of femur. Mesotarsus 2-segmented, segment | ca. 0.2 x length of 2, the 2 claws of equal length. Metacoxa and trochanter resembling those of mesothorac- ic leg in shape and relative proportions, fringed with long hairs on outer and inner margins, respectively. Meso- and metatib- iae and tarsi furnished with row of swim- ming hairs on outer surface, most developed on hindlegs. Abdomen, dorsally, brown with yellow- ish white markings; ventrally, yellowish white, covered by long hairs and greatly convex in middle 4; 7 pairs of spiracles evident but minute, Ist pair more medially placed. Paired ostioles of scent glands pres- ent dorsally between segments 3-4. Second instar (Fig. 5).—Length, 3.39 + 0.01; width, 2.13 + 0.03. Body widest at metanotum and 3rd abdominal segment. 293 Synthlipsis ca. 2x width of one eye. An- tennae with segment 2 ca. 3 x length of seg- ment | and 0.6 length of segment 3. Thoracic nota medially generally brown- ish with yellow markings or yellowish with brown markings, brown markings may be much reduced; lateral whitish areas of pronotum now often with few brown spots. Pronotum subequal to metanotum in length along midline, both longer than meso- notum. Posterior margin of mesonotum slightly arcuate medially, clearly arcuate lat- erally; ratio of mesonotal wing pad to meta- notum along lateral edge ca. 1:1. Double row of pegs of profemur reduced to single row except at base, area formerly occupied by remainder of 2nd peg row now occupied by short setae. Row of swimming hairs pres- ent on outer surface of mesotibia and tarsus, and on both inner and outer surfaces of metatibia and tarsus. Spiracles now more evident. Third instar (Fig. 6).—Length, 4.74 + 0.05; width, 2.97 + 0.02. Antenna with seg- ment 2 ca. 2x length of | and 0.5 x length of 3. Thoracic nota medially often with brown markings reduced or absent; lateral whitish areas of pronotum now with several brown spots. Pronotum longest of thoracic nota along m idline, slightly exceeding metanotum and much exceeding meso- notum. Mesonotum clearly arcuate medi- ally and laterally; ratio of wing pad to meta- notum along lateral edge ca. 4:3. Single row of pegs on profemur now surrounded on each side by setae, peg row may be incom- plete. Fourth instar (Fig. 7).—Length, 6.24 + 0.08; width, 3.93 + 0.06. Lateral area of pronotum now yellowish. Pronotum longest of thoracic nota, much exceeding either meso- or metanotum along midline. Ratio of mesonotal wing pad to metanotum along lateral edge ca. 2:1. Single row of pegs on profemur now incomplete, area covered by numerous setae. Fifth instar (Fig. 8).— Length, 8.17 + 0.09; width, 5.33 + 0.05. Wing pads of meso- 294 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 3-8. Immature stages of P. femoratus. 3, Egg. 4, First instar. 5, Second instar. 6, Third instar. 7, Fourth instar. 8, Fifth instar. | VOLUME 89, NUMBER 2 and metanota (latter present but covered by those of mesonotum) just extending to ab- dominal segment 3 laterally. Single row of pegs on profemur now almost absent, nu- merous setae present. ACKNOWLEDGMENTS We thank S. L. Keffer and T. E. Vogt, Department of Zoology, SIU-C, for their help with various aspects of this project, and Karen A. Schmitt, Scientific Photography and Illustration Facility, SIU-C, for the final illustrations and photographs of the various figures. We thank also R. H. Mohlenbrock, Department of Botany, SIU-C, for identi- fying the species of Ceratophyllum and Elo- dea mentioned in this study. LITERATURE CITED Blatchley, W. S. 1926. Heteroptera or True Bugs of Eastern North America with Especial Reference to the Faunas of Indiana and Florida. Nature Publ. Co., Indianapolis. 1116 pp. Bobb, M. L. 1974. The insects of Virginia: No. 7. The aquatic and semi-aquatic Hemiptera of Vir- ginia. Va. Polytech. Inst. State Univ. Res. Div. Bull. 87: 1-195. Ellis, L. L. 1952. The aquatic Hemiptera of south- eastern Louisiana (exclusive of the Corixidae). Am. Midl. Nat. 48: 302-329. Froeschner, R. C. 1962. Contributions to a synopsis of the Hemiptera of Missouri, Part V. Hydro- metridae, Gerridae, Veliidae, Saldidae, Ochteri- dae, Gelastocoridae, Naucoridae, Belostomatidae, Nepidae, Notonectidae, Pleidae, Corixidae. Am. Midl. Nat. 67: 208-240. Gonsoulin, G. J. 1973. Seven families of aquatic and 295 semiaquatic Hemiptera in Louisiana. Entomol. News 84: 83-88. Hungerford, H. B. 1927. Life history of the creeping water bug, Pelocoris carolinensis Bueno (Naucor- idae). Bull. Brooklyn Entomol. Soc. 22: 77-83. La Rivers, I. 1948. A new species of Pelocoris from Nevada, with notes on the genus in the United States (Hemiptera: Naucoridae). Ann. Entomol. Soc. Am. 41: 371-376. Lauck, D.R. 1959. The taxonomy and bionomics of the aquatic Hemiptera of Illinois. M.S. Thesis, University of Illinois, Urbana. Polhemus, J. T. 1979. Family Naucoridae/creeping water bugs, saucer bugs, pp. 131-138. Jn Menke, A. S., ed., The Semiaquatic and Aquatic Hemip- tera of California (Heteroptera: Hemiptera). Calif. Insect Surv. Bull. 21: 1-166. Sanderson, M. W. 1982. Aquatic and semiaquatic Heteroptera, pp. 6.1-6.94. Jn Brigham, A. R. et al., eds., Aquatic Insects and Oligochaetes of North and South Carolina. Midwest Aquatic Enterprises, Mahomet, Illinois. Slater, J. A. and R. M. Baranowski. 1978. How To Know the True Bugs (Hemiptera— Heteroptera). Wm. C. Brown Co., Dubuque, Iowa. 256 pp. Torre-Bueno, J. R. de la. 1903. Brief notes toward the life history of Pelocoris femorata Pal. B. with a few remarks on habits. J. N.Y. Entomol. Soc. 11: 166-173. 1905. Practical and popular entomology— No. 4. Notes on collecting, preserving and rearing aquatic Hemiptera. Can. Entomol. 37: 137-142. 1923. Family Naucoridae, pp. 402-404. Jn Britton, W. E., ed., Guide to the insects of Con- necticut. Part IV. The Hemiptera or sucking in- sects of Connecticut. Conn. State Geol. Nat. Hist. Surv. Bull. 34: 1-807. Uhler, P.R. 1884. Order VI—Hemiptera, pp. 204— 296. In Kingsley, J. S., ed., The Standard Natural History. Vol. II. Crustacea and Insects. S. E. Cas- sino and Company, Boston. 555 pp. Wilson, C. A. 1958. Aquatic and semiaquatic He- miptera of Mississippi. Tulane Stud. Zool. 6: 115- 170. PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 296-302 THREE NEW SPECIES OF STONEFLIES (PLECOPTERA) FROM THE OZARK-OUACHITA MOUNTAIN REGION BARRY C. POULTON AND KENNETH W. STEWART Department of Biological Sciences, North Texas State University, Denton, Texas 76203. Abstract.—Three new species of Plecoptera representing three families are described from the Ozark-Ouachita Mountain region: Allocapnia oribata (Capniidae), Alloperla caddo (Chloroperlidae), and Isoperla szczytkoi (Perlodidae). Morphological descriptions, illustrations, biological notes, and comparisons with other regional species are provided. All stages of Isoperla szczytkoi are described. Stark et al. (1983) and Ernst et al. (1986) have recently described new Alloperla and Neoperla species from the Ozark-Ouachita Mountain region of central North America. During our ongoing study of the regional Plecoptera fauna, three additional species representing three families were discovered. The following descriptions and morpholog- ical terms follow those of Ross and Ricker (1971), Surdick (1981), and Szczytko and Stewart (1976, 1979, 1984). Allocapnia oribata Poulton and Stewart, NEw SPECIES Figs. 1-3 Male.— Micropterous. Color dark brown in alcohol. Length of body 5.5-6.5 mm; wings reaching 6th tergum. Seventh tergum without a dorsal process, but with a distinct rounded membranous area connected to posterior margin (Fig. 2). Eighth tergum with anterior margin sloping abruptly upward from base forming a sclerotized process, ap- pearing as rounded knob in lateral view (Fig. 1), and wide, shallow, and U-shaped in dor- sal view (Fig. 2). Eighth tergum process sub- conical in posterior view (Fig. 3). Upper limb of epiproct narrow, and slightly wider apically in dorsal view (Fig. 2), with sub- equal proximal and distal sections, and cla- vate apically in lateral view (Fig. 1). Upper and lower epiproct limbs subequal in length in lateral view, and lower limb spatulate and abruptly curved downward at tip (Fig. 1). Female. — Unknown. Material examined. — Holotype 6, Arkan- sas, Searcy Co., Middle Fork Little Red River, Hwy 65 @ Shirley, 6-I-1985, B. C. Poulton; two additional 4 in poor condition, same locality and date, B. C. Poulton; 1 paratype 6, Arkansas, Van Buren Co., Ar- chey Creek, Hwy 254, 12.1 km NE Rupert, 6-I-1985, B. C. Poulton. Holotype depos- ited at United States National Museum, paratype deposited at North Texas State University museum. Etymology.—The species name is de- rived from the Greek root oribat-, meaning “mountain roaming.” Diagnosis.—The species represents the first discovery of a new Allocapnia from the Ozarks since A. warreni Ross and Yama- moto (1966). It is not clear which Ross and Ricker (1971) group this species belongs to, since: 1) the epiproct upper limb and the shape of the eighth dorsal process in dorsal view are typical of the pygmaea group; how- ever the process is unnotched, and 2) the arcuate elevated posterior ridge of the eighth dorsal process is characteristic of the recta VOLUME 89, NUMBER 2 Figs. 1-3. Allocapnia oribata. 1, Lateral terminalia, 6. 2, Dorsal terminalia, 4. 3, Posterior view, 8th dorsal process, 6. 4-7, Alloperla caddo. 4, Lateral (A) and anterior (B) view, ¢ epiproct tip. 5, Dorsal terminalia, 4. 6, Ventral, 2 subgenital plate. 7, Lateral, 2? abdominal segments 7-9. group (Figs. 1, 3). Even though 4. oribata is similar to 4. malverna Ross of the recta group, neither it nor 4. malverna have the thin blade-like epiproct upper limb typical of A. recta (Claassen) and other members of that group. Biological notes.—The type males were collected on bridges from permanent, 4th order streams which have rock-rubble sub- strate. Large numbers of A. mohri Ross and Ricker, A. rickeri Frison, and A. granulata (Claassen) were also collected at these lo- calities. Females of these species exhibit considerable variation in subgenital plate form; therefore we were unable to discern the female of A. oribata. Alloperla caddo Poulton and Stewart, New SPECIES Figs. 4-7 Male. — Macropterous. General color am- ber to white in alcohol. Forewing length 6- 7 mm; body length 5-7 mm. Abdominal dorsal stripe absent. Epiproct tip ca. 2x 298 longer than wide; lateral margins conver- gent near base (Fig. 5); anterior margin sharply upturned in lateral view and with a prominent anterior knob visible in dorsal view (Figs. 4A, 5). Epiproct tip with ventral groove visible in front view (Fig. 4B). Lat- eral and dorsal abdominal setae present, with longer brushes of setae on segments 7-9, and dorsally on segment 10. Posterior mar- gin of abdominal pleura 8 and 9 with 4-8 brown spinules. Basal cercal segments with thick setae ca. 1-2x segment width. Ae- deagus membranous. Female. — Macropterous. Color similar to male. Forewing length 6-7 mm; body length 5-7 mm. Subgenital plate little produced, with posterior margin rounded in ventral view (Fig. 6), and slightly protruding ven- trad in lateral view (Fig. 7). Cercal and ab- dominal setae similar to male. Fine setae on subgenital plate (Figs. 6, 7) ca. 2 length of lateral setae. Vagina membranous. Material examined.—Holotype ¢, allo- type 2, 7 paratype 6, and 4 paratype °, Ar- kansas, Garland Co., Middle Fork Saline River, Hwy 7 at Iron Springs Rec. Area, 6-VI-1984, B. C. Poulton; 4 paratype 46 and 1 paratype 2, Arkansas, Perry Co., Dry Fork Creek, Hwy 7, 14.3 km S. of Hollis, 6-VI- 1984, B. C. Poulton; 4 paratype 4, Arkansas, Perry Co., Bear Creek, Hwy 7, 4.4 km SE of Hollis, 12-V-1985, B. C. Poulton. Etymology.—This species is named after the Caddo mound builders, an American Indian tribe that inhabited parts of the Ouachita Mountains from 750-1200 AD. Diagnosis.— Recent descriptions and an illustrated key to Ozark-Ouachita Alloperla were provided by Stark et al. (1983), in- cluding the new species 4. ouachita Stark and Stewart, collected in the same region of the Ouachita Mountains as 4. caddo. The unique saddle-shaped epiproct tip of A. cad- do (Fig. 4) easily separates it from 4. /eon- arda Ricker and A. hamata Surdick. The two last species have flat, blade-like epi- proct tips, with either lateral serrations or PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON points (Surdick, 1981). Alloperla ouachita and A. caudata Frison have broadened epi- proct tips, but the lateral horns of A. ouachi- ta and the appressed hairs of A. caudata (Hitchcock, 1974; Stark et al., 1983) are ab- sent in A. caddo. The membranous basal lobes of the epiproct cowl in A. caddo are relatively smaller than those of A. ouachita and A. hamata (Stark et al., 1983). A. caddo represents the only Ozark-Ouachita A/lo- perla with an evenly rounded female subgenital plate; the other species all possess pointed subgenital plates similar to those described by Surdick (1981) for A. hamata and A. furcula Surdick. Biological notes.—The type localities are all in the Ouachita Mountains and comprise first or second order, rock-rubble, intermit- tent streams. Adults were collected by sweeping riparian vegetation. Tsoperla szczytkoi Poulton and Stewart, NEw SPECIES Figs. 8-20 Male.— Macropterous. Forewing length 10-11 mm, body length 9-10 mm. Body color yellow in life, light brown to amber in alcohol, with darker pattern on head, pronotum, and thorax. Head with brown circular patch enclosing light spot, con- necting median ocellus with anterior frons; lateral and median ocelli connected with an inverted U-shaped dark band with lighter peripheral patches, posterior ones extending to near eyes and back to occiput (Fig. 15). Pronotum with median light stripe; disks medium brown with dark brown rugosities. Pronotum stripe continues through meso- notum; rest of mesonotum and metanotum light brown. Cerci and antennae medium brown. Legs light brown, tibiae darker than femora, both with darker brown bands near their articulation. Cercal segments with short plumose setae and a single, long, dark brown posterioventral seta on each segment. Wings light brown with dark brown veins. Eighth sternum with vesicle 1.5—2 x wide as long VOLUME 89, NUMBER 2 Figs. 8-10. Jsoperla szczytkoi egg. 8, 700.9, 200. 10, 700~. (Fig. 13), with setae present along lateral and anterior margins. Paraprocts medium length, curving upward and inward, narrow- ing to a sharp point, and overlapping pos- terior margin of 10th tergum (Fig. 11). Ninth pleura with a single brown spot. Aedeagus membranous with bulb-like posteriodorsal section 1.5—2 = wider than aedeagal base; posterior portion with a row of 6-7 scler- otized, digited golden brown teeth sur- rounded by an unsclerotized triangular area; blunt spinulae present on elevated circular region posteriodorsal to sclerotized digited teeth; ventral portion of aedeagus with el- evated region containing sparse blunt spi- nulae, and a round patch with anterior half containing stout brown spinulae of medium density (Fig. 13). Basal portion of aedeagus with circular bands containing stout, blunt spinulae and short, rounded spinulae; lat- eral regions with patches of sclerotized punctures and sharp golden brown spinulae (Fig. 12). Posterodorsal tip of aedeagus without spinulae; separate membranous dorsal lobe between cerci unsclerotized and without spinulae (Fig. 12). Female.— Macropterous. Forewing length 11-12 mm, body length 9-11 mm. Body coloration and external morphology similar to male. Subgenital plate broadly rounded and produced posteriorly to anterior '3 of 9th sternum (Fig. 14). Vagina membranous. Nymph.— Body length 10-13 mm. Gen- eral color brown in alcohol. Abdomen with 2 faint dorsal longitudinal stripes with light borders, appearing slightly diagonal on each segment, with a median light blotch be- tween them; a row of 6-8 dark spots, 2 al- ways positioned dorsally inside median light blotch, and others dorsolaterally (Fig. 16). Head pattern with dark inverted U-shaped band connecting ocelli, with light area be- tween lateral ocelli, extending to posterior portion of head, and light oval area anterior to median ocellus (Fig. 17). Pronotum vari- able with median longitudinal brown band surrounded by lighter borders; irregular dark markings on discs and light lateral margins (Fig. 17). Lacinia with 2 teeth; subapical tooth 3-4 length of apical tooth and par- tially hidden behind | of 2 incomplete rows of stout hairs that continue along entire length of inner margin (Fig. 20). Labrum with median swelling (Fig. 17). Right man- dible deeply cleft, with 5 short, stout apical teeth, the outer 2 with inner serrated ridges. Inner margin of mandible with a dense brush of long setae (Fig. 18). Glossae produced and broadly pointed upward at apex; para- glossae slender, curved, with length ca. 1.5- 300 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 11-20. Jsoperla szczytkoi. 11, Dorsal, 6 segment 10. 12, Lateral terminalia, 4. 13, Ventral terminalia, 6. 14, Ventral terminalia, 2. 15, Head and pronotum, adult ¢. 16, Abdominal tergites, nymph. 17, Head and pronotum, nymph. 18, right mandible, nymph. 19, Nymphal labium. 20, Right lacinia, nymph. VOLUME 89, NUMBER 2 2 x width of base (Fig. 19). Posterior margin of abdominal segments with continuous fringe of hairs. Cercal segments with a dor- sal row of long setae, and whorles of short hairs on posterior margins. Egg.— Outline oval, cross section circular. Length 0.35 mm; width 0.27 mm. Collar absent. Chorion sculptured with numerous small punctations, and a few raised knobs, possibly marking a micropylar ring (Figs. 8-10). Material examined.— Holotype ¢ and al- lotype 2 (reared), Arkansas, Logan Co., Gut- ter Rock Creek, 33 km SE of Paris on side road near Hwy 309, 20-IV-1985, B. C. Poulton; 6 paratype 6 and 3 paratype 2 (reared), and 6 nymphs, same locality and date, B. C. Poulton; | paratype ¢ (reared), and 4 nymphs, same locality, 13-IV-1985, B. C. Poulton. Holotype, allotype, and 2 nymphs deposited at USNM; paratypes de- posited at NTSU Museum. Etymology.—This species is named in honor of Stanley W. Szczytko, who is pres- ently working on the revision of eastern Nearctic Jsoperla and has greatly contrib- uted to knowledge of Jsoperla in North America. Diagnosis.— During our intensive sam- pling in the Ozark and Ouachita Mountain region, our collections have revealed a total of nine /soperla species. The most recent descriptions of regional Jsoperla are pro- vided by Szczytko and Stewart (1976) and Stark and Stewart (1973). Stanley W. Szczytko (personal communication), has in- dicated that J. szczytkoi belongs to the J. decepta Frison-I. mohri Frison complex. Common characters of this complex include the following: 1) outer nymphal mandibular teeth deeply cleft and serrated, 2) nymphal lacinial shelf reduced, 3) nymphal subapical tooth of lacinia reduced or absent, 4) nymphal head pattern dark, 5) aedeagus with sclerotized patch-like process bearing teeth, and 6) ovum without a collar. Based on these characters, J. szczytkoi most closely resem- 301 bles J. decepta, another Ozark-Ouachita mountain species. Nymphs of these two species have a similar head pattern, but J. decepta lacks a median pronotal stripe and has a much weaker apical lacinial tooth. Adult J. decepta lack a contrasting head pat- tern as in J. szczytkoi (Fig. 15). Biological notes.— This species is known only from the type locality, an intermittent first-order stream with large rubble substra- tum, that flows down the north side of Mag- azine Mountain, having the highest eleva- tion in the Ozark-Ouachita region. ACKNOWLEDGMENTS We thank S. W. Szczytko for the SEM photography and helpful suggestions in di- agnosing the new /soperla species. This study was supported in part by National Science Foundation Grant BSR 8308422 and the Faculty Research Fund of North Texas State University. LITERATURE CITED Ernst, M. R., B. C. Poulton, and K. W. Stewart. 1986. Neoperla (Plecoptera: Perlidae) of the southern Ozark and Ouachita mountain region, and two new species of Neoperla. Ann. Entomol. Soc. Am. 79: 645-661. Hitchcock, S. W. 1974. Guide to the insects of Con- necticut, Pt. VII. The Plecoptera or stoneflies of Connecticut. Bull. State Geol. Nat. Hist. Surv. Conn. 107: 1-262. Ross, H. H. and W. E. Ricker. 1971. The classifi- cation, evolution, and dispersal of the winter stonefly genus Allocapnia. Ill. Biol. Monogr. 45: 166 pp. Ross, H.H. and T. Yamamoto. 1966. Two new sister species of the winter stonefly genus Allocapnia (Plecoptera: Capniidae). Entomol. News. 77: 265— 267. Stark, B. P. and K. W. Stewart. 1973. New species and descriptions of stoneflies (Plecoptera) from Oklahoma. Entomol. News. 84: 192-197. Stark, B. P., K. W. Stewart, and J. Feminella. 1983. New records and descriptions of A//operla (Ple- coptera: Chloroperlidae) from the Ozark-Ouachita region. Entomol. News. 94: 55-59. Surdick, R. F. 1981. New Nearctic Chloroperlidae (Plecoptera). Great Basin Nat. 41: 349-359. Szczytko, S. W. and K. W. Stewart. 1976. Three new 302 species of nearctic Jsoperla (Plecoptera). Great Ba- sin Nat. 36: 211-220. 1979. The genus Jsoperla (Plecoptera) of western North America; holomorphology and sys- tematics, and a new stonefly genus Cascadoperla. Mem. Am. Entomol. Soc. 32: 1-120. BOoK Foundations for a National Biological Survey. Edited by K. C. Kim and Lloyd Knutson. Association of Systematic Col- lections, Washington, D.C. 1986, x1 + 215 pp. $18.00. This important book is primarily a result of a symposium presented at the annual meeting of the Association of Systematic Collections held in May of 1985 in Victoria, British Columbia. A total of twenty-four contributors examine and comment on vir- tually all aspects of the formation, funding, and perceived results and effects of a National Biological Survey. After a brief foreword by E. O. Wilson and a preface by the editors, the heart of the book is pre- sented in six sections. An introduction by K. C. Kim and Lloyd Knutson addresses the scientific bases for such a Survey. This introduction provides an excellent descrip- tion of what a National Biological Survey might encompass and what might be ex- pected as a direct result of such a Survey. The introduction also provides in brief form and with well-chosen words what is ex- panded upon in the following sections of the volume. The second section consists of papers summarizing the relations of such a Survey on ecological and environmental consider- ations. Especially cogent are the articles on the relation of systematics to long-range ecologic research and the role of a National Biological Survey on environmental pro- tection, food production and plant protec- tion. This is followed by a section on bio- logical survey information in which the form of the data, how it is to be managed and how and to whom it is to be disseminated, is discussed. A fourth section dealing with REVIEW PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON . 1984. Descriptions of Calliperla Banks, Rick- era Jewett, and two new western nearctic [soperla species (Plecoptera: Perlodidae). Ann. Entomol. Soc. Am. 77: 251-263. PROC. ENTOMOL. SOC. WASH. 89(2), 1987, p. 302 legislative and historical perspectives for a National Biological Survey examines both federal and state legislation of the past with comments on possible future legislation. A fifth section describes the ambitious biolog- ical survey programs of Australia and Can- ada and the results of these surveys, and discusses the formation of these surveys and their possible use as models for such a sur- vey in this country. A final brief section consists of two arti- cles. First, a brief article by L. I. Nevling, Jr. summarizes the conference and provides a list of nine statements in which the con- tributors, representing highly diverse view- points, were in agreement. Second, R. M. West and W. D. Duckworth list five rec- ommendations distilled from the presen- tations of the participants. A final brief epi- logue by the editors completes the volume. This reviewer found considerable varia- tion in quality of the presentations, as is usually the case in symposium volumes. However, the editors did a remarkable job in finding excellent speakers and writers and a fine job of editing and introducing the results. Some ideas expressed in this book are of general interest, as well as of interest in relation to a possible National Biological Survey. For instance, this reviewer found Barry Chernoff’s discussion of the three main relationships between ecology and system- atics to be especially well expressed and thought provoking. The volume, in short, is well worth examining and should be on the shelf of anyone doing systematic, eco- logic, or survey studies involving the fauna or flora of North America. Wallace E. LaBerge, J/linois Natural His- tory Survey, Champaign, Illinois 61821. PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 303-321 A REVISION OF THE GENUS PHRYGANIDIA PACKARD, WITH DESCRIPTION OF A NEW SPECIES (LEPIDOPTERA: DIOPTIDAE) JAMES S. MILLER Postdoctoral Fellow, Department of Entomology, Smithsonian Institution, Washington, DC. 20560. Abstract.—The genus Phryganidia Packard (Lepidoptera: Dioptidae) is revised and a new species, Phryganidia chihuahua, is described. Following a brief review of the Diop- tidae, the three species of Phryganidia and the placement of the genus are discussed. Six autapomorphies for Phryganidia are identified. Relevant morphological characters are illustrated and distributional data are presented. When I began studying the systematics of the Dioptidae, John Burns presented me with a copy of a poem he had written en- titled “In Temperate Spring” (1986). The last two lines read, “Like the California oak- moth, I’m a mindless Phryganidia.”’ As the ironies of life would have it, I now find my- self revising the genus. To most North American entomologists, Phryganidia californica, the California Oak Moth, is the only member of the obscure family Dioptidae with which they are fa- miliar. However, the family contains nearly 400 species (Bryk, 1930), all of which, with the exception of P. californica, occur from Mexico south to Uruguay and Argentina. Most dioptids are relatively small, light- bodied, brightly colored moths that appear to mimic members of groups as disparate as the Pericopinae and Lithosiinae in the Arctiidae, and the Ithomiinae in the Nym- phalidae. The 39 other dioptid genera will be the subject of future publications; this paper concerns the species of Phryganidia. Because the Dioptidae are so poorly under- stood, it is necessary to review briefly the taxonomic history of the family and de- scribe some of the morphological characters that previous authors have discussed. Ever since the family was erected by Walker in 1865, its phylogenetic position has been controversial. After examining the larva and pupa of Phryganidia californica, it was Packard’s (1895) opinion that the Dioptidae were closely related to the Geo- metridae. Dyar (1896) suggested that the group was associated with the Pericopinae. Forbes (1916) argued that dioptids belonged to the Noctuid/Arctiid lineage within the Noctuoidea, but subsequently (1922) fol- lowed Fracker (1915) and Mosher (1916) in moving them to a position close to the No- todontidae, a relationship with which all re- searchers have since agreed. Not until Minet (1983) was a significant change in the phy- logenetic position of the Dioptidae pro- posed. Minet argued that the group should be recognized as a tribe, the Dioptini, within the Notodontinae, but he was uncertain about the tribe’s affinities within that subfamily. Minet based his proposal on a single character complex, the structure of the last pair of larval prolegs. Several character complexes have figured 304 prominently in theories concerning dioptid relationships. The structure of the dioptid tympanum indicates a close relationship with the Notodontidae. The Noctuoidea (sensu Hodges et al., 1983; Watson et al., 1980) can be defined by the presence of a thoracic tympanum (Richards, 1933; Brock, 1971: Minet, 1983), and only in the Diop- tidae and Notodontidae is the tympanal membrane directed ventrally (Richards, 1933: Sick, 1940; Kiriakoff, 1950a, b, c, 1963; Minet, 1983). Variation in tympanal structure within the Dioptidae has also re- ceived considerable attention (Sick, 1940; Kiriakoff, 1950a). Since Fracker (1915), many authors have recognized a similarity between the larvae of Phryganidia and those of notodontids. Minet (1983) described the anal prolegs of dioptids as being ‘reduced or absent’ and claimed that this apomorphic character state is shared with the Notodontinae. I have ex- amined larval specimens representing the dioptid genera Phaeochlaena, Phryganidia, Zunacetha, and Josia. The anal prolegs are sometimes elongate (e.g., Phaeochlaena gyon, figured in Bastelberger, 1908), but they are not “reduced or absent” as Minet sug- gested. His hypothesis is therefore in doubt. Wing venation has provided the majority of characters previous authors have used to define dioptid genera (Prout, 1918; Hering, 1925; Bryk, 1930), but offers little infor- mation concerning higher-level relation- ships. All species are “‘trifid’’ in the fore and hind wings (M, being closer to M, than to M;), a plesiomorphic character state shared with the Notodontidae and many lepidop- teran families (see Forbes, 1948). Two char- acters that are frequently observed in the Notodontidae, the presence of an ‘“‘acces- sory cell’ in the fore wing (Forbes, 1942, 1948), and Sc in the hind wing being parallel with R for two-thirds the length of the discal cell (Franclemont, 1970), do not occur in dioptids. As this discussion indicates, the phylo- genetic position of the Dioptidae remains PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON unresolved. Available evidence strongly supports a relationship with the Notodon- tidae but is inadequate to allow a more pre- cise hypothesis. Until more becomes known it seems better to retain family-level status for the group. No autapomorphies for the Dioptidae are known, but a character of the larvae appears promising. The larval skin of all species I have examined is covered with microscopic cuticular projections, described by Fracker (1915) for Phryganidia californica. This condition has also been termed “minutely rugose”” (Forbes, 1942) or “shagreened” (Peterson, 1965). The presence of these cu- ticular projections may ultimately prove to be a synapomorphy for the Dioptidae, but it is important that the immatures of very few species have been collected. Phryganidia has historically included only two species. Packard (1864) described the genus in the subfamily Psychinae (Bom- bycidae), with P. californica as the type species. He chose the name to note conver- gent similarities between these moths and trichopteran species in the Phryganidae. Phryganidia was subsequently referred to the Zygaenidae (Stretch, 1872), and finally to the Dioptidae (Kirby, 1892). Like P. cal- ifornica, P. naxa was originally placed with the psychids. Druce (1885) described P. naxa in the psychid genus Typhonia Bruand. Prout (1918) united both species in the Dioptidae as members of Phryganidia but expressed doubt as to the correct affinity of P. naxa. The only other taxon that has been named, fasciata, was described by Hering (1928) as a form of P. naxa. This paper treats these three taxa and describes a new species, P. chihuahua. METHODS Morphological terminology of the head and antennae follows Forbes (1923) and Hodges (1971). Names for the structures of the genitalia follow Klots (1970) and Hodges (1971). However, the homology of lepidop- teran genitalic structures needs careful study. VOLUME 89, NUMBER 2 For example, it is uncertain whether the terms socli, uncus, and transtilla used here to describe dioptids are homologous with structures in other groups to which the same names have been applied. I will use the terms while recognizing that future research is es- sential. The color guide of Smithe (1975) was followed as closely as possible in de- scriptions of vestiture and wings. Genitalia and adomens were prepared by soaking the entire abdomen in 10% KOH for 12-15 hours. Specimens were placed in 20% ethanol and the abdomen was cut lon- gitudinally along the pleural membrane on the right side. The terminalia were then sep- arated from the rest of the abdomen and scales and soft tissues were removed with a brush. The aedoeagus was removed, and the vesica was inflated using fluid pressure from a syringe inserted in the anterior (proximal) end of the aedoeagus. The bursa was simi- larly inflated using a syringe inserted in the ostium. Preparations were then stained for 10 to 20 seconds in Chlorazol Black (ICN Pharmaceuticals), followed by one hour in Safranin O (Wards’s Natural Science Estab- lishment), and mounted on slides. Line drawings were made using a camera lucida attached to a Wild MS dissecting micro- scope. Lateral views of genitalia were drawn from specimens in ethanol. Drawings of pal- pi, legs, anal views of male genitalia, ae- doeagi, and tergites/sternites were made from specimens mounted on slides in Can- ada Balsam. Scanning electron micrographs were tak- en with a Cambridge Scan 100 microscope. Specimens were mounted on stubs and sputter-coated with gold/palladium. Before being mounted, antennae were cleaned by rinsing them for several minutes in 95% eth- anol. The thorax of dried specimens was carefully brushed free of scales for micro- graphs of the tympanum. Preserved pupae were removed from ethanol and air dried before being mounted and sputter-coated. Acronyms for museums are as follows: American Museum of Natural History, New 305 York, (AMNH); British Museum (Natural History), London, (BMNH); Canadian Na- tional Collections, Ottawa, (CNC); Cornell University Insect Collections, Ithaca, (CU); Carnegie Museum of Natural History, Pitts- burgh, (CMNH); Essig Museum of Ento- mology, Berkeley (UCB); Natural History Museum of Los Angeles County, Los An- geles, (LACM); Museum of Comparative Zoology, Cambridge, (MCZ); National Mu- seum of Natural History, Washington, (USNM). Genus Phryganidia Packard Phryganidia Packard, 1864: 348 [type species: Phryganidia californica Packard by monotypy]. Diagnosis.—Small to medium-sized moths, fore wing length 12 to 22 mm. Most- ly light brown to gray-brown. Adult.— Head: Labial palpi upturned to just above clypeus, segment 2 slightly longer than segment | (Fig. 1). Antennae of male widely bipectinate (Fig. 4), those of female narrowly bipectinate (Fig. 5). Thorax: Pleural region and tegulae cov- ered with long, hair-liked scales. Epiphysis long and slender (Fig. 2), reaching to distal end of tibia or beyond. Fore wing (Fig. 3) with R, arising from discal cell, other radials in the pattern R,+[(R,;+R,)+R,]; M, and CuA, stalked; posterior angle of discal cell oblique. Hind wing with M, and CuA, stalked, posterior angle of discal cell oblique. Tympanal membrane not enclosed by met- epimeron (Figs. 6 and 7). Male abdomen (Figs. 10, 15, 18, 20): Ter- gite VIII arched dorsally, anterior margin slightly emarginate; posterior margin weak- ened medially, with numerous membra- nous folds and 2 lateral excavations. Pos- terior margin of sternite VIII excavated medially, with a pair of sclerotized lateral pockets; a single broad, blunt apodeme on anterior margin. Genitalia with uncus broad and hood-like, ventral surface covered with a mat of short setae; socii elbowed, with 306 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON igsseleeos no. JSM 110; scale line = 0.5 mm). 2, Left fore leg of Phryganidia chihuahua male in lateral view (Mexico, CNC slide no. JSM 115, scale line = 1.0 mm). short, spine-like setae; anal tube kinked dis- tally; valvae with a small lateral patch of androconial scales, a short process at dorsal margin of apex; sacculus small; arms of transtilla narrow, meeting above aedoeagus anteriorly. Aedoeagus short, with a ventral tooth and small dorsal sclerite distally; vesi- ca with 15 to 25 spine-like cornuti. Female abdomen (Figs. 11, 19): Genitalia with ostium simple; ductus bursae short; corpus bursae with small spicules at base and a pair of small signa; posterior margin of tergite VIII with a pair of long, slender projections. Posterior margin of sternite VII excavated medially. Immatures.—Larvae with skin sha- greened; anal prolegs not elongate, crochets present. Pupae with hook-shaped setae on dorsum of abdominal segments 7-10 and on cremaster (Figs. 8 and 9). Natural history.— Within the genus only the immatures of P. californica are known. The larvae feed on many oak species (Tietz, 1972), often causing severe defoliation 1, Right labial palpus of Phryganidia californica male in lateral view (California, AMNH slide (Brown and Eads, 1965). Its primary host is Quercus agrifolia (Volney et al., 1983; Puttick, 1986). Largely as a result of its eco- nomic importance, P. californica has been the subject of ecological (Harville, 1955), physiological (Volney et al., 1983; Puttick, 1986), and bio-control (Milstead et al., 1980) research. Hochberg and Volney (1984) found evidence of a female produced pheromone. The parasites of P. californica have been studied by Young (1982, and references therein). Apomorphies.—Of the morphological characters described above, the following appear to be autapomorphies for Phrygan- idia: 1) Epiphysis long and slender; 2) Pos- terior margin of male tergite VIII with nu- merous folds and two lateral excavations; 3) Posterior margin of male sternite VIII with a pair of sclerotized, lateral pockets; 4) Uncus broad and hood-like; 5) anal tube with a distal kink; and 6) Posterior margin of female tergite VIII with a pair of long, slender projections. VOLUME 89, NUMBER 2 307 3 Fig. 3. Fore and hind wing venation of Phryganidia chihuahua (Mexico, CNC slide no. JSM 114). A, anal vein; C, costal vein; CuA, cubito-anal vein; M, medial vein; R, radial vein; Rs, radial sector; Sc, subcostal vein. KEY TO THE SPECIES OF 2. Ground color light brown; tegulae buff-yellow; PHRYGANIDIA PACKARD fore wing length 112 to 8 mm -22>......... Ie Ground' color offore wines gray-brown, hind =e ee ee cee esse eee P. californica Packard wings charcoal gray ............ P. naxa Druce -— Ground color gray-brown; tegulae gray-brown; — Ground color of fore and hind wings concol- fore-wing length: 191022 mmr"... 2... 2... GEOUS PE te eet eee 8 Sa EA Te ryt 2 As ee ee ee P. chihuahua, new species 308 Phryganidia californica Packard Figs. 1, 4-14 Phryganidia californica Packard, 1864: 348. Diagnosis.— Fore wing length 12 to 18 mm. Ground color lighter than other species in the genus. Head: Scales on first two segments of la- bial palpus buff-yellow, third segment light gray-brown; front tan to light gray-brown. Antennae black ventrally, light gray-brown scales dorsally. Thorax: Legs, pleural region, and dorsum gray-brown; patagia buff to light brown; teg- ulae buff-yellow. Wings translucent, light brown; dorsal surface of male fore wing with a cream colored fascia beyond discal cell, fascia fainter below. Abdomen: Light brown. Male terminalia as in Fig. 10; sternite VIII with a large, membranous “window”; a small patch of androconial scales on lateral surface of val- vae. Female terminalia as in Fig. 11; sternite VII with short, lateral processes on anal margin, largely membranous medially; ter- gite VIII with a membranous region me- dially; corpus bursae with two small patches of spicules at base and two indistinct, spi- culate signa. Type material.—The male lectotype, herein designated, bears three labels: “‘San Mateo, California, [no date], A. Agassiz”; a red label stating ““MCZ Type 23388”; and an “A.S. Packard Type” label. It is in fair condition and is in the MCZ. A second male with the same label data is here designated as a paralectotype. Other specimens examined.—64 6, 32 2 (USNM); 82 6, 21 2 (AMNH); 32 4, 4 2 (CMNH); 9 pupae, 25 larvae (USNM). Dis- Sected)5 6) 512: Distribution. — Oregon. Two localities in Lane County (Wickman and Kline, 1985). California. Happy Camp, Siskiyou County (CMNH); Overwintering from southern Mendocino County south to San Diego, in- land only in the Riverside area (Fig. 14). Also recorded from the Channel Islands PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (Anacapa, Santa Cruz, and Santa Rosa Is- lands), and Catalina Island. A single spec- imen in the USNM labeled “Cn[Canyon] de las Cruces, Baja California, Mexico.”’ Two specimens (one from the AMNH and one from the LACM) are recorded as being from Arizona. Phryganidia chihuahua J. Miller, NEw SPECIES Figs. 2, 3, 15-17 Diagnosis.—Fore wing length 19 to 22 mm. This is the largest of the three Phry- ganidia species. It is darker than most spec- imens of P. californica, the species with which it is superficially most similar. Fe- male specimens are not known. Head: Scales on first segment of labial palpus light yellow, second and third seg- ments gray-brown; front light yellow, gray- brown at outer margins; antennae black ventrally, dark gray-brown scales dorsally. Thorax: Legs, pleural region, dorsum, and tegulae gray-brown; patagia light yellow. Wings gray-brown, translucent, without markings either above or below; (a single specimen with a faint fascia beyond discal cell). Abdomen: Gray-brown. Male terminalia as in Fig. 15; anterior apodeme of sternite VIII broad; valvae narrow, a small patch of androconial scales on lateral surface. Type material.—The holotype male (Fig. 16) bears two labels. The first has the data: ‘““Mexico. Chifhuahua], Mesa del Huracan, 108°15'30°4’, 7400’, VIII-21-25-1964, J. E. H. Martin.”’ The second is a “holotype” la- bel with red lettering. The specimen is in excellent condition and is deposited in the Canadian National Collections, Ottawa. Paratypes: 9 4, all with the same label data as the holotype. Dissected 3 4. Other specimens examined.— Mexico. Durango, El Salto, 28 mi E, 8000’, VII- 22-64, J. Powell (1 6, UCB). Distribution (Fig. 17).—Known only from two localities; the type-locality, and El Sal- to, Durango. VOLUME 89, NUMBER 2 Figs. 4-9. Scanning electron micrographs of Phryganidia californica. 4, Male antenna (ventral view, scale line = 100 wm). 5, Female antenna (ventral view, scale line = 100 um). 6, Meso- and metathoracic segments plus first two abdominal segments (anterior at left, scale line = 0.5 mm). Al, abdominal tergite 1; A2, abdominal tergite 2; C, metacoxa; E, metepimeron; M2, mesomeron; M3, metameron; S, metepisternum; SL, metascutellum; ST2, mesoscutum, T, tympanum. 7, Metathoracic and first abdominal segments (anterior at left, scale line = 250 um). E, metepimeron; TM, tympanal membrane. 8, Terminal segments of pupa (dorsal view, scale line = 0.5 mm). C, cremaster; H, hook-shaped setae on segment 10. 9, Hook-shaped setae on segment 10 of pupa (scale line = 100 um). 310 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 10. Male terminalia of Phryganidia californica (California, AMNH slide no. JSM 110; scale line = 1.0 mm). 10a, Genitalia in anal view with aedoeagus removed. A, anal tube; P, apical process; S, socius; Sc, sacculus; T, transtilla; U, uncus. 10b, Aedoeagus in lateral view (anterior at left). 10c, Tergite VIII in dorsal view. 10d, Sternite VIII in ventral view. VOLUME 89, NUMBER 2 311 Fig. 11. mm). | la, Genitalia in lateral view (anterior at left). 1 1b, Tergite VIII in dorsal view. 1 1c, Sternite VII in ventral view. Etymology.—A noun in apposition. The name comes from the type locality. Remarks.—Phryganidia chihuahua and P. californica are more similar in habitus than either is to P. naxa, but without fe- males of P. chihuahua there is insufficient character information to resolve relation- ships within the genus. Phryganidia naxa Druce Figs. 18-26 Typhonia naxa Druce, 1885: 229. Phryganidia naxa: Prout, 1918: 412. Diagnosis.— Fore wing length 12 to 18 mm. The fore wings are narrower and the Female terminalia of Phryganidia californica (California, USNM slide no. 29171; scale line = 1.0 hind wings less full than in the other Phry- ganidia species. In addition, the wings are not concolorous and are only slightly trans- lucent (not at all in males). Head: Scales on labial palpus and front uniformly gray. Antennae black ventrally, dark gray scales dorsally. Thorax: Legs, pleural region, patagia, dorsum, and tegulae gray. Males (Figs. 21, 23-26) with fore wing ground color gray- brown, ventral surface with a yellow fascia beyond discal cell; some color forms (fas- ciata of Hering, Fig. 21) with an equivalent fascia on dorsal surface as well. Males with hind wings charcoal gray above and below. 312 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Bigs sl Zeal: Females with fore wings gray-brown, hind wings charcoal gray; slightly translucent; without markings either above or below. A single male specimen from Guatemala (Figs. 20, 25, 26) with a light yellow longitudinal stripe on ventral surface of fore wing from base stopping short of fascia; a longitudinal orange-yellow stripe on both surfaces of hind wing from base stopping short of outer mar- gin; fringe scales on anal margin yellow. Abdomen: Gray, lighter ventrally. Male genitalia as in Figs. 18 and 20; costa of valve expanded; uncus bulbous. Female genitalia as in Fig. 19; corpus bursae with a sclero- tized region and two patches of spicules at base; two more or less distinct, spiculate signa. Type material.—The female lectotype (Fig. 22), herein designated, is labeled “Guatemala City, 5000 feet, [no date], Champion.” It bears two additional labels: “B.C.A. Lep. Het. Typhonia naxa,” and “Godman-Salvin Coll. 98.-40.” A_ third handwritten label by D. Goodger reads ‘probable syntype of naxa Druce.”’ It is in good condition and is in the British Mu- seum (Natural History). A female with the data: ““S. Geronimo, [no date], Champion,” and bearing a blue, circular ““syntype”’ label is here designated as a paralectotype. It has no abdomen and is in rather poor condition. Other specimens examined.—Guatema- la. [18]87-88, Conradt, det. Martin Hering Phryganidia californica (dorsal view). 12, Male; 13, Female. (1 2, BMNH); [no date], B. Bruckner S.G., (form “‘fasciata’’) det. Martin Hering (1 2, BMNH); Guatemala City, Sept. & Nov. [no year], Schaus and Barnes (2 6, USNM); Vol- can de Santa Maria, Nov. [no year], Schaus and Barnes (1 2, USNM). Mexico. Nuevo Leon, Chipinque Mesa, 4300’, IX-19-20- 1975 (at light), J. Powell, J. Chemsak, and T. Friedlander (14 4, 42, UCB); Nuevo Leon, 4 mi W Iturbide, 5500’, [IX-25-1975 (black- light trap), J. Powell, J. Chemsak, T. Fried- lander (1 6, UCB); Nuevo Leon, 6.4 km W Iturbide, 24°44'N-99°56'W, 1800 M, VII- 16-79, D.C. Darling (1 6, CMNH). Dis- sected 6 4, 3 2. Distribution (Fig. 17).—Known from Nuevo Leon, Mexico and Guatemala. I have been unable to verify a locality reported by Prout (1918) for Panama. Collecting in higher elevation sites between the Mexican and Guatemalan localities will likely yield additional specimens. Remarks.—The wing pattern of P. naxa is extremely variable. The specimen in Fig. 21 is indistinguishable from one figured by Hering (1928) as “‘form fasciata.”’ I found no genitalic differences between specimens with either this color pattern, or that of the specimen in Figs. 23 and 24, in which there is only a ventral fascia. The specimen most distinct from others, placed in the USNM collection as “form fasciata,” is shown in Figs. 20, 25 and 26. It is essentially identical VOLUME 89, NUMBER 2 313 CALIFORNIA INSECT SURVEY Department of Entomology and Parasitology -@ &- UNIVERSITY OF CALIFORNIA ORAFT 1955 Yarc0on 1 fe} 114 Fig. 14. Overwintering range of Phryganidia californica. in male genitalic morphology to other spec- imens of P. naxa dissected (compare Figs. 18 and 20). The color forms are sympatric at both localities but never differ morpho- logically. Although the number of available speci- mens was small, morphological differences between Mexican and Guatemalan speci- mens were observed: 1) The vestiture is sil- ver-gray in Mexican specimens and brown- gray in those from Guatemala. 2) The male socil are somewhat more sharply elbowed 314 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 15. Male terminalia of Phryganidia chihuahua (Mexico, CNC slide no. JSM 112; scale line = 1.0 mm). 15a, Genitalia in anal view with aedoeagus removed. 15b, Aedoeagus in lateral view (anterior at left). 15c, Tergite VIII in dorsal view. 15d, Sternite VIII in ventral view. in Mexican specimens than in Guatemalan specimens. 3) In females from Guatemala M, is short-stalked with R,.;, whereas it arises from the discal cell in examples from Mexico. Rather than recognize these as dis- tinct species, I have chosen to wait and see whether collecting in localities between Nuevo Leon and Guatemala yields additional VOLUME 89, NUMBER 2 16 Fig. 16. Phryganidia chihuahua holotype male (Mexico, dorsal view). specimens. Their discovery may indicate that morphological differences between the northern and southern populations form continuous clines. 315 DISCUSSION Most of the character states described for Phryganidia are shared with species in one or more additional dioptid genera. Some are widespread in the family. The combination of wing venational characters found in Phryganidia, including the arrangement of the radial veins and stalking of M; and CuA, in the fore and hind wings, is characteristic of over 270 dioptid species belonging to 14 genera (approximately 68% of the family). This venational pattern does not occur else- where in the Noctuoidea and provides per- haps the most convenient means for im- mediately recognizing the majority of species in the Dioptidae. The only large genus with different venation is Myonia (57 species), in CONIC PROJECTION | 110 105 95 90 saa i 80 west voncituse | Fig. 17. Geographical distributions of P. chihuahua (a) and P. naxa (@). 316 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 18. Male terminalia of Phryganidia naxa (Guatemala, USNM slide no. 29174; scale line = 1.0 mm). 18a, Genitalia in anal view with aedoeagus removed. 18b, Aedoeagus in lateral view (anterior at left). 18c, Genitalia in lateral view (anterior at left). 18d, Tergite VIII in dorsal view. 18e, Sternite VII in ventral view. VOLUME 89, NUMBER 2 317 Fig. 19. Female terminalia of Phryganidia naxa (Guatemala, USNM slide no. 29176; scale line = 1.0 mm). 19a, Genitalia in lateral view (anterior at left). 19b, Tergite VIII in dorsal view. 19c, Sternite VII in ventral view. which R, and M, of the fore wing arise from the discal cell. Sick (1940) and Kiriakoff (1950a) argued that the tympanum of Phryganidia is ple- siomorphic within the Dioptidae. In diop- tids with a derived tympanal configuration such as Josia and related genera, the meta- thoracic epimeron forms a “pocket” that almost completely encloses the tympanal membrane (figures in Sick and Kiriakoff). The hook-shaped setae on the abdomen of Phryganidia pupae were first noted by Mosher (1916: plate XX VII, fig. 116). I ob- served identical structures on the abdomen of pupae in other dioptids, including Zu- nacetha annulata and Tithraustes demades. The complete taxonomic distribution of this and other pupal traits will provide invalu- able phylogenetic information. The labial palpi in many dioptid genera are upturned to the dorsal portion of the front (Sometimes far beyond), and segment 2 in these cases 1s extremely long. The pres- ence of relatively short palpi in Phryganidia is likely a plesiomorphic character state. In many Neotropical notodontid genera the sacculus of the male valve is greatly ex- panded, consisting of a series of “pleats” 318 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON sf i jallaebat! Fig. 20. Male terminalia of Phryganidia naxa “form fasciata” (Guatemala, USNM slide no. 29175; scale line = 1.0 mm). 20a, Genitalia in anal view with aedoeagus removed. 20b, Aedoeagus in lateral view (anterior at left). 20c, Tergite VIII in dorsal view. 20d, Sternite VIII in ventral view. which are associated with long androconial scales (Forbes, 1942; Barth, 1955). Similar valvae occur throughout the Dioptidae. The absence ofa pleated sacculus in Phryganidia may be a derived state within the family. The transtilla does not join medially in the notodontids I have examined whereas it does in virtually all dioptids. It is not known whether the character state in dioptids is plesiomorphic or apomorphic relative to the Notodontidae. There is variation in the structure of the transtilla within the Diop- tidae as well. Prout (1918) and Hering (1925) suggested that Phryganidia is structurally most similar to species in the genus Polypoetes. I ob- served female genitalic similarities between members of these two genera. However, my research suggests that Po/ypoetes is not monophyletic. A complete understanding VOLUME 89, NUMBER 2 319 25 26 Figs. 21-26. Phryganidia naxa. 21, P. naxa male similar to form fasciata of Hering (Mexico, UCB; dorsal view). 22, P. naxa female lectotype (Guatemala, BMNH; dorsal view). 23, P. naxa male (Mexico, UCB; dorsal view). 24, Same specimen (ventral view). 25, P. naxa “form fasciata’’ male (Guatemala, USNM; dorsal view; Genitalia, Fig. 20). 26, Same specimen (ventral view). of Phryganidia’s relationship to the other genera will require cladistic analyses, in- cluding a thorough investigation of char- acter state distributions throughout the Dioptidae and Notodontidae. ACKNOWLEDGMENTS I am extremely grateful to J. Donahue (LACM), who first noticed the specimens of P. chihuahua in the collection at the CNC. He also provided valuable distributional data for P. californica. J. Powell was par- ticularly helpful in providing the map of California, as well as distributional data for P. californica and several references on its biology. J. Donahue, B. Poole, S. Weller, J. Franclemont, J. Rawlins, and R. Gagné kindly offered comments on the manu- script. The photographs were taken by V. Krantz (USNM). I thank the following in- dividuals and their institutions for the loan of material: F. Rindge (AMNH), A. Watson 320 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (BMNH), D. Lafontaine (CNC), J. Rawlins (CMNH), D. Bowers (MCZ), J. Powell (UCB), J. Liebherr (CU), B. Poole (SEL- USDA at USNM). This research was sup- ported by a postdoctoral fellowship from the Smithsonian Institution. LITERATURE CITED Barth, R. 1955. Maennliche Duftorgane brasilian- ischer Lepidopteren. 10. Mitteilung: Hemiceras proximata Dogn. (Notodontidae). An. Acad. Bras. Ciéne. 27: 539-544. Bastelberger, M. 1908. Geho6rt die Gattung Phaeo- chlaena zu den Geometridae? Entomol. Z., Stutt- gart 22: 66-67. Brock, J. P. 1971. A contribution towards an under- standing of the morphology and phylogeny of the ditrysian Lepidoptera. J. Nat. Hist. 5: 29-102. Brown, L. R. and C. O. Eads. 1965. California Oak Moth. Jn A technical study of insects affecting the oak tree in southern California. Calif. Agric. Exp. Stn. Bull. 810: 44-47. Bryk F. 1930. Dioptidae. Jn Strand, E., ed., Lepi- dopterorum Catalogus (42). Junk, Berlin. 65 pp. Burns, J. M. 1986. In temperate spring. BioScience 36: 294. Druce, H. 1885. Insecta, Lepidoptera-Heterocera, Vol. 1. Biologia Centrali-Americana. 490 pp. Dyar, H. G. 1896. On the probable origin of the Pericopidae: Composia fidelissima. J. N.Y. Ento- mol. Soc. 4: 68-70. Forbes, W. T. M. 1916. On the tympanum of certain Lepidoptera. Psyche 23: 183-192. 1922. The position of the Dioptidae (Lepi- doptera). J. N.Y. Entomol. Soc. 30: 71. 1923. Lepidoptera of New York and neigh- boring states, part 1. Cornell Agric. Exp. Memoir 68. 729 pp. 1942. Family Dioptidae, pp. 318-322. The Lepidoptera of Barro Colorado Island, Panama. Bull. Mus. Comp. Zool. 85: 99-322. 1948. Family Notodontidae, pp. 203-237. Lepidoptera of New York and Neighboring States, Part 2. Cornell Agric. Exp. Memoir 274. 263 pp. Fracker, S. B. 1915. The classification of lepidopter- ous larvae. Ill. Biol. Monogr. 2: 1-161. Franclemont, J. G. 1970. Dioptidae, pp. 9-11. Jn Dominick, R. B. etal., eds., Perspectus of the Moths of America North of Mexico. E. W. Classey Lim- ited and the Wedge Entomological Research Foundation, London. Harville, J. P. 1955. Ecology and population dynam- ics of the California Oak Moth Phryganidia cali- fornica Packard (Lepidoptera: Dioptidae). Mi- croentomology 20: 83-166. Hering, E.M. 1925. Dioptidae, pp. 501-534. Jn Seitz, A., ed., Die Gross-schmetterlinge der Erde, Vol. 6. Alfred Kernen, Stuttgart. 1928. Neue und alte Heteroceren aus dem Zoologischen Staatsmuseum Berlin. Dtsch. Ento- mol. Z. “Iris” (Dresden). 42: 268-282. Hochberg, M. E. and W. J. A. Volney. 1984. A sex pheromone in the California Oakworm Phrygan- idia californica (Dioptidae). J. Lepid. Soc. 38: 176—- 178. Hodges, R. W. 1971. The Moths of America North of Mexico, Fascicle 21, Sphingoidea. E. W. Classey Limited and R.B.D. Publications Inc., London. 158 pp. Hodges, R. W., et al., eds. 1983. Check List of the Lepidoptera of America North of Mexico. E. W. Classey and the Wedge Entomological Research Foundation, London and Washington, D.C. 284 pp. Kirby, W. F. 1892. A Synonymic Catalogue of Lep- idoptera Heterocera (Moths), Vol. 1, Sphinges and Bombyces. Gurney and Jackson, London. 951 pp. Kiriakoff, S.G. 1963. The tympanic structures of the Lepidoptera and taxonomy of the order. J. Lepid. Soc. 17: 1-6. 1950a. Recherches sur les organes tympa- niques des Lépidoptéres en rapport avec la clas- sification. III. Dioptidae. Bull. Ann. Soc. Entomol. Belg. 86: 67-86. 1950b. Recherches sur les organes tympa- niques des Lépidoptéres en rapport avec la classi- fication. IV. Notodontidae. Biol. Jaarb. 17: 66- WiWile 1950c. Sur la classification et la phylogenie de la superfamille Notodontoidea (F. d’Almeida) Kiriakoff. Bull. Ann. Soc. Entomol. Belg. 86: 23- ZS: Klots, A. B. 1970. Lepidoptera, pp. 115-130. Jn Tux- en, S. L., ed., Taxonomist’s Glossary of Genitalia in Insects. Munksgaard, Copenhagen. 359 pp. Milstead, J. E., D. Odom, and M. Kirby. 1980. Con- trol of early larval stages of the California Oak- worm by low concentrations of Bacillus thurin- giensis applied to lower leaf surfaces. J. Econ. Entomol. 73: 344-345. Minet, J. 1983. Eléments sur la systematique des No- todontidae et nouvelles données concernant leur étude faunistique 4 Madagascar (Lepidoptera: Noctuoidea). Bull. Soc. Entomol. Fr. 87: 354-370. Mosher, E. 1916. A classification of the Lepidoptera based on characters of the pupa. Bull. Ill. St. Lab. Nat. Hist. 12: 14-159. Packard, A. S. 1864. Synopsis of the Bombycidae of the United States. Proc. Entomol. Soc. Philad. 3: 331-396. 1895. A clew to the origin of the geometrid moths. J. N.Y. Entomol. Soc. 3: 30-32. VOLUME 89, NUMBER 2 Peterson, A. 1965. Larvae of Insects, Part 1, Lepi- doptera and Plant Infesting Hymenoptera. Fifth Edition. Ohio State University, Columbus. 315 pp. Prout, L. B. 1918. A provisional arrangement of the Dioptidae. Novit. Zool. 25: 395-429. Puttick, G. M. 1986. Utilization of evergreen and deciduous oaks by the California Oak Moth Phry- ganidia californica. Oecologia (Berl.). 68: 589-594. Richards, A. G. 1933. Comparative skeletal mor- phology of the noctuid tympanum. Entomol. Am. 13: 1-43. Sick, H. i940. Beitrag zur Kenntnis der Dioptidae, Notodontidae und Thaumetopoeidae und deren Verwandtschaftsbeziehungen zueinander. Zool. Jahrb. (Anat.) 66: 263-290. Smithe, F. B. 1975. Naturalist’s Color Guide. Amer- ican Museum of Natural History, New York. Stretch, R. H. 1872. Illustrations of the Zygaenidae and Bombycidae of North America, vol. 1, part 1-9, pp. 90-93. Tietz, H. M. 1972. An Index to the Described Life Histories, Early Stages and Hosts of the Macrolep- 321 idoptera of the Continental United States and Canada, vol. 1. Allyn Museum of Entomology, Sarasota, Florida. 536 pp. Volney, W. J. A., J. E. Milstead, and V. R. Lewis. 1983. Effects of food quality, larval density, and photoperiod on the feeding rate of the California Oakworm (Lepidoptera: Dioptidae). Environ. Entomol. 12: 792-798. Walker, F. 1865. List specimens Lepid. Colln. Br. Mus. 31: 146. Watson, A., D. S. Fletcher, and I. W. B. Nye. 1980. Noctuoidea (part), Jn Nye, I. W. B., ed., The Ge- neric Names of Moths of the World. Vol. 2. British Museum (Natural History), London. 243 pp. Wickman, B. E. and L. N. Kline. 1985. New Pacific Northwest records for the California Oakworm Phryganidia californica. Pan.-Pac. Entomol. 61: 1152: Young, L. C. 1982. Key to the pupal parasites of California oakworm, Phryganidia californica (Lepidoptera: Dioptidae), based on larval exuviae. Pan-Pac. Entomol. 58: 42-47. PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 322-324 A NEW SPECIES OF NOTIPHILA (NOTIPHILA) (DIPTERA: EPHYRIDAE) FROM OHIO ALEXANDER D. HURYN Department of Entomology, University of Georgia, Athens, Georgia 30602. Abstract. — Notiphila (Notiphila) kentensis n. sp. is described from a freshwater marsh near Kent, Ohio. Notiphila kentensis is a member of the adjusta species group and is most closely related to Notiphila mathisi. While examining a series of Notiphila spp. collected from a marsh near Kent State Uni- versity, Portage County, Ohio (Todd, 1985), I encountered an undescribed species of the subgenus Notiphila, Notiphila (N.) kentensis n. sp. Specimens were collected using de- tergent pan traps placed in an area domi- nated by the emergent macrophyte, Nuphar luteum (L.) Sibthorp & Smith (Todd, 1985). Other members of Notiphila, e.g. Notiphila (N.) bella Loew, Notiphila (N.) mathisi Hu- ryn, and Notiphila(N.) theonae Huryn, have also been collected almost exclusively in as- sociation with foliage and flowers of Nuphar (personal observation; Huryn, 1984; Todd, 1985). Although use of Nuphar for ovipo- sition and resting sites has been reported in the literature, association of the larvae of Notiphila with the roots of these plants has not been reported (cf. Mathis, 1979; Van Der Velde and Brock, 1980). At the type locality of N. kentensis, larvae of the sub- genus Notiphila were collected in associa- tion with the root systems of Nuphar (B. A. Foote, personal communication). Although specific identification is not possible at pres- ent, the association of the immature stages of Notiphila (Notiphila) with the yellow pond-lily deserves further study. In the description below, numerical char- acters follow Mathis (1979) and are based on male specimens. Unless otherwise des- ignated, other character states utilized are based upon examination of both male and female specimens. Notiphila (Notiphila) kentensis Huryn, NEw SPECIES Description. —Shore flies of medium size [males 3.5—4,0 mm (n = 6); females 4.6-4.9 mm (n = 3)]. Ground color blue-grey; ex- treme dorsolateral margins of mesonotum bordered by pair of distinct dark-brown stripes. Head: Eye ratio 1:0.71-1:0.78 (n = 6); eye to cheek ratio 1:0.15-1:0.22; postfrons ratio 1:1.25-1:1.43; prefrons ratio 1:0.62-1:0.75. Frons generally concolorous throughout, blue-grey. Paravertical bristles medium in size, noticeably more robust than postocel- lar setae. Single fine proclinate fronto orbital seta present. First and second antennal seg- ments brown, third segment variable, either: (1) light brown proximally becoming darker distally, or (2) entirely dark brown; arista with 10-12 dorsal branches, usually 11. Face microtomentose, variable: (1) generally yel- low near antennal bases becoming silver to- ward oral margin, or (2) yellow throughout. Facial setae fine; gena blue-grey; genal bris- tle similar in dimensions to paraverticle bristle; maxillary palps orange. Thorax: Mesonotum and pleural regions concolorous, blue-grey; lateral margins with VOLUME 89, NUMBER 2 Figs. 1-8. 1, 2, 4-8, Notiphila kentensis. 3, Notiphila mathisi. 1, Epandrium, ventral. 2, Epandrium, lateral. 3, Basiphallus, dorsal. 4, Same. 5, Aedeagal apodeme, lateral. 6, Basiphallus, lateral. 7, Hypandrial process, lateral. 8, Hypandrial receptacle, lateral. a, Scale for Figs. 5-8. b, Scale for Figs. 1-4. distinctive brown stripes extending poster- iad from area anterior of presutural bristle, across the extreme dorsal region of noto- pleuron, terminating near base of supra-alar bristle. Anepisternum with variable dark brown region consisting of either: (1) dark- ened region about prothoracic spiracle, (2) two darkened regions, one about spiracle 324 and one located posterodorsally on pleurite, or (3) an elongate, rectangular darkened re- gion extending posteriorly from spiracle to posterior margin of pleurite. Lateral mar- gins of scutellum nearly black with pigmen- tation extending anteriad onto mesonotum to form short stripes terminating near bases of intra-alar bristles. Femora light-grey, yel- low apically; tibia and tarsi yellow; setal fas- cicle of hind basitarsus yellow. Abdomen: Abdominal ratio 1:0.60—1:0.66 (n = 5); tergum V/IV ratio 1:0.64—-0.79; ter- gum V ratio 1:0.41-0.66. Ground color blue- grey with dark-brown geminate fascia on segments III-IV (e.g., fig. 2, Huryn, 1984). Male genitalia: epandrium generally rect- angular in shape (Fig. 1) with extreme an- terior tapered and bilobed, produced into anteriorly directed projection extending ventrad of epandrial processes (Fig. 2); epandrial processes narrow, parallel, form- ing lateral boundary of narrow emargina- tion (Fig. 1). Aedeagal apodeme as in Fig. 3. Basiphallus (Fig. 6) strongly sclerotized with apical % strongly recurved; in dorsal view, parallel sided proximally with lateral margins converging distally (Fig. 8). Hy- pandrial process (Fig. 4) considerably longer than wide (width: length ratio ca. 1:7), par- allel with no indication of club on apical portion; apical third sparsely invested with fine spinules; hypandrial receptacle reduced to 2 elongate sclerotized strips (Fig. 5). Specimens examined.— Holotype é. Ohio, Portage County, 1.3 km E of Kent State University. 13 September 1984. Julie L. Todd, deposited in National Museum of Natural History, Smithsonian Institution. Paratypes: 5 6, 3 2, same data as holotype except 19 July 1984; 9 4, same data as ho- lotype except 31 August 1984; 1 6, same data as holotype. Deposited in USNM (5 2, 1 2), KSU (5 4, 1 2), and University of Geor- pia (od, 9): PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Etymology.— Through the efforts of Ben- jamin A. Foote and his colleagues, the wet- land areas surrounding the Kent State Uni- versity campus have been the site of numerous studies of the Ephydridae. It is in recognition of these accomplishments and the type locality that I name the new species N. kentensis. REMARKS Notiphila kentensis is a member of the adjusta species group as defined by Mathis (1979) and is apparently most closely relat- ed to Notiphila mathisi. These species are readily distinguished by characters of the basiphallus. The lateral margins of the bas- iphallus of N. kentensis converge distally to form an acutely angled structure (Fig. 4), whereas those of N. mathisi diverge to form a spoon-shaped structure (Fig. 3). N. math- isi 1s known only from the Okefenokee Swamp, Georgia (Huryn, 1984). ACKNOWLEDGMENTS All specimens utilized in this study were obtained through the efforts of J. L. Todd and B. A. Foote. W. N. Mathis examined and offered opinion on specimens of N. ken- tensis. LITERATURE CITED Huryn, A. D. 1984. New Notiphila (Diptera: Ephy- dridae) from the Okefenokee Swamp, Georgia. Proc. Entomol. Soc. Wash. 86: 942-945. Mathis, W. N. 1979. Studies of the Notophilinae (Diptera: Ephydridae), I: Revision of the Nearctic species of Notiphila Fallen, excluding the caudata group. Smithson. Contrib. Zool. 287: 1-111 + iv. Todd, J. L. 1985. The community organization of acalypterate Diptera in a freshwater marsh. M.S. thesis. Kent State University, Kent, Ohio. Van Der Velde, G. and Brock, Th. C. M. 1980. The life history and habits of Notiphila brunnipes Ro- bineau-Desvoidy (Diptera: Ephydridae), an aut- ecological study on a fly associated with nym- phaeid vegetations. Tijdschr. Entomol. 123: 105- 127. PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 325-328 VADUM, A NEW GENUS OF NEARCTIC BRACONIDAE (HYMENOPTERA) W. R. M. MASON Biosystematics Research Centre, Agriculture Canada, Research Branch, Ottawa, Ontario K1A 0C6, Canada. Abstract.—Vadum volatum, a new genus and species of Diospilini (Braconidae), is de- scribed from southern Missouri and eastern Texas. It is distinguished from its nearest relatives, Diospilus and Taphaeus, by the long, narrow first metasomal tergum, the trun- cately toothed clypeus and the four-segmented labial palpus. Diospilini are a small but widely distrib- uted tribe of Braconidae placed in Helcon- inae and believed to be parasitic on Co- leoptera larvae with few or no exceptions. There are about 10 described genera, the majority of which contain only one or two tropical species of uncertain status. Generic names representing tropical groups that I believe should be placed in Diospilini are: Austrodolops Blanchard, Eudiospilus Sze- pligeti, Parabaeacis Granger, Repetiodio- spilus Shenefelt, and Westwoodiella Szepli- geti. There is an obvious need for considerably more taxonomic work, espe- cially on the rich tropical forest faunas, and one should not lightly add new genera. However, the Holarctic fauna is much bet- ter known, containing only a few genera: many species of Diospilus Haliday, a few of the doubtfully different Taphaeus Wesmael, and one or two each placed in Baeacis Foerster and Aspigonus Wesmael. The new genus here described differs radically from all Diospilini known to me by its extremely long and untapered first metasomal tergite and by the median elevated bilobate pro- jection of the clypeus that appears like a truncate tooth (most Diospilini have the clypeal margin flat and evenly curved, sometimes, e.g. Aspigonus and Baeacis, bearing a small single median point). In ad- dition, the new genus has four-segmented labial palpi and the second flagellomere about '3 longer than the first, features ap- parently unique within the tribe, where the labial palpi are usually three-segmented and the first flagellomere is normally the longest. Vadum Mason, NEw GENUS Type species.—Vadum volatum Mason, new species. Head transverse and large (Fig. 1), about 50% wider than mesoscutum between teg- ulae; clypeus (Fig. 3) rather flat and wide, about 3x as wide as long, central quarter elevated and bilobately protruding, forming a truncate tooth-like elevation; labrum about half as wide as clypeus, its margin arcuate but flattened medially; anterior tentorial pits very large, with a deep groove between them; laterally clypeus separated from face by a distinct fine groove; mandibles not twisted, teeth subequal; maxillary palpi with 6 ar- ticles, length about equal to height of head; labial palpi short, with 4 articles; hyposto- mal and occipital carinae complete, meeting at a distance above mandibular opening about equal to basal width of mandible. Pronotum (Fig. 2) medially with a U-shaped carina, which is concave ante- 326 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-4. riorly and about *4 as wide as ocellar tri- angle, its arms enclosing a shallow, for- ward-facing cavity; propleuron with a ven- trolateral lobe overlapping pronotum. Mesothorax smooth, notauli strong, rugose; transscutal groove visible medially; axillae broad, roundedly sloping down behind, lat- eral margin defined by a strong carina; pre- scutellar scrobe very wide and deep, with a few transverse costae; post scutellar groove Vadum volatum, n. sp. 1, Dorsal aspect of 2, wings drawn following conventions of Mason (1986). 2, Pronotum, dorsal aspect. 3, Head. 4, Abdomen of 2. broad, transcostate; epicnemial carina strong, extending far dorsally; metapleuron and propodeum mostly rugose and bearing vague carinae. Wings (Fig. 1) with full venation; in fore- wing vein M reaching R proximad to origin of Rs, i.e. Ist discoidal cell sessile; A, with a strong dip at which a sclerotized, tubular stump of A, occurs, vein a absent; veins 2RS and 2r-m weakly converging anteriorly. Vein VOLUME 89, NUMBER 2 A, of hind wing absent. Legs long and slen- der; mid and hind tibial spurs shorter than apical depth of tibia; tarsal claws simple. Abdomen (Figs. 1, 4) long and slender, about '4 as wide as thorax and a little less than twice as long. Metasoma I in dorsal view about 4 times as long as its minimum (subbasal) width; base and apex of equal width, about 10% wider than minimum; spiracles prominent, subbasal; dorsal and dorsolateral carinae conspicuous basally; ventral edges of tergite widely separated; complete first sternite about half as long as first tergite; basal strongly sclerotized part of sternite I smooth, pointed posteriorly and about '4 as long as tergite I; sternite 2 lying half beneath tergite I and half beneath ter- gite II; sculpture of tergite I completely ru- gose-reticulate. Remainder of metasoma smooth, apical terga extensively telescoped, metasomal terga 1-5 occupying basal 0.8- 0.9 of length. Ovipositor (Fig. 4) very long and slender, about 1.6 times length of fore- wing or twice length of metasoma. Valvulae 2 with a subapical dorsal notch. Cerci half- discoid, articulated. Life-history and immature stages un- known. The generic name is neuter, referring to shallow water or a shoal; the specific name means turning about or rolling. The epithet commemorates the type locality, the Becker Farm, known as “Rolling Shoals.” Vadum yvolatum Mason, NEw SPECIES Figs. 1-4 Clypeus (Fig. 3) confused punctate ba- sally; median bilobate elevation shining, impunctate; apicolateral margins strongly depressed and narrowly decurved. Face and genae smooth, with sparse, fine punctures; vertex very sparsely punctate, frons im- punctate, shining. Antennae long and slen- der, about 25% longer than forewing; basal flagellomeres about 4 times longer than wide, apical ones about 1.5 times longer than wide, very little taper; flagellum broken at article 327 29; first flagellomere about 75% as long as second. Sides of pronotum broadly smooth above and below with a broad central rugulose area. Scutum and scutellum shining, punctate; notauli broad, rugulose near pronotum, deep and transcostulate centrally, merging into a large rugose area behind; mesopleuron mostly smooth but bearing a broad, sinuate rugulose groove from upper epicnemium to middle coxa. Color.— Thorax and legs fulvous with pi- ceous to castaneous prothorax, propodeum, hind tibia and tarsus. Head and abdomen black; mouthparts fulvous. Wings hyaline with brown veins and stigma. Variation.—Very little in the few speci- mens available; flagellum of 28-32 articles; length of forewing 3.3—4.5 mm, of head and body 4-6 mm, of ovipositor sheath 5-7.5 mm; metapleuron and notauli sometimes piceous. Both males have forewing about 4 mm long and broken antennae, otherwise very similar to female in all somatic char- acters. Specimens. — 2 4, 5 2, as follows: Holotype ?, Missouri, Williamsville, J. T. Becker, Malaise Trap. June 1-16, 1969. CNC No. 19357. Paratypes: 1 6, 2 2°, same data but 1 2 dated 16-26 June (CNC). Texas, Brazos Co., College Stn., R. Wharton, Malaise Trap. 1 6, May 4, 1983; 2 29, April 11-15 and 12- 25, 198) (ix2A.M.U)): RELATIONSHIPS Vadum resembles the cosmopolitan gen- era Diospilus and Taphaeus in most of its features and appears closely related. Vadum differs from the many species of Diospilus and Taphaeus by the following four fea- tures, each of which appears derived in Vad- um: (character state for Diospilus and Tap- haeus in parentheses) 1, apical margin of clypeus (Fig. 3) depressed on the lateral 0.45 so that there appears to be an elevated me- dian bilobate tooth (margin of clypeus not depressed and bearing no such median 328 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tooth), 2, second flagellomere distinctly longer than the first (first longer than sec- ond), 3, metasomal tergite I unusually elon- gated, the base and apex equally wide, length/apical width = 3.0-4.0 (metasomal tergite I broadening apically, length/apical width = 1.0-1.5), 4, in forewing vein a ab- sent (vein a present). Vadum might be re- garded as no more than a derived species of Diospilus were it not for the labial palpus which has four articles of subequal length, whereas all the species of Diospilus (as well as all other Diospilini known to me, in- cluding Aspigonus, Baeacis and Taphaeus) have only three articles in the labial palpus, clearly a derived condition. Thus I believe the evidence indicates Vadum to be a sister group to Diospilus and the other three gen- era named above. I thank my colleagues M. J. Sharkey and J. R. Barron for reviewing the manuscript and making useful suggestions. LITERATURE CITED Mason, W. R. M. 1986. Standard drawing conven- tions and definitions for venational and other fea- tures of wings of Hymenoptera. Proc. Entomol. Soc. Wash. 88: 1-7. PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 329-343 A NEW NEARCTIC GLYPHIDOCERA WITH DESCRIPTIONS OF ALL STAGES (LEPIDOPTERA: BLASTOBASIDAE: SYMMOCINAE) D. ADAMSKI AND R. L. BROWN Department of Entomology, Drawer EM, Mississippi State University, Mississippi State, Mississippi 39762. Abstract.—A new species of Blastobasidae, Glyphidocera juniperella Adamski, from southeastern United States is described. This species feeds on Juniperus and is a pest on ornamental varieties. Illustrations of the adult, male and female genitalia, and larval chaetotaxy are provided. Scanning electron micrographs of egg, larva, pupa, and abdom- inal sex scales of male adult are included. Glyphidocera and its type-species, G. au- dax, were described from Saint Vincent Is- land in the West Indies by Walsingham (1892). Thirty-nine species occur in the New World tropics and 9 species occur in Amer- ica north of Mexico. Host plants and biol- ogy of these species are unknown. Many Glyphidocera are uniformly brown and usually possess relatively broad and truncate forewings. The genitalia are char- acterized by the following: male with valvae narrowed basally, abruptly broadened dis- tally, apex protracted; costa with fingerlike projection at base; gnathos projecting from beneath tuba analis; aedeagus with medium to large cornutus; female usually with duc- tus bursae sclerotized and apically expand- ed; ductus seminalis spiralled and originat- ing from posterior end of corpus bursae; corpus bursae with two patches of denticles at the anterior and posterior ends, and usu- ally with a sclerotized plate near constricted end of ductus bursae. This study resulted from the discovery of larvae of a new species of Glyphidocera feeding on nursery stock varieties of Junip- erus horizontalis Moench, in Mississippi and Florida. The biology of this species has been studied by Schiffhauer and Mizell (accepted for publication), and insecticides have been evaluated for its control by Mizell and Schiffhauer (accepted for publication). MATERIALS AND METHODS The adult, egg, larva, and pupa were ex- amined with an incandescent light source (reflected light). The Methuen Handbook of Colour (Kornerup and Wanscher, 1978) was used as a color standard for the description of the adult. Genitalia were dissected as de- scribed by Clarke (1941), except Mercuro- chrome and chlorazol black were used as stains. All preparations were examined with dissecting and compound microscopes. The ultrastructure of egg, larva, pupa, and abdominal sex scales of males was studied with a Hitachi HH-S-2R scanning electron microscope at an accelerating voltage of 20 kV. For SEM examination, immature spec- imens were fixed in 3% glutaraldehyde in 0.1 M potassium phosphate buffer (pH 7.3), rinsed in phosphate (pH 7.3), and postfixed in 2% osmium tetroxide in 0.1 M potassium phosphate (pH 7.3). After dehydration in ethyl alcohol, specimens were critical point dried, mounted on stubs with silver paint 330 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. and paste, and coated with gold-palladium in a Polaron E5100 sputter coater. Abbreviations used in this paper are: ANT, antenna; AX TB, axillary tubercle; C, cremaster; FR CL S, frontal clypeal suture; GL, galea; L3 (Fig. 28), metathoracic leg; LB, labium; LBP, labial palp; MD, man- dible; MX, maxilla; MXP, maxillary palp; ?, pit (chordotonal organ, pore, other); PIV A, pivot area; PR SC, proleg scars; SP, spi- racle; SPIN, spinneret; TON FIB PL, tonofi- brillary platelets. Glyphidocera juniperella Adamski, NEw SPECIES Adult.— Head: Scales light-brown or greyish-orange basally, greyish-orange api- cally; frons usually lighter than vertex; an- tenna dorsally with dark-brown and grey- ish-orange scales intermixed, ventrally with greyish-orange; labial palpus medially light greyish-orange, or light greyish-orange scales intermixed with dark-brown scales, later- Holotype of Glyphidocera juniperella Adamski (male) (9.2 x). ally dark-brown, apex of second segment usually greyish-orange. Thorax: Tegulae, mesonotum, and meta- scutum dark-brown or with dark-brown scales intermixed with greyish-orange scales; color of legs similar to labial palpi. Forewing (Fig. 1): 7.0-8.5 mm long; dor- sal surface with greyish-orange scales inter- mixed with dark-brown scales, appearing uniformly light-brown without microscope; one to two small dark-brown spots present in discal cell near distal end and near mid- dle; some specimens with dark-brown spots near base and on Cu near middle of wing; fringe scales brownish-grey. Ventral surface brownish-grey. Hindwing: Dorsal surface light orange- grey at base, becoming greyish-orange api- cally; ventral surface light brownish-grey, intermixed with dark-brown on costa. Abdomen: Greyish-orange scales inter- mixed with dark-brown scales; male with irregular, transverse row of sex scales on VOLUME 89, NUMBER 2 % 08905 &.b..—- fA Tage 8g 08a. Pore 2 — Figs 2. Ww Ww — en Dorsal view of abdominal terga 1-4 of male Glyphidocera juniperella. Sex scales are indicated by arrows pointed towards anterior margins of terga 3-4. Line scale = 1 mm. anterior margins of terga III and IV (Fig. 2); each scale with sub-parallel sides, obtuse apex, with base abruptly curved to short pedicel (Fig. 3); interridge grooves deep, fe- nestrae Oval, irregularly spaced, each cir- cumscribed by a wide rim; some cross ribs medially fused and parallel with longitudi- nal ridges (Fig. 4). Male genitalia (Fig. 5): Tegumen with heavily sclerotized anterior margin, scaled laterally below gnathos; uncus without scales or setae, dorsally sclerotized and confluent with tegumen, ventrally sclerotized to near base; gnathos fused and projecting dorsally from below tuba analis, sparsely setose; ae- deagus long, cornutus medium sized; juxta dorsally rolled around base of aedeagus, ventrally narrowed and elongated; valva heavily sclerotized except for ventral half of cucullus and medial groove from base to cucullus; costa sparsely setose, basally with fingerlike, apically setose projection; cucul- lus with ventral half densely setose, dorsal half relatively bare; sacculus with single row of 5—7 setae near middle and numerous se- tae on ventral margin. Female genitalia (Fig. 6): Sternum and tergum VII lightly sclerotized and evenly scaled; sternum and tergum VIII with nu- merous microtrichia that increase in density toward posterior margins, with several long, simple setae restricted to posterior half of sclerites; papillae anales with two types of setae: type I setae long, simple, directed pos- teriorly and posterolaterally; type II setae (visible at high magnification) minute, re- curved, and directed laterally; antrum with two internal, basal prongs, each directed posterolaterally; corpus bursae with smooth, sub-circular plate near antrum, anterior cluster of denticles near inception of ductus seminalis, and posterior cluster of denticles 332 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 3, 4. SEM of abdominal sex scales of male Glyphidocera juniperella. 3, 4000x. 4, 20,000 x. VOLUME 89, NUMBER 2 333 Fig. 5. Scale = 1 mm. at the posterior end; ductus seminalis spi- ralled, bearing cluster of denticles near base. Egg.—Eggs laid singly or in imbricate clusters on scales of branchlets of Juniperus spp. Egg (Figs. 7-8) width nearly two-thirds the length; broadly raised along the median longitudinal axis and flattened towards the outer margin. The surface is characterized by slightly raised convolutions. Aeropyles Genitalia of male G/lyphidocera juniperella (aedeagus is detached). D. Adamski Gen. Slide no. 2854. appear to be distributed nonrandomly over the dorsal surface. A distinct micropylar area has not been detected. Color changes of the embryo (ranging from yellow-orange to dark-brown) are readily observable through the translucent chorion. Larva.—Length 12.0-15.6 mm [10n]. Body greyish-brown with a velvety textured appearance due to slightly raised stellate cu- 334 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Genitalia of female Glyphidocera juniperella. D. Adamski Gen. Slide no. 2855. Line scale = 1 mm. VOLUME 89, NUMBER 2 : be ‘ q . Z Figs. 7, 8. SEM of egg of Glyphidocera juniperella. 7, 150.8, 750 (arrows point to aeropyles). Ww nN 336 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ticular projections on the surface; head cap- sule, prothoracic shield, anal plate, pina- cula, and legs dark brown. Head (Figs. 9- 10): hypognathous; epicranium with ridges arranged polygonally, ridges on mouthparts indistinct; adfrontal sclerites broad (Fig. 17), delimiting frons dorsolaterally; frons closed; F1, F2, and C3 sub-equal in length; F1 clos- er to C3 than F2, in straight line with C2; P1 long, closer to P2 than A2; A1 closer to A2 than to A3; Cl arising from small depression on ventral lateral margin of clyp- eus, C2 closer to Cl than C3; labrum with distal margin modified into two lobes pro- jecting medially, with 12 setae, two medial pairs, with outer pair at least four times length of inner pair, one pair on each lateral margin, and two pairs on distal margin; mandibles (Figs. 10, 15-16) slightly asym- metrical, with pair of setae on outer surface (basal seta longer than distal seta (Fig. 10)); labium strongly ridged distally, with micro- trichia proximally; submental pit absent; spinneret bulbous at base, narrowed distal- ly; labial palpus two segmented, each seg- ment with a dorsally directed apical seta; maxilla (Figs. 10-11, 13) prominent; sen- silla types and arrangement on medial lobe and apex of palpus (Figs. 11, 13), similar to those of Choristoneura fumiferana (Clem.) (Albert, 1980), Heliothis zea (Boddie), (Avé, 1981), and other ditrysian Lepidoptera (Grimes and Neunzig, 1986a, b). Stemmata III and IV approximate, associated setae as in Fig. 21. Sensilla types on antenna (Fig. 14) similar to other Lepidoptera (Schoon- hoven and Dethier, 1966). Prothorax (Fig. 18): prothoracic shield medially bissected by ecdysial line; D1 and D2 parallel or sub- parallel, D1 slightly more than half the length of D2; LI at least twice the length of L2, closer to L2 than L3, and slightly below line between L2 and L3, L2 and L3 sub-equal in length; SV1 and SV2 widely divergent, SV2 slightly shorter than SV1 and pointed posteriorly; distance between SV1 and SV2 less than distance between L1 and L2. Me- sothorax (Fig. 18): D1 anterodorsal to D2, on same pinaculum; D1 slightly more than half the length of D2, SD2 slightly more than half the length of SD1; L2 and L1 di- vergent, on same pinaculum; L2 more than twice length of L1; L3 posterodorsal to L1; MV1 on small pinaculum, located within fold on anteroventral margin of segment be- tween L and SV groups. Metathorax as de- scribed for mesothorax. Abdomen (Figs. 18— 20, 22-26): prolegs on A3-6 and A10 and of equal size; crochets uniserial and bior- dinal (Fig. 26); D1 and D2 parallel or sub- parallel on Al-9; D1 anterior to and widely separate from D2 on A1-8, and anteroven- tral and closer to D2 on A9; SD1 and SD2 dorsal to spiracle on Al-8, SD2 minute, detectable only at high magnification (Figs. 22-23); SD1 and SD2 on Al-8 approximate to an invagination on pinaculum; L1 and L2 in nearly straight line with spiracle and SD1 on Al, anterior to line between spiracle and SD1 on A2-8, L3 pinaculum postero- ventral to L1 and L2 pinaculum on A1-8, ventral to L1 and L2 pinaculum on AQ, L3 in straight line with L2 and SV3 on A9; SV bisetose on Al and A7, trisetose on A2-6, unisetose on A8-9; V1 pinacula in or near straight line between each pair of SV’s on A7-8, slightly anterior to line between SV’s on A9, V1’s slightly closer on A9 than A7-— 8; AlO with D1’s athwart (Fig. 20); D2’s stout, recumbent, and directed ventrally (Figs. 25, 27). Small, slightly pigmented, ir- regularly shaped and sized, tonofibrillary “platelets” are located in transverse folds of T2-3, on intersegmental membrane of T2- A9 on two irregular lines, one between D and SD setal groups, and one between spi- racles and L setal groups (Figs. 22, 24). Pupa.—Pupa (Figs. 28-34): scabrous, an- teriorly truncate, widened in thoracic re- gion, narrowed posteriorly to a bifurcate cremaster; ecdysial line extending from pos- terior margin of metathorax to anterior margin of vertex; axillary tubercles (cocoon cutter) located on each anterolateral margin VOLUME 89, NUMBER 2 Figs. 9-14. SEM of larva of Glyphidocera juniperella. 9, Frontal view of head capsule, 55x. 10, Ventral view of head capsule, 150. 11, Maxillae, 850 x. 12, Labium, 1000. 13, Sensilla on apex of maxillary palpus, 5000 x. A2 = sensillum styloconicum; Al, A3, M1, M2, L1, L2, and L3 = sensilla basiconica; SD = sensillum digitiform; 14, sensilla on apical portion of antenna, 2500, 1 = sensilla basiconica, 2 = sensilla chaetica, 3 = sensillum styloconicum, 4 = sensillum trichodeum. 338 A10 AQ Figs. 15-21. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Larva of Glyphidocera juniperella. 15, Right mandible. 16, Left mandible. 17, Frons and associated setae. 18, Setal map of thorax and Al-7. 19, Setal map of A8-10. 20, Setal map of A10. 21, Arrangement of stemmata with associated setae. of mesothorax (Figs. 28-29, 31); epicranial suture V-shaped; fronto-clypeal suture pres- ent; antenna, maxilla, legs 1, 2, and apex of 3 exposed ventrally, femur of leg 1 exposed as short, narrow sclerite, leg 3 extending be- yond margin of forewing; terminal six ab- dominal segments, pivoting as a unit from intersegmental area between fourth and fifth segments, pivotal area with narrow ridges separated by areas with irregular rows of small pits (Figs. 29-32); AS5—6 with scars of abdominal prolegs present; pupal setae long and straight, setae on cremaster are apically recurved (Figs. 30, 33-34). Holotype.—¢, Miss.[issippi], Monroe Co., Hamilton, Amfac Nursery, 25 Aug. 1981, VOLUME 89, NUMBER 2 339 Figs. 22-27. SEM of larvae of Glyphidocera juniperella. 22, Portion of A6 showing positional relationship between spiracle and SD group and tonofibrillary platelets, 200 x. 23, SD1 and SD2 on Al. Note invagination (labeled ‘*?”’), 100 x. 24, Tonofibrillary platelets and stellate integument (Al), 1000 x. 25, A8-10, 50x. 26, Left proleg on A5, 500. 27, Anal plate with large recumbent D2 setae and A10 prolegs, 100~. 340 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ery a oe i ‘ i a » » . % FRCL SC Figs. 28-30. SEM of pupa of Glyphidocera juniperella. 28, Lateral view, 30x. 29, Dorsal view, 30x. 30, ventral view, 30. David Tatum [Coll.]; iss. 14-[X-81, ex Gold Coast Juniper. The holotype is deposited in the U.S. National Museum on indefinite loan from the Mississippi Entomological Mu- seum (MEM). Data are given as on labels except for bracketed information. Paratypes.— Mississippi: Monroe Co., Hamilton, 25 Aug. 1981, R. L. Brown [Coll.], ex Juniperus, 2 6, David Tatum [Coll.], ex Gold Coast Juniper, em. 26-28 Aug. 1981, 2 6, D. Adamski genitalia slide no. 577; 5 2, D. Adamski gen. sl. nos. 542, 580, 582, and 2855; em. 29 Aug. 1981, 3 4, D. Adamski gen. sl. nos. 579, 585, and 2888; VOLUME 89, NUMBER 2 Figs. 31-34. SEM of pupa of Glyphidocera juniperella. 31, Auxillary tubercle (= cocoon cutter), 400 x. 32, Area demarcating pivotal area of pupa, 450 x. 33, Ventral view of cremaster, 150 x. 34, Dorsal view of cremaster, 150x. 3 2, D. Adamski gen. sl. no. 581; em. 3 Sept. 1981, 6 6, D. Adamski gen. sl. nos. 587, 2853, 2854, 2887, USNM gen. sl. nos. 11426, 11427; 3 2, D. Adamski gen. sl. no. 2856, USNM gen. sl. no. 11429; em. 5 Sept. 1981, 4 6, D. Adamski gen. sl. nos. 578, 584, and 586; 1 2, USNM gen. sl. no. 11428; iss. 12-IX-81, 4 6, 2 2. Paratypes of male and female are deposited in the British Mu- seum (Natural History). Other paratypes are deposited in the Mississippi Entomological Museum (MEM). Other specimens examined. — Florida: Gadsden Co., Quincy, larva coll. 1 Apr. 1981, Juniperus horizontalis “‘wiltoni,” R. F. Mizell 2 6. Maryland: Prince George’s Co., Adelphi, VIII-8-1970, R. W. Hodges 2 2; USNM gen. sl. no. 81436. Mississippi: Hinds Co., Clinton, 20 Jul. 1963, and 24 Jul. 1963, Bryan Mather 2 2; Clay Co., West Point, Gold Coast Juniper, 5-23-80, J. D. Solomon; Hopkins no. $1705, 1 2. Larvae— Florida, Gadsden Co., Quincy, ex Juniperus horizontalis *‘wiltoni,” Imperial Nursery, | Apr. 1984, R. F. Mizell, 10 fifth instars, 14 early instars. DIAGNOSIS Based upon the examination of type- specimens of all Glyphidocera species at the United States National Museum and British Museum, G. juniperella is most sim- 342 ilar in wing coloration and pattern to G. barythyma Meyrick, described from For- estburg, Texas and G. rhypara Walsingham, described from Guerrero, Mexico. How- ever, specimens are paler in the latter two species. Males of G. barythyma can be dis- tinguished from those of G. juniperella by the presence of abdominal sex scales only on tergum 3. G. juniperella has sex scales on terga 3 and 4. The male genitalia of G. barythyma have a cucullus that is broader at the apex, a shorter fingerlike projection at the base of the costa, and a larger and toothed cornutus. Males of G. rhypara can be distinguished from G. juniperella by presence of a narrow valval with a rounded apex and a dense setal cluster on the inner surface of the cucullus. Female rhypara have a more narrow antrum and broader eighth sternum. In addition, the denticles within the corpus bursae in rhypara are stouter and are not basally attached to each other as in G. juniperella. DISCUSSION Clarke (1969) was the first to illustrate abdominal sex scales and scale tufts on male Glyphidocera. G. juniperella males lack ab- dominal scale tufts; however, they do pos- sess tergal sex scales. These scales normally cannot be seen on pinned specimens be- cause they are covered by folded interseg- mental membrane, but they are easily de- tected on dissected specimens. The characteristic exterior rims of fenestrae in these sex scales also occur at various sex scales of Tortricidae (Brown and Miller, 1983). The invagination or pit proximal to SD1 and SD2 on the larval abdomen may demarcate muscle attachments of dorsal chordotonal organs (Kristensen, pers. comm.), a pore, or some other structure of unknown function. ACKNOWLEDGMENTS We thank David Tatum, Mississippi Di- vision of Plant Industry, for calling our at- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tention to this species through his initial collection; Russell F. Mizell, of the Agri- cultural Research Center, University of Florida, for providing us with immatures as well as reared material; Ronald W. Hodges, Systematic Entomology Laboratory, Agri- cultural Research Service, at the U.S. Na- tional Museum, and Klaus Sattler of the British Museum (National History) for their cooperation provided during the examina- tion of type material; and Greta E. Tyson and Michael Sullivan, of the Electron Mi- croscopy Center, Mississippi State Univer- sity, for their help in the preparation of spec- imens and photographic plates. This research was supported in part by grants from NSF Grant BSR85-01212 and Sigma Xi. LITERATURE CITED Albert, P. J. 1980. Morphology and innervation of mouthpart sensilla in larvae of the spruce bud- worm, Choristoneura fumiferana (Clem.) (Lepi- doptera: Tortricidae). Can. J. Zool. 58: 842-851. Avé, D. A. 1981. Induction of changes in the gus- tatory response by individual secondary plant compounds in larvae of Heliothis zea (Boddie) (Lepidoptera, Noctuidae). Ph.D. dissertation. De- partment of Entomology, Mississippi State Uni- versity. 89 pp., 4 tables, 37 figs. Brown, R. L. and P. R. Miller. 1983. Studies of Lep- idoptera hindwings with emphasis on ultrastruc- ture of scales in Cydia caryana (Fitch) (Tortrici- dae). Entomography 2: 261-295. Clarke, J. F. G. 1941. The preparation of slides of the genitalia of Lepidoptera. Bull. Brooklyn Ento- mol. Soc. 36: 149-161. 1969. Catalogue of the Type Specimens of Microlepidoptera in the British Museum (Natural History) described by Edward Meyrick. Vol. VII. British Museum (Natural History), London. 531 pp. Grimes, L. R. and H. H. Neunzig. 1986a. Morpho- logical survey of the maxillae in last stage larvae of the suborder Ditrysia (Lepidoptera): Palpi. Ann. Entomol. Soc. Am. 79: 491-509. . 1986b. Morphological survey of the maxillae in last-stage larvae of the suborder Ditrysia (Lep- idoptera): Mesal lobes (laciniogaleae). Ann. Ento- mol. Soc. Am. 79: 510-526. Kornerup, A. and J. H. Wanscher. 1978. Methuen VOLUME 89, NUMBER 2 Handbook of Colour. 2nd ed. Methuen and Co., Ltd., London. 243 pp. Mizell, R. F. II] and D. E. Schiffhauer. Evaluation of insecticides for control of Glyphidocera juniperella Adamski, (Lepidoptera: Blastobasidae: Symmo- cinae). Fl. Entomol. (accepted for publication). Schiffhauer, D. E. and R. F. Mizell III]. Bionomics of Glyphidocera juniperella Adamski, (Lepidoptera: Blastobasidae: Symmocinae) a newly discovered 343 pest of container-grown juniper. Fl. Entomol. (ac- cepted for publication). Schoonhoven, L. M. and Dethier, V. G. 1966. Sen- sory aspects of host-plant discrimination of lepi- dopterous larvae. Arch. Neerlandaises Zool. 16(4): 497-530. Walsingham, Lord (Thomas de Grey). 1892. On the Micro-lepidoptera of the West Indies. Proc. Zool. Soc. Lond. 1892: 492-549. PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 344-350 A NEW SPECIES OF PHERBELLIA FROM NORTH AMERICA WITH RANGE EXTENSIONS FOR P. HACKMANI AND P. GRISEICOLLIS (DIPTERA: SCIOMYZIDAE) R. E. ORTH Department of Entomology, Division of Biological Control, University of California, Riverside, California 92521. Abstract. —Pherbellia fisheri is described from Northwest Territories, Canada. New distributional information is recorded for P. griseicollis (Becker) and P. hackmani Roz- kosny is reported for the first time from North America. The male genitalia of all three species are illustrated and maps of the North American distribution records are given. In America north of Mexico the genus Pherbellia consists of 41 species including the new species described here, and P. hack- mani Rozkosny, recorded here from North America for the first time. Not among the above 41 species is P. fusca (Cresson), which recently has been considered a nomen du- bium (Knutson et al., 1986). This study (1) recognizes Pherbellia fish- eri, new species, (2) extends the known dis- tribution of P. hackmani, described from Europe, to North America, (3) expands the known distribution of P. griseicollis (Beck- er) in North America, (4) provides illustra- tions of the terminalia of the above three species, and (5) maps the localities in North America where the species have been re- corded. Seven North American species including one subspecies of Pherbellia are now known to be Holarctic: P. albocostata (Fallén), P. argyra Verbeke, P. griseicollis, P. griseola (Fallén), P. hackmani, P. nana nana (Fal- lén), and P. obscura Ringdahl. In the Palae- arctic, these seven species have essentially a Fennoscandian distribution; some of them extend into other parts of Europe, and to North Africa, Soviet Central Asia, Siberia and China. Several attempts have been made to sep- arate the genus Pherbellia into subgenera. The most recent division was by Rozkosny (1964). The species discussed herein are in the subgenus Chetocera Robineau-Des- voidy, as defined by Rozkosny. Rozkosny (1984) further alludes to a P. griseicollis group in which he listed P. griseicollis and P. sordida (Hendel). Pherbellia fisheri is a member of that group and perhaps P. hack- mani may also belong there. Furthermore, the Pherbellia fuscipes group (species with a row of hairs on the posterior margin of the anepisternum, as designated by Steyskal (1961), belongs to Chetocera, as defined by Rozkosny (1964). Pherbellia fisheri, P. griseicollis, P. hack- mani and P. sordida share the following combination of characters: 1. Midfrontal stripe less than '2 distance from anterior ocellus to frontal margin. 2. Two fronto-orbital bristles. 3. Wing not patterned. 4. First vein surpassing level of anterior crossvein. 5. Anepisternum bare. 6. Anepimeron with 2 (rarely 3) large bris- tles and several fine setae. 7. Katepisternum with only fine setae dor- sally. VOLUME 89, NUMBER 2 Pherbellia fisheri Orth, NEw SPECIES Figs. 1-3, 8 Holotype male. — Height of head *4 width. Medifacies yellowish, pruinose; facial grooves subshiny; parafacies and cheeks pruinose, pale yellow. Frons testaceous, yel- lowish anteriorly. Midfrontal stripe extend- ing less than '2 distance from anterior ocel- lus to frontal margin. Ocellar triangle and orbital plates tannish grey, tomentose. Or- bital plates tapered anteriorly, extending be- yond midfrontal stripe. Orbito-antennal spot lacking; narrow white tomentose stripe along upper orbital margin. Two pairs fronto-or- bital bristles; anterior bristle approximately */; as long as posterior bristle; ocellars, post- ocellars, and inner and outer verticals well developed. Occiput tannish grey, tomen- tose. Short black setae on lower 7 of cheeks, on anterior '4 of frons, between ocellar and postocellar bristles, on orbital plates, and in mid-cervical patch. Lateral occipital mar- gins with somewhat stronger setae and bris- tles. Antenna testaceous; aristal hairs ap- proximately as long as width of first segment of arista. Thorax greyish brown, tomentose, slight- ly mottled, with 4 indistinct brown longi- tudinal stripes. Pleura tannish grey, tomentose. Anepi- sternum bare, with upper posterior surface more brownish. Anepimeron with cluster of small bristles and 2 much longer, stronger bristles situated mid-anteriorly. Katepister- num with fine setae over much of surface and well-developed setae ventrally. Pro- sternum bare. Coxae greyish white, tomentose. Forefe- mur brownish, infumated. Mid- and hind- legs tawny, slightly infumated; hindfemur slightly darkened distally. Tarsal segments tawny except brown 4th and 5th segments on fore- and midlegs. Wing length 4.2 mm. Membrane greyish yellow, hyaline, costal margin and veins brownish yellow, area around crossveins clouded. No stump veins; first vein sur- passing level of anterior crossvein; anal vein 345 reaching wing margin. Halter, squama, and squamal cilia pale yellow. Abdominal segments greyish brown, slightly infumated dorsally; terminalia as in Figs. 2 and 3. Allotype female.—Similar to holotype except for reproductive structures. Wing length 5.0 mm. Diagnosis.— Among Pherbellia species, P. fisheri appears to be most closely related to P. griseicollis and P. sordida. Identification should be made by examining male termi- nalia and using the locality maps. Pherbellia fisheri is set apart from both species by the smaller, less deeply emarginated ventral posterior lobe on the anterior surstylus (for illustrations of P. sordida see Rozkosny (1984)). Holotype.—¢, CANADA, Northwest Termtones, Aklavik, July 27, 1931), ©: Bryant, Lot 234. Deposited in United States National Museum of Natural History. Allotype.—@, same data as holotype ex- cept, July 27, 1932, Lot 306. Deposited with holotype. Paratypes.—CANADA. Alberta: Banff, July 19, 1915 (1 2), N. B. Sansome, May 25, 1922 (1 2, 3 4), May 26, 1922 (2 2), June ls 1922783). June 21¥ 1922 (12), CoB: Garrett; Lancaster Park, July 28, 1963 (1 6), J. R. Vockeroth; Waterton Lakes Na- tional Park, July 7-12, 1980 (1 3), H. J. Teskey. Manitoba: Churchill, June 25, 1930 (1 4), collector ? Northwest Territories: Ak- lavik, Sept. 12, 1930 (1 4) Lot 142, Sept. 18, 1930 (1 2) Lot 153, O. Bryant; USA. Alaska: Umiat, July 3, 1959 (1 9, 1 3), July 7, 1959 Go 3:6) J. E2H.Martin: Etymology.—I take pleasure in naming this species in honor of T. W. Fisher, whom I have enjoyed working with for many years. Distribution.—This boreal species is known from widely separated areas in Alas- ka and Canada. The northernmost collect- ing site is Umiat, Alaska, 69°25’, and the southernmost is Waterton Lakes National Park, Alberta, 49°06’. Discussion. —The illustrations of the ter- 346 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 0.4 mm Figs. 1-3. Pherbellia fisheri, new species. 1, Head, holotype male. 2, Terminalia, ventral view, paratype male. 3, Terminalia, sinistral view, inverted, paratype male. minalia of Palaearctic Pherbellia sordida by Rozkosny (1984) resemble P. fisheri. A specimen of P. sordida collected in Wus- tung, Poland, by O. Duda, May 24, 1921, was borrowed from R. Rozko$&ny, J. E. Pur- kyne University, Brno, Czechoslovakia, for examination. Comparison of the terminalia of P. sordida with P. fisheri revealed them to be distinct. Pherbellia fisheri was independently rec- ognized as undescribed by L. Knutson sev- eral years ago. Upon receipt of my manu- script for review he informed me of his findings and suggested that I proceed with its description. He also provided most of the specimens used in this paper. I gratefully acknowledge his contribution. The northernmost collection sites of P. VOLUME 89, NUMBER 2 fisheri (Umiat, Alaska, and Aklavik, North- west Territories) lead one to speculate that, as with other similarly distributed species, P. fisheri may also occur in Fennoscandia. Closer study of material labeled P. grisei- collis from northern Europe may reveal the presence of the new species there. Pherbellia griseicollis (Becker) Figs. 4, 6, 8 Externally, Pherbellia griseicollis and P. fisheri are so similar that I cannot separate them with a great degree of confidence. In general, the dorsum is greyer and the legs are slightly darker than P. fisheri. Distribution.—In the Palaearctic P. gris- eicollis is known to occur in Fennoscandia, the Murmansk region, the Karelian ASSR of the USSR, and in Western Siberia (Roz- kosny, 1984). Prior to this study, P. gris- eicollis was known only from Prudhoe Bay, Alaska, in North America (Knutson et al., 1986). It is now known to be rather wide- spread with the southern limit of its distri- bution near Lima, Montana, approximately 45°N latitude. Collection records.—CANADA. Alberta: Banff, July 25, 1924 (1 8), E. Hearle; Banff National Park, June 4, 1955 (1 2), June 9, 1955 (1 2), G. E. Shewell, July 9, 1955 (2 2), J. R. McGillis. British Columbia: Buck- inghorse Provincial Campground, Alaska Hwy. DC-175 (281.6 km), June 27, 1978 (1 6), P. H. Arnaud, Jr.; Lac le Jeune, June 25, NOS2(2°2)~ June 27, W973 eo 1S). a Teskey. Manitoba: Warkworth Creek, nr. Churchill, June 10, 1952 (1 8), G. E. Shew- ell. Northwest Territories: Aklavik, June 7, 1931 (1 8), July 25, 1931 (1 84), O. Bryant; Yellow Knife, Kam Lake, June 20, 1966 (1 6), G. E. Shewell; 21 mi E of Tuktoyaktuk, June 17, 1971 (1 9), June 21-25, 1971 (1 8), 8-12 July, 1971 (1 9), 17-21 July, 1971 (2 2), 20-25 July, 1971 (1 2, 1 6), D. M. Wood. Yukon: Herschel Island, 24-28 July, 1971 (1 2, 14), D. M. Wood. USA. Alaska: Circle, July 26, 1971 (1 4), B. A. Foote; N. Coast, Prudhoe Bay, June 8, 1971 (1 2), June 16, 347 1971 (1 8), June 20, 1971 (1 8), July 4, 1971 (1 2, 1 3), M. Deyrup; Umiat, Aug. 7, 1959 (16), R. Madge. Idaho: Idaho Co., Lolo Pass, June 18, 1965 (1 3), R. L. Westcott. Mon- tana: Beaverhead Co., 9 miS of Lima, June 26, 1966 (1 4), B. A. Foote. Pherbellia hackmani Rozkosny ISS 5: ies Externally, P. hackmani is set apart from P. fisheri, P. griseicollis and P. sordida by a more intense clouding of the membrane bordering the anterior and posterior cross- veins and a yellowish brown gross aspect. The male terminalia are as in Figs. 5 and 7. The anterior surstyli are similar to P. argyra Verbeke (see Fisher and Orth, 1983). Distribution.—In the Palaearctic this species is known only from the type series (Rozko&sny, 1984). The holotype male was collected at Mukkavuoma, Torne Lappmark, Sweden. There are ten additional paratypes from Sweden, Finland, and western Siberia. This study presents the first records of Pherbelli hackmani in North America. It is a boreal species, and has been recorded south of 60°N latitude only at King Salmon, Alas- ka (58°40'N, 156°40’W). Collection records.—CANADA. North- west Territories: Aklavik, June 15, 1930 (1 2, 1 3), O. Bryant; 21 mi E of Tuctoyaktuk, July 17-21, 1971 (1 4), D. M. Wood; Tun- unuk, Aug. 15, 1930 (1 9, 1 4), O. Bryant. Yukon: Dickson Lake, Mt. Mye, 133°08’- 62°21",-5000 ft. June14> 1960 (1, 9). Je E: H. Martin; Firth River, Aug. 3, 1956 (2 9), R. E-Leech: Herschel Iss. July 22, 1953 (1 6), J. S. Waterhouse, (1 2), C. D. Bird; North Fork Pass, Ogilvie Mts., 4100 ft., June 11, 1962 (1 4), June 12, 1962 (1 2), June 20, 1962" (isc) RsEseech June 18. 1962*(1 2), July 1, 1962 (1 9), P. J. Skitsko. USA. Alaska: Cape Thompson, June 10, 1960 (sex ?), W. C. Hanson; Cape Thompson, Crow- bill Mountain, June 6, 1960 (1 2), W. C. Hanson; Gulkana, Paxon Lodge, Aug. 4, 1951 (2 9, 1 6), W. R. M. Mason; Isabella Pass, Mi. 206 Richardson Highway, 2900 0.4 mm Figs. 4-7. 4, 6, Pherbellia griseicollis, male, 21 mi E of Tuktoyaktuk, Northwest Territories. 4, Terminalia, ventral view. 6, Terminalia, sinistral view, inverted. 5, 7, Pherbellia hackmani, male, Summit Lake, Isabella Pass, Alaska. 5, Terminalia, ventral view. 7, Terminalia, sinistral view, inverted. VOLUME 89, NUMBER 2 349 NORTH AMERICA GRISEICOLLIS P, EISHERI ieee: iS 120 800 1000 MILES 1 GOODE BASE MAP SERIES 200 1400 KILOMETERS DEPARTMENT OF GEOGRAPHY THE UNIVERSITY OF CHICAGO HENRY M LEPPARD, EDITOR LAMBERT AZIMUTHAL EQUAL-AREA PROJECTION Fig. 8. Collection sites for Pherbellia griseicollis, P. hackmani, and P. fisheri. 350 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ft., July 17, 1962 (1 2), P. J. Skitsko; Isabella Pass, Summit Lake, July 9, 1951 Q 9), P. J. Skitsko, Aug. 3, 1951 (1 9, 1 36), Mason- McGillis; King Salmon, Naknek River, July 9, 1952 (1 8), J. B. Hartley, July 13, 1952 (1 4), Aug. 4, 1952 (1 3), W. R. Mason; Kotzebue, June 24, 1951 (1 9), R. I. Sailer; Nome, July 9, 1951 (1 4), D. P. Williams; Umiat, June 10, 1947 (1 9), C. Smith, July 3, 1959 (72); Julye10) 1959 (Ged) aE. Ft: Martin, July 10, 1959 (1 4), July 23, 1959 (1"2)5 Aug. 3.1959 (2:95 2:6); Aug: 7, 1959 (1 2), Aug. 12, 1959 (1 2), R. Madge; Una- lakleet, June 11, 1961 (2 9), June 18, 1961 (2 3), R. Madge. ACKNOWLEDGMENTS I thank the following individuals and in- stitutions for their assistance or loan of ma- terial: H. J. Teskey, Biosystematics Re- search Centre, Agriculture Canada, Ottawa, Ontario; L. Knutson, Biosystematics and Beneficial Insects Institute, USDA, Belts- ville, Maryland; A. D. Bratt, Calvin College, Grand Rapids, Michigan. A special thanks to T. W. Fisher, University of California, Riverside, and L. Knutson for reviewing the manuscript. LITERATURE CITED Fisher, T. W. and R. E. Orth. 1983. The Marsh Flies of California (Diptera: Sciomyzidae). Bull. Calif. Insect Surv. 24, Univ. Calif. Press. vii + 117 pp. Knutson, L., R. E. Orth, T. W. Fisher, and W. L. Mur- phy. 1986. Catalog of Sciomyzidae (Diptera) of America north of Mexico. Entomography 4: 1-53. Rozko&sny, R. 1964. Zur Taxonomie der Gattung Pherbellia Robineau-Desvoidy (Diptera, Scio- myzidae). Acta Soc. Entomol. Cechoslov. 61(4): 384-390. Rozkosny, R. 1984. The Sciomyzidae (Diptera) of Fennoscandia and Denmark. Fauna Entomol. Scand. 14: 1-224. Steyskal, G. C. 1961. The North American Scio- myzidae related to Pherbellia fuscipes (Macquart) (Diptera: Acalyptratae). Pap. Mich. Acad. Sci. Arts Lett. 46: 405-415. PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 351-355 BIOLOGY OF A WOOD-NESTING WASP, MIMUMESA MIXTA (W. FOX) (HYMENOPTERA: SPHECIDAE), AND ITS PARASITE, ELAMPUS VIRIDICYANEUS NORTON (HYMENOPTERA: CHRYSIDIDAE) Jay A. ROSENHEIM AND J. KENNETH GRACE Department of Entomological Sciences, 201 Wellman Hall, University of California, Berkeley, California 94720; JKG Current address: Faculty of Forestry, University of Toronto, Toronto, Ontario M5S 1A1, Canada. Abstract.—An aggregation of Mimumesa mixta nests was located near the end of a decayed Douglas-fir timber in a second-story porch deck in Berkeley (Alameda Co.), California. Cells were excavated in the rotting wood in approximately linear series parallel to the grain and separated from one another by plugs of macerated wood fragments. Nests were provisioned with several adult and nymphal cicadellid and delphacid homopterans. Both M. mixta and its chrysidid parasite Elampus viridicyaneus Norton were reared from the nests. The sex ratio of M. mixta did not differ significantly from 1:1, while all emerging E. viridicvaneus were female. The cocoons of both wasps are described. The genus Mimumesa (Hymenoptera: Sphecidae) is distributed throughout the Nearctic, Neotropical, Palearctic, and Ori- ental Regions (Bohart and Menke, 1976). These wasps excavate multicellular nests in level or sloping ground, decaying wood, and plant stems and provision them with del- phacid or cicadellid prey (Bohart and Menke, 1976; Spooner, 1948; Tsuneki, 1959; Petit, 1979). Gurney (1951) provided the only de- scription of the nesting biology of a North American species of Mimumesa, that of Mi- mumesa nigra (Packard). The discovery of a large aggregation of Mimumesa mixta (W. Fox) nests provided an opportunity to study the biology of this species as well as that of its parasite, Elampus viridicvaneus Norton (Hymenoptera: Chrysididae). METHODS The timber containing the nesting aggre- gation was collected in Berkeley, Alameda County, California on 15 July, 1985. To simulate natural conditions the timber was maintained in an exterior screened cage in a shaded exposure at the Oxford Agricul- tural Tract, University of California, Berke- ley until the adult emergence occurred dur- ing May, 1986. Cell dimensions were measured to the nearest millimeter, and host and parasite cocoons were measured with a calipers accurate to 0.05 mm. Voucher spec- imens of both wasp species have been de- posited in the R. M. Bohart Museum of Entomology, Department of Entomology, University of California, Davis. RESULTS Nest location and architecture.— Nests were located near one end (approximately 0.6 m) of a large timber made by glue-lam- inating seven Douglas-fir, Pseudotsuga menziessi (Mirb.) Franco, 2 x 4 (inch) boards together. The timber was found in a stack of lumber that had been removed dur- ing the previous week from a second story 352 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-4. 1, Mimumesa mixta nesting aggregation in decayed end of a9 x 30 cm timber. 2, Linear series of M. mixta cells. 3, Cocoons of M. mixta (left) and Elampus viridicyaneus (right). Cocoons pictured in upper row are intact, while those below have been sectioned lengthwise to reveal internal structure. Small divisions of scale are in mm. 4, Cocoons of E. viridicyaneus (above) and M. mixta (below), showing emergence holes. Same scale as previous figure. porch deck. No active wasps were observed at the time of collection, the entire aggre- gation being composed of overwintering in- dividuals. The timber was located between 2.5 and 4 m above ground. At the time of collection the end of the timber containing the wasp nests was dry but already decayed by brown-rot fungi (Basidiomycetes). MM. mixta was apparently able to excavate the decay-softened wood. The nest aggregation consisted of over 400 cells which extended throughout the de- cayed portion of the timber but not into the sound wood (Fig. 1). Cells were arranged end to end in approximately linear series parallel to the grain (Fig. 2), each being sep- arated from the next by a plug of macerated wood fragments. Nests appeared to be in- terconnected, and we were unable to deter- mine where an individual nest began or end- ed. Measurements of 30 randomly selected cells gave a mean cell length of 9.3 + 1.8 mm and mean width of 4.0 + 0.6 mm (range 5-12 mm by 3-5 mm). No cell contained more than a single cocoon. Host and parasite populations.—After adult emergence the wood was dissected to provide an estimate of the sizes of the host and parasite populations (Table 1). A total of 44 M. mixta emerged successfully. An additional 119 M. mixta larvae, pupae, and adults were found inside cocoons; some of these were alive, and it is unclear to what extent additional emergence might have oc- curred. Two host cocoons, one containing a dismembered adult wasp and the other a mass of insect frass, had apparently been attacked by fly larvae of the family Sciari- VOLUME 89, NUMBER 2 Table 1. No. ofspecimens of Mimumesa mixta and Elampus viridicyaneus reared or dissected from a nest- ing aggregation. Species and Stage No. Mimumesa mixta Emerged adults @ 19 3 25 Pupae/adults within cocoons @¢ 333) 3 31 Pupae within cocoons, unsexable! 12 Larvae within cocoons 43 Emerged cocoons 131 Cocoons with fly feeding holes only 2 Elampus viridicyaneus Emerged adults ° 22 3 0 Pupae/adults within cocoons @ 12 3 0) Pupae within cocoons, unsexable' 5 Larvae within cocoons 18 Emerged cocoons 192 Cocoons with fly feeding holes only 3 ' Pupae that died before sclerotization. dae, which left a single, small (ca. 0.7 mm diameter), irregular hole in each cocoon. These fly larvae were common throughout the nest aggregation and appeared primarily to be fungivores, only incidentally attacking cocoons. The sexes of 63 of the sphecid pu- pae could be determined, which when com- bined with the data from the adults yielded a sex-ratio of 52 females: 56 males or 1: 1.08, which was not significantly different irom lln(Z-="—0, 39, P > 0.5). The ls! host cocoons with emergence holes exceed- ed the number of emerged adults by 87, indicating that some of the cocoons may have remained from one or more previous generations. Twenty-two adult EF. viridicyaneus emerged successfully. Twenty-five larvae, pupae, and adults were found inside co- coons; some of these appeared to be alive and might have emerged later. Three empty parasite cocoons had apparently been at- tacked by fly larvae. The 22 adults and 12 pupae whose sex could be determined were 353 Table 2. Species of Homoptera recovered from cells of Mimumesa mixta. Taxon No. Adults/Nymphs Delphacidae Delphacodes sp. 1 adult Cicadellidae Sorhoanus helvinus (Van Duzee) 59 adults Tiaja californica (Ball) 8 adults Euscelidius variegatus (Kirshbaum) 8 adults Amblysellus grex (Oman) 5 adults Endria lassus (Ball) 2 adults Calanana rubralineata (Beamer) 1 adult Reticopsis nubila (Van Duzee) 1 adult Deltocephalinae, undetermined 10 adults Deltocephalinae, undetermined 5 nymphs all females. Barring strongly sex-specific mortality in the immature stages, FE. viri- dicyaneus showed a female-biased sex ratio (Z-= 5.8.9P = 0,001). .mhe 192" E. wirdiey- aneus cocoons with normal emergence holes exceeded the number of observed emerged adults, suggesting the existence of one or more earlier generations of parasites in the same nesting aggregation. Nest provisions.—M. mixta provisioned cells with adult and nymphal Homoptera (Table 2). Nine species of cicadellids and one delphacid were recovered; 59 of the 100 prey individuals recovered represented a single species of cicadellid, Sorhoanus hel- vinus (Van Duzee). Intact prey were found both in cells in which the wasp larva had died and in cells containing cocoons. Two apparently complete cells contained 18 and 19 prey items, respectively. Mimumesa mixta cocoons.—Cocoons of M. mixta (Figs. 3, 4) were oblong, spongy in texture, and generally covered with wood and fragments of devoured provisions. Co- coons averaged 8.5 + 0.7 mm long by 3.3 + 0.3 mm wide (range 6.7-9.7 mm by 2.8- 3.8 mm, n = 60). The inner surface of the cocoon was light brown and very smooth; 354 the end opposite that with the generally off- center exist hole was coated thinly with the dark brown meconium. Elampus viridicvaneus cocoons.—Co- coons of EF. viridicyaneus (Figs. 3, 4) were superficially similar to those of M. mixta, being oblong and covered with wood frag- ments and prey debris. However, they were shorter, and the cocoon walls were crisp, paper-like, and had a varnished appearance. The inner cocoon surface was darker than that of M. mixta and was roughly textured. The exit hole was centered and represented the removal of the end of the cocoon. Mean cocoon dimensions were 6.7 + 0.8 mm long by 3.2 + 0.2 mm wide (range 4.9-9.0 mm by 2.6-3.88 mm, n = 42). DISCUSSION Mimumesa mixta. —Several Mimumesa species are known to nest in the soil, and Mimumesa dahlbomi (Wesmael), Mimu- mesa dahlbomi pacifica (Tsuneki), and Mi- mumesa nigra (Packard) are known to nest in decayed wood (Bohart and Menke, 1976; Spooner, 1948; Tsuneki, 1959; Petit, 1979). Spooner (1948) suggested that Mimumesa unicolor (Vander Linden) and Mimumesa littoralis (Bondroit) (as Mimesa celtica Spooner) may nest in plant stems, but /. unicolor has subsequently been found nest- ing in the ground (Petit, 1979) and the bi- ology of M. /ittoralis has not been described. The short description by Gurney (1951) of a M. nigra nest is the only description of the biology of a North American species. Gurney (1951) reared a single M. nigra male from a cocoon taken from a nest con- taining “‘a dozen or more cells” in the de- cayed end of a 2 x 4 (inch) fence board ca. 30 cm above ground. Although abandoned beetle borings are used by M. dahlbomi and M. dahlbomi pacifica to gain entry into the wood (Bohart and Menke, 1976; Spooner, 1948; Tsuneki, 1959), Gurney (1951) did not mention any evidence of beetle infes- tation in the board containing the M. nigra nest. Likewise, we found no evidence of PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON wood-boring beetle activity in the timber containing the M. mixta nests. M. mixta may either have excavated directly into the soft, decayed end-grain of the board or en- tered through cracks in the decayed wood. Species of the genus Psen, the only other genus in the subtribe Psenina known to nest in decayed wood (Bohart and Menke, 1976), apparently excavate directly into stumps (Iwata, 1938; Tsuneki, 1959). The number of empty host and parasite cocoons exceeded the number of newly emerged adults, suggesting that both M. mixta and its parasite were active at the nesting site for one or more previous gen- erations. The complete history of the nest- ing aggregation cannot, however, be in- ferred from our single collection due to the possible re-use of old cells by M. mixta. Mimumesa unicolor, the only Mimumesa species known to be attacked by chrysidid parasites, has been reported as a host of Chrysis succincta L., Hedychridium ardens Cocquebert, and Omalus auratus (L.) (Bo- hart and Menke, 1976; Moczar, 1967; Spoon- er, 1948). Elampus_ viridicvaneus.—The genus Elampus contains six species in North America (Bohart and Kimsey, 1982) for which the biology is almost completely un- known (Huber and Pengelly, 1977). The only published host record for a North American species is for E. viridicyaneus parasitizing Hoplisoides costalis (Cresson) (as Psam- maecius costalis (Cr.)) (Krombein, 1958). Huber and Pengelly (1977) relate that this record is based upon a single specimen in the U.S. National Museum bearing the label “Bred from nests of Gorytes (s.1.) from Hun- tington, L.I. Cocoon March 24, 1924. em. April 30, 1924. S. C. Bridwell.” Hoplisoides costalis is a ground-nesting, membracid- provisioning nyssonine (Evans, 1966), while M. mixta, as presented above, nests in de- caying wood, provisions with cicadellids and delphacids, and is in the subfamily Pem- phredoninae. Thus Krombein’s (1958) re- cord combined with that presented here im- VOLUME 89, NUMBER 2 plies a host range for FE. viridicyaneus that is broad in both the systematic and behav- ioral/ecological senses. Observations of sand grains trapped in the coarse integumentary punctations of curated specimens of E/am- pus spp. (Huber and Pengelly, 1977; Kur- czewski and Kurczewski, 1970) support the suggestion that the host pool of North American Elampus spp. includes at least some ground-nesting species. Spooner (1948), summarizing the European litera- ture, and Moczar (1967) together listed three species in the genus Mimesa as hosts of two European E/ampus species. These three Mi- mesa species, and indeed all the members of the genus for which biological informa- tion exists, construct nests in the ground (Bohart and Menke, 1976). Thus, world- wide, the genus Elampus does parasitize both wasps that excavate nests in rotting wood and wasps that nest in the ground. The genus Mimesa is closely related to Mi- mumesa, both being members of the sub- tribe Psenina (Bohart and Menke, 1976). The significance of the female-biased sex ratio of emerging FE. viridicyaneus is unclear. Museum collections of this and other E/am- pus species generally include approximately equal numbers of both sexes (Bohart and Kimsey, 1982; Huber and Pengelly, 1977). Highly female-biased sex-ratios for single batch rearings of chrysidid parasites are not uncommon (e.g. Krombein, 1967; Medler, 1964) and sex ratios may vary between sites (Krombein, 1967). ACKNOWLEDGMENTS We thank the following taxonomists for their identifications: R. M. Bohart (Spheci- dae), L. S. Kimsey (Chrysididae), Depart- ment of Entomology, University of Cali- fornia, Davis; J. P. Kramer (Homoptera), Systematic Entomology Laboratory, Agri- cultural Research Service, USDA; E. I. Schlinger (Sciaridae), Department of En- tomological Sciences, University of Cali- fornia, Berkeley. We are grateful also to R. 355 M. Bohart and G. W. Frankie (Department of Entomological Sciences, University of California, Berkeley) for critical readings of earlier drafts of the manuscript. This ma- terial is based in part upon work supported under a NSF Graduate Fellowship to JAR and by the Division of Entomology and Par- asitology, U.C. Berkeley (JKG). LITERATURE CITED Bohart, R. M.and A.S. Menke. 1976. Sphecid Wasps of the World. Univ. Calif. Press, Berkeley. 695 pp. Bohart, R. M. and L. S. Kimsey. 1982. A synopsis of the Chrysididae in America north of Mexico. Mem. Am. Entomol. Inst. 33: 1-266. Evans, H. E. 1966. The Comparative Ethology and Evolution of the Sand Wasps. Harvard University Press, Cambridge, Mass. 526 pp. Gurney, A. B. 1951. The nesting habits of Mimesa (Mimumesa) nigra (Packard). Proc. Entomol. Soc. Wash. 53: 280. Huber, J. T. and D. H. Pengelly. 1977. A revision of the genus E/ampus Spinola (Notozus Auctt.) (Hy- menoptera: Chrysididae) in America north of Mexico. Proc. Entomol. Soc. Ont. 108: 75-137. Iwata, K. 1938. Habits of some Japanese pemphre- donids and crabronids (Hymenoptera). Mushi 11: 20-41. Krombein, K. V. 1958. Hymenoptera of America North of Mexico. Synoptic Catalogue. First sup- plement. U.S. Dept. Agric., Agric. Monogr., Washington, D.C. 305 pp. 1967. Trap-nesting Wasps and Bees: Life Histories, Nests and Associates. Smithsonian Press, Washington, D.C. 570 pp. Kurczewski, F. E. and E. J. Kurczewski. 1970. An annotated list of cuckoo-wasps from Erie County, Pennsylvania (Hymenoptera: Chrysididae). Proc. Entomol. Soc. Wash. 72: 190-201. Medler, J.T. 1964. Parasitism of Eumeninae by cuck- 00 wasps in trap-nests in Wisconsin. Proc. Ento- mol. Soc. Wash. 66: 209-215. Moczar, L. 1967. Hymenoptera III. Chrysidoidea. Fauna Hungariae 86: 1-118. Petit, J. 1979. Note sur Mimumesa sibiricana R. Bo- hart (Hym. Sphecidae). Lambillionea 79: 9-14. Spooner, G. M. 1948. The British species of psenine wasps (Hymenoptera: Sphecidae). Trans. R. Ento- mol. Soc. Lond. 99: 129-172. Tsuneki, K. 1959. Contributions to the knowledge of the Cleptinae and Pseninae faunae of Japan and Korea (Hymenoptera, Chrysididae and Spheci- dae). Mem. Fac. Lib. Arts, Fukui Univ. (2, Nat. Sci.) 9: 1-78. PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 356-358 SYNONYMIC NOTES ON THE BETHYLIDAE DESCRIBED BY V. bE MOTSCHULSKY (HYMENOPTERA: ACULEATA) KARL V. KROMBEIN Department of Entomology, National Museum of Natural History, Smithsonian Insti- tution, Washington, D.C. 20560. Abstract.—The Motschulsky types of Bethylidae were studied. Do/us Motschulsky is synonymized with Epyris Westwood by designation of politus Motschulsky as type-species; the Egyptian politus and apicalis Motschulsky are new combinations in Epyris; the Cey- lonese subnitidus Motschulsky and opacicollis Motschulsky are new combinations in Ho- lepyris Kieffer, and the former species is synonymized under the latter. Lectotypes are designated for politus and opacicollis. The Ceylonese Homalus? amplipennis Motschulsky is a new combination in Holepyris. The Ceylonese Goniozus montanus Kieffer, replace- ment name for Bethylus distigma Motschulsky not Thomson, is confirmed as a species of Goniozus. The Russian coleopterist Motschulsky published a catalog of Ceylonese insects in three parts and a supplement (1861-65). He listed therein previously known species and described a number of new insects. Most of the new species were Coleoptera but he also included some species of Hymenoptera and other orders. The type series are in the Zoo- logical Museum, Moscow State University, USSR. Motschulsky described one genus and six species of Bethylidae in this catalog of Cey- lonese species, four species from Ceylon and two from Egypt. The Ceylonese insects came from J. Nietner, a coffee planter and ama- teur coleopterist. The specimens are not in good condition, some having suffered dam- age from dermestids or breakage so that ap- pendages are often lacking. The specimens are glued on small cards. One specimen of each species bears a small hand-written la- bel “Type” and a larger label bearing Mot- schulsky’s hand-written identification label. The identification labels of the Ceylonese species also bear a notation such as “‘I. or. Ceyl. Mt. N.E.” indicating that this speci- men came from the Oriental island of Cey- lon from ‘‘Montagnes de Nura-Ellia.”’ Spec- imens that I consider to be syntypes do not bear identification labels but were placed in the collection next to the specimen bearing the “‘type”’ label. Motschulsky did not des- ignate types in his paper. In the two species containing several syntypes, I have selected as lectotype the specimen bearing the “type” label. Motschulsky listed the localities from which the bethylids were described as ‘““Montagnes de Nura-Ellia” and as “Mont Patannas.’’ Nuwara Eliya is a town at an altitude of some 1800 m. Two mountains nearby are Hakgala and Pidurutalagala at- taining elevations of about 2100 and 2500 m respectively. Patannas is not a mountain; patanas are areas of montane grassland oc- curring at altitudes above 500 m. I am publishing the following synonymic notes in advance of a revisionary study of VOLUME 89, NUMBER 2 the Ceylonese Bethylidae so the informa- tion can be included in a world catalog of Bethylidae in preparation by Gordon Gordh. Subfamily Epyrinae Dolus Motschulsky The genus Dolus (1863: 27) has been a puzzle since its description. Dalla Torre (1898: 536) assigned it to the Dryininae; (Motschulsky placed all his bethylids under the heading Dryinides). Ashmead (1902: 272) placed it with question as a synonym of Mesitius sensu Ashmead not Spinola. Kieffer (1908) omitted it in his treatment of Bethylidae in Genera Insectorum, but in- cluded it as a questionable [zweifelhafte] ge- nus in his monograph of the family in Das Tierreich (1914). Motschulsky described Dolus for four new species, two from Ceylon, subnitidus and opacicollis, and two from Egypt, politus and apicalis. The first two species are congeneric with species placed in Holepyris Kieffer, 1904, and the last two with species placed in Epyris Westwood, 1832. In the interest of nomenclatorial stability I designate po/- itus as the type-species of Dolus, thus sink- ing that genus as a synonym of Epyris. Dolus politus and apicalis are new com- binations in Epyris. The former species was described from three conspecific females, one of them on a single card bearing “‘type”’ and identification labels, the other two fe- males on a single card without labels. I des- ignate as lectotype the specimen bearing the “type” label. Dolus apicalis is based on a unique female. Neither species is known to occur in Sri Lanka. It should be noted that Epyris apicalis Smith, 1874, becomes a secondary hom- onym with the transfer of apicalis Mot- schulsky to Epyris. I shall not propose a substitute name inasmuch as a junior syn- onym may be available for apicalis Smith. Holepyris opacicollis (Motschulsky), NEw ComMBINATION Dolus opacicollis Motschulsky, 1863: 28. 35¥7 Dolus subnitidus Motschulsky, 1863: 27-28. New SYNONYM. Dolus opacicollis was described from three conspecific females from “‘Montagnes de Nura-Ellia et Patannas.” One specimen bears a label “‘type” and another label on which Motschulsky wrote his identification and “I. or. Ceyl. Mt. Pat.” Two unlabeled females standing next to this specimen are syntypes; we know that Motschulsky had a series of opacicollis for he gave the length as 1%-1% lines. I designate as lectotype the specimen bearing the “type”’ and identifi- cation labels. Dolus subnitidus was de- scribed from a single female from ‘‘Mon- tagnes de Nura-Ellia’’; it has suffered some dermestid damage. The minor differences between the two species noted by Motschulsky fall within the normal range of variation of this taxon. The specimen of subnitidus has the head some- what less dull than is typical so I place it as a synonym of opacicollis. Ihave two females collected in Udawattakele Sanctuary, Kan- dy, by the Smithsonian’s Ceylon Insect Project, and a third labeled just Ceylon, col- lected by G. H. K. Thwaites, the first Di- rector of the Royal Botanic Garden at Per- adeniya near Kandy, and accessioned by the British Museum in 1867. These three spec- imens are very similar to the type of opa- cicollis. Holepyris amplipennis (Motschulsky), NEw COMBINATION Homalus? amplipennis Motschulsky, 1863: 26. Perisemus amplipennis Dalla Torre, 1898: 549. The unique type is a male from “‘som- mités du Mont Patannas.” It is not the op- posite sex of opacicollis (Motschulsky). Subfamily Bethylinae Goniozus montanus Kieffer Bethylus distigma Motschulsky, 1863: 26- 27. (Preoccupied in Goniozus by distig- mus Thomson, 1861.) 358 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Goniozus montanus Kieffer, 1908: 15 (New name for distigma Motschulsky.) The unique type is a female from ‘“‘Mon- tagnes de Nura-Ellia.”” Motschulsky wrote bistigma rather than distigma on his label. I can confirm Kieffer’s transfer of this species to Goniozus. ACKNOWLEDGMENTS I am grateful to my colleague Alexander V. Antropov, Moscow State University, USSR, for lending the Motschulsky mate- rial for study. I thank also Curtis W. Sa- brosky, Systematic Entomology Labora- tory, U.S. Department of Agriculture (ret.), for reviewing the manuscript and making helpful suggestions. LITERATURE CITED Ashmead, W.H. 1902. Family XX XII.—Bethylidae. In Classification of the fossorial, predaceous and parasitic wasps, or the superfamily Vespoidea. Can. Entomol. 34: 268-273, 287-290. Dalla Torre, K. W. von. 1898. Chalcididae et Proc- totrupidae. Catalogus Hymenopterorum 5: 1-598. Kieffer, J. J. 1908. Bethylidae. Jn Wytsman, Genera Insectorum 76: 1-50. 1914. Bethylidae. Das Tierreich 41: 1-595. Motschulsky, V. de. 1863. VI: Hyménoptéres. Jn Es- sai d’un catalogue des insectes de l’ile Ceylan. Bull. Soc. Imp. Nat. Moscou 36: 11-73. PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 359-362 HABITS AND BIOLOGY OF THE BEECH MEALYBUG, PELIOCOCCUS SERRATUS (FERRIS) (COCCOIDEA, PSEUDOCOCCIDAE) LouIsE M. RUSSELL Systematic Entomology Laboratory, BBII, Agricultural Research Service, USDA, BARC- West, Beltsville, Maryland 20705. Abstract. —The beech mealybug, Peliococcus serratus (Ferris), lives on Fagus grandifolia J. F. Ehrh. in eastern North America. In Maryland the species has two generations a year. Adult females settle on the bark of tree trunks and form a covering ovisac in which eggs are deposited from June until August and October through November. Eggs laid in summer hatch in about 7-14 days while those deposited in the fall overwinter. Mealybugs and eggs are destroyed by adverse weather conditions, parasitoids, and predators. Annotated citations are given to literature on the species. This article portrays, for the first time, the biology and habits of the beech mealy- bug, Peliococcus serratus (Ferris), in North America. It supplements the meager data available on a scale insect that lives on American beech, Fagus grandifolia J. F. Ehrh. (= F. americana Sweet), a valued for- est and shade tree. The mealybug merits study because of general interest in the ecol- ogy of indigenous insects and because it adds to the little that is known of the life cycles of most mealybugs native to the Nearctic fauna. The beech mealybug occurs sparingly and would achieve pest status only under greatly changed conditions. Although P. serratus apparently is restricted to Fagus (Fagaceae) in North America, it was re- ported from Corylus avellana L. (Betula- ceae) in Italy by Tranfaglia (1976). Literature on P. serratus is limited to the following annotated citations: Phenacoccus serratus Ferris, 1925: 231-232 (description, 3 collections); Trimble, 1928: 43 (recorded as rare); Friend, 1932: 596 (ovisacs described); Britton, 1933: 375 (recorded); Rau, 1942: 124 (location of immatures, attended by ants, reared parasites Homalotylus sp. and Leptomas- tidea sp.); Herting and Simmonds, 1972: 118 (rerecorded parasites listed by Rau). Peliococcus serratus (Ferris), Ferris, 1950: 118 (redescription, 2 additional collec- tions); Baker, 1972: 99 (erroneously re- corded from birch); Tranfaglia, 1976: 134-136 (redescribed from Corylus ay- ellana in Italy, biology, habits, attended by ants, Crematogaster sp.), 1981: 9 (re- ferred to 1976 report); Drooz, 1985: 94 (distribution in U.S.). METHODS My study of Peliococcus serratus extend- ed from October 1977 through September 1986 in a forested area in Silver Spring, Montgomery County, Maryland. About 50 trees were scrutinized to a height of 4.5 m with the unaided eye and to 9.1 m with binoculars. Tree trunks harboring mealy- bugs or ovisacs measured 15.2—20.3 cm in diameter at base. Insects or ovisacs were located 0.3-2.4 m from the ground on tree trunks. Mealybugs without ovisacs were ob- 360 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. served on leaves and twigs 2.4 m above ground although they unquestionably were present at a much greater height. Trees were examined monthly all year and, if infested, usually were scrutinized weekly or daily April through November. Mealybugs or ovisacs were found on only four trees, one of which was located | km from the others and on which ovisacs were found only in 1986. The other trees were in close prox- imity, one being 3.6 m from two that were 1.5 m apart. Mealybugs or ovisacs were present on one tree each of the 9 years, on one in 1981, 1983, 1985, 1986, and on the other only in 1985. Tops of the two closest trees intertwined. Specimens were not seen from October 1983 to November 1984, the longest period in which mealybugs or ovi- sacs were not observed. After examination of the trees, immatures, adult females, and Ovisacs were brought into the laboratory for further study. Ovisacs of Peliococcus serratus in knothole on bark of tree trunk. RESULTS Ovisacs are formed gradually in 3-6 days and when complete are subrectangular, 5— 8 mm long, and pure white (Fig. 1). Those formed in the summer tend to be smaller than those made in the fall. The outside is rather feltlike, the inside is fluffy and fila- mentous, and they are thicker dorsally than ventrally. When females are ready to oviposit, they crawl down the tree trunk from the tree can- opy and usually settle in a knothole, scar, crevice or other rough, protecting place. Oc- casionally, however, they settle on smooth bark or move to a second location even after the ovisac is formed. At this time the fe- males are deep pink or dark purple and have a thin, white, waxy substance arranged in transverse rows across the body. A large fe- male in this condition on 6 October 1985 had a little white fuzzy material on the body, VOLUME 89, NUMBER 2 Adult Females Eggs Fig. 2. Seasonal history of Peliococcus serratus in Maryland. Solid line denotes first generation, broken line second generation. more posteriorly than anteriorly, on 7 Oc- tober. The buildup continued through 11 October and on 12 October the ovisac was complete. This female started laying eggs 3- 4 days after the ovisac was completed. On 30 October the ovisac was removed and brought to the laboratory. The shrivelled, but soft body of the female was located at one end and 225 eggs filled the remainder of the ovisac. The female died, and a few eggs hatched within 3 days but most of them collapsed. The development of this female, Ovisac, and eggs was analogous to that of other specimens. Eggs deposited in October and November overwinter and hatch from late April to ear- ly May of the following year (Fig. 2). Crawl- ers travel up tree trunks and onto the branches where they and later stages occur on the lower surface of leaves and on bark in twig axils. In this location adults develop in approximately 4 weeks. From late June into July and occasionally into August, fe- males come down the tree trunks, form ovi- sacs, and lay eggs which hatch in 7-14 days. Only 55-60 eggs were found in summer ovi- sacs and only 60 eggs were counted in | ovisac formed in October. However, 200- 225 eggs were found in most ovisacs formed in the fall. Initially the eggs are bright yellow and some remain yellow, but the majority turn pink, and some, presumably dead, be- come black. Descending and ovisac-forming females were not found in September, but second stage specimens and adult females were present on leaves and twigs at this time. Owing to the location of the insects in the tree canopy, I was unable to determine the length of the first and second stages. Adult males were not found, but 6 male cocoons, possibly of P. serratus because no other co- coon-forming species was present, were col- lected in July and September. Rau (1942) noted that P. serratus was oc- casionally attended and shielded by ants, and Tranfaglia (1976) wrote that the species is habitually cultured by a formicid for the production of honeydew. I have not ob- served these activities although I have seen ants on the same tree trunks as the mealy- bugs. P. serratus exists under hazardous con- ditions. Insects and ovisacs are dislodged by strong winds and heavy rains almost in- stantly, and they are attacked by hymenop- terous parasites (Rau, 1942; Herting and Simmonds, 1972). I have observed adult hymenopterous parasites, coccinellid and chrysopid larvae on tree trunks near mealy- bugs, syrphid larvae near and inside mealy- bugs, and both hibernating and predaceous 362 mites in ovisacs. Ovisacs also are some- times covered with fungus. Ovisacs, females and eggs may be totally destroyed by one or more of these agents within 8 hours. The distribution of P. serratus as indi- cated by literature citations and/or the pres- ence of specimens in the National Collec- tion of Coccoidea at Beltsville, Maryland is as follows: Canada.— Ontario (Ferris, 1925, 1950). United States.—Connecticut: (Friend, 1932), (Britton, 1933), USNM; District of Columbia: USNM; Maryland: (Ferris, 1950), USNM; Massachusetts: USNM; New Hampshire: USNM; New Jersey: USNM; New York (Ferris, 1925, 1950), USNM; Ohio: (Ferris, 1925, 1950), USNM; Penn- sylvania: (Trimble, 1928), USNM; Tennes- see: USNM; Virginia: (Drooz, 1985), USNM. Europe.—Italy: Campania (Tranfaglia, 1976). ACKNOWLEDGMENTS I gratefully acknowledge the assistance of members of the Systematic Entomology Laboratory, BBII, Agricultural Research Service as follows: J. M. Kingsolver for re- viewing the manuscript, D. R. Miller for identifying the mealybugs, and M. B. Sto- etzel for photographing the ovisacs and giv- ing helpful suggestions. I thank J. A. Da- vidson, University of Maryland, College Park, Maryland, and Michael Kosztarab, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Virginia Polytechnic Institute and State University, Blacksburg, Virginia for review- ing the manuscript. LITERATURE CITED Baker, W. L. 1972. Eastern Forest Insects. U.S. Dept. Agr. For. Serv. Misc. Publ. 1175. 642 pp. Britton, W. E. 1933. Entomological features of 1932. Conn. Agr. Exp. Sta. Bull. 349: 369-381. Drooz, A. T. 1985. Insects of Eastern Forests. U.S. Dept. Agr. For. Serv. Misc. Publ. 1426. 608 pp. Ferris, G. F. 1925. Notes on Coccidae XI (Hemip- tera). Can. Entomol. 57(9): 228-234. 1950. Atlas of the Scale Insects of North America. (ser. 5) V: vii + 278 pp. Stanford Uni- versity Press, Stanford, California. Friend, R. B. 1932. A new scale insect on beech. Conn. Agr. Exp. Sta. Bull. 338: 596. Herting, B. and F. F. Simmonds. 1972. A Catalogue of Parasites and Predators of Terrestrial Arthro- pods. Commonwealth Inst. Biol. Control. 210 pp. Rau, G. J. 1942. The Canadian apple mealybug, Phenacoccus aceris Signoret, and its allies in northeastern America. Can. Entomol. 74(7): 118- 2S. Tranfaglia, A. 1976. Studi sugli Homoptera Coccoi- dea IV. Su alcune cocciniglie nuove 0 poco con- osciute per |’Italia (Coccidae, Eriococcidae, Pseu- dococcidae). Boll. Lab. Entomol. Agr. “Filippo Silvestri’? Portici 33: 128-143. 1981. Studi sugli Homoptera Coccoidea V. Notizie morphosistematiche su alcune specie di cocciniglie con descrizione di tre nuove specie di Pseudococcidi. Boll. Lab. Entomol. Agr. “Filippo Silvestri’? Portici 38: 3-28. Trimble, F. M. 1928. Scale insects of Pennsylvania (Homop.: Coccidae). Entomol. News 39(2): 42- 47. PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 363-366 A NEW SUBFAMILY OF NOGODINIDAE (HOMOPTERA: FULGOROIDEA) WITH THE DESCRIPTION OF A NEW SPECIES OF GASTRINIA R. G. FENNAH Commonwealth Institute of Entomology, % British Museum (Natural History), London SW7 5BD, United Kingdom. Abstract. —The genus Gastrinia Stal (Homoptera: Fulgoroidea) is redefined and trans- ferred from Issidae to Nogodinidae, and a new subfamily is proposed for its reception. Supplementary characterization is provided for the type species, G. vaginata Stal, and a new species, G. phidon, is described from Minas Gerais, Brazil. Stal (1866a) referred his genus Gastrinia to his new subfamily Tropiduchida, and Melichar (1914) placed it in his new tribe Hiraciini. In 1982, the present writer ex- cluded both Hiracia and Gastrinia from the Tropiduchidae and transferred them to the Issidae. Further study of these genera has confirmed the conclusion that Hiracia is an issid, but has revealed that Gastrinia differs from both Issidae and Nogodinidae in the proportions of the abdominal laterotergites and pleurites, the shape of the spiracles, and the structure of the aedeagus and the ovi- positor. However, the number and arrange- ment of the post-tibial and post-tarsal spines, the relationship of the basal sclerite of the tegmen with the base of the clavus, the fre- quency of branching and direction of the wing-veins and the proportions of the gen- ital styles are broadly similar to correspond- ing features to be found in Nogodinidae, though not in combination. In view of the magnitude of the differences that separate Gastrinia from all other Nogodinidae, a new subfamily is now erected for its accom- modation, characterized as follows. Family Nogodinidae Gastriniinae, NEw SUBFAMILY Habitus broad and depressed. Pronotum much wider than head, with 2 carinae be- tween eye and tegula. Tegmen with basal cell absent, basal sclerite narrow and straight. Post-tibia with 10-12 long teeth apically; basal metatarsal segment rather long, with coarse teeth apically. Abdominal spiracles elongate, narrow, embedded in lateroter- gites strongly obliquely to lower margin; la- terotergites IV-VI not much longer than broad, and much broader than pleurites. Genital style elongate, with narrow sides and a short process dorsally at apex. Aedeagus with a short basal collar. With the exception of the number of api- cal teeth on the post-tibia, none of the above characters occurs in the nominate subfam- ily. Gastrinia Stal Gastrinia Stal, 1859: 319. Type species, Gastrinia vaginata Stal. Vertex pentagonal. Frons strongly de- flexed ventrocaudad, median disc promi- nent at base, sublateral carinae curving mesad basally and not meeting anterior margin of vertex. Post-clypeus with sides very narrow, and visible in anterior view. Antennae with second segment globose or nearly so, infuscate except at apex, with pal- lid disc sensilla, microsetae minute, mac- rosetae apparently absent. Ocelli present. Rostrum distally overlapping post-coxae. 364 Dt pl s Figs. 1-13. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Vee lyre Z iy Lily OF y 1-12, Gastrinia phidon. 1, Head and thorax, dorsal view. 2, Head, thorax and base of tegmen, side view. 3, Frons and clypeus, anteroventral view. 4, Basal part of third antennal segment. 5, Tegmen. 6, Apex of post-tibia and basal 2 segments of metatarsus, ventral view. 7, Laterotergite with spiracle (It), pleurite (pl) and lateral part of sternite (s) of fourth abdominal segment. 8, Ninth abdominal segment and anal segment of male, anterodorsal view. 9, Male genitalia, right side. 10, Aedeagus, ventral view. 11, Apex of aedeagus, left side, with spinose process slightly displaced outward. 12, Phallobase (suspensorium), posteroventral view, with basal part of phallus emerging from orifice. 13, Gastrinia vaginata. Female genitalia, right side. Pronotum much wider than head, and broader than long in middle (about 4:1), disc slightly produced anteriorly, posterior margin shallowly concave; 2 carinae be- tween eye and tegula. Mesonotum almost flattened, with lateral carinae strongly con- verging cephalad. Tegula large, exposed, and abruptly decurved. Post-trochanters rock- ing mesad-laterad. Post-tibia with 4 spines laterally, 10-12 apically. Basal metatarsal segment with 10-13 coarse teeth an a tract of sparse long setae; second segment with 2 teeth; apical margin between them shallow- ly convex. Tegmen coriaceous, distally sub- tectiform, costal and commissural margins parallel, and apical margin angulate, scarce- ly surpassing abdomen, Sc+R scarcely bent at junction with M basally, M and Cu, sim- ple to beyond level of apex of clavus, an oblique row of veinlets from apex of clavus to M near apical margin. Basal cell obsolete, basal sclerite narrow, straight. Wing-tuck- ing process absent. Wing ample, with apical margin not deeply cleft, and costal margin straight. Venation regular, not bent ante- riorly in distal half, M with 2-3 branches apically, Cu, with 3-5 branches, apical cells almost parallel-sided, much longer than broad. Abdomen strongly dorsoventrally compressed; tergites narrowly divided by membrane along middle line, sternites not divided medially, laterotergites large, little longer than broad, and several times broad- er than pleurites; spiracles greatly elongated, embedded in laterotergites strongly obliquely to lower margin. Pygofer rela- tively short, with a small angulate medio- ventral process. Aedeagus long, straight and tubular, supported in basal quarter by a short tubular suspensorium. Genital style elon- VOLUME 89, NUMBER 2 gate, with narrow sides and a short process dorsally at apex. Anal segment of female elongate, with lateral margins deep and de- cumbent. Ovipositor elongate, slender and porrect. Gastrinia vaginata Stal Fig. 13 Gastrinia vaginata Stal, 1859: 319. Hiracia lacerdae Signoret, 1861: 57. (syn.) Stal, 1866b: 393. Female (supplementary description).— Length with tegmen, 15 mm; tegmen, 11.8 mm. Vertex broader than long (almost 1.2:1). Frons longer medially than broad (1.2:1), medially ecarinate. Rostrum with apical segment longer than broad in side view (6.6: 1). Basal metatarsal segment with 13 teeth api- cally, longer dorsally in middle line than wide between tips of outermost teeth (1.9:1), and than length of outer apical spine mea- sured from level of apex of mid-dorsal mar- gin (2.1:1). Second metatarsal segment lon- ger dorsally than wide between apical spines (about 1.2:1). Tegmen longer than broad (3.0:1), cell PCu longer than common claval vein (about 1.3:1). Tegmen with 2 narrow oblique bands from Sc to basal angle of clavus and from Sc to apex of clavus, reddish brown. Ninth abdominal segment very short dor- sally, overlapped by seventh and eighth ter- gites. Anal segment in side view longer than deep dorsoventrally (about 4.5:1), narrow- ing and weakly ascending distad and nar- rowly rounded apically; sides decumbent, ventral surface deeply hollowed to ensheath Ovipositor; anal foramen situated at middle. Ovipositor slender, much longer than deep in side view (about 18:1), of almost equal width throughout, and weakly ascending distad. Seventh sternite short, not quite overlapped by sixth, with hind margin weakly concave; area between this and base of ovipositor apparently consisting of tough membrane, and with a weak median ridge. 365 Material examined. | 2, Brazil, Bahia (coll. Signoret). This specimen, determined by Stal as be- longing to his G. vaginata, (Melichar 1914: 210) agrees with the description and illus- tration of Hiracia lacerdae given by Sig- noret, and also agrees with the description of Gastrinia vaginata Stal based on a female from Bahia, and with the figure of G. (= Am- fortas) vaginata Stal given by Melichar (1914: fig. 30). It is one of two specimens from Bahia in the Signoret Collection in the Naturhistorisches Museum, Vienna, and the only one that agrees with the original de- scription of H. lacerdae. It is here desig- nated as the lectotype, and has been labelled as such. The status of the second female from Ba- hia in the Signoret Collection, labelled ““Ba- hia, Coll. Signoret, det. Signoret,” and con- sidered to be specifically distinct from G. vaginata by Melichar must remain unset- tled until further material is available for study. It is not mentioned by Signoret, and its size and tegminal marking are not cov- ered by the description given for H. /acer- dae. The length/greatest breadth ratios of the frons, apical segment of the rostrum, basal metatarsal segment, second metatarsal seg- ment and tegmen are 1.2:1,5.1:1, 1.6:1, 1.2:1 and 2.9:1, respectively, and the ratio of lengths of tegminal cell PCu/common claval vein, about 1.3:1. Gastrinia phidon Fennah, NEw SPECIES Figs. 1-12 Male. — Length with tegmen, 9.0 mm; teg- men, 7.0 mm. Vertex broader than long (slightly less than 1.1:1). Frons longer me- dially than broad (1.2:1), feebly medially carinate in distal half. Rostrum with apical segment longer than broad in side view (4.9:1). Basal metatarsal segment with 12 teeth apically, longer dorsally in middle line than wide between tips of outermost teeth (1.5:1), and than length of outer apical spine 366 measured from level of apex of mid-dorsal margin (1.9:1). Second metatarsal segment longer dorsally than wide between apical spines (1.1:1). Tegmen longer than broad (2.6:1), cell PCu longer than common claval vein (2.6:1). Wing with Sc 2-3 branched, R simple, M and Cu, each with 3 branches, PCu forked basad of level of Cu, fork. Dorsally yellowish brown, mottled with paler spots. Mesonotum with a suffusion in each lateral field and a small round spot near each posterolateral margin of disc, piceous. Lower surface of thorax and legs more or less reddish brown, mottled with pale round spots; abdomen ventrally pale brown with fuscous spots at base of setae. Tegmen light yellowish brown, with light reddish brown markings in depressions; venation coarse, with supernumerary irregular veinlets, mostly concolorous, but with a few spots on main veins in corium and clavus and a larg- er spot close to M, ,, subapically, dark red- dish brown or piceous. Wing uniformly dilute fuscous, apical veins almost concol- orous, but orange brown on margin at apices of R, M and Cu;; longitudinal veins slightly darker. Anal segment of male relatively large, in dorsal view widest near base, with lateral margins curving to deeply rounding apex; anal orifice in distal half; anal style very short. Pygofer with lateral margins sinuate, dorsolateral angles obscure; medioventral process broader at base than long. Aedeagus porrect caudad, with lateral margins of phallobase meeting at an obtuse angle below phallus in basal quarter; a pair of delicate spinose processes ventrolaterally near apex of aedeagus, each directed cephalad and lying close against ventrolateral surface. Genital style in side view about 3.5 times as long as wide, and widest near base, stiffened with a shallow ridge internally, extending from base to apex; dorsal and ventral margins gradually converging distad in basal three- quarters, thence subparallel; apical margin shallowly rounded, shortly produced dor- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON sally in a peg-like process. Length, 9.0 mm; tegmen, 7.0 mm. Holotype ¢.— Brazil: Minas Gerais, Pedra Azul, xii. 1970, (F. M. Oliviera) in British Museum (Natural History). This species differs from G. vaginata and the second specimen of Gastrinia in the Sig- noret collection in the ratio of the basal width of the frons to the width at the frontoclypeal suture (about 1.1:1 in G. phidon and 1.3:1 in the other two), the ratio of length to width in the tegmen (2.6:1 in G. phidon and 3.0:1 and 2.9:1 in the others) and the relative lengths of cell PCu and the common claval vein in the tegmen (2.6:1 in G. phidon and about 1.3:1 in the others). The name phidon is a classical personal name, and is used in apposition. ACKNOWLEDGMENTS I thank W. J. Knight (British Museum (Natural History)) for the privilege of study- ing the Fulgoroidea in his charge, A. Kal- tenbach (Naturhistorisches Museum, Vi- enna) for the loan of the female of Hiracia lacerdae Signoret referred by Stal to Gas- trinia vaginata Stal and for supplying in- formation on a second female in the Sig- noret collection, and two anonymous reviewers for helpful comments on the manuscript. LITERATURE CITED Fennah, R. G. 1982. A tribal classification of the Tropiduchidae (Homoptera: Fulgoroidea), with the description of a new species on tea in Malaysia. Bull. Entomol. Res. 72: 631-643. Melichar, L. 1914. Monographie der Tropiduchiden. Verh. Naturforsch. Ver. Brunn 53: 1-145. Signoret, V. 1861. Description de quelques Hémip- téres nouveaux. Ann. Soc. Entomol. Fr. (4)1: 55- 58. Stal, C. 1859. Novae quaedam Fulgorinorum formae speciesque insigniores. Berl. Entomol. Zeit. 3: 313- B27 . 1866a. Hemiptera Homoptera Latr. Hemip- tera Africana 4: 1-276. . 1866b. Analecta hemipterologica. Berl. Ento- mol. Z. 3: 319: PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 367-368 NOTE The Prevalence of Icosta americana (Diptera: Hippoboscidae) on Ruffed Grouse (Bonasa umbellus) in Wisconsin Little has been published regarding the distribution of Jcosta americana (Leach) from ruffed grouse, Bonasa umbellus L., in Wisconsin. MacArthur (1948. Bull. Public Mus. Mil. 8: 367-440) reported americana from a “partridge,” locality unknown and Bequaert (1955. Entomol. Am. 34—35(N:S.): 1-611) reported infected grouse from three counties. Because of the lack of information from other areas of the state, our study was initiated. From 1968 through 1985, 206 ruffed grouse were collected by the senior author and his father from seven Wisconsin coun- ties and examined for americana. Preva- lence of infection in the samples was: 11/ 80 Richland Co.; 1/98 Portage Co.; 0/17 Adams Co.; 0/4 Chippewa Co.; 0/2 Forest Co.; 0/4 Monroe Co.; and 0/1 Waushara Co. Seven grouse from Richland Co. harbored one americana each and four birds carried two flies each, while the infected Portage Co. grouse had one. Infection rate in south- western Richland Co. (13.7%) was consid- erably higher than the approximately 1% in the more central Portage Co. In the 1984 and 1985 seasons Wisconsin hunters were asked to collect any flies seen on ruffed grouse. Six additional hippobos- cids were collected: two from Tremapealeau Co., and one each from Grant, Richland, Jackson, and Rusk counties. All of these are in the lower two-thirds of Wisconsin with the exception of Rusk which is in the upper one-third. The reason for this distribution is un- known and may be related to one or more ecological factors. Bequaert (op. cit.) sug- gested americana may not occur north of 48°10’ latitude, approximately the Cana- dian border in central United States. Ben- nett (1961. Can. J. Zool. 39: 379-406) found only a “few” specimens of americana from more than 400 ruffed grouse in Ontario and 63 ruffed grouse and spruce grouse (Can- achites canadensis L.) taken between July and November in Algonquin Park, Ontairo just slightly south of 48°10’ north latitude. In Wisconsin there is a tension zone di- viding the state into two floristic provinces (Curtis, 1959, The Vegetation of Wisconsin. The University of Wisconsin Press, Madi- son, WI). The southwest third of the state contains southern hardwood forests with some prairie elements, and the northern half features coniferous-hardwood forests with a few boreal elements. The narrow tension zone separating each province contains members of each. Bequaert (op. cit.) records americana from ruffed grouse in Clark, Marathon, and Rusk counties. Although these counties are just north of the tension zone, our data suggest this fly is more common on grouse in or below the tension zone. Bequaert (op. cit.) believed that native galliform birds were the original hosts of americana, and that raptors later acquired the fly by preying on game birds. Converse- ly, Bennett (op. cit.) believed americana is primarily a parasite of birds of prey and occur only accidentally on galliformes. By definition, accidental parasites are rare, do not breed on their host, and remain with the host only fora short period of time. That 13.7% of ruffed grouse from Richland Co. have this parasite argues against its being accidental on grouse. Also, in three in- stances female americana were captured alive and viable. When placed in a jar, they 368 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON larviposited. Female americana carrying larvae and a male were collected as late as January 28. These data suggest americana is common on grouse in southern Wisconsin. This species most likely breeds on the host, and female americana may overwinter with lar- vae in utero. Voucher specimens of two americana from ruffed grouse are deposited in the Na- tional Museum of Natural History in Wash- ington, D.C. The remainder of the speci- mens are in the Museum collection at the University of Wisconsin-Stevens Point. We thank D. J. Taft for his help in col- lecting americana. Stephen J. Taft, Department of Biology, University of Wisconsin—Stevens Point, Ste- vens Point, Wisconsin 54481 and Susan Marcquenski, Wisconsin Department of Natural Resources, Box 7921, Madison, Wisconsin 53707. PROC. ENTOMOL. SOC. WASH. 89(2), 1987, p. 368 Book NOTICE The Insect and Spider Collections of the World, by Ross H. Arnett, Jr. and G. A. Samuelson, assisted by John B. Heppner, Gordon M. Nishida, J. Charles Watt, and Robert E. Woodruff. E. J. Brill Publishers, New York, NY. 1986. 220 pp. Cost: $19.95 (paper with plastic comb binding). Part I has a list and description of 918 public collections known to exist; 377 of the major collections are described in detail. They are arranged alphabetically by coun- try, state/province, and city. Every country in the world is listed—those without known collections are indicated. The mailing ad- dress of each museum is cited, and a four- letter coden is assigned to each. If a ques- tionnaire was returned, the names of cura- tors, phone numbers, details about the size and content of the collection, and data about primary types and special collections are given. Part II has a list and descriptions of 211 private collections, arranged alphabetically by owner’s name, with pertinent data. A coden is assigned to each collection. To systematists who borrow specimens for research and to collectors who would like to have their collections identified and used in research, this list will be most wel- come. (E. J. Brill, founded in 1683 in The Netherlands, now has an office in USA: Suite 404, 225 W. 57th St., NYC 10019.) T. J. Spilman, Systematic Entomology Laboratory, BBII, Agricultural Research Service, U.S.D.A., % U.S. National Mu- seum NHB 168, Washington D.C. 20560. PROC. ENTOMOL. SOC. WASH. 89(2), 1987, p. 369 NOTE New Distributional and Rearing Records for Neotropical Flower Flies (Diptera: Syrphidae) The biology of most Neotropical flower flies is poorly known. In this paper, I report new information on the distribution, flower visitation, or larval habitat of 4 syrphids. Copestylum sexmaculatum (Palisot de Beauvois) (= C. pallens of authors). — First state record: Illinois, Pope County, Bell Smith Springs Rec. Area, 19 July 1975, 14 feeding on blossoms of Pycnanthemum ten- uifolium Schrad. (Labiatae). Wirth et al. (1965. In Stone et al., Cat. Diptera America North of Mexico, Agric. handbook 276: 602) and Thompson (1981. Mem. Entomol. Soc. Wash. 9: 138-139) reported this fly from North Carolina, Florida to Arizona, and the West Indies. Meromacrus panamensis Curran.— First United States record: Texas, Cameron County, Sabal Palm Grove Sanctuary, 14 December 1984, 1 4 feeding on blossoms of Xylosma flexuosa (H.B.K.) O. Ktze. (Fla- courtiaceae). Thompson (ibid.: 157; 1985, pers. comm.) previously noted that this species is distributed from Mexico to Brazil. Meromacrus pratorum (Fabricius). — First rearing record: Puerto Rico, El Yunque Rec. Area, | é reared from larva collected in fluid and debris of upright flower bract of Heli- conia sp. (Musaceae) on 29 May 1981. Sack (1921. Arch. Naturgesch. (Abt. A) 87(3): 144-145) reported that rat-tailed larvae of M. pratorum live in sediment of wet rot pockets of trees. His identification was ap- parently incorrect because Thompson (1981: 157) found M. pratorum to be endemic to the West Indies. Snow (1958. Ecology 39: 83-88) observed larvae of M. acutus (Fa- bricius) in decaying stumps filled with water. The larval habitat of M. pratorum in Puerto Rico more closely resembles that of Qui- chuana angustiventris (Macquart) reared from Heliconia bracts by Seifert and Seifert (1976. J. N.Y. Entomol. Soc. 94: 233-242; 1979. Ecology 60: 462-467) than that of other Meromacrus species. Additional study is needed to determine the diversity of lar- val habitats used by species of Meromacrus and related genera. Polybiomyia macquarti Shannon. —First rearing record: Texas, Cameron County, Sa- bal Palm Grove Sanctuary, 2 2? reared from larvae collected from sap-soaked detritus in small rot pocket in trunk of Leucaena pul- verulenta (Schlecht.) Benth. (Leguminosae) on 14 December 1984. Other cerioidine species reared by Johnson (1893. Entomol. News 4: 91), Banks (1902. Proc. Entomol. Soc. Wash. 5: 310) and Maier (1982. Proc. Entomol. Soc. Wash. 84: 603-609) have similar larval habitats. I thank F. Christian Thompson (System- atic Entomology Laboratory, Agricultural Research Service, USDA) for identifying most of the syrphids. The staff of the Sabal Palm Grove Sanctuary, Brownsville, Texas provided valuable biological information. Chris T. Maier, Department of Ento- mology, The Connecticut Agricultural Ex- periment Station, P.O. Box 1106, New Ha- ven, Connecticut 06504. PROC. ENTOMOL. SOC. WASH. 89(2), 1987, p. 370 NOTE Anomalous Tubercle Patterns Found on Lymantria dispar (L.) Caterpillars in the Field in Maryland (Lepidoptera: Lymantriidae) The gypsy moth caterpillar is conspicu- ous by having a double row of colored spots along its dorsal surface during and after the fourth instar. The anterior 5 pairs are blue, while the posterior 6 pairs are a dull-to- bright red. These spots are dorsal tubercles that form the basal portion of hair tufts, and they represent outgrowths of the integument bearing numerous setal sockets (Traxler, J. LOTT AN YoEntomol soc: 853 7197). In the spring of 1986, I had occasion to observe gypsy moth caterpillars from near- ly 50 locations in Northern Maryland, amounting to several thousand collected, and many thousands more observed 1n the field either by myself, or by collaborators Kathleen Tatman, Kevin Boyd, and Anne Wieber. From these, 17 individual cater- pillars were collected having anomalous tu- bercle patterns. Eight had only the first 4 anterior spot-pairs blue while the 7 poste- rior spot-pairs were red. Four individuals were missing one red spot (never the same one); three individuals were missing a pair of red spots (never the same pair), while one individual was missing 3 red spots. The final oddity was a caterpillar whose dorsal integ- ument was albino along its entire length on the left side, while the right side was a nor- mal mottled black. All dorsal tubercles were present, and of normal pigmentation, on this individual. All specimen were collected from 6 locations in northern Baltimore County, Maryland. All of the above caterpillars appeared normal except for their color patterns. The 11 individuals not parasitized or diseased produced apparently normal pupae and adult females. If any of these patterns prove to be heritable, it could prove the basis of a marked strain of gypsy moths of potential utility in sterile-male release programs or for release-recapture ecological studies. Ralph E. Webb, Florist and Nursery Crops Laboratory, HSI, Agricultural Research Service, U.S.D.A., Beltsville, Maryland 20705. SociETY MEETINGS 925th Regular Meeting— October 2, 1986 The 925th Regular Meeting of the Ento- mological Society of Washington was called to order by President E. M. Barrows in the Naturalist Center, National Museum of Natural History, at 8 p.m. on October 2, 1986. Twenty-one members and four guests were present. Minutes of the previous meet- ing were read. Membership Chairman, G. White, read the names of the following ap- plicants for membership: P. Adler, Clemson, South Carolina; C. Agnew, College Station, Texas; D. Haile, Reading, Pennsylvania; K. Hoffman, Clemson, South Carolina; E. Lip- pert, Guelph, Ontario, Canada; J. Mackley, Laredo, Texas; P. Newhouse, Silver Spring, Maryland; A. Zuccaro, Jr., Natchez, Mis- sissippi; A. Ritchie, Hamden, Connecticut; R. K. Robbins, Washington, D.C. Editor R. Gagné displayed the recently rediscovered Society seal embosser. This embosser, which may have been used in the past to emboss the seal on portfolios of greetings and other correspondence, will re- main with the Editor’s office. R. Gagné showed slides of various ceci- domyiid galls on hickory including husk swellings caused by “‘Cecidomyia”’ nucicola. The discovery of this species, which has not been reported since its description in 1870, except for one additional record in 1906, allowed him to place the species in the genus to which it properly belongs. G. Steyskal showed a recent publication by Spencer and Steyskal on the agromyzid leaf miners of the United States. E. Barrows displayed an unusual day- flying trichopteran with dark wings and or- ange body. The speaker of the evening was F. Eugene Wood, University of Maryland who pre- sented a talk and video tape entitled ‘“‘Den- izens of the Dark: Observations on Termite Biology and Management.” PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 371-374 Following the introduction of visitors, the meeting was adjourned at 9:10 p.m. Paul M. Marsh, Recording Secretary 926th Regular Meeting— November 6, 1986 The 926th Regular Meeting of the Ento- mological Society of Washington was called to order by President Edward M. Barrows in the Naturalist Center, National Museum of Natural History, at 8 p.m. on November 6, 1986. Twenty-two members and thirteen guests were present. Corresponding Secre- tary R. G. Robbins read the names of the following applicants for membership: Paul E. Blom, Department of Entomology, Uni- versity of Idaho, Moscow; Paul Courneya, Harlingen, Texas; and Eugene G. Munroe, Ottawa, Ontario, Canada. President Barrows summarized the latest meeting of the Executive Committee, held on the 23rd of October. Among the topics covered were a proposed increase in dues for life memberships, an invitation to the Maryland Entomological Society to cospon- sor the annual banquet, use of name tags at all ESW meetings, revision and republica- tion of the Bylaws, and efforts to encourage young entomologists to become more active in the Society. The Nominating Committee Chairman, D. M. Anderson, presented the following nominations for 1987 officers: President-Elect— F. Eugene Wood Treasurer— Norman E. Woodley Corresponding Secretary—Robert G. Robbins Recording Secretary— Paul M. Marsh Editor—Raymond J. Gagné Associate Editor—no nominee Membership Chairman—Geoffrey B. White Program Chairman— Michael J. Raupp Custodian— Victor L. Blackburn Nominations will remain open until the 372 December meeting when elections will take place. R. Gagné noted that there was no nom- inee for Associate Editor in the report of the Nominating Committee. He reminded the Nominating Committee that 1987 would be his last year as Editor and that, according to a recently enacted bylaw (see Minutes for January 5, 1984, PESW 86: 974), the Nom- inating Committee is responsible for finding an Associate Editor, who would begin pro- cessing manuscripts about June in prepa- ration for the January 1988 issue of the Pro- ceedings. T. J. Spilman displayed a newly published reference, Insect and Spider Collections of the World, compiled by Ross H. Arnett, Jr., and colleagues. The speaker for the evening was Jonathan A. Coddington, Associate Curator, Depart- ment of Entomology, Smithsonian Institu- tion. His talk was entitled “Evolution in Orb-Weaving Spiders.” Refinements in web construction and prey capture strategies were discussed and illustrated. Visitors were introduced and the meeting was adjourned at 9:30 p.m., after which re- freshments were served. Richard G. Robbins, Corresponding Sec- retary 927th Regular Meeting— December 4, 1986 The 927th Regular Meeting of The En- tomological Society of Washington was called to order by President E. M. Barrows in the Naturalist Center, National Museum of Natural History, at 8 p.m. on December 4, 1986. Twenty-two members and six guests were present. Minutes of the previous two meetings were read. Membership Chair- man, G. White, read the names of the fol- lowing applicants for membership: G. T. Baker, Mississippi State, Mississippi; J. Johnson, Moscow, Idaho; J. Rosenheim, Berkeley, California; M. Sharkey, Ottawa, Canada; D. Wahl, Gainesville, Florida; D. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Wood, Ottawa, Canada; S. Larcher, Wash- ington, D.C. Annual reports of officers were given by the Treasurer, Editor, Membership Chair- man, Corresponding Secretary, and Custo- dian. Program Chairman, M. Raupp, an- nounced that the next regular meeting would be held on the second Thursday of next month, January 8, 1987. Nominating Committee Chairman, D. Anderson, presented the slate of nominees for 1987 officers. Wayne N. Mathis was nominated as Associate Editor. President Barrows called for further nominations of which there were none. A motion was made and seconded that the slate be accepted as presented. The motion was unanimously passed. President Barrows discussed the Execu- tive Committee recommendation that life membership dues be raised from $150 to $200. After some discussion of procedure, a motion to amend the bylaws to raise life membership dues was made, seconded, and passed. President Barrows also discussed the Ex- ecutive Committee recommendation to in- vite the Maryland Entomological Society to cosponsor the annual June banquet. It was decided to table any action until the Wash- ington Pest Science Society was contacted. E. Bickley presented two notes in the area of medical entomology. The first concerned Aedes albopictus and its movement north and east; it is now established in 12 states. The second concerned Lyme disease of which there were 74 human cases in Cali- fornia during 1985 and more expected in 1986. Ixodes pacificus is thought to be the most important vector. R. Robbins displayed a brochure an- nouncing a new book on insect dormancy from the Biological Survey of Canada. D. Nickle exhibited several live and un- usual katydids collected during his recent Earthwatch trip to Peru. The speaker of the evening was Douglas W. Tallamy, University of Delaware, who VOLUME 89, NUMBER 2 presented a talk entitled “The Ecology of Maternal Behavior in Insects.” Following the introduction of visitors, President Barrows thanked the Society of- ficers individually for their help to him and their services to the Society. He thanked also the Hospitality Committee composed of Margaret Collins, Mignon Davis, and David Nickle for preparing the fine refresh- 373 ments that are served following each meet- ing. The gavel is usually passed to the new President at this time, but T. Wallenmaier did not attend the meeting because of a con- flict with his honeymoon. The meeting was adjourned at 9:50 p.m. Paul M. Marsh, Recording Secretary REPORTS OF OFFICERS Treasurer’s Report SUMMARY FINANCIAL STATEMENT FOR 1986 Special General Publications Total Fund Fund Assets Assets: November 1, 1985 $21,235.76 $60,137.63 $81,373.39 Total Receipts for 1986 35,317.46 4,771.80 60,089.26 Total Disbursements for 1986 6100151 0.00 61,001.51 Assets: October 31, 1986 ley you leah 64,909.43 80,461.14 Net Changes in Funds $ 5,684.05— 4,771.80 912.25- Norman E. Woodley, Treasurer CORRESPONDING SECRETARY'S SUMMARY OF Major ACTIVITIES FOR CALENDAR YEAR 1986 Letters of welcome were sent to 38 new members. Ten members who had earlier been dropped from our rolls for nonpay- ment of dues were readmitted. The mem- bership list (now computerized) tonight stands at 592. Two of our members were welcomed to Emeritus status: Clyde F. Smith of Raleigh, North Carolina and John V. Thompson of Linwood, New Jersey. Eight (after this evening, 9) letters were written thanking our guest speakers: Some two doz- en thank-you letters were written to mem- bers who contributed to our Special Publi- cation Fund when paying their dues. Fifteen letters (about the same number as in pre- vious years) were written to members who persist in paying the institutional subscrip- tion rate ($35/U.S., $40/foreign) rather than regular member’s dues. The postage costs of this Office amounted to $25.00. Richard G. Robbins, Corresponding Sec- retary EDITOR’S REPORT Beginning with the January, 1987 issue, the Proceedings will begin a two-column format. The Publications and Executive Committees agree that this measure will save some expense without sacrificing utility or beauty. Arly Allen of Allen Press, our print- er, suggested the change to us as a cost- effective measure and to reduce the size of future volumes. The main space-saving fea- ture comes from reducing the width of some tables and plates to one column instead of spreading them across the width of a whole one-column page. 374 Next year the Society will publish a mem- oir, an identification manual to the North American genera of Braconidae. It will be about 100 pages long, half of them filled with drawings and SEM photographs. We think it will be a good seller. The memoir will be spiral bound. This will allow the book to lie perfectly flat while in use, a good feature when both hands are occupied turn- ing a specimen or focussing a dissecting mi- croscope. Respectfully submitted, Raymond J. Gagné, Editor List oF NEw MEMBERS FOR 1986 Warren G. Abrahamson Peter H. Adler Charles W. Agnew Paul E. Blom Tim A. Christiansen Atilano Contreras-Ramos Paul Courneya Paul S. Cwikla Nathan Erwin David L. Haile John M. Heraty Steve Heydon Kevin M. Hoffman Gregory A. Hoover Edward Lippert James W. Mackley PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Jonathan Mawdsley Mary Jo Molineaux Eugene Munroe Paul Newhouse Yuen-Shaung Ng John D. Plakidas Peter W. Price Alasdair J. Ritchie Bob Robbins Michael D. Schwartz Barbara J. Sheffer Joseph D. Shorthouse John R. Spence Margery G. Spofford Terry A. H. Stasny Michael R. Wagner Gwendolyn W. Waring P. Juliette Weinstein James B. Wooley Wu Yar David Z. Zeigler A. Edward Zuccaro, Jr. TOTAL: 38 TOTAL MEMBERSHIP: 592 (Dec. 1986) (Was 632 in Feb. 1986) Current membership reflects names dropped from list for nonpayment of dues as well as 10 members reinstated after payment of dues in arrears. Geoffrey B. White, Membership Chair- man PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 375-383 OBITUARY Glen Milton Kohls 1905-1986 Glen M. Kohls, an internationally rec- ognized authority on ticks and tickborne diseases, died August 3, 1986 at his home in Hamilton, Montana, following a battle with cancer that lasted several months. He was born October 23, 1905, in Vesta, Minnesota. During his early life, he moved to Montana and graduated from high school in Kalispell. Following high school, Glen enrolled at Montana State College (now Montana State University), and was intro- duced to Entomology by Dr. Robert A. Cooley, who was head of the department at that time. During his junior year, Dr. Cool- ey convinced him to leave school and work at the Rocky Mountain Laboratory (RML) in Hamilton on a project concerned with growing hymenopterous tick parasites. Af- ter a year at RML, Glen returned to school and received his Bachelor of Science degree 376 in 1929. Following graduation, he returned to the RML and this association lasted for nearly 40 years. Glen’s service at the RML was inter- rupted on two occasions. First, he enrolled in graduate school at the University of Min- nesota, and received a Master of Science degree in 1937. His return to school was prompted by the threat of a forced vacation without pay that was common at the Rocky Mountain Lab during the depression. The second interruption was for service with the U.S.A. Typhus Commission dur- ing World War II. Before going overseas, Glen served as an instructor at Walter Reed Army Medical School in Washington, D.C. In September of 1943, he left for New Guinea. His research there centered on de- velopment and testing of repellents for the chiggers biting service personnel. Through a program using clothing impregnated with repellents, Kohls and his team convincingly showed a reduction in the transmission rate of scrub typhus and these preventive mea- sures were introduced throughout the areas of scrub typhus endemicity. Later in the war, Glen was transferred to Burma, where he was deputy commander of the typhus unit and worked with four other scientists from the RML. He ended his military career as a lieutenant colonel and returned to Ham- ilton in 1946, where he continued his work on ticks and tickborne diseases. In 1949, Glen was commissioned in the U.S. Public Health Service. He was awarded an honorary doctorate of science degree from Montana State University in 1966 and, shortly before his retirement in 1969, he received the Meritorious Service Medal from the U.S. Public Health Service. Throughout his career, he worked on a variety of projects, and these efforts resulted in over 180 publications from 1927-1973. These manuscripts included work on dis- eases such as tularemia, scrub typhus, Rocky Mountain spotted fever, Colorado tick fe- ver, and ectoparasites such as fleas and ticks. Although Glen worked on many projects, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON his main interest centered on tick taxono- my. Starting from Dr. Cooley’s time, Glen built up the RML tick collection until it was the largest and best curated in the world. He was recognized as an authority on ticks from several areas including the United States, South America and the Near and Far East. Glen was known as a meticulous tax- onomist and nearly all his work is as valid today as when published. He has been hon- ored by several of his associates, who have named 12 species of insects and acari kohlsi. Glen was a quiet family man who went about his everyday tasks without much fan- fare. His legacy to science is demonstrated by the list of publications given below. He had a lot to offer all those who crossed his path and will be missed by friends and col- leagues alike. Glen is survived by his wife, Jeannette, of Hamilton, Montana, whom he married in 1931, and three daughters, Carol Bang- hart, of Arizona; Margy Cotter, of Cali- fornia; and Glenda Kohls-Chase, of Wash- ington. Glen M. Kohls (1906-1986) A Bibliography 1928 Cooley, R. A. and G. M. Kohls. Egg laying of [xodi- phagus caucurtei du Buysson in larval ticks. Sci- ence 67: 656. 1930 Kohls, G. M. A summary of parasite liberations. Mon- tana State Board Entomol., 8th Bien. Rept.: 26- 34. 1934 Cooley, R. A. and G. M. Kohls. A summary of tick parasites. Proc. 5th Pac., Sci. Cong. 5: 3375-3381. 1936 Jellison, W. L. and G. M. Kohls. Distribution and hosts of the human flea, Pulex irritans L., in Montana and other western states. Public Health Repts. 51: 842-844. Kohls, G. M. and R. A. Cooley. Notes on the occur- rence and host relationships of the tick Ornithodo- VOLUME 89, NUMBER 2 ros talaje in Arizona. Public Health Repts. 51: 512-513. 1937 Davis, G. E. and G. M. Kohls. /xodes ricinus califor- nicus (Banks) a possible vector of Bacterium tu- larense. Public Health Repts. 52: 281-282. Kohls, G. M. Tick rearing methods with special ref- erence to the Rocky Mountain wood tick, Der- macentor andersoni Stiles, pp. 246-256. In Galt- Solty 22) S20 Ee BP lutz. PS: Welch, and’ J. G. Needham, eds., Culture Methods for Invertebrate Animals. Comstock Publ. Co., Inc., Ithaca. Kohls, G. M. Hosts of the immature stages of the Pa- cific Coast tick Dermacentor occidentalis Neum. (Ixodidae). Public Health Repts. 52: 490-496. Kohls, G. M. A new species of bat flea (Siphonaptera) from Arizona. J. Parasitol. 23: 300-302. Kohls, G. M. and R. A. Cooley. North American rec- ords of the tick Ixodes ricinus californicus (Banks). Public Health Repts. 52: 282-284. 1938 Cooley, R. A. and G. M. Kohls. Two new species of ticks (Ixodes) from California. Public Health Repts. 53: 1616-1621. Cooley, R. A. and G. M. Kohls. /xodes marmotae—a new species of tick from marmots (Acarina: Ixodi- dae). Public Health Repts. 53: 2174-2181. Jellison, W. L. and G. M. Kohls. Tick-host anemia: A secondary anemia induced by Dermacentor an- dersoni Stiles. J. Parasitol. 24: 143-154. Kohls, G. M. Two new species of Meringis Jordan (Siphonaptera). Public Health Repts. 53: 1216- 1220: 1939 Cooley, R. A. and G. M. Kohls. Amblyomma philipi— a new tick from Texas and Mexico, with a key to known species of Amb/yomma in the United States (Acarina: Ixodidae). Public Health Repts. 54: 44—- 47. Jellison, W. L. and G. M. Kohls. Siphonaptera: A list of Alaskan fleas. Public Health Repts. 54: 2020- 2023. Kohls, G. M. Siphonaptera: Notes on synonymy of North America species of the genus Hoplopsyllus Baker. Public Health Repts. 54: 2019-2020. Parker, R. R., G. M. Kohls, G. W. Cox, and G. E. Davis. Observations on an infectious agent from Amblyomma maculatum. Public Health Repts. 54: 1482-1484. 1940 Cooley, R. A. and G. M. Kohls. Two new species of 377 Argasidae (Acarina: Ixodoidea). Public Health Repts. 55: 925-933. Kohls, G. M. Siphonaptera: A study of the species infesting wild hares and rabbits of North America north of Mexico. Nat. Inst. Health Bull. No. 175. 34 pp. Parker, R. R., W. L. Jellison, G. M. Kohls, and G. E. Davis. Tularemia infection found in streams. Pub- lic Health Repts. 55: 227. 194] Cooley, R. A. and G. M. Kohls. Ornithodoros viguerasi, a new species of tick from bats in Cuba (Acarina: Ixodoidea). Public Health Repts. 56: 396-399. Cooley, R. A. and G. M. Kohls. Three new species of Ornithodoros (Acarina: Ixodoidea). Public Health Repts. 56: 587-594. Cooley, R. A. and G. M. Kohls. Further new species of Ornithodoros from bats (Acarina: Argasidae). Public Health Repts. 56: 910-914. 1942 Cooley, R. A. and G. M. Kohls. Antricola new genus, Amblyomma gertschi new species, and notes on Ixodes spinipalpis (Acarina: Ixodoidea). Public Health Repts. 57: 1733-1736. Cooley, R. A. and G. M. Kohls. /xodes baergi, a new species of tick from Arkansas (Acarina: Ixodidae). Public Health Repts. 57: 1869-1872. Cooley, R. A. and G. M. Kohls. [xodes mexicanus n. sp. e Ixodes tancitarius n. sp. dos nuevas garra- patas Mexicanas (Acarina: Ixodidae). Rev. Soc. Mex. Hist. Nat. 3: 149-154. Jellison, W. L., G. M. Kohls, W. J. Butler, and J. A. Weaver. Epizootic tularemia in the beaver, Castor canadensis, and the contamination of stream water with Pasteurella tularensis. Am. J. Hyg. 36: 168- 182. Kohls, G. M. Siphonaptera: Ptilopsylla dunni, a new species of bat flea from Panama. J. Parasitol. 28: 361-362. Steinhaus, E. A. and G. M. Kohls. Isolation of an acid- fast bacillus from a hawk. J. Am. Vet. Med. Assoc. 101: 502. 1943 Cooley, R. A. and G. M. Kohls. [xodes californicus Banks, 1904, Ixodes pacificus n. sp., and Ixodes conepati n. sp. (Acarina: Ixodidae). Pan-Pacific Entomol. 19: 139-147. Foote, H. B., W. L. Jellison, E. A. Steinhaus, and G. M. Kohls. Effect of chlorination on Pasteurella tularensis 1n aqueous suspension. J. Am. Water Works Assoc. 35: 902-910. Jellison, W. L., G. M. Kohls, and H. B. Mills. Siphon- 378 aptera: Species and host list of Montana fleas. Montana State Board Entomol., Misc. Pub. No. 2 22spp: Kohls, G. M. and E. A. Steinhaus. Tularemia: Spon- taneous occurrence in shrews. Public Health Repts. 58: 842. Parker, R. R. and G. M. Kohls. American Q fever: The occurrence of Rickettsia diaporica in Amblyomma americanum in eastern Texas. Public Health Repts. 58: 1510-1511. Parker, R. R., G. M. Kohls, and E. A. Steinhaus. Rocky Mountain spotted fever: Spontaneous infection in the tick Amblyomma americanum. Public Health Repts. 58: 721-729. Parker, R. R., E. A. Steinhaus, and G. M. Kohls. Tu- laremia in beavers and muskrats and the contam- ination of natural waters and mud by Pasteurella tularensis in the northwestern United States. J. Bacteriol. 45: 56-57. 1944 Cooley, R. A. and G. M. Kohls. The Argasidae of North America, Central America and Cuba. Am. Mid- land Nat. Monogr. No. 1. The University Press, Notre Dame, Indiana. 152 pp. Cooley, R. A. and G. M. Kohls. The genus Amblyom- ma (Ixodidae) in the United States. J. Parasitol. 30: 77-111. 1945 Blake, F. G., K. F. Maxcy, J. F. Sadusk, Jr., G. M. Kohls, and E. J. Bell. Studies on tsutsugamushi disease (scrub typhus, mite-borne typhus) in New Guinea and adjacent islands: Epidemiology, clin- ical observations and etiology in the Dobadura area. Am. J. Hyg. 41: 243-373. Blake E.G Ke Es Maxcy, J; Es sadusk, Jr. .G. M. Kohls, and E. J. Bell. Tsutsugamushi disease (scrub typhus, mite-borne typhus) in New Guinea. Am. J. Public Health 35: 1121-1130. Blake, F. G., K. F. Maxcy, J. F. Sadusk, Jr., G. M. Kohls, and E. J. Bell. Trombicula fletcheri Wom- ersley and Heaslip 1943, a vector of tsutsugamushi disease (scrub typhus) in New Guinea. Science 102: 61-64. Cooley, R. A. and G. M. Kohls. The genus /xodes in North America. Nat. Inst. Health Bull. No. 184. 246 pp. Kohls, G. M., C. A. Armbrust, E. N. Irons, and C. B. Philip. Studies on tsutsugamushi disease (scrub typhus, mite-borne typhus) in New Guinea and adjacent islands: Further observations on epide- miology and etiology. Am. J. Hyg. 41: 374-396. Philip, C. B. and G. M. Kohls. Studies on tsutsuga- mushi disease (scrub typhus, mite-borne typhus) in New Guinea and adjacent islands: Tsutsuga- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON mushi disease with high endemicity on a small south sea island. Am. J. Hyg. 42: 195-203. 1946 Mackie, T. T., G. E. Davis, H. S. Fuller, J. A. Knapp, M. L. Steinacker, K. E. Stager, R. Traub, W. L. Jellison, D. D. Millspaugh, R. C. Austrian, E. J. Bell, G. M. Kohls, W. Hsi, and J. A. Girsham. Observations on tsutsugamushi disease (scrub ty- phus) in Assam and Burma. Preliminary report. Am. J. Hyg. 43: 195-218. Mackie, T. T., G. E. Davis, H. S. Fuller, J. A. Knapp, M. L. Steinacker, K. E. Stager, R. Traub, W. L. Jellison, D. D. Millspaugh, R. C. Austrian, E. J. Bell, G. M. Kohls, W. Hsi, and J. A. Girsham. Observations on tsutsugamushi disease (scrub ty- phus) in Assam and Burma. Preliminary report. Trans. Roy. Soc. Trop. Med. Hyg. 40: 15-46. 1947 Kohls, G. M. Vectors of rickettsial diseases. Ann. Int. Med. 26: 713-719. Kohls, G. M. Notes on the tick, Ixodes howelli Cooley and Kohls, with descriptions. J. Parasitol. 33: 57- 61. Kohls, G. M. Ixodes radfordi, a new species of tick from rats in eastern India (Acarina: Ixodidae). J. Parasitol. 33: 497-498. Kohls, G. M. and J. M. Brennan. The Lone Star Tick Amblyomma americanum (Linnaeus). Rocky Mountain Lab. Circ. No. 14. 5 pp. Womersley, H. and G. M. Kohls. New genera and species of Trombiculidae from the Pacific islands. Trans. Roy. Soc. S. Australia 71: 3-12. 1948 Kohls, G. M. Vectors of rickettsial diseases, pp. 83- 96. In Proc. Rickettsial Diseases of Man Sym- posium A. A. A. S., Boston, Mass., Dec. 26-28, 1946. Kohls, G. M. Haemaphysalis ratti, a new species of tick from rats in New Guinea and Haemaphysalis krijgsmani, new name for Haemaphysalis novae- guineae Krijgsman and Ponto, 1932, pre-occu- pied. J. Parasitol. 34: 154-157. Kohls, G. M. and W. L. Jellison. Ectoparasites and other arthropods occurring in Texas bat caves. Nat. Speleological Soc. Bull. No. 10, 116-117. Kohls, G. M. and R. R. Parker. Occurrence of the brown dog tick in the western states. J. Econ. Ento- mol. 41: 102. Philip, C. B. and G. M. Kohls. Mites and scrub typhus. Proc. IV Int. Cong. Trop. Med. Malaria 2: 1656- 1663. VOLUME 89, NUMBER 2 1949 Kohls, G. M. Haemaphysalis centropi, a new species of tick from birds in the Far East. J. Parasitol. 35: 388-390. 1950 Brown, J. H. and G. M. Kohls. The ticks of Alberta with special reference to distribution. Canad. J. Res. 28: 197-205. Jellison, W. L., D. C. Epler, E. Kuhns, and G. M. Kohls. Tularemia in man from a domestic rural water supply. Abstracts of papers, p. 82. Fifth Int. Cong. Microbiol., Rio de Janeiro. Jellison, W. L., D. C. Epler, E. Kuhns, and G. M. Kohls. Tularemia in man from a domestic rural water supply. Public Health Repts. 65: 1219-1226. Jellison, W. L. and G. M. Kohls. Persistence of agglu- tinins against Pasteurella tularensis in serums of naturally infected sheep. J. Am. Vet. Med. Assoc. 117: 405-408. Kohls, G. M. Description of the male of Ixodes woodi Bishopp (Acarina: Ixodidae). J. Parasitol. 36: 65- 66. Kohls, G. M. Note on the occurrence of the flea Nearc- topsylla hyrtaci (Roth.) in the United States. J. Parasitol. 36: 311. Kohls, G. M. Two new species of ticks from Ceylon (Acarina: Ixodidae). J. Parasitol. 36: 319-321. Kohls, G. M. Ticks (Ixodoidea) of the Philippines. Nat. Inst. Health Bull. No. 192. 28 pp. 195i Jellison, W. L., G. M. Kohls, and C. B. Philip. Tula- remia—muskrats as a source of human infection in Utah. Rocky Mountain Med. J. 48: 594-597. Parker, R. R., E. A. Steinhaus, G. M. Kohls, and W. L. Jellison. Contamination of Natural Waters and Mud with Pasteurella tularensis and Tularemia in Beavers and Muskrats in the Northwestern United States. Nat. Inst. Health Bull. No. 193. 61 pp. Philip, C. B. and G. M. Kohls. Elk, winter ticks, and Rocky Mountain spotted fever: A query. Public Health Repts. 66: 1672-1675. Wharton, G. W., D. W. Jenkins, J. M. Brennan, H. S. Fuller, G. M. Kohls, and C. B. Philip. The ter- minology and classification of trombiculid mites (Acarina: Trombiculidae). J. Parasitol. 37: 13-31. 1952 Gregson, J. D. and G. M. Kohls. The male of [xodes soricis Gregson (Acarina: Ixodidae). Canad. Ento- mol. 84: 185-188. Kohls, G. M. A record of the occurrence of the tick Ixodes muris Bishopp and Smith on muskrats in Utah. Great Basin Nat. 12: 65-66. Kohls, G. M. and H. T. Dalmat. The male of Der- 379 macentor dissimilis Cooley (Acarina: Ixodidae). J. Parasitol. 38: 140-142. Kohls, G. M. and N. J. Kramis. Tick paralysis in the American buffalo, Bison bison (Linn.). Northwest Sci. 26: 61-64. 1953 Jellison, W. L. and G. M. Kohls. Tularemia in sheep and sheep industry workers in the western United States. Riass. Comun. Sixth Congr. Int. Microbiol. 2: 593, Rome, Sept. 6-12. Kohls, G. M. Notes on the ticks of Guam with the description of Amblyomma squamosum n. sp. (Acarina: Ixodidae). J. Parasitol. 39: 264-267. Kohls, G. M. Ixodes venezuelensis, a new species of tick from Venezuela, with notes on /xodes minor Neumann, 1902 (Acarina: Ixodidae). J. Parasitol. 39: 300-303. Kohls, G. M. and A. J. Rogers. Notes on the occurrence of the tick Ixodes affinis Neumann in the United States. J. Parasitol. 39: 669. 1954 Kohls, G. M. The order Acarina (ticks and mites), pp. 661-679. In Mackie, T. T., G. W. Hunter, III, and C. B. Worth, eds., A Manual of Tropical Medicine, 2 ed. W. B. Saunders Co., Philadelphia. Kohls, G. M. New distributional records for Trichobius corynorhini (Diptera: Streblidae). Wasmann J. Biol. 12: 27-28. Kohls, G. M. and B. Locker. Isolation of Pasteurella tularensis from the tick [xodes ricinus in Norway. Nord. Vet.-Med. 6: 883-884. 1955 Eklund, C. M., G. M. Kohls, and J. M. Brennan. Dis- tribution of Colorado tick fever virus-carrying ticks. J. Am. Med. Assoc. 157: 335-337. Jellison, W. L. and G. M. Kohls. Tularemia in Sheep and Sheep Industry Workers. Public Health Monogr. No. 28. 17 pp. Kohls, G. M. Colorado tick fever discovered in Cali- fornia. Calif. Vector Views 2: 1. Kohls, G. M. Two new species of ticks from North Borneo (Acarina: Ixodidae). J. Parasitol. 41: 312- 315). 1956 Kohls, G. M. Eight new species of xodes from Central and South America (Acarina: Ixodidae). J. Par- asitol. 42: 636-649. Kohls, G. M. Concerning the identity of Amblyomma maculatum, A. tigrinum, A. triste, and A. ovatum of Koch, 1844. Proc. Entomol. Soc. Wash. 58: 143-147. Kohls, G. M. The identity of Ixodes boliviensis Neu- 380 mann 1904 and J. bicornis Neumann 1906 (Ixodi- dae). Proc. Entomol. Soc. Wash. 58: 232-233. 1957 Kohls, G. M. Malaysian parasites— X VIII. Ticks (Ixo- doidea) of Borneo and Malaya, pp. 65-94. Jn Audy, J. R., ed., Malaysian Parasites XVI-XXXIV. Studies from the Institute for Medical Research, Federation of Malaya, No. 28. The Caxton Press, Ltd., Kuala Lumpur. Kohls, G. M. Insects of Micronesia. Acarina: Ixodoi- dea. Insects of Micronesia 3: 85-104. Kohls, G. M. Ixodes downsi, a new species of tick from a cave in Trinidad, British West Indies (Acarina— Ixodidae). Proc. Entomol. Soc. Wash. 59: 257- 264. 1958 Eklund, C. M., G. M. Kohls, and W. L. Jellison. Iso- lation of Colorado tick fever virus from rodents in Colorado. Science 128: 413. Jellison, W. L., D. C. Epler, E. Kuhns, and G. M. Kohls. Tularemia in man from a domestic rural water supply. Arq. V Cong. Int. Microbiol. (Rio de Ja- neiro, August 17-24, 1950), 1: 301-304. Kohls, G. M. Amblyomma imitator, a new species of tick from Texas and Mexico, and remarks on the synonymy of A. cajennense (Fabricius) (Acarina— Ixodidae). J. Parasitol. 44: 430-433. 1959 Hoogstraal, H. and G. M. Kohls. The Haemaphysalis ticks (Acarina: Ixodidae) of birds. I. H. ornitho- philan. sp. from Burma and Thailand. J. Parasitol. 45: 417-420. 1960 Hoogstraal, H. and G. M. Kohls. Observations on the subgenus Argas (Ixodoidea, Argasidae, Argas). 1. Study of A. reflexus reflexus (Fabricius, 1794), the European bird argasid. Ann. Entomol. Soc. Am. 53: 611-618. Hoogstraal, H. and G. M. Kohls. Observations on the subgenus Argas (Ixodoidea, Argasidae, Argas). 3. A biological and systematic study of A. reflexus hermanni Audouin, 1827 (revalidated), the Afr- can bird argasid. Ann. Entomol. Soc. Am. 53: 743- 155: Kohls, G. M. Ixodides, pp. 298-299. In McGraw-Hill Encylopedia of Science and Technology. Kohls, G. M. Ixodes (Endopalpiger) zaglossi, n. sp. from the long-beaked echidna of New Guinea (Ac- arina, Ixodidae). Acarologia 2: 447-452. Kohls, G. M. Ticks. A Monograph of the Ixodoidea. Part V. On the genera Dermacentor, Anocentor, Cosmiomma, Boophilus, and Margaropus, by Don PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON R. Arthur. (Book review.) Ann. Entomol. Soc. Am. 53: 855-856. Kohls, G. M. Records and new synonymy of New World Haemaphysalis ticks, with descriptions of the nymph and larva of H. juxtakochi. J. Parasitol. 46: 355-361. Kohls, G. M. and H. Hoogstraal. Observations on the subgenus Argas (Ixodoidea, Argasidae, Argas). 2. A. cooleyi, new species, from western North Amer- ican birds. Ann. Entomol. Soc. Am. 53: 625-631. 1961 Clifford, C. M. and G. M. Kohls. A new distribution and host record for Ixodes muris Bishopp and Smith, 1937 (Acarina: Ixodidae). Proc. Entomol. Soc. Wash. 63: 210. Jellison, W. L., C. O. Owen, J. F. Bell, and G. M. Kohls. Tularemia and animal populations: Ecology and epizootology. Wildl. Dis. 17: 1-22. Kohls, G. M. Rediscovery of Haemaphysalis mjobergi Warburton, 1926 (Acarina: Ixodidae). Pacific In- sects 3: 305-306. Kohls, G. M. and C. M. Clifford. A new species of Ixodes (Lepidixodes) from bats in Malaya, North Borneo, and the Congo (Acarina: Ixodidae). Acar- ologia 3: 285-290. Kohls, G. M. and H. Hoogstraal. Observations on the subgenus Argas (Ixodoidea, Argasidae, Argas). 4. A. neghmei, new species, from poultry houses and human habitations in northern Chile. Ann. Ento- mol. Soc. Am. 54: 844-851. Kohls, G. M., H. Hoogstraal, and C. M. Clifford. Ob- servations on the subgenus Argas (Ixodoidea, Ar- gasidae, Argas). 5. Study of A. brevipes Banks, 1908, from birds in Arizona and California, U.S.A., and Baja California, Mexico. Ann. Entomol. Soc. Am. 54: 869-877. Osgood, S. B., W. L. Jellison, and G. M. Kohls. An episode of pseudo-pediculosis. J. Parasitol. 47: 985— 986. Philip, C. B. and G. M. Kohls. Dermacentor andersoni Stiles, 1908 (Acarina): Proposed validation under the plenary powers. Revision of Opinion 78. Z. N. (S.) 260. Bull. Zool. Nomencl. 18: 316-318. 1962 Clifford, C. M. and G. M. Kohls. Description of the female of Dermacentor latus Cooley and Am- blyomma albopictus Neumann (Acarina: Ixodi- dae). J. Parasitol. 48: 486-489. Eklund, C. M., G. M. Kohls, W. L. Jellison, W. Burg- dorfer, R. C. Kennedy, and L. Thomas. The clin- ical and ecological aspects of Colorado tick fever. Proc. Sixth Int. Cong. Trop. Med. Malar. (Lisbon, September 5-13, 1958), 5: 197-203. (dated 1959). Eklund, C. M., G. M. Kohls, and R. C. Kennedy. Lack of evidence of transovarial transmission of Col- VOLUME 89, NUMBER 2 orado tick fever virus in Dermacentor andersonsi, pp. 401-406. Jn Libikova, H., ed., Biology of Vi- ruses of the Tick-borne Encephalitis Complex. Czechoslovak Acad. Sci., Prague. Eklund, C. M., H. G. Stoenner, and G. M. Kohls. Rocky Mountain spotted fever, pp. 433-445. In Harvey, J. C., ed., Vol. 4 (Revised). Tice-Harvey Practice of Medicine. W. F. Prior Co., Inc., Hagerstown, Maryland. Hoogstraal, H. and G. M. Kohls. Bat ticks of the genus Argas (Ixodoidea, Argasidae). 5. Description of larvae from Australia and New Guinea Carios- group populations. Proc. Linn. Soc. New South Wales 87: 275-280. Kohls, G. M. A new species of /xodes from Malaya (Acarina: Ixodidae). Proc. Entomol. Soc. Wash. 64: 103-105. Kohls, G. M. and C. M. Clifford. Ixodes tiptoni, a new species of tick from Panama. J. Parasitol. 48: 182- 184. Kohls, G. M. and H. Hoogstraal. Bat ticks of the genus Argas (Ixodoidea, Argasidae). 4. A. (Carios) aus- traliensis n. sp. from Australia. Ann. Entomol. Soc. Am. 55: 555-559. Kohls, G. M. and R. E. Ryckman. New distributional records of ticks associated with cliff swallows, Pet- rochelidon spp., in the United States. J. Parasitol. 48: 507-508. Ryckman, R. E. and G. M. Kohls. The desert tortoise, Gopherus agassizi, a host for the tick Ornithodoros turicata in California. J. Parasitol. 48: 502-503. Sonenshine, D. E., C. M. Clifford, and G. M. Kohls. The identification of larvae of the genus Argas (Acarina, Argasidae). Acarologia 4: 193-214. 1963 Bell, E. J., G. M. Kohls, H. G. Stoenner, and D. B. Lackman. Nonpathogenic rickettsias related to the spotted fever group isolated from ticks, Derma- centor variabilis and Dermacentor andersoni from eastern Montana. J. Immunol. 90: 770-781. Brinton, E. P. and G. M. Kohls. New distributional and host data for the tick Dermacentor hunteri Bishopp. Great Basin Nat. 23: 166. Clifford, C. M. and G. M. Kohls. Observations on the life cycle of the tick Argas brevipes Banks, 1908. J. Parasitol. 49: 527. Hoogstraal, H. and G. M. Kohls. Observations on the subgenus Argas (Ixodoidea: Argasidae, Argas). 6. Redescription and biological notes on A. /ageno- plastis Froggatt, 1906, of Australian fairy martins, Hylochelidon ariel (Gould). Ann. Entomol. Soc. Am. 56: 577-582. Kohls, G. M. and C. M. Clifford. Ornithodoros sparnus sp. n., a parasite of wood rats, Neotoma spp. and deer mice, Peromyscus spp. in Utah and Arizona (Acarina: Agarsidae). J. Parasitol. 49: 857-861. Nadchatram, M. and G. M. Kohls. Discovery of the 381 tick Argas boueti Rouboud and Colas-Belcour (Ac- arina: Argasidae) in Malaya. J. Parasitol. 49: 350. 1964 Clifford, C. M., G. M. Kohls, and D. E. Sonenshine. The systematics of the subfamily Ornithodorinae (Acarina: Argasidae). I. The genera and subgenera. Ann. Entomol. Soc. Am. 57: 429-437. Kaiser, M. N., H. Hoogstraal, and G. M. Kohls. The subgenus Persicargas, new subgenus (Ixodoidea, Argasidae, Argas). 1. A. (P.) arboreus, new species, an Egyptian persicus-like parasite of wild birds, with a redefinition of the subgenus Argas. Ann. Entomol. Soc. Am. 57: 60-69. Kohls, G. M. Rocky Mountain spotted fever, pp. 349- 356: Jn’ Coates, J: B; Jr., EC. Hoff, and P.M: Hoff, eds., Preventive Medicine in World War II., Vol. VII. Washington, D.C. Kohls, G. M. and C. M. Clifford. Ixodes eadsi, a new species of tick from rodents in southern Texas (Acarina: Ixodidae). J. Parasitol. 50: 466-470. Kohls, G. M. and C. M. Clifford. Ornithodoros (Alec- torobius) boliviensis sp. n. (Acarina: Argasidae) from bats and houses in Bolivia. J. Parasitol. 50: 792- 796. Tonn, R. J. and G. M. Kohls. Ectoparasites of birds and mammals of Costa Rica. 2. Ticks. Rev. Biol. Trop. 11: 217-220 (1963). 1965 Hoogstraal, H. and G. M. Kohls. Bat ticks of the genus Argas (Ixodoidea, Argasidae). 6. The female and larva of A. (Carios) australiensis. Ann. Entomol. Soc. Am. 58: 816-820. Hoogstraal, H. and G. M. Kohls. Descriptions, hosts, and ecology of Ixodes kuntzi n. sp., Kuntz’s Tai- wan flying squirrel tick (Ixodoidea, Ixodidae). J. Med. Entomol. 2: 209-214. Hoogstraal, H. and G. M. Kohls. Southeast Asian Hae- maphysalis ticks (Ixodoidea, Ixodidae). H. ban- dicota sp. n. from bandicoot rats in Taiwan, Thai- land, and Burma. J. Parasitol. 51: 460-466. Hoogstraal, H., G. M. Kohls, and H. Trapido. Hae- maphysalis (Rhipistoma) eupleres sp. n. from a Madagascar carnivore and definition of the sub- genus Rhipistoma Koch (resurrected) (Ixodoidea, Ixodidae). J. Parasitol. 51: 997-1000. Hoogstraal, H., H. Trapido, and G. M. Kohls. Studies on Southeast Asian Haemaphysalis ticks (Ixodoi- dea, Ixodidae). H. atherurus sp. n., and redescrip- tion of type material of H. birmaniae Supino, 1897. J. Parasitol. 51: 114-125. Hoogstraal, H., H. Trapido, and G. M. Kohls. Studies on Southeast Asian Haemaphysalis ticks (Ixodoi- dea, Ixodidae). H. (Kaiseriana) papuana nadcha- trami ssp. n. and a redescription of H. (K.) se- mermis Neumann. J. Parasitol. 51: 433-451. Hoogstraal, H., H. Trapido, and G. M. Kohls. Studies 382 on Southeast Asian Haemaphysalis ticks (Ixodoi- dea, Ixodidae). The identity, distribution, and hosts of H. (Kaiseriana) hystricis Supino. J. Parasitol. 51: 467-480. Hoogstraal, H., H. Trapido, and G. M. Kohls. Studies on Southeast Asian Haemaphysalis ticks (Ixodoi- dea, Ixodidae). H. (Kaiseriana) celebensis sp. n. from wild boar in Celebes. J. Parasitol. 51: 1001- 1003. Kohls, G. M. Die Zeckenfauna Mitteleuropas, by San- dor Babos. (Book review.) J. Parasitol. 51: 381. Kohls, G. M., D. E. Sonenshine, and C. M. Clifford. The systematics of the subfamily Ornithodorinae (Acarina: Argasidae). II. Identification of the lar- vae of the Western Hemisphere and descriptions of three new species. Ann. Entomol. Soc. Am. 58: 331-364. 1966 Fairchild, G. B., G. M. Kohls, and V. J. Tipton. The ticks of Panama (Acarina: Ixodoidea), pp. 167— 219. In Wenzel, R. L., and V. J. Tipton, eds., The Ectoparasites of Panama. Field Museum of Nat- ural History, Chicago. Hoogstraal, H. and G. M. Kohls. Ornithodoros (Pav- lovskyella) macmillani, new species (Ixodoidea: Argasidae), a nest parasite of Australian cockatoos (Kakatoe roseicapilla). Ann. Entomol. Soc. Am. 59: 86-92. Hoogstraal, H. and G. M. Kohls. Argas (Microargas) transversus Banks (new subgenus) (Ixodoidea, Ar- gasidae), a diminutive parasite of the Galapagos giant tortoise: Redescription of the holotype male and description of the larva. Ann. Entomol. Soc. Am. 59: 247-252. Hoogstraal, H., H. Trapido, and G. M. Kohls. Studies on Southeast Asian Haemaphysalis ticks (Ixodoi- dea, Ixodidae). Speciation in the H. (Kaiseriana) obesa group: H. semermis Neumann, H. obesa Larrouse, H. roubaudi Toumanoff, H. montgo- meryi Nuttall, and H. hirsuta sp. n. J. Parasitol. 52: 169-191. Kohls, G. M. A new sea bird tick, Jxodes amersoni, from Phoenix Island. J. Med. Entomol. 3: 38-40. Kohls, G. M. and C. M. Clifford. Three new species of Ixodes from Mexico and description of the male of I. auritulus auritulus Neumann, J. conepati Cooley and Kohls, and J. /asallei Mendez and Or- tiz (Acarina: Ixodidae). J. Parasitol. 52: 810-820. Sonenshine, D. E., C. M. Clifford, and G. M. Kohls. The systematics of the subfamily Ornithodorinae (Acarina: Argasidae). III. Identification of the lar- vae of the Eastern Hemisphere. Ann. Entomol. Soc. Am. 59: 92-122. 1967 Hoogstraal, H. and G. M. Kohls. Bat ticks of the genus Argas (Ixodoidea, Argasidae). 7. A. (Chiropterar- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON gas) cordiformis, new species, a heart-shaped par- asite from South-West Africa. Ann. Entomol. Soc. Am. 60: 63-67. Hoogstraal, H., G. M. Kohls, and D. W. Parrish. Stud- ies on Southeast Asian Haemaphysalis ticks (Ixo- doidea, Ixodidae). H. (Kaiseriana) psalistos sp. n., a parasite of deer and boars in Luzon, Philippines. J. Parasitol. 53: 1096-1102. Hoogstraal, H., G. M. Kohls, and H. Trapido. Studies on Southeast Asian Haemaphysalis ticks (Ixodoi- dea, Ixodidae). H. (Kaiseriana) anomala Warbur- ton: Redescription, hosts, and distribution. J. Par- asitol. 53: 196-201. Kohls, G. M. and C. M. Clifford. Ixodes (Haemixodes) uruguayensis, new subgenus, new species (Acari- na: Ixodidae) from small rodents in Uruguay. Ann. Entomol. Soc. Am. 60: 391-394. Kohls, G. M. and C. M. Clifford. The male and larva of Ixodes laysanensis Wilson with notes on rearing (Acarina: Ixodidae). J. Med. Entomol. 4: 83-86. 1968 Clifford, C. M., G. M. Kohls, and H. Hoogstraal. Ixodes walkerae, n. sp., from a bird in Kenya (Acarina: Ixodidae). J. Med. Entomol. 5: 513-514. Clifford, C. M., L. A. Thomas, L. E. Hughes, G. M. Kohls, and C. B. Philip. Identification and com- parison of two viruses isolated from ticks of the genus Ornithodoros. Am. J. Trop. Med. Hyg. 17: 881-885. Hoogstraal, H., M. N. Kaiser, and G. M. Kohls. The subgenus Persicargas (Ixodoidea, Argasidae, Ar- gas). 4. Argas (P.) robertsi, new species, a parasite of Australian fowl, and keys to Australian argasid species. Ann. Entomol. Soc. Am. 61: 535-539. Hoogstraal, H., M. N. Kaiser, and G. M. Kohls. The subgenus Persicargas (Ixodoidea, Argasidae, Ar- gas). 5. A. (P.) zumpti, new species, a parasite of the South African cape vulture. Ann. Entomol. Soc. Am. 61: 744-751. Hoogstraal, H. and G. M. Kohls. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). Redescription of type material of H. (Rhipistoma) heinrichi Schulze, a parasite of the ferret-badger in Burma and Vietnam. J. Parasitol. 54: 1057- 1062. Hoogstraal, H., G. M. Kohls, and D. W. Parrish. Stud- ies on Southeast Asian Haemaphysalis ticks (Ixo- doidea, Ixodidae). H. (Kaiseriana) susphilippensis sp. n., a parasite of Luzon and Mindanao boars. J. Parasitol. 54: 616-621. Hoogstraal, H., F. H. S. Roberts, G. M. Kohls, and V. J. Tipton. Review of Haemaphysalis (Kaiseriana) longicornis Neumann (resurrected) of Australia, New Zealand, New Caledonia, Fiji, Japan, Korea, Northeastern China and USSR, and its parthe- nogenetic and bisexual populations (Ixodoidea, Ixodidae). J. Parasitol. 54: 1197-1213. VOLUME 89, NUMBER 2 Kohls, G. M. Scientific names of important ticks in the United States. Vector Control Briefs No. 21: 26. Kohls, G. M. and C. M. Clifford. The subgenus Per- sicargas (Ixodoidea, Argasidae, Argas). 6. A. (P.) giganteus, n. sp., from wild birds in western United States and Sonora, Mexico. Ann. Entomol. Soc. Am. 61: 1113-1116. Kohls, G. M. and C. M. Clifford. Ixodes (Lepidoxides) kopsteini Oudemans, 1925, valid name for J. (L.) paradoxus Kohls and Clifford, 1961, with notes on distribution and hosts (Acarina: Ixodidae). J. Med. Entomol. 5: 503-506. Wilson, N., H. Hoogstraal, and G. M. Kohls. Studies on Southeast Asian Haemaphysalis ticks (Ixodoi- dea, Ixodidae). Redescription of H. (Rhipistoma) bartelsi Schulze (resurrected), the Indonesian flying squirrel haemaphysalid. J. Parasitol. 54: 1223- 12272 1969 Kohls, G. M. New records of ticks from the Lesser Antilles, pp. 126-134. Jn Hummelinck, P. W., ed., Studies of the Fauna of Curacao and Other Carib- bean Islands. Vol. 28. The Hague. Kohls, G. M. Obituary, Robert Allen Cooley 1873- 1968. J. Econ. Entomol. 62: 972. Kohls, G. M. Ixodes taglei n. sp. (Acarina: Ixodidae) a parasite of the deer, Pudu pudu (Wol.), in Chile. J. Med. Entomol. 6: 280-283. Kohls, G. M. A new species of amblyomma from igua- nas in the Caribbean (Acarina: Ixodidae). J. Med. Entomol. 6: 439-442. Kohls, G. M., C. M. Clifford, and H. Hoogstraal. Two new species of Ornithodoros from the Galapagos Islands (Acarina: Argasidae). J. Med. Entomol. 6: 75-78. Kohls, G. M., C. M. Clifford, and E. K. Jones. The systematics of the subfamily Ornithodorinae (Ac- arina: Argasidae). IV. Eight new species of Orni- thodoros from the Western Hemisphere. Ann. Entomol. Soc. Am. 62: 1035-1043. Kohls, G. M., D. E. Sonenshine, and C. M. Clifford. Ixodes (Exopalpiger) jonesae sp. n. (Acarina: Ix- odidae), a parasite of rodents in Venezuela. J. Par- asitol. 55: 447-452. Sonenshine, D. E., G. M. Kohls, and C. M. Clifford. Ixodes crenulatus Koch, 1844. Synonymy with J. kaiseri Arthur, 1957 and redescriptions of the male, female, nymph, and larva (Acarina: Ixodidae). Acarologia 11: 193-206. 1970 Keirans, J. E. and G. M. Kohls. [xodes philipi n. sp. (Acarina: Ixodidae), a sea bird parasite in Japan. J. Med. Entomol. 7: 725-726. 383 Kohls, G. M., C. M. Clifford, and H. Hoogstraal. Ixodes (Scaphixodes) mitchelli, n. sp. (Acarina: Ixodidae), a tick parasitizing pheasants and partridges in high mountains of Nepal. J. Med. Entomol. 7: 348- 352: Kohls, G. M., H. Hoogstraal, C. M. Clifford, and M. N. Kaiser. The subgenus Persicargas (Ixodoidea, Argasidae, Argas). 9. Redescription and New World records of Argas (P.) persicus (Oken), and resur- rection, redescription, and records of A. (P.) ra- diatus Railliet, A. (P.) sanchezi Dugés, and A. (P.) miniatus Koch, New World ticks misidentified as A. (P.) persicus. Ann. Entomol. Soc. Am. 63: 590- 606. 1971 Clifford, C. M., H. Hoogstraal, and G. M. Kohls. Lxodes hyatti, n. sp., and I. shahi, n. sp. (Acarina: Ixodi- dae), parasites of pikas (Lagomorpha: Ochotoni- dae) in the Himalayas of Nepal and West Pakistan. J. Med. Entomol. 8: 430-438. 1972 Jones, E. K., C. M. Clifford, J. E. Keirans, and G. M. Kohls. The ticks of Venezuela (Acarina: Ixodoi- dea) with a key to the species of Amblyomma in the Western Hemisphere. Brigham Young Univ. Sci. Bull. Biol. Ser. 17: 1-40. Oliver, J. H., Jr., P. R. Wilkinson, and G. M. Kohls. Observations on hybridization of three species of North American Dermacentor ticks. J. Parasi- tol. 58: 380-384. 1973 Clifford, C. M., D. E. Sonenshine, J. E. Keirans, and G. M. Kohls. Systematics of the subfamily Ixodi- nae (Acarina: Ixodidae). |. The subgenera of Ixodes. Ann. Entomol. Soc. Am. 66: 489-500. Jellison, W. L. and G. M. Kohls. Cornelius Becker Philip: An appreciation. Exper. Parasitol. 33: 407— 423. Carleton M. Clifford, P.O. Box 1230, Ham- ilton, Montana 59840 and James E. Keirans, De- partment of Health and Human Services, Public Health Service, National Institutes of Health, National Institute of Allergy and Infectious Dis- eases, Department of Entomology, Museum Support Center, Smithsonian Institution, Wash- ington, D.C. 20560. PROC. ENTOMOL. SOC. WASH. 89(2), 1987, pp. 384-388 OBITUARY Jon Lamar Herring 1922-1985 Jon Lamar Herring, born December 30, 1922, died on February 25, 1985 after sev- eral years of illness. Jon graduated from Crescent City High School, Crescent City, Florida in 1942. He enlisted in the Army in May 1943 and gained specialized surgical training. This prepared him for overseas duty as a surgical techni- cian caring for war casualities. After the ser- vice, Jon enrolled at the University of Flor- ida at Gainesville (1946-1955) where he obtained his B.S. as an honor student and inductee into Phi Kappa Phi and an M.S. under R. F. Shirley. While at Gainesville he met Roland F. Hussey who became a life- long friend and kindled his interest in the Heteroptera, especially in its aquatic mem- bers. One of the highlights of this associa- tion was the rediscovery of Thomas Say’s “long-lost species” of giant water bug, Be- lostoma (now Abedus) immaculata (Belo- stomatidae). From Florida, he entered a VOLUME 89, NUMBER 2 Ph.D. program (1955-1958) under R. L. Usinger at the University of California, Berkeley; he continued his interest in aquat- ic Heteroptera with a dissertation, pub- lished in 1961, comprising a world revision of the marine water strider genus Halobates (Gerridae). For this latter project he spent five months in Hawaii studying field and laboratory populations. Shortly after receiving his degree, Jon was hired by the Government of American Sa- moa at Pago Pago as Staff Entomologist in charge of all entomological affairs. Those days, he said, were among his happiest. His duties included identification of insects, in- vestigations of their pest status on principal crops, inspection of plantations and incom- ing commerce, training of local farmers on methods of insect and disease control, and involvement in the rearing and release of parasites and predators. After two years in the South Pacific, Jon returned to the United States to take a po- sition with the U.S. Department of Agri- culture in Beltsville, Maryland to work on biological control of insects. Within two years, however, he moved on to become the Heteropterist for the Systematic Entomol- ogy Laboratory (Agricultural Research Ser- vice, U.S.D.A.) in Washington, D.C. where he spent the remainder of his career. Al- though his taxonomic speciality had been aquatic Heteroptera, Jon developed an in- terest in the predatory family Anthocoridae and wrote papers on this group, as well as on several other terrestrial families of ag- ricultural importance. Among his finer con- tributions were his revision of Halobates, key to the genera of North American An- thocoridae, and his role in initiating a cat- alog of the Heteroptera of Canada and the United States. He published 45 entomolog- ical papers and described as new 6 genera, 62 species, and 3 subspecies during his ca- reer, Jon was respected by his fellow heter- opterists who often sought his opinion and help through letters or personal visits. Such 385 help was freely given. He was companion- able, a good story teller, and a good listener. His broad interest in the world around him was evidenced in his reading matter which often found its way into conversations. Pos- sessor of real artistic talent, he illustrated many of his own papers and often dabbled in creative art. He was editor of the Pro- ceedings of the Entomological Society of Washington from 1963-1967; during that time he used his artistry to redraw the water strider Rheumatobates rileyi Bergroth for the Society seal that has appeared on the cover of the Proceedings from 1894-1920 and from 1937 to date (see Herring, 1964a: 1). He was also adept at macramé weaving and made many pieces for colleagues and friends. His skill with foreign languages allowed him to help colleagues with French, German, and Spanish. He also avidly collected and had a broad knowledge of U.S. coins. His in- terest in geneology enabled him to trace his ancestry into pre-Revolutionary times in the southern part of the present United States. Jon retired in August 1979 and moved to Ocala, Florida. He is survived by his wife Katherine (Kate) and three daughters, Kitt, Tiara, and Trina. INSECT TAXA DESCRIBED BY J. L. HERRING Family Anthocoridae: Genera: Alofa Herring, 1976c: 150 Dolostethus Henry and Herring, 1978: 520 Kitocoris Herring, 1967: 396 Tiare Herring, 1967: 400 Species: Blaptostethus pacificus Herring, 1967: 395 Dolostethus pubescens Henry and Herring, 1978: 522 Kitocoris omura Herring, 1967: 395 Lasiochilus ashlocki Herring, 1966b: 128 Lasiochilus ather Herring, 1967: 410 Lasiochilus campylus Herring, 1967: 409 Lasiochilus mesostenus Herring, 1967: 408 Lasiochilus palauensis Herring, 1967: 407 Nidicola aglaia Drake and Herring, 1964: 59 Nidicola engys Drake and Herring, 1964: 61 Nidicola etes Drake and Herring, 1964: 59 Nidicola mazda Herring, 1966b: 129 386 Nidicola mitra Drake and Herring, 1964: 61 Orius alcides Herring, 1966c: 1101 Orius candiope Herring, 1966c: 1098 Orius championi Herring, 1966c: 1095 Orius diespeter Herring, 1966c: 1098 Orius euryale Herring, 1966c: 1103 Orius florentiae Herring, 1966c: 1095 Orius harpocrates Herring, 1966c: 1097 Orius ixionides Herring, 1966c: 1100 Orius jasiones Herring, 1966c: 1108 Orius lesliae Herring, 1966c: 1096 Orius niobe Herring, 1967: 399 Orius pele Herring, 1966c: 1096 Orius thyestes Herring, 1966c: 1099 Tiare nesiotis Herring, 1967: 401 Xylocoris dybasi Herring, 1967: 413 Family Belostomatidae: Species: Abedus cantralli Hussey and Herring, 1950a: 85 (a junior synonym of Abedus immaculatus (Say), 1832) Family Coreidae: Species: Chelinidea staffilesi Herring, 1980: 239 Family Gerridae: Species: Halobates browni Herring, 1961: 270 Halobates bryani Herring, 1961: 286 Halobates calyptus Herring, 1961: 285 Halobates darwini Herring, 1961: 278 Halobates eschscholtzi Herring, 1961: 254 Halobates fijiensis Herring, 1958a: 10 Halobates galatea Herring, 1961: 294 Halobates katherinae Herring, 1958a: 8 Halobates kelleni Herring, 1961: 266 Halobates nereis Herring, 1961: 272 Halobates panope Herring, 1961: 295 Halobates peronis Herring, 1961: 278 Halobates poseidon Herring, 1961: 287 Halobates salotae Herring, 1961: 260 Halobates tethys Herring, 1961: 273 Halobates trynae Herring, 1964: 85 Hermatobates bredini Herring, 1965a: 124 Hermatobates tiare Herring, 1965Sa: 126 Rheumatobates crinitus Herring, 1949: 160 Subspecies: Metrobates anomalus comatipes Hussey and Her- ring, 1949: 169 Metrobates hesperius depilatus Hussey and Her- ring, 1949: 168 Metrobates hesperius ocalensis Hussey and Her- ring, 1949: 167 Family Miridae: Species: Corticoris infuscatus Henry and Herring, 1979: 85 Corticoris mexicanus Henry and Herring, 1979: 87 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Neoborella canadensis Kelton and Herring, 1978: 779 Neoborella pseudotsugae Kelton and Herring, 1978: 779 Neoborella xanthenes Herring, 1972: 9 Rhinacloa callicrates Herring, 1971: 449 Trynocoris lawrencei Herring, 1976a: 93 Family Saldidae: Species: Omania nauruensis Herring and Chapman, 1967: 355 Family Veliidae: Genus: Husseyella Herring, 1955: 21 Species: Halovelia danae Herring, 1958a: 12 Microvelia alachuana Hussey and Herring, 1950b: 117 Microvelia starmuehlneri Polhemus and Herring, 1970a: 181 Rhagovelia pidaxa Polhemus and Herring, 1970a: 183 ENTOMOLOGICAL PAPERS PUBLISHED BY J. L. HERRING Dailey, P. J., R. C. Graves, and J. L. Herring. 1978. Survey of Hemiptera collected on common milk- weed, Asclepias syrica, at one site in Ohio. Ento- mol. News 89: 157-162. Drake, C. J. and J. L. Herring. 1964. The genus Ni- dicola (Hemiptera: Anthocoridae). Proc. Biol. Soc. Wash. 77: 53-64. Henry, T. J. and J. L. Herring. 1978. A new genus and species of Cardiastethini from Peru (Hemip- tera: Anthocoridae). Proc. Entomol. Soc. Wash. 80: 520-523. Henry, T. J. and J. L. Herring. 1979. Review of the genus Corticoris with descriptions of two new species from Mexico (Hemiptera: Miridae: Iso- metopinae). Proc. Entomol. Soc. Wash. 81: 82- 96. Herring, J. L. 1948. Taxonomic and distributional notes on the Hydrometridae of Florida (Hemip- tera). Fla. Entomol. 31: 112-116. Herring, J. L. 1949. A new species of Rheumatobates from Florida (Hemiptera: Gerridae). Fla. Ento- mol. 32: 160-165. Herring, J. L. 1950a. The aquatic and semiaquatic Hemiptera of northern Florida. Part |: Gerridae. Fla. Entomol. 33: 23-33. Herring, J. L. 1950b. The aquatic and semiaquatic Hemiptera of northern Florida. Part 2: Veliidae and Mesoveliidae Fla. Entomol. 33: 145-150. Herring, J. L. 195la. The aquatic and semiaquatic Hemiptera of northern Florida. Part 3: Nepidae, VOLUME 89, NUMBER 2 Belostomatidae, Notonectidae, Pleidae and Co- rixidae. Fla. Entomol. 34: 17-29. Herring, J. L. 1951b. The aquatic and semiaquatic Hemiptera of northern Florida. Part 4: Classifi- cation of habits and keys to the species. Fla. Ento- mol. 34: 146-161. Herring, J. L. 1955. A new American genus of Ve- liidae (Hemiptera). Fla. Entomol. 38: 21-25. Herring, J. L. 1958a. The marine water-striders of the “Dana” Expeditions (Insecta: Hemiptera). Carlsberg Foundation’s Oceanographic Expedi- tion Round The World 1928-30 And Previous “‘Dana’’—Expeditions. Rept. No. 44. Andr. Fred. Host & Son, Copenhagen. 13 pp. Herring, J. L. 1958b. Evidence for hurricane trans- port and dispersal of aquatic Hemiptera. Pan-Pac. Entomol. 34; 174-175. Herring, J. L. 1961. The genus Halobates (Hemip- tera: Gerridae). Pac. Ins. 3(2—3): 223-305. Herring, J. L. 1964a. The official seal of the Ento- mological Society of Washington. Proc. Entomol. Soc. Wash. 66: 1. Herring, J. L. 1964b. A new species of Halobates from the Bay of Bengal (Hemiptera: Gerridae). Proc. Entomol. Soc. Wash. 66: 85-86. Herring, J. L. 1965a. Hermatobates, a new generic record for the Atlantic Ocean, with descriptions of new species (Hemiptera: Gerridae). Proc. U.S. Nat. Mus., No. 3510. 117: 123-130. Herring, J. L. 1965b. The status of Amphiareus Dis- tant, Buchaniella Reuter and Poronotellus Kir- kaldy. Proc. Entomol. Soc. Wash. 67: 202-203. Herring, J. L. 1966a. The correct name for an an- thocorid predator of the Cuban laurel thrips (He- miptera: Anthocoridae). Proc. Entomol. Soc. Wash. 68: 93. Herring, J. L. 1966b. The Anthocoridae of the Ga- lapagos and Cocos Islands (Hemiptera). Proc. Entomol. Soc. Wash. 68: 127-130. Herring, J. L. 1966c. The genus Orius of the Western Hemisphere (Hemiptera: Anthocoridae). Ann. Entomol. Soc. Am. 59: 1093-1109. Herring, J. L. 1967. Insects of Micronesia. Heter- optera: Anthocoridae. B. P. Bishop Mus. Ins. Mi- cronesia, Honolulu, Hawaii 7(8): 391-414. Herring, J.L. 1971. A new species of Rhinocloa from palo verde and ocotillo in the western U.S. Proc. Entomol. Soc. Wash. 73: 449. Herring, J.L. 1972. Anewspecies of Neoborella from dwarf mistletoe in Colorado. Proc. Entomol. Soc. Wash. 74: 9-10. Herring, J. L. 1973. Insects not known to occur in the United States. South African grain bug (Mac- chiademus diploterus (Distant)). U.S. Dept. Agric., Coop. Econ. Ins. Rpt. 23(43): 733-734. Herring, J. L. 1974. Insects not known to occur in the continental United States. New Caledonian 387 orchid pest (Nysius caledoniae Distant). U.S. Dept. Agric., Coop. Econ. Ins. Rpt. 24(30): 590-591. Herring, J. L. 1976a. A new genus and species of Cylapinae from Panama (Hemiptera: Miridae). Proc. Entomol. Soc. Wash. 78: 91-94. Herring, J. L. 1976b. Pests not known to occur in the United States. Pacific rice chinch bug, Dimpor- hicus pilosus (Barber). Hemiptera: Lygaeidae. U.S. Dept. Agric., Coop. Plant Pest Rpt. 1(48-52): 897- 899. Herring, J. L. 1976c. Keys to genera of Anthocoridae of America north of Mexico, with description of a new genus (Hemiptera: Heteroptera). Fla. Ento- mol. 59: 143-150. Herring, J. L. 1980. A review of the cactus bugs of the genus Chelinidea with the description of a new species (Hemiptera: Coreidae). Proc. Entomol. Soc. Wash. 82: 237-251. Herring, J. L. and P. D. Ashlock. 1971. A key to the nymphs of the families of Hemiptera (Heterop- tera) of America north of Mexico. Fla. Entomol. 54: 207-212. Herring, J. L. and H. C. Chapman. 1967. A new species of Omania from Micronesia (Hemiptera: Saldidae). Proc. Entomol. Soc. Wash. 69: 354— 359. Herring, J. L. and P. S. Dale. 1960. Pest Control in Samoa. Bulletin of the Board of Agriculture, For- estry and Fisheries. Apia, Western Samoa. Herring, J. L. and H. G. Dowling. 1947. Observation on Dolomedes albineus Hentz. Fla. Entomol. 30: 42. Hussey, R. F. and J. L. Herring 1949. Notes on the variation of the Metrobates of Florida (Hemiptera, Gerridae). Fla. Entomol. 33: 32: 166-170. Hussey, R. F. and J. L. Herring. 1950a. A remarkable new belostomatid (Hemiptera) from Florida and Georgia. Fla. Entomol. 33: 84-89. Hussey, R. F. and J. L. Herring. 1950b. A new Mi- crovelia from Florida (Hemiptera: Veliidae). Fla. Entomol. 33: 117-120. Hussey, R. F. and J. L. Herring. 1950c. Rediscovery of a belostomatid named by Thomas Say (He- miptera). Fla. Entomol. 33: 154-156. Kelton, L. A.and J. L. Herring. 1978. Two new species of Neoborella Knight (Heteroptera: Miridae) found on dwarf mistletoe, Arceuthobium spp. Can. Ento- mol. 110: 779-780. Mead, F. W. and J. L. Herring. 1974. A cactus bug, Chelinidea vittiger aequoris McAtee, in Florida (Hemiptera: Coreidae). Florida Dept. Agric. & Consumer Serv., Div. Plant Industry, Entomol. Circ. No. 149. 2 pp. Polhemus, J. T. and J. L. Herring. 1970a. Ergebnisse der Osterreichischen Neukaledonien Expedition. Aquatic and semiaquatic Hemiptera. Proc. Ento- mol. Soc. Wash. 72: 179-187. 388 Polhemus, J. T. and J. L. Herring. 1970a. Etudes hydrobiologiques en Novelle-Calédonie (Mission 1965 du premier Institut de Zoologie de lUni- versité de Vienne) (Suite). X. Aquatic and semi- aquatic Hemiptera of New Caledonia. Cah. O.R. S. T. O. M., sér. Hydrobiol. 4(2): 3-12. Polhemus, J. T. and J. L. Herring. 1979. A further description of Hermatobates bredini Herring and a new record for Cuba (Hemiptera: Hermatoba- tidae). Proc. Entomol. Soc. Wash. 81: 25 3-254. Usinger, R. L. and J. L. Herring. 1957. Notes on marine water striders of the Hawaiian Islands (He- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON miptera: Gerridae). Proc. Haw. Entomol. Soc. 16: 281-283. Wheeler, A. G., Jr. and J. L. Herring. 1979. A po- tential insect pest of azaleas. Q. Bull. Am. Rho- dodendron Soc. 33: 12-14, 1 color fig. Thomas J. Henry, Systematic Entomology Laboratory, BBII, Agricultural Research Service, U.S.D.A., % USNM, Washington, D.C. 20560, and Richard C. Froeschner, Department of Ento- mology, Smithsonian Institution, Washington, DIE; 20560: PUBLICATIONS FOR SALE BY THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MISCELLANEOUS PUBLICATIONS Cynipid Galls of the Eastern United States, by Lewis FH. Weld ec sceececeeneseneeeeteceteeeentnnnnnnnenee $ 5.00 Cynipid Galls of the Southwest, by Lewis H. Weld... ES Ne 2 3.00 MOEMpADELS ON CyNIDIGl AAllk Ge eA UtL eM UAT ee MER) BOM ACU AC SY aA DR ER 6.00 Identification of Alaskan Black Fly Larvae, by Kathryn M. 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The Nearctic Leafhoppers, a Generic Classification and Check List, by Paul Wilson Oman. : DSS) (V0). INO Ae UN 8 AL RU Ee A AR Ok UP a AA SUA 15.00 No. 4. A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 195220000 15.00 No. 5. A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp. os MAO CEE CE es ee eT CRD aE 15.00 No. 6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi ABaAASI 2S Ope VOOM eeu ene we NUE wt My Te Std ei BOY Ba 0 ge Wa BS oe Be SN 15.00 No.7. Ant Larvae: Review and Synthesis, by George C. Wheeler and Jeanette Wheeler. 108 pp. / emer AWA UND TUR PINE ie Ly eR CU VCE Pee NE SUA Pace 11.00 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 0 12.00 No.9. The Flower Flies of the West Indies (Diptera: Syrphidae), by F. 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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. 4 : j CONTENTS (Continued from front cover) McPHERSON, J. E., R. J. PACKAUSKAS, and P. P. KORCH, III—Life history and laboratory rearing of Pelocoris femoratus (Hemiptera: Naucoridae), with descriptions of immature © stages 1), Fe SUA ON TAU A ee ae ee Ce Hl ee ORTH, R. E.—A new species of Pherbellia from North America with range extensions for P. hackmani and P. griseicollis (Diptera: Sciomyzidae) ..............0.0. 0002 c cee cece ee POULTON, B. C. and K. W. STEWART—Three new species of stoneflies (Plecoptera) from fi the/Ozark-Ouachita: Mountain Region yy Oe) eat ea ed a ee RITCHIE, A. J. and J. D. SHORTHOUSE-—A review of the species of Synergus from Guatemala, | with notes on Cynips guatemalensis Cameron (Hymenoptera: Cynipidae) .............. ROSENHEIM, J. A. and J. K. GRACE—Biology of a wood-nesting wasp, Mimumesa mixta (W. Fox) (Hymenoptera: Sphecidae), and its parasite, E/ampus viridicyaneus Norton (Hy- menoptetayG@hrysididae) a's YR Cw FS Woe A Le eed ee OIC at acm ee ET aa RUSSELL, L. M.—Habits and biology of the beech mealybug, Peliococcus serratus (Ferris) ; (Caccnidea. Pseudococeidde) Ao ke FAT a ed A a 359 SABROSKY, C. W.—A new species of Leptometopa (Diptera: Milichiidae) from Madagascar — pollinating |\Ceropesia’ (Asclepiadaceae) i hd h Ae ak ce ee a) Lees ree SPANGLER, P. J.—A new species of water penny beetle, Pheneps cursitatus, from Cerro de la Neblina, Venezuela (Coleoptera: Dryopoidea: Psephenidae) ................... PET RRR WEBB, D. W.—A revision of the Nearctic species of Arthroceras (Diptera: Rhagionidae) .... WHEELER, A. G., JR., and J. E. FETTER—Cilacis typhae (Heteroptera: Lygaeidae) and the subfamily Artheneinae new to| North America .°.)..05 5) 506 Dba de fea 244 NOTES MAIER, C. T.—New distributional and rearing records for Neotropical flower flies (Diptera: _ Syrphidae) iy iy Py a has MO AA AU TT ea FO) ie a a ' TAFT, S. J. and S. MARCQUENSKI—The prevalence of Jcosta americana (Diptera: Hippo- boscidae) on ruffed grouse (Bonasa umbellus) in Wisconsin ....................2.+25-- WEBB, R. E.—Anomalous tubercle patterns found on Lymantria dispar (L.) caterpillars in the field\in' Maryland, (Lepidoptera: Lymantriidae)\\ i Soa ae ae a eat) ea BOOK NOTICE SPILMAN, T. J.— The Insect and Spider Collections of the World ...........0..6.00 000.000 BOOK REVIEW LABERGE, W. E.— Foundations for a National Biological Survey ...........00 0.00 eevee OBITUARIES CLIFFORD, C. M. and J. E. KEIRANS—Glen Milton Kohls 1905-1986 .................. HENRY, T. J. and R. C. FROESCHNER—Jon Lamar Herring 1922-1985 ................ SOCIETY MEETINGS AND REPORTS OF OFFICERS FOR 1986 .2....05.0.0.0) 0.0.00: VOL. 89 JULY 1987 NO. 3 tone | (ISSN 0013-8797) PROCEEDINGS of the IN TOMOLOGICAL SOCIETY = ee Sew -'s < . - a x eee Sa cepa hi, | CONTENTS Ne il i, idae) a beat former on the ostrich fern, Matteuccia struthiopteris, with notes on othe insect- REGEN ASSOCIATES pea ela eR ee Leh Me EN DNL LOI AR RG Doe OM A RE aM i MUR Be) 532 g _ BEYER, W.N., G. W. MILLER, and W. J. FLEMING— Populations of trap-nesting wasps near : Nf "3 it _-a major source of fluoride emissions in western Tennessee ................0000-2 220055 478 We BLANC, F. L. and R. H. FOOTE—Taxonomic observations on United States Tephritidae iy mvMiptera), with descriptions of mew species) 1. /.)) 05.) CA We SEO Ee ae ie by 425 i) By BORKENT, A., W. W. WIRTH, and A. L. DYCE—The newly discovered male of Austroconops #5 rt _ (Ceratopogonidae: Diptera) with a discussion of the phylogeny of the basal lineages of the : , aa Oe) DE Ce Oy RU ME ATL MUA LEE 587 1 WE DA AVIS, H. G., L. M. MCDONOUGH, and D. C. FERGUSON-—Sex attractant for Scoparia Bit) ai mamacials Keenideptera: Peyralidaey 205.0) Ee See eB Pel 500 i 0 DAVIS, J. R.—A new species of Farrodes (Ephemeroptera: Leptophlebiidae: Atalophlebiinae Rem pomincrn Texas 7 MMe ib lek Ae a ee ei ee Oe 407 r Ns! | - DODSON, G.—Host-plant records and life history notes on New Mexico Tephritidae | ae PRETAU VCMT AURA Li iA be Ce UR A eae mee ME RAC CPE ONLY t Beiiig By Di loo asf incre Be 607 __ DOWNES, W.L., Jk.—The impact of vertebrate predators on early arthropod evolution .... 389 i EVANS, H. E.—A new species of Jrenangelus from Costa Rica (Hymenoptera: Pompilidae: Hy) iy a re eo AOU AT eG Renae SCT PS a ae A MI SP Tt AE 559 GILES, F. E. and W. W. WIRTH—New species and records of New Caledonia Alluaudomyia i Mt pnserlrnteoparanidacy).)). cee. tk a EM ee Be he OEY a eh 458 it _ GOEDEN, R. D., T. D. CADATAL, and G. A. CAVENDER-— Life history of Neotephritis finalis i} _ (Loew) on native Asteraceae in southern California (Diptera: Tephritidae) .............. 552 ‘\ eas, S. A.—A review of the Holarctic genus Terrilimosina (Diptera: Sphaeroceridae) i i : _ with descriptions of new species from Nepal and Japan ............0..6 00000000 ce eee. 502 (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ORGANIZED MARCH 12, 1884 OFFICERS FOR 1987 THOMAS E. WALLENMAIER, President MICHAEL J. RAupp, Program Chair an F. EUGENE Woop, President-Elect GEOFFREY B. WuiTE, Membership Chairman PAUL M. MarsH, Recording Secretary VicTOR L. BLACKBURN, Custodian RICHARD G. RosBINS, Corresponding Secretary MANYA B. STOETZEL, Delegate, Wash. Acad. Sci. NORMAN E. WoopDLEY, Treasurer RAYMOND J. GAGNE, Editor Publications Committee DAvID R. SMITH THEODORE J. SPILMAN GEORGE C. STEYSKAL — Associate Editor my HIRAM G. LAREW Honorary President C. F. W. MUESEBECK Honorary Members : FREDERICK W. Poos _ ASHLEY B. GURNEY THEODORE L. BISSELL — All correspondence concerning Society business should be mailed to the appropriate officer at the follov rin g address: Entomological Society of Washington, c/o Department of Entomology, NHB 168, Smithsonian Insti- tution, Washington, D.C. 20560. i 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 publish i regularly in the Proceedings. ; MEMBERSHIP.— Members shall be persons who have demonstrated interest in the science of entonialae py. Annual dues for members are $20.00 (U.S. currency) of which $18.00 is for a subscription to the Procee of the Entomological Society of Washington for one year. PROCEEDINGS. — The Proceedings are published quarterly beginning in January by The Entomological Soci of Washington, % Department of Entomology, NHB-168, Smithsonian Institution, Washington, D.C. Member ; in good standing receive the Proceedings of the Entomological Society of Washington. Nonmember subscription: sat) are $35.00 per year, domestic, and $40.00 per year, foreign (U.S. currency), payable in advance. Foreign deliver yo 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. a é Title of Publication: Proceedings of the Entomological Society of Washington. . ’ Frequency of Issue: Quarterly (January, April, July, October). i Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of Was h- 2 ington, c/o Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash- ington, D.C. 20560. Editor: Raymond J. Gagné, Systematic Entomology Laboratory, c/o U.S. National Museum NHB 168, Wash: ington, D.C. 20560. At Managing Editor and Known Bondholders or other Security Holders: none. Please see p. 730 of the July 1984 issue for information regarding preparation of manuscripts. STATEMENT OF OWNERSHIP This issue was mailed 23 July 1987 Second Class Postage Paid at Washington, D.C. and additional mailing office. PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA PROC. ENTOMOL. SOC. WASH. 89(3), 1987, pp. 389-406 THE IMPACT OF VERTEBRATE PREDATORS ON EARLY ARTHROPOD EVOLUTION WILLIAM L. Downes, JR. Department of Entomology, Michigan State University, East Lansing, Michigan 48824. Abstract.—The extinctions of several “‘orders”’ of insects near the close of the Paleozoic resulted from increasingly effective predation by visually hunting tetrapods. Forms living on the surface of the ground were the first to be affected. The larger immatures of the early Pterygota were especially vulnerable because they could not yet fly, nor could they enter crevices easily because they were encumbered with large, laterally projecting wing pads. Surviving “orders” had immatures capable of avoiding diurnal tetrapod predators. Some immatures escaped by living in water (Ephemeroptera, Odonata, Plecoptera); some hid in crevices during the day (Blattodea, Coleoptera, etc.), or were too small to be attractive as prey (Collembola, Psocoptera). Endopterygote larvae lived out of sight in soil litter; mecopteroid larvae lived there permanently, neuropteroid larvae hid there during the day, but foraged above ground at night. Intrinsic advantages in exploiting divergent larval and adult habitats did not induce the complete metamorphosis of endopterygotes; vertebrate predators coerced the diver- gence between larval and adult modes of life. The adaptive radiation of the Endopterygota also did not result from direct competitive advantages of their metamorphosis. The larvae simply moved into new niches created directly or indirectly by the adaptive radiations of vascular plants. There were probably no fewer exposed niches for exopterygotes, but tetrapod predators severely restricted the exploitation of those niches. The neopterous wing-flexing mechanism was originally an adaptation for settling into open crevices. It eliminated the silhouette that revealed the insect’s position, especially on tree trunks. It was evidently very effective until beaked birds began fishing insects from crevices. Aerial predation by Odonata and Protodonata deterred early insects from flying freely during the day. For this reason the adults of most orders came to rely primarily on means other than flight to escape tetrapods. Early terrestrial arthropods played an in- dispensible role in the origin and early evo- lution of terrestrial vertebrates: they were the food that sustained vertebrates as they became terrestrial. Apparently, only the Ar- thropoda developed terrestrial populations sufficiently large to support the widescale adaptive radiations that occurred among the early terrestrial vertebrates. Terrestrial ver- tebrate herbivores, capable of supporting significant populations of vertebrate pred- ators, did not appear until the Permian. In- deed, the rapid evolutionary expansion of early reptiles seems to have occurred pri- marily in response to the rapid increase in numbers and kinds of insects (Olson, 1976). Many later adaptive radiations among ter- restrial vertebrates similarly appear to have depended on an “insect”? food base (Roh- dendorf, 1970). 390 The earliest known terrestrial vertebrates were found in Carboniferous deposits. Their dentition consisted of small, sharp-pointed teeth. Since such teeth generally function in seizing small arthropod prey and swallow- ing them whole in modern species, then the early terrestrial vertebrates must have been similarly “‘insectivorous” (Carroll, 1982; Gregory, 1950; Olson, 1976; Milner, 1980). The preceding ideas have been widely ac- cepted, and are not in question here. The references cited provide access to additional discussion on this matter. MISSING HYPOTHESES The early dependence of terrestrial ver- tebrates upon arthropods has not lacked in interest, nor was the idea suggested only re- cently. But, curiously, the reciprocal impact of early tetrapods on insects and their rel- atives has never received comparable atten- tion. The primary focus has almost invari- ably been on vertebrate evolution, although Rolfe (1980) made several references to pos- sible coevolutionary relationships between tetrapods and terrestrial or amphibious ar- thropods, and Kukalova-Peck (1978) spelled out anumber of significant but seldom men- tioned features of Paleozoic insects, consid- ered mainly in relation to wing evolution and metamorphosis. Obviously, the advent of reptiles was a critical development for terrestrial arthro- pods. The reptiles were not a minor group, and terrestrial arthropods were their main food. It is inconceivable that early terrestrial vertebrate predators did not seriously affect the course of arthropod evolution on land. The purpose of this paper is to point out that the extinctions of the Palaeodictyoptera and several comparable “‘orders’’ of insects in the late Paleozoic resulted from early ver- tebrate predation. Further, many conspic- uous features of the surviving arthropod or- ders were shaped originally, and have continued to be shaped, in no small measure by terrestrial vertebrate predators. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON These hypotheses and their derivatives differ from previous considerations in that they do not have vertebrate evolution as their subject; they concern particular changes that occurred among early arthropods as a result of terrestrial vertebrate predation. Ex- tinction, of course, is one of the more sig- nificant and drastic changes. SOURCES OF THE EXPLANATIONS The ideas presented here are hypotheses extrapolated from published ideas or ideas stemming from common field observations of modern species. The key hypothesis de- rives (in part) from the following: verte- brates (presumably birds foremost, but also lizards and diurnal amphibia) have had a staggering impact on insects in the tropics. Elton (1973) reported finding widespread feeding damage on the foliage of Neotrop- ical trees, but, paradoxically, virtually no sign during the day of any insect that could have caused the damage. Evidently, neither cryptic nor warning coloration nor any or- dinary escape mechanisms provide sufh- cient protection under the highly competi- tive conditions of the tropics. The insects have resorted to hiding physically from their vertebrate predators during the day. Adaptive coloration in insects 1s one of the strongest indications of the profound impact of visually-hunting vertebrate pred- ators. All insects of any size exposed during the day are colored. This is so obvious that it is seldom mentioned—in such general terms. Yet, the fact that cave insects, fly maggots, certain beetle larvae, etc., all of which lead continuously concealed lives, are not colored points to the all-pervasive effect of visually-hunting vertebrate predators. The arthropod compound eye is not known to provide sufficient visual acuity to account for the elaborate colors and structural pat- terns of contemporary insects, which leaves vertebrates as the primary selective agents for most of these patterns. Some coloration is not related to verte- VOLUME 89, NUMBER 3 brate predation, such as that employed in courtship or heat regulation or the blacks or browns incidental to strong sclerotizations. None of these (as color patterns) are as prev- alent or basic among insects as cryptic or aposematic patterns that relate to vertebrate predation. The eye-spot patterns on the wings of cer- tain Paleozoic insects (Carpenter, 1971) suggests that the use of color patterns in insects as a response to tetrapod predators was determined that long ago, no doubt largely by the fundamental structure and ca- pabilities of the vertebrate eye and brain. According to Blest (1957), the eye-spot pat- terns of Lepidoptera wings may serve two functions: 1) they may simulate a larger predator and so frighten off an attacker, or 2) they may, as a salient target, deflect an attack to a less vulnerable part of the lepi- dopteran body. The last, presumably, de- flects bird pecks from the vulnerable body to the less vulnerable wing extremities. In either case, eye spots function as defenses against vertebrate attack. But birds did not exist in the Paleozoic! EXTINCTION BY VERTEBRATE PREDATION It is not possible to say exactly when any major type of terrestrial arthropod became extinct. The fossil record does not reveal when the last palaeodictyopteran disap- peared from the earth. Nevertheless, certain groups occur regularly up to a certain point, but are consistently absent from later de- posits. There is always the possibility that the early and later deposits are not equiv- alent, and that later representatives simply were not fossilized. However, the total ab- sence of Palaeodictyoptera and various oth- er orders from all records after the Permian indicates that a major change had taken place, whether catastrophic or gradual, even if this change was completed in the Triassic. For the purposes of this paper, such dis- appearances from the record are treated simply as extinctions. If some relicts lasted 391 much longer than they appear to have from the fossil record, it is of little consequence for the hypotheses developed here. The hypothesis that early vertebrate predators reduced populations of certain types of insects, such as the Palaeodictyop- tera, but not others, to the point of extinc- tion accords far better than any alternatives with conventional evolutionary thought and the data of the fossil record. As already mentioned, tetrapods began by eating ter- restrial arthropods. It is unreasonable to as- sume that terrestrial arthropods were unaf- fected, since vertebrates were becoming increasingly numerous and better adapted for life on land. Moreover, the extinction of major arthropod types correlates in geolog- ical time with the early adaptive radiations among terrestrial vertebrates, the reptiles in particular. Vertebrates are (and were) normally larg- er than their arthropod prey—in vertebrate- arthropod encounters it is the arthropod that would ordinarily end up as prey. While some mammals are nocturnal insectivores, the early tetrapods appear to have been diurnal and visually oriented hunters. Conceivably, arthropods and their tetra- pod predators might enter into coevolu- tionary progressions without major extinc- tions. No doubt many did. However, extinctions are possible under certain con- ditions. For example, New Zealand mayflies evolved in streams in which they were safe from predatory fish. Their immature stages lived an exposed life, and did not hide like their relatives in the rest of the world. Some swam freely through open water. Trout, 1n- troduced early in this century, ate them in such numbers that in 20 years the once ex- tremely abundant mayflies had become rare if not extinct (Tillyard, 1926; Burks, 1953). It does not matter that the New Zealand mayflies may not be completely extinct. The drastic reduction in their numbers effec- tively removed their evolutionary potential in New Zealand from them. The Paleozoic 392 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tetrapods are equivalent to the trout in New Zealand. EXTINCTIONS: ALTERNATIVE EXPLANATIONS A few alternative explanations have been suggested for the early arthropod extinc- tions. Major climatic changes, a large me- teorite impact, or simple replacements by more efficient types have usually been re- garded as the more plausible. Climatic changes in combination with replacements by new types have been suggested as the primary cause of the late Paleozoic extinc- tions (Ross, 1965, for example). Hennig (1981) suggested essentially the same idea, but emphasized replacements by more vig- orous types of insects. The Permian has been characterized as a time of tectonic upheaval and great climatic change. It has been called “‘a crisis point in the history of life’? (Schuchert and Dunbar, 1941). Both Newell (1982) and Raup (1979) discuss the peak of extinctions of major groups that occurred near the end of the Permian. This is superficially consistent with the idea that climatic changes were respon- sible for the extinctions of various arthro- pod groups, which were replaced by more adaptable types later on. However, the Appalachian Revolution with its widespread uplifts and more varied, drier climates did not abruptly terminate the Permian Period. The Revolution began with and continued during the Period (Col- bert, 1980). In such case the extinctions ought to have occurred mainly at the be- ginning of the Permian when arthropods would have been most seriously stressed in their first encounters with the new climates. On the contrary, Carpenter (1977) point- ed out that the Permian insect fauna was obviously much more diverse than the Pennsylvanian—“‘In terms of diversity of form and the association of generalized and specialized species, the fauna of the Perm- ian was probably the most diverse in the history of the Insecta.” If anything, the widespread uplift with the increased diver- sity of climates and habitats had an effect opposite to the one called for: it seems to have been more effective in producing new orders and adaptive radiations rather than extinctions among arthropods. Later tec- tonic upheavals and the accompanying cli- matic changes also do not seem to have pro- duced comparable major extinctions among arthropods as, for example, during the Lar- ramide Revolution at the end of the Me- sozoic. The latter revolution is as much as- sociated with adaptive radiations as with extinctions. From these considerations alone there is no reason to assume that any or- dinary climatic changes caused the late Pa- leozoic arthropod extinctions. A serious drawback to the climatic change hypothesis 1s that it is extremely vague. Ex- actly how did Permian climatic changes af- fect the evolution of any given species or group? Most of the known insect fossils are from only a few tropical swamp forest de- posits, although some are from temperate deposits (Wootton, 1981). And for what reason would one order, such as the Pa- laeodictyoptera, be affected disastrously by climatic changes, but not another, such as the Blattodea. In its present loose formu- lation the climatic change hypothesis does not permit deducing accurate enough con- sequences to permit practical observational tests. More plausibly, extinctions might have resulted from a climatic disaster as might have been caused by a large meteorite 1m- pact. This has been suggested as the reason for the extinctions of the dinosaurs at the close of the Mesozoic (Alvarez et al., 1982, is a general consideration of this topic and is part of a major symposium on the subject, q.v.). This proposal has the advantage of yielding several precise consequences which are that the extinctions should be abrupt, extensive and synchronized, and that they should be followed by a slow recovery from an impoverished fauna. The geological record for the closing pe- riod of the Paleozoic fits this pattern in some VOLUME 89, NUMBER 3 respects: the later Permian mass extinction has been regarded as “‘the single most dev- astating collapse of the marine ecosystem” we know of (Sepkoski, 1982). Numerous ex- tinctions appear to have occurred at that time (Raup, 1979). However, the arthropod extinctions were not all simultaneous. Some occurred earlier or later (diagram in Smart and Hughes, 1973). Most of the extinctions were not sud- den (Tappan, 1982). Further, the record of the extinctions is essentially a marine re- cord. There are few fossils of terrestrial ar- thropods to compare from the early Trias- sic; the apparently synchronous extinctions of several major arthropod groups near the close of the Permain might be an artifact of the fossil record. What evidence there is, including much from the vertebrate record, ‘““makes a catastrophic event at the end of Permian time difficult to support’? (Olson, 1982) (speaking of terrestrial environ- ments). No one has been able to confirm an iridium anomaly near the end of the Perm- ian (Silver, 1982; the iridium anomaly has been intepreted as a result of a major me- teorite impact in the Cretaceous). But could the extinctions reflect replace- ments by better adapted types, as Hennig (1981) seems to favor? At least from the standpoint of major types of insects, there does not appear to be any support for this view either. Which order supplanted the Palaeodictyoptera (or any other group), or which order was transformed into the Dip- tera (or any other order)? Virtually none of the relevant major phylogenetic questions have been answered (Wootton, 1981). Pres- ent information indicates that a number of orders of both insects and arachnids dis- appeared in the late Paleozoic with no known descendents and no recognizable replace- ments. Another possibility is that the extinction of a major plant group would result in the extinctions of insect dependents. The diffi- culty with this hypothesis is that this pre- supposes that a major arthropod group was 393 entirely dependent on a particular plant group that then became extinct near the end of the Paleozoic. Some palaeodictyopteroid insects may have fed on the endosperm of paleozoic megaspores, but this is far from suggesting that all of them did, or that none were capable of evolving along with plants that survived. The extinction of the plant group itself would still beg explanation. Tappan (1982) developed an interesting plant dependency hypothesis. She related the marine extinctions at the end of the Permian to the development of a more ad- vanced land vegetation. The increased bio- mass and more extensive soils which ac- companied this vegetation retained much of the nutrient material that formerly washed into the sea. Presumably this effect was ex- aggerated by the widespread uplift that seems to have characterized the Permian (because it resuted in a more extensive terrestrial vegetation). As a result, the marine phyto- plankton became increasingly starved, and eventually whole food chains collapsed. The collapse of marine food chains, ac- cording to Tappan’s hypotheses, does not entail a terrestrial catastrophe. They imply exactly the opposite: a greater development of terrestrial plant life and, presumably, an enriched arthropod evolution there. It should be pointed out, however, that a ma- rine catastrophe of the magnitude recog- nized must surely have affected life on land to some degree. If nothing else, marine pho- tosynthesis would probably have been af- fected, which is no minor matter. However, no one seems to have detailed such conse- quences, nor have any of the major terres- trial extinctions been related specifically to the marine catastrophe. Another possibility is that predatory arachnids rather than vertebrates caused the major arthropod extinctions. This also is not plausible. The fossil record for terres- trial arthropods is very poor until the Car- boniferous. At that point both arachnids and insect appear as rich coexisting faunas, which became more diverse together during the 394 Carboniferous and Permian (Savory, 1977). Major extinctions caused by arachnida must have occurred prior to the development of a significant fossil record, perhaps in the Devonian. The Devonian myriapods (Peach, 1889), Collembola (Scourfield, 1940), and arachnids (Rolfe, 1980) point to an earlier inland differentiation of which we have scant record. But, since we know so little, it is not possible to develop precise ideas about such early events. Another possibility 1s to assume that there is no one explanation, but various unknown reasons for the extinctions. To take this po- sition precludes developing precise expla- nations, and it leaves untouched the prob- lem of why certain orders should become extinct rather than others. Why not some ofeach? The fact that it was not simply some of each, and that many of the extinctions occurred in the same general time period indicates that the extinctions were not ran- dom or “pseudo-random” but according to some pattern that affected certain arthropod types more seriously than others. Hence, general explanations should be possible. A MobE oF LIFE PROBLEM Schmalhauzen (1957 —discussed in Roh- dendorf, 1970) argued for a view that is not compatible with the vertebrate predation hypothesis developed here. They suggested that the first terrestrial vertebrates were ini- tially ““passive,”’ feeding on insects when the latter crawled into their mouths. Glandular areas in the amphibian mouth were evi- dently taken as the supporting evidence, ap- parently on the assumption that such glands must have been present originally because the “salivary” glands of contemporary am- phibia must be derivatives of some such precursors. The glands were said to function in the “fixation” of the food insects. While not mentioned by Rohdendorf, nocturnal habits might be regarded as likely for early amphibia becoming reptiles. Their moist skin would render them susceptible to desiccation; nocturnal habits would have PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON them active when relative humidities would ordinarily be higher than during the day. Nocturnal habits would not be especially conducive to active visual predation, and, hence, these ideas are more compatible with Schmalhauzen’s and Rohdendorf’s ideas than with the hypotheses developed here. But a host of implications have been ig- nored. There are simply no grounds for as- suming that whatever glands may have been present functioned as sticky traps in propped-open mouths. Clamping the mouth shut should have “‘fixated”’ any ordinary in- sect prey. It is also difficult to conceive of any features of an aquatic ancestor that would predispose them or their amphibian descendants to such a mode of prey capture. That 1s, no origin or pathway can be pointed to that might lead to the situation depicted by Rohdendorfand Schmalhauzen, and nei- ther the hypothesis nor its implications are supported by any observational data I have been able to find. Had the first terrestrial vertebrates started as postulated, selection might be expected to have produced ever better methods of baiting and sticky-trap- ping. So far as we know, it produced neither; it produced active hunters. Neither fossil nor modern amphibian structures suggest the ‘“‘passive’’ feeding hypothesized by Schmalhauzen and Rohdendorf. Even the nocturnal habits of modern Am- phibia may not be primarily adaptations for avoiding desiccation. They are as easily ex- plained as the result of predation by larger vertebrates. Frogs, which ambush insect prey and might be regarded as the closest modern approximation to the Schmalhauzen pro- totype, depend on their comparatively large eyes for prey capture; these indicate earlier diurnal habits. The highly poisonous har- lequin frogs (Bufonidae) of Central America are neither retiring nor nocturnal (Martha L. Crump, personal communication, 1985); but they are apparently too poisonous to be attractive as prey for most predators. By way of contrast, the simplest transition from an aquatic life to one on land entails VOLUME 89, NUMBER 3 supposing only that certain amphibians moved from seizing prey and swallowing it whole in water to doing the same on land, as their teeth suggest. This is a direct, simple and obvious pathway that began with na- scent amphibia making temporary forays onto land for insect prey. It implies active searching rather than passive trapping, which is consistent with the continued de- velopment of eyes and legs in the subse- quent adaptive radiations of terrestrial ver- tebrates. The eyespot patterns on the wings of a Paleozoic insect (Carpenter, 1971) make little sense, except in the context of visually hunting tetrapods. OBSERVATIONAL TESTS OF THE THEORY The vertebrate-predation hypothesis de- veloped here, when conjoined with various generally accepted or fairly obvious ideas, yields numerous implications (deductions or predictions) that fit available observa- tional data extremely well. Several exam- ples are discussed below. The earliest vertebrate incursions into the terrestrial arthropod world probably began in the Devonian in the vicinity of bodies of water where the early vertebrates lived. As mentioned above, the habit of seizing prey and swallowing it whole would suffice for prey capture on land as well as in water. No major structural or behavioral changes would be called for aside from those related to coping with the physical problems of an aquatic animal attempting to live on land. The essentially untapped terrestrial arthro- pod fauna would have been a major entice- ment for the earliest tetrapods to forage there, and no comparable predators had preempted their predatory role on land. The arachnids were not equivalent competitors, since the tetrapods could eat them also. The vertebrate emergence from water onto land was no minor transition, since virtually every organ system would need to be re- organized for the vastly different conditions of life out of water. Consequently, the first amphibian tetrapods must have been com- 395 paratively inefficient and clumsy predators on land; their impact on arthropods must have been minimal until reptiles more fully adapted for life on land evolved. At that point, it would no longer be safe for arthro- pods to roam freely over the surface of the ground during the day; arthropods active on the ground surface would be especially vul- nerable to early reptilian predation. In general, sensory-nervous mechanisms are slow to evolve (Smith, 1952), apparently because of their extreme complexity. Ac- cordingly, we should expect the early ter- restrial arthropods to be limited to rela- tively simple mechanisms for avoiding vertebrate predation. The following seem most likely: 1) The early arthropods might not “re- spond”’ at all, but simply succumb to ver- tebrate predation and become extinct. 2) They might avoid vertebrates, as by crawling out of sight into the soil litter. 3) They might become too small to be attractive as prey. 4) They might evade predators by jump- ing. 5) They might utilize distinctive color patterns to frighten vertebrates (the eye- spots mentioned above). 6) They might develop noxious sub- stances. 7) They might survive by living up on tree trunks out of reach of the early verte- brates rather than on the ground. The last, a variant of option 2, may not be obvious. Tetrapod surface gaits are not suitable for climbing trees; for them, tree climbing requires the evolution of consid- erable ‘‘eye-hand” coordination, that is, it requires new, well-coordinated sensory- neural-muscular mechanisms that primi- tive tetrapods would not have had time to develop. Climbing otherwise would be haz- ardous, since falls from any height would injure animals of their size. Fossils of the earliest land vertebrates do not show ad- aptations for tree-climbing, as would be ex- pected, although insectivorous, gliding rep- 396 tiles appear in the Upper Permian (Evans, 1982). Insects and myriapods could climb easily because of their clawed legs and small size. Falls pose little threat, even to wingless in- sects, since most reach a low terminal ve- locity so quickly, that they are rarely injured in a fall, regardless of height. Epiphytic al- gae, fungi and lichens would enable them to feed on tree trunks as readily as on the ground. From this it follows that some early ar- thropods living on tree trunks would sur- vive early tetrapod predation longer than ground-surface species. Winged insects would best fit the epiphytic mode of life, since their adults could move the species between trees or stands (which would give easier access to their food and enable them to maintain larger outcrossing populations so important for their evolution). But trunk- dwelling arthropods would eventually face the same basic problems their predecessors had faced earlier on the ground. Simply climbing and feeding on tree trunks would, accordingly, be of only temporary utility, and epiphytic insects would eventually be limited to the first six options. In summary, we should predict that forms dwelling on the surface of the ground would be the first to disappear (if of sufficient size to be attractive prey) followed by those that lived on tree trunks. As a corollary, all sur- viving groups should exemplify one or more of the first six responses to predators, as listed above. These predictions are realized abundantly among tracheates. Example 1.—The recent and Paleozoic diplopod faunas do not resemble each other (Kraus, 1974). For this reason it had been assumed that the Paleozoic diplopods must not be closely related to modern groups. But, as Kraus pointed out, the modern clas- sification is based on characters seldom pre- served in Paleozoic forms, and there is good reason for believing that at least some mod- ern diplopod groups have ancient roots. Significantly, the conspicuous differences PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON between modern and Paleozoic diplopods relate to mode of life: the large eyes, long bifurcated spines, etc., indicate that the Pa- leozoic forms lived exposed on surfaces, perhaps on plants. The large eyes suggest diurnal habits. The majority of modern Di- plopoda live below the soil surface, at least during the day. Is it an accident that the conspicuous differences between Paleozoic and modern millipedes are precisely those that would result from the elimination of surface-dwelling species by vertebrate pred- ators? Example 2.— Manton (1965) pointed out that the long-legged house centipede, Scu- tigera, appears never to have been adapted for burrowing or for living in shallow crev- ices, as have most modern centipedes. Fur- ther, it and its long-legged relatives have compound eyes. Obviously, the Chilopoda, like the Diplopoda, were not originally bur- rowers; they were adapted for a life on sur- faces where their long legs and compound eyes were functional. Such a mode of life has not been open to them since the Paleo- zoic, and only the hidden burrowers and a few retiring scutigeromorphs have sur- vived, which is again readily explained as a consequence of predation by visually hunt- ing terrestrial predators. Example 3.—The Collembola do not pro- vide compelling data, but the available data are in accord with the hypotheses as pre- sented. They jump, and they are too small to be attractive prey to most vertebrate predators. Jumping does not make sense in confined spaces, and the more deeply “bur- rowing” species usually lose the furcula and become white. This group was obviously adapted originally for a surface life, and this is supported by the elaborate color patterns of many species. Interestingly enough, some earlier Col- lembola were much larger than the species we are familiar with. Matthew (1895) de- scribed a collembolan from the “Little Riv- er Group” of New Brunswick, a deposit identified as either Silurian or Devonian by VOLUME 89, NUMBER 3 Wilmarth (1938). This animal had an ab- domen of “about” seven segments, and a strong “spring,” but it was 20 mm long— large enough to be attractive prey for an insectivorous tetrapod. No Collembola this large have survived. Example 4.—A rather involved body of theory indicates that Symphyla and Diplura were also originally surface-dwellers, but the account 1s too lengthy to develop here. It can be pointed out, though, that their rel- atively long, many-segmented antennae are incongruous with a burrowing life in the soil. Both groups appear to be crawlers-in- small-spaces, and, according to Scheller (1982), the Symphyla are incapable of bur- rowing. Example 5.—Individuals of some species became so small as to be hard to see and perhaps too small to be worth chasing by most vertebrate predators. The Collembola exemplify this response on the ground; the Psocoptera exemplify it on tree trunks. So do the lice orders, although only deriva- tively. The Hemiptera and Homoptera probably also belong in this class, although less obviously. I postulate this: the hemip- teroid precursors became small, so small that their jaws (especially in their early in- stars) could no longer gape widely enough for effective biting of the now relatively larg- er plant cells. It became easier to puncture plant cells and suck out the contents. These are the circumstances that would select for stylet-like mouthparts. It is the obvious pathway to entognathy. This explains the small size of the entognaths (in a functional, not a phyletic sense, to include tardigrades, Symphyla, Collembola, Diplura, Thysa- noptera, and, incipiently, the Psocoptera). Once an entognathous mode of feeding developed, it is conceivable that an insect might retain piercing mouthparts while evolving to a larger body size— providing that stylet feeding led to tapping adequate food sources along the way. It did, and yielded a variety of Homoptera in the Perm- ian and the Heteroptera later on. 397 There are several hints that the Hemip- tera were originally smaller than most are today. They are usually thought to have evolved from a psocopteroid ancestor, and the Psocoptera are small. The Sternorrhyn- cha are small. The hemipteran antennae are comparatively short and usually have few segments, and the tarsi have fewer than the primitive five segments. In Scutigera (Chilopoda), and presum- ably elsewhere, long antennae are important in sensing a leg track and the many-seg- mented tarsi (with claws) are important as automatic grappling devices in species which run too fast to select footholds (Manton, 1952). In very small individuals a secure foothold is usually more advantageous than running speed, and species of small body size have often lost tarsal segments and an- tennal length. Many Hemiptera or Homoptera have reattained a larger body size and have ac- quired additional means for avoiding their predators. The Auchenorrhyncha, for ex- ample, are able to jump, or they live beneath the ground surface (cicadas). Example 6.—The implications for the Pterygota are somewhat different than for primitively wingless tracheates, since the winged adults would conceivably fly to es- cape; their immature stages could not. Ac- cordingly, we should postulate that, 1) ex- tinct pterygote orders characteristically had exposed, terrestrial immatures that were es- pecially susceptible to tetrapod predation, and 2) the immature stages of surviving or- ders will have effective means for avoiding visually-hunting tetrapod predators. Both predictions are fulfilled among Pa- leoptera: the only surviving orders are the Ephemeroptera and Odonata, and both have aquatic immature stages. Some fossils in- dicate that the Paleozoic Ephemeroptera were also aquatic (Hubbard and Kukalova- Peck, 1980). The aquatic habits of these or- ders effectively removed their immature stages from terrestrial reptile predators. But would an aquatic life secure the 1m- 398 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON matures from vertebrate predation? Fish, many of which would eat insects, were al- ready present, and would not the amphib- ians and littoral reptiles pursue insects into the water? The answer is that pterygote insects had two significant advantages: they were small- er, and their adults flew. Flight enabled the adults to colonize smaller, less permanent and more widely scattered bodies of water. Their larvae would be able to utilize food in bodies of water or portions of them not suitable for or even accessible to fish or any vertebrate specialized for an aquatic mode of life. The small size of these insects en- abled more individuals per body of water (for a given productivity), and, hence, rel- atively larger breeding populations and a relatively larger probability of at least some adults finding another body of water. The early aquatic paleopterans retained their evolutionary potential because of the size of their breeding populations and adult mi- grations. While not paleopterous, the Plecoptera also appear to have survived the Paleozoic because their immature stages had become adapted to living in water beyond reach of tetrapod predators. The paleopterous orders that became ex- tinct would be expected to have had terres- trial immature stages. This is suggested by the long ovipositors of many of these species (Carpenter, 1971), that usually correlate with oviposition in soil, various crevices, or plant tissues rather than in water. Fossil 1mma- tures are very scarce for these groups, which is probably a consequence of terrestrial hab- its that seldom placed them in sites suitable for fossilization (Lemche, 1940). No im- matures of these orders have been found yet that exhibit aquatic adaptations, and con- vincing information has been accumulating to show that the immature stages of at least some Palaeodictyoptera and Megasecoptera were terrestrial (Lemche, 1940; Carpenter, 1977; Carpenter and Richardson, 1971; Wootton, 1972). Carpenter and Richardson (1968) men- tion some crucial information: the later in- stars of the Plaeozoic Megasecoptera and Palaeodictyoptera had large, posterolater- ally projecting wing pads. It is these large, developing wings that would cause serious problems in avoiding vertebrate predators. In immatures not yet able to fly, large pads would interfere with burrowing into ground litter or entering crevices to hide. These im- matures must have been easy prey for ar- thropod-eating reptiles; it is not surprising that they became extinct. Carpenter (1977) thought that the Prot- odonata must have been aquatic, apparent- ly because they are so similar to the Odo- nata, as he had mentioned earlier (1947). However, he also mentioned (1947) that August Krogh had stated that the immature stages of the giant meganeurids would have been unable to respire via caudal tracheal or rectal gills. The vertebrate predation hy- potheses developed here, are in accord with Krogh’s statement, and it seems much more likely that the Protodonata were simply the terrestrial branch of the Odonata. They be- came extinct because their immature stages were exposed on the ground where they were vulnerable to vertebrate predators, al- though it may be that their early instars were passed in water. Their near-adult imma- tures may have been sufficiently large to re- sist many tetrapod predators. The Paleoptera did not originate as aquat- ic animals. Both adults and immature stages (Odonata and Ephemeroptera) are abun- dantly supplied with tracheae that belie their terrestrial origins. The idea that wings orig- inated as aquatic adaptations is beset with serious weaknesses, inasmuch as movement in water and in air would impose vastly different demands on lateral flaps. More- over, at least some Paleoptera were clearly not aquatic as immatures, as already men- tioned. The immatures of the Odonata ev- idently became aquatic independently of the Ephemeroptera, since the immatures of the two have developed different respiratory adaptations for their aquatic lives. In general, Ephemeroptera show more VOLUME 89, NUMBER 3 primitive features than Odonata, and their gill covers seem to be serially homologous with wings (see Kukalova-Peck, 1978). However, gill covers may just as easily be adaptations of preexisting lateral flaps (whatever their origin) to an aquatic life as new structures resulting from selection in an aquatic environment. Since there is no very feasible pathway from an epiphytic mode of life to an aquatic one, the Odonata and Ephemeroptera are necessarily interpreted as remnants of lin- eages that originally lived on the ground surface. Their terrestrial relatives are all now extinct. Several endopterygotes (Neuroptera, Me- coptera, Trichoptera, and Coleoptera) ap- pear in Lower Permian deposits (Carpenter, 1977). Odonata appear about the same time. The Ephemeroptera appear in the Carbon- iferous as only a single species, but they became abundant in the Permian. The Palaeodictyoptera show the reverse pattern of many species in the Carboniferous, but only a sparse representation in the Permian (Carpenter, 1976). These data suggest that vertebrate predation on the ground became critical about the time of the Carboniferous- Permian transition, that is, about the time that the reptiles began their first explosive adaptive radiation. The data are also con- sistent with the idea that some species, in this case the few Permian Palaeodictyop- tera, may have survived in arboreal refugia until reptiles became efficient climbers. Example 7.—Endopterygotes evolved from species in which the immature stages had entered the soil litter. Projecting wing pads impede back-and-forth maneuverings in subsurface environments, and were se- lected against in burrowing pterygote im- matures; the endopterygote condition is thus a result of entry upon a burrowing life (Hin- ton, 1948, 1977). It is also a result of selec- tion by visually hunting vertebrate preda- tors (the two ideas are compatible). Tetrapod predators eliminated groups that had 1m- mature stages exposed on the surface of the ground. 399 Hennig (1981, p. 278) objected to Hin- ton’s views, and asserted that thysanuroid or campodeiform larvae were primitive for endopterygotes, and that these were not suited for burrowing in firm substrates. Hence, the endopterygote condition arose for reasons other than as an adaptation for burrowing life: it may have contributed to the subsequent evolution of endopterygotes but not to their origin. However, it 1s un- likely that Hinton implied anything more in his general statement than that the en- dopterygote condition resulted from selec- tion in a subsurface environment. Obvious- ly, the first larvae could not have been very different from the immatures of other in- sects. They did not begin as burrowers in plant tissues or firm soil; they merely en- tered spaces in the ground litter. A thysanu- riform larva, as occurs in many neuropter- oid insects, is a reasonable model for an early endopterygote larva, as Hennig noted and as Hinton probably implied. Because about 88% of all insects are en- dopterygotes (Hinton, 1977) it has often been assumed that the internalization of the developing wings in endopterygote larvae preadapted them to invade environments not available to immatures with external wing pads (see Hinton, 1948, 1963, 1977; Hennig, 1981; Clements, 1968). For the first time the distributive and reproductive adult stage was completely dissociated from the feeding larval stages. According to this view, the adult-larval dissociation conferred ad- vantages in utilizing a new, hidden type of food from which the less-suited exoptery- gote immatures were excluded (Hinton, 1977). This implies that selection was ba- sically for an ability to obtain hidden, larval foods by burrowing, while the adults could still disperse. Once larvae had gained effec- tive access to such foods they underwent a major adaptive radiation on them. The hypotheses developed here require that these ideas be modified. Visually hunt- ing, terrestrial vertebrate predators selected severely against species that had surface- dwelling immature stages. Endopterygote 400 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON larvae were not affected, since they lived out of sight under litter, and their adults prob- ably spent much of their time up on vege- tation out of reach of early tetrapod pred- ators. The ensuing adaptive radiation of endo- pterygotes occurred in groups having larvae that lived in litter—to which they were lim- ited during the day by the larger tetrapod predators. It did not occur because larvae gained sudden access to a previously un- tapped, major food resource. That would have been reflected in a sudden “explosion” of the endopterygotes. The radiations oc- curred over a longer period of time because the increasing complexity of land environ- ments provided more hidden “‘niches”’ into which endopterygote larvae could move from time to time. The adaptive radiations of the angiosperms appear to have resulted in the greatest complexity and the most niches for endopterygote larvae. There is no reason to assume that there were fewer ex- posed niches for exopterygotes. Exoptery- gotes have been unable to radiate freely in exposed niches since the end of the Paleo- zoic because of the ever-present vertebrate predators. The divergence of the Endopterygota into the neuropteroid and mecopteroid orders seems to have been initiated by compara- tively simple specializations for two diver- gent modes of larval life. Mecopteroid lar- vae found food in the soil and fed there day or night out of sight of potential vertebrate predators. The neuropteroids hid in soil lit- ter during the day, but emerged to forage at night. The nocturnal wanderings of some early neuropteroid larvae led them into crevices beneath bark where they found an ample supply of fungi. Their adults could enter the same crevices, and these became the Coleoptera. The longer antennae and legs of the larvae of primitive Neuroptera (sensu lato) and Coleoptera at first appear somewhat incon- gruous with the invaginated wing pads and lack of compound eyes (the first suggest an active life on surfaces, the second a burrow- ing mode of life). These are, however, just the features required for insects that burrow during the day but forage over surfaces at night. The wandering habit also explains how the Neuroptera got to their extremely diverse larval habitats (in water, soil, under bark, stones, on plants). Mecopteroid larvae did not leave the soil to wander at night, and became more worm-like as befits a bur- rowing mode of life and, accordingly, de- veloped very short antennae and legs. The mecopteroid adults could not remain ex- posed near their underground larvae be- cause of vertebrate predators, nor could they specialize like their larvae for burrowing, since that would require sacrificing the adult adaptations for flight and dispersal. In this sense vertebrate predators coerced the adult- larval dissociation of endopterygotes. More accurately, they ate the comparatively vul- nerable surface-dwellers out of existance, leaving behind larval burrowers that had arboreal and aerial adults. Before the advent of reptiles, such species would probably have appeared as minor evolutionary side issues. Example 8.—Neoptera are able to flex their wings backward over or alongside their abdomens when not flying. The flexing mechanism originated very early in the evo- lution of insects. It was present among the first known winged insects of the Carbon- iferous (Carpenter, 1977; Kukalova-Peck, 1978: Rolfe, 1980). That vertebrate pred- ators were involved in selecting for the wing- flexing mechanism is at least implied by statements that it enabled winged adults to enter crevices and hide (Carpenter, 1977; Kukalova-Peck, 1978; and others). Presum- ably, they hid from their larger tetrapod predators. Some authors mention that flexing the wings backwards would facilitate crawling through vegetation with less risk of damage to the wings (Kukalova-Peck, 1978; and others). If so, the greater facility in moving through vegetation might also be related, at least in part, to predator escape. Wootton VOLUME 89, NUMBER 3 (1981) suggested that the ability to flex the wings backwards provided three advantages for Paleozoic insects: it enabled a greater mobility with less risk of wing damage, as just mentioned; it reduced their wind resis- tance and the likelyhood of the insect being blown off vegetation; and it enabled the adults to enter crevices. Neither mayflies nor dragonflies have moved toward a wing-flexing mechanism even though they typically perch in exposed sites that would seem to render them es- pecially vulnerable to wind gusts. Damsel- flies ordinarily operate in less exposed hab- itats where winds are seldom a problem. Yet, it is they that are characterized by an analog of the wing-flexing mechanism. Their thorax is so extremely tilted that the wings extend backwards over the abdomen even though they are only simply folded dorsally from a morphological perspective. Winds would seem to be among the less plausible agents selecting for a wing-flexing mecha- nism, but there is no very obvious way to assess earlier roles of wind selection. There is some evidence for believing that mobility in vegetation was important; ant and termite adults discard or chew off their wings once they have completed their nup- tial flight—that is, after the wings are no longer needed. There are two difficulties to be resolved if neoptery is to be explained as a conse- quence of entering crevices. Difficulty 1. The first Neoptera could fly. If so, for what reason would winged insects that could fly to escape predators resort to hiding in crevices? Theoretically, flight should have been a most effective means of escape, and the large wings would certainly cause major problems in entering crevices. This difficulty results from an uncritical extrapolation from too simple a basis. The early Neoptera were very primitive insects. They had essentially no metamorphosis (Kukalova-Peck, 1978). A distinct meta- morphosis, as characterizes most modern winged insects, may not have developed ful- 401 ly until after the Paleozoic (Kukalova-Peck, 1978). The primitive adults developed only gradually from immature stages from which they probably differed little in either mor- phology or mode of life. In this light, entering crevices would pre- sumably have been one of the most im- mediately available means for escaping ver- tebrate predators. It would not require new behavior patterns, since even before wings arthropods must have entered crevices reg- ularly to avoid desiccation during dry pe- riods. The arthropods could remain in the crevices during the day, but emerge to feed at night when vertebrates would be unable to see and pursue them effectively. Since their development was so simple, basic escape patterns would necessarily be carried from instar to instar. But, since there was no significant metamorphosis, the es- cape patterns of the immatures would al- most certainly be carried into adulthood. This is consistent with Kukalova-Peck’s idea (1978) that neoptery, among other events, developed first in the immature stages, al- though, strictly speaking, the changes would have developed simultaneously in both im- mature and adult stages. It is not unreason- able that the adults should escape via the same means as their immatures, particular- ly in primitive insects. It is still the rule in the jumping Collembola, Saltatoria, or Ho- moptera, in the cryptically colored orthop- teroid insects, or in Blattodea—essentially all insects that have not developed widely divergent adult and immature modes of life. Difficulty 2. Neoptera appear to have dif- ferentiated along two incompatible lines, with but few persisting, undifferentiated rel- icts. The Blattodea exemplify one type. These are fitted characteristically, and no doubt primitively, for a life in crevices, such as under the bark of trees. In such confined spaces the wings are subject to abrasion, and the front wings became thickened, pro- tective covers for the hind wings. The prothorax is also characteristically and primitively rather loosely joined to the 402 mesothorax in these insects, probably be- cause in shifting the burden of flight to the metathorax there was no longer any great need for a large, fixed phragma between the pro- and mesothorax for the attachment of the mesothoracic, indirect flight muscles. A second type of Neoptera relies primar- ily on the mesothorax rather than the meta- thorax for flight. The prothorax is always more or less fused with the mesothorax in these groups, because in this case a well- developed phragma between these segments is essential for the operation of the indirect flight muscles. Ifa pair of wings is modified, it is the hind pair, principally by reduction, as in Diptera or Hymenoptera. But these Neoptera never show any hint of thickened mesothoracic wings, nor does their behav- ior suggest that they ever lived in crevices. Thus the difficulty: “‘crevice-entering” behavior that resulted in thickened front wings seems to have affected the evolution of only some Neoptera and only after the wing-flexing mechanism had already origi- nated, apparently for reasons other than en- tering crevices. The hypothesis consistent with the most data is that the wing-flexing mechanism first evolved as a facet of “‘crevice-settling”’ be- havior, in which the winged adult simply settled into open crevices in bark or on the ground. Since they were only partly con- cealed, immobility during the day would be almost essential to the behavior. Even if crevice-settling insects were not completely concealed from view, it would not be easy for a much larger vertebrate to remove them from small crevices. Perhaps more crucial, especially on trees, settling into crevices and flexing the wings back along the abdomen eliminated the silhouette that otherwise would so readily disclose the insect’s posi- tion to potential tetrapod predators. One reason for thinking that crevice-set- tling rather than running through vegetation was the more significant feature of early winged insects is that crevice-inhabiting in- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON sects appear to have supported major adap- tive radiations among beaked birds and per- haps the pterodactyls earlier. The loss of teeth and development of a relatively long beak are adaptations for fishing insects out of crevices. The beak developed in winged animals, as would be expected: the ability to move from tree to tree (or from stand to stand) would be essential for survival, since single trunk surfaces would not ordinarily support sufficiently large insect populations for predators as large as birds or pterodac- tyls. Insectivores, such as bats, shrews, and lizards, that do not pick their prey from crevices, have not developed beaks. THE PROBLEM WITH FLYING DURING THE DAY By and large the neopterous orders seem primitively to have had nocturnal adults. Many adults are still nocturnal. In general, these are singularly reluctant to fly to escape diurnal predators. Modern representatives most often rely primarily on physical con- cealment, cryptic coloration, or means oth- er than flight to escape diurnal predators, as in moths or Trichoptera that are concealed by their coloration on tree bark, or in Blat- todea, Coleoptera or Dermaptera that seem primitively to have hidden in crevices with their immatures. This reluctance to fly (during the day) is understandably adaptive in modern envi- ronments. To fly out in the open in the day is to be conspicuous and to risk capture by birds. However, the basic nocturnal pat- terns for all these orders appear to be very ancient; they were laid down long before birds evolved. This is not so puzzling as it might seem. While not often considered in this regard, modern Odonata are significant aerial pred- ators. In the tropics they are evidently more effective than birds in excluding Ephemer- optera from diurnal activity over the streams in which they breed (Edmunds and Ed- munds, 1980). Presumably, the Protodo- VOLUME 89, NUMBER 3 nata earlier augmented the depredations of the Odonata. Insects flying or swarming during the day would be conspicuous targets for such aerial predators. This surely ac- counts for the early avoidance of diurnal flight in many insects. Of course, flight is also costly in terms of energy, an added reason to depend first on means other than energy-expensive flight for escaping predators. Nevertheless, the Odo- nata, Ephemeroptera and Diptera seem to have been fundamentally diurnal and ae- rially oriented. The Odonata and Ephemeroptera have short, setaceous antennae. They do not feel the substrate with them as they move about at night (they do not seem to walk much at all). Diptera have also repeatedly developed comparable antennae (stylate and aristate types), and they do not use them to test the substrate like Blattodea or other active noc- turnal insects. Most Diptera also fly readily to escape during the day. Many Ephemeroptera swarm at dusk or at night, although, like the diptera, they do so at visual markers, which betrays their earlier diurnal orientation. The nocturnal habits of tropical species at sites remote from breeding sources (Edmunds and Edmunds, 1980) are probably the extreme effects of severe selection pressure on primitively diurnal insects. They have obviously not been able to cope well as adults, and have survived mainly because of advantages their immature stages developed in their long his- tory in aquatic environments. Their lack of a wing-flexing mechanism prevents practi- cal crevice-entering behavior, and it reduces the effectiveness of cryptic coloration be- cause of their conspicuous silhouette. As developed elsewhere (Downes and Dahlem, 1987), the labellum of Diptera gave flies early access to sugar fuels in the thin, dried films of Homoptera honeydews. Dip- tera alone among insects had consistent ef- fective access to naturally occurring sugars until flowers appeared in the Cretaceous. 403 Thus, they almost alone among insects could afford the high cost of flight, or, perhaps, the cost of flight swift enough to evade early odonate predators. The early endopterygote adults probably spent much of their lives on trees. As rep- tiles became efficient climbers, the endo- pterygotes could respond to this new threat in several ways. They might become very small so as to become unattractive as prey. This is not a practical response for endo- pterygotes, since moving from their larval life in litter or soil to an adult life on tree trunks would present major transportation difficulties for such small insects. Only the exopterygote Psocoptera, in which both im- matures and adults live together on tree trunks, appear to have pursued this option. Visually hunting vertebrates must often have selected continuously for another re- sponse — better cryptic coloration—as seems to have developed in Trichoptera or Lepi- doptera. Another possibility would be for the adults to move out to the tips of branch- es where the heavier vertebrates would have difficulty in pursuing them. The Diptera seem to have taken this tactic, and it is one that would be more likely to place them on leaf surfaces bearing honeydew than the al- ternative of hiding in crevices on trunks. Crevice-settling would not be possible at branch tips, and this may have contributed to the dipteran tendency to remain basically aerial in their orientation. CONCLUSIONS Vertebrate predators brought about the extinction of several major arthropod types near the close of the Paleozoic. They were major shapers of the conspicuous adaptive features of every major surviving terrestrial arthropod group. They indirectly account for the adaptive radiation of Endopterygota, simply because they inhibited the adaptive radiations of related groups having com- paratively exposed immature stages. For adults, flight would seem to be a first 404 choice and the most effective means of es- caping tetrapod predators. It may have been an effective escape in the earliest Pterygota, but by the Permian winged insects that flew to escape tetrapod predators risked capture by Odonata or Protodonata. For this reason most adult Pterygota opted for various al- ternatives to flight as their primary means for escaping tetrapod predators. In many an escape response of the immatures was car- ried into adulthood, and adults and im- matures pursued similar lives in the same concealed places. The Neoptera simply continued a very elementary escape pattern of their 1mma- tures; they settled into open crevices. The ability to flex their wings backwards enabled them to enter crevices, and flexing served the larger immatures as well as the adults. The flexing mechanism also eliminated the insect’s characteristic silhouette that so readily gave away its position. Elytra and thickened front wings developed after some winged Neoptera joined their immature stages in larger crevices out of sight of tet- rapod predators, at least during the day. Odonate adults continued to escape tet- rapod predators by flying: but they were the aerial predators, and they have remained strong fliers. Because they could not flex their wings, the Ephemeroptera adults could not settle into or enter crevices as Neoptera, nor could they join their aquatic immatures. Flight seems to have remained as their prin- ciple means of escape, but they have sur- vived mainly on the strengths of their im- mature stages in aquatic environments. The Diptera are the third surviving order in which the adults have retained an essen- tually aerial orientation. They evidently left their original crevice-settling behavior very early in their evolution to assume a more thoroughly aerial life as they gained access to the sugar-fuels of Homoptera honeydew. They do not seem to be derived from stock that had nocturnal adults. Their early access to fuels enabled a faster flight, and their hal- teres probably conferred an agility that al- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON lowed them to fly more freely during the day with less risk of capture than in other Pterygota. Efficient modern predators, such as birds or Odonata, have probably selected for the nocturnal or crepuscular habits of certain groups, although many “lower” Diptera still swarm in the open during the day, at least in temperate regions. Their small size rendered Collembola and Psocoptera unattractive as prey for most tetrapod predators. Only a few of the implications of tetra- pod-predation hypothesis are developed above. Thus far, all fit available observa- tional data. No other hypotheses about the major features of terrestrial arthropod evo- lution currently account for such a wide va- riety of observational data. LITERATURE CITED Alvarez, W., L. W. 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The importance of the nervous system in the evolution of animal flight. Evolution 6: 127-129. Tappan, H. 1982. Extinction or survival: Selectivity PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON and causes of phanerozoic crises. Geol. Soc. Am., Spec. Pap. 190: 265-275. Tillyard, R. J. 1926. The Insects of Australia and New Zealand. Sidney, N. S. W., Angus and Rob- ertson, Ltd. xv + 560 pp. + 44 Pls. Wilmarth, M. G. 1938. Lexicon of geological names of the United States (including Alaska). Part 1, A- L. U.S. Dept. Interior, Geol. Surv. Bull. 896: 1- 1244. Wootton, R. J. 1972. Nymphs of Palaeodictyoptera (Insecta) from the Westphalian of England. Pa- laeontology 15: 662-665. Wootton, R. J. 1981. Palaeozoic insects. Ann. Rev. Entomol. 26: 319-344. PROC. ENTOMOL. SOC. WASH. 89(3), 1987, pp. 407-416 A NEW SPECIES OF FARRODES (EPHEMEROPTERA: LEPTOPHLEBIIDAE: ATALOPHLEBIINAE) FROM SOUTHERN TEXAS JAcK R. DAvis Texas Water Commission, Water Quality Division, P.O. Box 13087, Capitol Station, Austin, Texas 78711. Abstract. — Farrodes texanus n. sp. from southern Texas is described from nymphs and male and female imagos. This report includes the first specific records of the genus from outside the West Indies. The new species resembles the three previously described species of Farrodes in a number of morphological respects. However, the degree of difference in male genitalia structure and the distinctive abdominal color pattern indicate that F. texanus 1s not closely related to West Indian species, but rather represents a separate evolutionary line. The new species is probably derived from a South American ancestor that invaded Texas during the early Pleistocene. Farrodes texanus is relatively uncommon, known thus far only from a few scattered localities in southern Texas, where it apparently is restricted to small, rocky-bottomed, clean-water streams. Collections of benthic macroinverte- brates during water quality investigations conducted since 1981 have resulted in the discovery of a new leptophlebiid mayfly from southern Texas, described herein as Farrodes texanus n. sp. These constitute the first specific records from outside the Ca- ribbean area, the genus formerly having been reported from the West Indian islands of Cuba, Jamaica, and Grenada (W. L. Peters, 1971). METHODS AND MATERIALS Nymphs were collected using a Surber square foot sampler or a d-frame aquatic net, or by handpicking from overturned rocks. All imagos were obtained through rearing of mature nymphs. Rearing was ac- complished in a polystyrene container with water agitated by air stones, and in an air powered, artificial stream modified after Lawson (1982). In each instance, stones from the reference stream were placed in the rear- ing chamber to provide food material for nymphs. Both rearing techniques produced excellent results. Physicochemical data used to character- ize the nymphal habiiat are available for all seven collection localities, as all collecting has been in conjunction with intensive water quality surveys conducted by the Texas Water Commission. Each survey was diur- nal in nature, with dissolved oxygen, tem- perature, pH, and specific conductance measured 3 or 4 times from predawn to sunset. These data, therefore, represent physiochemical extremes for a particular date and locality. Data for other physico- chemical parameters are based on compos- ite samples, with equal aliquots composited on each sampling run. Intensive survey methodologies are described in detail in Buzan (1982). Farrodes texanus Davis, NEw SPECIES Figs. 1-10 Homothraulus sp.: Edmunds, Jensen, and Berner, 1976: 224; Edmunds, 1978: 79. 408 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-7. Farrodes texanus n. sp. 1, Schematic adult fore wing, showing abbreviations of venational ter- minology used in this paper. 2, Same, hind wing. 3, Adult fore wing. 4, Adult hind wing. 5, Hing wing enlarged. 6, Dorsal view, abdominal segments 6-10, male imago. 7, Lateral view, abdominal segments 6 and 7, male imago. VOLUME 89, NUMBER 3 409 Figs. 8,9. Farrodes texanus n. sp. Genitalia of male imago. 8, Ventral view; 9, Lateral view. Farrodes sp.: Edmunds, 1982a: 243, 1982b: 372,984: 97, 125, Male imago (in alcohol).— Length: body, 4.6-5.0 mm; fore wings, 4.6-5.1 mm; ter- minal filament, 8.4-9.7 mm; caudal cerci, 5.1-5.9 mm. Upper portion of eyes buff, lower portion dark gray. Head light yellow- ish-brown; carinae and mouthparts washed with black. Antennae light brown, flagellum pale yellow. Base of ocelli black, remainder white. Thorax yellowish-brown, carinae darker, sutures lighter; prothorax washed with brown, pronotum with dark brown markings and black margins; pleural areas dorsal to base of legs and near anterior base of fore and hind wings washed with brown. Legs pale yellow; coxae suffused with black; apical half of prothoracic femora variably washed with light brown, apex of mesotho- racic femora light brown, and apex of meta- thoracic femora brown. Ratios of segments in fore legs, 0.54:1.00 (1.78 mm):0.07:0.33: 0.25:0.14:0.09. Tarsal claws of a pair dis- similar, one apically hooked, the other ob- tuse, padlike. Wings (Figs. 1-5): longitudi- nal veins of fore and hind wings buff, cross veins paler; membrane of fore and hind wings hyaline, except distal third of cells C and Sc of fore wings translucent, and base of wings buff. Fork of vein MP of fore wings symmetrical; base of vein ICu, of fore wings not attached to vein CuA or CuP. Abdomen (Figs. 6, 7): segments 1-6 semihyaline, seg- ments 7-9 opaque. Each tergum washed with dark reddish-brown, with pale yellow lateral margins, and with five principal areas of pale yellow lightening: (1) a median, ante- riorly-directed, triangular mark gradually increasing in width from tergum | through tergum 6 or 7, then narrowing posteriorly, (2) two anterior, submedian, oval marks, and (3) two anterior, sublateral, rectangular marks. Sterna 1-7 pale yellow; sterna 8 and 9 slightly darker, washed with light brown. Spiracles black, tracheae semihyaline washed variably with black. Genitalia (Figs. 8, 9): apical one-third of penes divided, a ventral appendage arising from apex of each penis lobe; posterolateral margins of styliger plate produced, extending posteriorly dorsal to forceps; basal one-third of forceps seg- ment | light brown, remainder of forceps and penes pale yellow. Caudal filaments semihyaline, white to pale yellow. Female imago (in alcohol).— Length: body, 4.4-4.9 mm; fore wings, 4.7—5.3 mm; 410 Fig. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON roo a , J = Farrodes texanus n. sp., dorsal view, female nymph. VOLUME 89, NUMBER 3 terminal filament, 7.3-8.7 mm; caudal cer- ci, 4.7-6.0 mm. Eyes dark gray. Head buff, carinae and mouthparts washed with black. Antennae light brown. Color of thorax, legs, and wings, and aspects of wing venation, essentially as in male imago. Abdomen: ter- ga 1-9 washed with dark reddish-brown, generally darker than in male imago, espe- cially in anterior segments. Lightening pat- tern of terga relatively similar to that in male imago (Figs. 6, 7), except median triangular marks not increasing gradually in width posteriorly from tergum 1, but rather uni- formly very narrow or absent on terga 1-4, suddenly becoming distinctive on tergum 5, and attaining maximum width on 6 or 7 before narrowing posteriorly. Spiracles and tracheae as in male imago (Fig. 7). Sterna pale yellow, lateral margins of sterna 1-4 or 5 often washed with light brown. Apex of ninth sternum produced, entire, and bluntly rounded. Caudal filaments semihyaline, light brown. Mature nymph (in alcohol) (Fig. 10).— Length: body, 4.3-5.6 mm; terminal fila- ment, 9.5 mm; caudal cerci, 5.2-5.3 mm. Head prognathous; dorsum brown with pale yellow markings as in Fig. 10; venter pale yellow. Antennae 2'2 times maximum length of head, brownish-yellow in color. Thorax light brown with brown and pale yellow markings as in Fig. 10. Legs pale yellow; femora with a brown, transverse, apical band; tibiae with brown, transverse, basal and subapical bands; tarsi with a brown, transverse, basal band. Tarsal claws with 14-18 denticles becoming progressively larger apically, except apical denticle much larger. Abdominal terga washed with brown, with pale yellow markings essentially as de- scribed for imagos; sterna pale yellow. Pos- terolateral spines present on abdominal seg- ments 8 and 9, that on 9 larger. Abdominal gills present on segments 1-7, all alike, each portion of gills long, slender, and gradually tapering to apex. Length-to-width ratio of middle abdominal gills 10:1 to 12:1; gill membrane translucent, gray, tracheae black. 411 Caudal filaments brownish-yellow, with darker annulations at articulations through- out length of filament. Etymology.—The species 1s named for the state of Texas, to which it apparently is en- demic. Type material.— Holotype, 4 imago, Tex- as: Caldwell County, Plum Creek at FM 86 SE Lockhart, 7 September 1984; allotype, 2 imago, same data as for holotype; paratypes, 19 6 imagos, 22 2 imagos, 2 6 subimagos, 2 2 subimagos, same data as for holotype. Ad- ditional paratypes, 9 6 imagos, 9 2 imagos, 1 6 subimago, Texas: Bexar County, Salado Creek at Goliad Road in SE San Antonio, 27 August 1984; 16 nymphs, Texas: Bexar County, Salado Creek at Goliad Road in SE San Antonio, 29 July 1981; 12 nymphs, Texas: Bee County, Aransas River at Ryan Ranch E Papalote, 29 April 1982. All types were collected by J. R. Davis and are in alcohol. The holotype, allotype, 16 6 para- types, 19 2 paratypes, and 12 nymphal para- types are deposited in the collections of the United States National Museum of Natural History; the remaining types are deposited in the collections of Florida A&M Univer- sity. Additional material.—Two 6 imagos, 2 2 imagos, 15 nymphs, Texas: Bexar County, Salado Creek at Southside Lion’s Park in San Antonio, 26 July 1984; 2 nymphs, Tex- as: Hays County, Blanco River at old Austin Highway N San Marcos, 4 June 1985; | nymph, Texas: Travis County, Barton Creek at Camp Craft Road near Rollingwood, 23 May 1985; 2 nymphs, Texas: Travis Coun- ty, Gilleland Creek 3.0 km SE Pflugerville, 27 August 1985. All specimens were col- lected by J. R. Davis and are in alcohol in the author’s personal collection. Remarks.—Farrodes texanus conforms to W. L. Peters’ (1971) generic account, ex- cept for two characteristics of the male ima- go: (1) the fore tibiae are reduced by about 0.5 mm, with femora and tibial segments 1, 4, and 5 ofrelatively greater proportion, and tibial segments 2 and 3 of slightly lesser 412 proportion; and (2) only the apical one-third of the penes is divided, versus the apical two-thirds as previously described for the genus. The new species, Farrodes texanus, 1s dis- tinguishable from the three previously-de- scribed species of Farrodes by the following characters. (1) In the male imago, the size and shape of the ventral appendages arising from the penis lobes are unique, and the penes are only shallowly divided apically (Figs. 8, 9). (2) The abdominal color pattern, which is relatively uniform in males, fe- males, and nymphs, is characteristic (Figs. 6 and 10). (3) In the male and female ima- gos, the spiracles, combined with the vari- ation in wash on the tracheae, are very dis- tinctive (Fig. 7). (4) In the nymphs, the brown banding on the tibiae and tarsi 1s unique (Fig. 10), and the caudal filaments bear distinct annulations at the articulations (Fig. 10). In contrast, the caudal filaments of previously-described species are uni- formly pale. PHYLOGENY Farrodes texanus resembles the three pre- viously-described West Indian species in a number of morphological respects, partic- ularly in the nymphs. However, the new species obviously represents a separate evo- lutionary line with a considerable history of geographical isolation, judging from the de- gree of difference in male genitalia structure and the distinctive abdominal color pattern. The Caribbean and Texas representatives are probably both derived from a South American ancestor. W. L. Peters (1971) stated that the relationships of the West In- dian Leptophlebiidae clearly appear to be with those of Central and South America, and many biotic elements of southern Texas have Neotropical affinities and apparently are of Central or South American ancestry (Blair, 1950). In addition, recent phyloge- netic studies indicate that Farrodes evolved in continental South America (W. L. Peters, 1986). On the basis of Halffter’s (1974) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON model of dispersion, Farrodes probably in- vaded Texas during the early Pleistocene, with dispersal through the Mexican Tran- sition Zone occurring along the eastern coastal plain. DISTRIBUTION The first published account of the genus Farrodes consisted of descriptions of three species from the West Indian islands of Cuba, Jamaica, and Grenada by W. L. Pe- ters (1971). At the time of Peters’ report, no closely-related taxa were known even from circum-Caribbean continental areas. Sub- sequent collecting efforts have demonstrat- ed, however, that Farrodes is widespread throughout tropical areas of continental South America, and the range is now known to extend from Argentina northward to southern Mexico, the West Indies, and southern Texas (W. L. Peters, 1986). Re- ports of leptophlebiids in Edmunds et al. (1976: 224) (as a segregate of Homothrau- lus, from Argentina, Guatemala, and Texas) and Allen and Brusca (1978: 424) (as Thrau- lodes sp. F, from Guatemala and Vera Cruz, Mexico) are now known to be referrable to Farrodes (R. K. Allen and H. M. Savage, personal communication). In addition, specimens of Farrodes from Venezuela and Peru have recently been reared by H. M. Savage and from Panama by R. W. Flowers, and W. L. Peters has recently collected the genus in Trinidad (H. M. Savage and W. L. Peters, personal communication). Farrodes texanus 1S apparently a disjunct, isolated unit of the genus, as years of collecting by R. K. Allen along the east coast of Mexico between Vera Cruz and Texas have failed to produce Farrodes. The invasion of southern Texas by Far- rodes has been promoted by the favorable climate together with the presence of small, rocky-bottomed streams. The genus has not colonized other climatically-suitable U.S. regions such as Florida, in spite of the prox- imity of Caribbean epicenters, probably due to physical habitat limitations imposed by VOLUME 89, NUMBER 3 the predominance of unfavorable, lowland, sandy-bottomed streams (see Berner, 1950). Farrodes texanus is thus far known only from seven scattered localities in southern Texas: two in the San Antonio River basin, two in the Guadalupe River basin, two in the Colorado River basin, and one in the Aransas River basin. Evidently, the genus also occurs in the Trinity River basin in northcentral Texas. As mentioned by Ed- munds et al. (1976: 225), many years ago two adult male leptophlebiids were sub- mitted to Edmunds for identification, with the following collection data: Texas: Dallas County, Elm Fork Trinity River below Car- rollton Dam, 20 November 1949, Louis E. Moore (G. F. Edmunds, Jr., personal com- munication) (i.e. collecting locality #16 of Moore, 1950). The specimens were identi- fied as a probable new genus and returned at the sender’s request. The material sub- sequently has been lost (W. L. Peters, per- sonal communication), so the actual iden- tity will never be known. However, through recollection of the characters, Edmunds be- lieves that the specimens were Farrodes. All F. texanus collection localities, except that on the Aransas River, are proximal to the Balcones Escarpment, the northern boundary of neotropical influence (Blair, 1950). In this region, winter temperatures are ameliorated by warm, moist air moving inland from the Gulf of Mexico, and small, rocky-bottomed streams are numerous. To the north and west, winter temperatures probably are limiting, while to the east, un- favorable sandy-bottomed streams become increasingly predominant. Farrodes texanus is seldom encountered, and is generally not abundant where it oc- curs. Its uncommon occurrence is evl- denced by the failure of Henry (1981) to record it during an extensive survey of the mayflies of the Concho River basin, a single locality record during a similar survey of the Guadalupe River basin by M. S. Peters (1977) (not reported in Peters, but discussed by Edmunds et al., 1976: 225), and by the 413 author’s rare encounters with it during 11 years of collecting throughout Texas. HABITAT AND BIOLOGY Farrodes texanus nymphs have been found only in small, warmwater streams having an abundance of rocky-bottomed, riffle habitat, at an altitudinal range of 24 to 177 m above msl. These streams are al- kaline, well-oxygenated, and relatively clear, with low concentrations of oxygen-de- manding substances, ammonia nitrogen, and nitrate nitrogen (Table 1). Total phosphorus was elevated at three sites located in recov- ery zones below small sewage treatment plant discharges (Plum Creek, Gilleland Creek, Aransas River). The species was ab- sent at a physically suitable site further up- stream on the Aransas River, closer to the effluent source, which suggests a high sen- sitivity to organic pollution. Its occurrence has always been associated with diverse macrobenthic communities (Table 1), a fur- ther indication that its presence reflects clean water and healthy environmental condi- tions. Nymphs typically occur in shallow water near the stream banks, along the edges of rifles where current velocities are reduced and some sediment deposition occurs. They are usually found on the underside of cob- blestones that support small growths of periphytic algae. Specimens were occasion- ally found in swift current in the center of rifles at sites where other leptophlebiids were scarce or absent, which suggests a fac- ultative ability to exploit true erosional mi- crohabitats under conditions of low com- petition. From four to eight other mayfly species occurred sympatrically, and from zero to three other leptophlebiids (Thraulodes gon- zalesi, Choroterpes (Neochoroterpes) mexi- canus, Traverella presidiana). Farrodes texanus was generally spatially segregated from other species. When other leptophle- biids were abundant, they predominated in swift-current areas, with F. texanus restrict- 414 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table |. Biological and physicochemical parameters for water quality sampling stations at which Farrodes texanus n. sp. was collected. Numbers in parentheses indicate the number of determinations comprising the data base. Biological Parameters Community ’ ; Physicochemical Parameters** Diversity F. texanus Relative Locality (H)* Abundance D.O. Temp. pH Aransas R. @ Ryan Ranch 4.04 222/m? 6.8-11.1 21.9-27.2 7.8-8.3 E Papalote (4) (4) (4) Salado Cr. @ Southside Lion’s 4.15 108/m? 4.6-5.0 22.8-24.5 7.1-7.3 Park in San Antonio (4) (4) (4) Salado Cr. @ Goliad Rd. 3.97 544/m 7.1-7.2 22.5-24.1 7.4-7.6 In SE San Antonio (4) (4) (4) Barton Cr. @ Camp Craft Rd. nr ili 5/m? 6.9-9.3 23.9-25.8 8.0-8.1 Rollingwood (4) (4) (4) Plum Cr. @ FM 86 SE shed very common 4.8-5.6 26.3-26.9 7.4-7.7 Lockhart (4) (4) (4) Blanco R. @ old Austin Hwy. th occasional 6.8-8.6 27.8-30.0 8.0-8.2 N San Marcos (4) (4) (4) Gilleland Cr. 3.0 km SE ty occasional 5.1-6.9 24.1-25.9 7.7-8.0 Pflugerville (3) (3) (3) Cumulative ranges 4.6-11.1 21.9-30.0 7.1-8.3 (27) (27) (27) * Calculated according to the equation described by Shannon and Weaver (1963). ** Units for all parameters in milligrams per liter, except temperature (°C), pH (standard units), specific conductance (umhos/cm), and flow (cubic feet per second). *** No diversity index available due to lack of quantitative data. However, qualitative sampling revealed high community diversity in each case. ed to riffle margins. Conversely, when other leptophlebiids were less numerous (as in Sa- lado Creek at Goliad Road) or absent (as in the Aransas River), F. texanus exhibited its greatest abundance and was able to exploit erosional microhabitats. A degree of inter- specific spatial niche overlap did occur at some sites, particularly with 7. gonzalesi which occurred sympatrically at five sites. However, when the two species overlapped, they were seldom found together on the same rock, which seemingly reflects an avoidance mechanism. In addition to previously dis- cussed physiographic and climatic factors, competitive exclusion by other leptophle- biids may also be a factor limiting the range of F. texanus, whose northern and western range limits correspond to the southern and eastern range limits of a number of lepto- phlebiid species. Foregut contents of nymphs from Salado Creek and the Aransas River consisted al- most entirely of detritus and fine sand, along with small numbers of diatoms. Thus, the VOLUME 89, NUMBER 3 415 Table 1. Extended. Physicochemical Parameters** Spec. Cond. BOD, TSS NH,-N NO,-N T-P TDS Flow 2010-2030 0.5 <10 <0.02 0.88 2.19 1120 13 (4) (1) (1) (1) (1) (1) (1) (1) 635-673 1.0 33 0.10 0.87 0.17 366 22 (4) (1) (1) (1) (1) (1) (1) (1) 595-647 1.0 24 0.02 0.89 0.15 352 25 (4) (1) (1) (1) (1) (1) (1) (1) 455-461 1.0 7 0.05 0.04 0.04 260 9 (4) (1) (1) (1) (1) (1) (1) (1) 1109-1124 0.5 30 <0.02 3:72 1.45 670 3 (4) (1) (1) (1) (1) (1) (1) (1) 375-387 1.0 5 <0.02 0.02 0.02 196 37 (4) (1) (1) (1) (1) (1) (1) (1) 12711285 1.5 12 <0.02 0.15 4.10 704 l (3) (1) (1) (1) (1) (1) (1) (1) B75-2030° 105=1:5 < 5-33 <0102-0:10 0.02=3.72> 0102=4.10 196-1120 1-37 (27) (7) (7) (7) (7) (7) (7) (7) species apparently is a detritivore that feeds ACKNOWLEDGMENTS by gathering decaying fine particulate or- ganic matter. In rearing chambers exposed to natural photoperiods, subimagos emerged at dusk and molted at sunrise. Although not ob- served, mating probably occurs in the morning in full sunlight. Such a cycle has been reported for Farrodes hyalinus in Ja- maica (W. L. Peters, 1971). Mature nymphs were collected from April through Septem- ber, indicating that emergence occurs for at least six months of the year. Gratitude is extended to R. K. Allen, H. M. Savage, W. L. Peters, and G. F. Ed- munds, Jr. for providing information per- tinent to this study. Savage, Peters, and Ed- munds also reviewed the manuscript and contributed valuable, constructive criti- cism, and Peters provided loan specimens of the type species for comparative study. LITERATURE CITED Allen, R. K. and R. C. Brusca. 1978. Generic revi- sions of mayfly nymphs II. Thraulodes in North 416 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON and Central America (Leptophlebiidae). Can. Entomol. 110: 413-433. Berner, L. 1950. The mayflies of Florida. Univ. Flor- ida Studies, Biol. Sci. Ser., No. 4. 267 pp. Blair, W. F. 1950. The biotic provinces of Texas. Texas J. Sci. 2: 93-117. Buzan, D. 1982. Intensive survey of Salado Creek, Segment 1910. Rept. No. IS-42, Texas Water Commission, Austin. 34 pp. Edmunds, G. F., Jr. 1978. Ephemeroptera, pp. 57- 80. In R. W. Merritt and K. W. Cummins, eds., An Introduction to the Aquatic Insects of North America. Kendall/Hunt Publ. Co., Dubuque, Iowa. 441 pp. . 1982a. Ephemeroptera, pp. 242-248. Jn S. H. Hurlbert and A. Villalobos-Figueroa, eds., Aquatic Biota of Mexico, Central America and the West Indies. San Diego State Univ., San Diego, California. 1982b. Historical and life history factors in the biogeography of mayflies. Am. Zool. 22: 371- 374. 1984. Ephemeroptera, pp. 94-125. Jn R. W. Merritt and K. W. Cummins, eds., An Introduc- tion to the Aquatic Insects of North America, 2nd edition. Kendall/Hunt Publ. Co., Dubuque, Iowa. 722 pp. Edmunds, G. F., Jr.,S. L. Jensen, and L. Berner. 1976. The mayflies of North and Central America. Univ. Minnesota Press, Minneapolis. 330 pp. Halffter, G. 1974. Elements Anciens de l’Entomo- faune neotropicale: ses implications biogeogra- phiques. Quaest. Entomol. 10: 223-262. Henry, B.C., Jr. 1981. The mayflies (Ephemeroptera) of the Concho River system in central Texas. M.S. thesis, Angelo State University, San Angelo, Tex- as. 54 pp. Lawson, P. W. 1982. A simple air powered pump for laboratory streams. Freshwater Invertebr. Biol. 1: 48-52. Moore, L. E., Jr. 1950. Distribution of mayfly nymphs (Ephemeroptera) in streams of Dallas County, Texas. Field Lab. 18: 103-112. Peters, M. S. 1977. The mayfly nymphs (Insecta: Ephemeroptera) of the Guadalupe River Basin, Texas. M.S. thesis, Southwest Texas State Uni- versity, San Marcos. 84 pp. Peters, W. L. 1971. A revision of the Leptophlebiidae of the West Indies (Ephemeroptera). Smithson. Contrib. Zool., No. 62. 48 pp. . 1986. Origins of the North American Ephem- eroptera fauna, especially the Leptophlebiidae. Can. Entomol. (In press.) Shannon, C. E. and W. Weaver. 1963. The mathe- matical theory of communication. Univ. Illinois Press, Urbana. 117 pp. PROC. ENTOMOL. SOC. WASH. 89(3), 1987, pp. 417-424 WOOD-DECAY FUNGI ASSOCIATED WITH SUBTERRANEAN TERMITES (RHINOTERMITIDAE) IN LOUISIANA DEBORAH A. WALLER, JEFFERY P. LA FAGE, ROBERT L. GILBERTSON, AND MEREDITH BLACKWELL (DAW, JPL) Louisiana Agricultural Experiment Station, Louisiana State University Agricultural Center, Baton Rouge, Louisiana 70803; (RGL) Department of Plant Pa- thology, University of Arizona, Tucson, Arizona 85721; (MB) Department of Botany, Louisiana State University, Baton Rouge, Louisiana 70803. Abstract. —Wood-rotting fungi associated with the subterranean termites Coptotermes formosanus and Reticulitermes spp. in southern Louisiana were examined. Fruiting bodies of thirty basidiomycetes and one ascomycete were collected from termite-infested logs. All of the basidiomycete species caused white rots. These fungi previously have been reported to repel termites. In several cases we found Reticulitermes feeding on basidio- carps, and species of both termite genera were attracted to Ganoderma basidiocarps in choice tests in the laboratory. Although termites often augment their ni- trogen-poor diets by feeding on decayed wood containing fungal mycelia (Sands, 1969), the interactions between termites and wood-rotting basidiomycetes are more complex. Some fungal species attract ter- mites (Esenther et al., 1961), while others reportedly inhibit feeding (Amburgey, 1979). The observation that the foragers of Reticulitermes flavipes (Kollar) make trails to branches decayed by the basidiomycete Gloeophyllum trabeum (Pers.: Fr.) Murr. (Esenther et al., 1961) led to the discovery that extracts of this fungus are attractive to a number of subterranean termite species (Allen et al., 1964). One of the attractive chemicals is apparently identical to the trail pheromone of R. virginicus (Banks) (How- ard et al., 1976). Laboratory studies have since demonstrated that termite response to G. trabeum and other decay species is de- pendent on the fungal strain, the fungal sub- strate, and the bioassay technique used (Becker and Lenz, 1976; Esenther and Beal, 1979). The dynamics of termite-fungal associa- tions in nature are poorly known, and it has been suggested that these associations de- pend upon rot type. White rot basidiomy- cetes catabolize both cellulose and lignin, as do termites (Waller and La Fage, 1987), while brown rots leave a lignin residue (Gil- bertson, 1981). Preliminary field observa- tions suggest that termites avoid wood de- cayed by white rot fungi (Amburgey and Beal, 1977), but several white rot species have been shown to be phagostimulatory in laboratory bioassays (Gilbertson, 1984). The importance of brown and white rot fungi in termite feeding is therefore uncertain, as few natural termite-fungal associations have been identified. In the present study we examined the wood-rotting fungi associated in southern Louisiana with the subterranean termites Coptotermes formosanus Shiraki and Retic- ulitermes spp. Coptotermes formosanus is a pest species introduced into the area ap- proximately forty years ago (La Fage, 1986), and species of Reticulitermes are native. We 418 were interested in comparing fungal asso- ciations with these species in natural habi- tats. Previous studies of termite-fungal inter- actions have focused on termite response to fungal mycelia. During our study we found termites feeding on fruiting bodies of sev- eral species. Therefore we also tested ter- mite attraction to fruiting bodies of several fungal species to determine whether some were more attractive than others. METHODS Termite species.— The Formosan subter- ranean termite Coptotermes formosanus has large colonies numbering several million in- dividuals (Su et al., 1983) that build nests underground or in tree hearts. This species attacks dead wood and live trees (Lai et al., 1983) and feeds on both sound and decayed wood (Smythe and Carter, 1970; Ambur- gey, 1979). Coptotermes is readily identifi- able by its numerous distinct soldiers, which have oval heads with a large anteriorly di- rected fontanelle (Weesner, 1969). Reticulitermes colonies contain several hundred-thousand individuals (Howard et al., 1982) that live underground and forage on sound and decayed dead wood (Smythe and Carter, 1970; Amburgey, 1979), in- cluding dead branches on live trees. Soldiers of the most common Reticulitermes species in Louisiana, R. flavipes and R. virginicus, are difficult to distinguish; species identifi- cations can be made reliably only by using alates. Thus associations were recorded as Reticulitermes sp. for this genus. Because the workers of Coptotermes and Reticuli- termes are extremely similar morphologi- cally, we recorded only “termite workers” if no soldiers could be found. However, C. formosanus soldiers are very numerous and aggressive, while Reticulitermes soldiers are rare (Howard and Haverty, 1981); workers without soldiers were therefore probably Reticulitermes. The dry-wood termite, Kalotermes ap- proximatus Snyder (Kalotermitidae), was PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON found in one collection. This termite has small colonies of several thousand individ- uals, and nests are constructed entirely within dead wood (Weesner, 1965). Field associations with decay fungi.— We collected fungal basidiocarps from termite- infested logs and live trees in Baton Rouge, Lake Charles, and New Orleans, Louisiana. Identifications were made by MB and RLG; in some cases, basidiocarps were deterio- rated, and identifications could be made only to genus. Fungal presence was determined by basidiocarp presence. Because decay iso- lations were not made, additional species may have been undetected. It is not known if fungal mycelia of detected species were viable. Host woods were identified when possible. Bioassays with fungal fruiting bodies. — We tested the response of Coptotermes for- mosanus and Reticulitermes sp. to basidio- carps of Ganoderma lucidum (W. Curt.: Fr.) Karst. (Ganodermataceae), Phellinus gilvus (Schw.) Pat. (Hymenochaetaceae), Hexa- gonia hydnoides (Fr.: Sw.) M. Fidalg. (Po- lyporaceae) and Pycnoporus sanguineus (L.: Fr.) Murr. (Polyporaceae) by using a mod- ification of the technique of Allen et al. (1964). Although laboratory bioassays do not necessarily reveal field preferences, they are important in providing preliminary data for more comprehensive studies. Basidio- carps were collected from logs without ter- mites, and were dried at 60°C for at least 48 h prior to the bioassay. We tested six colonies of both Coptotermes and Reticu- litermes and used two replications per col- ony (only one replication for one of the Re- ticulitermes colonies) for each test. One hundred termites per test were placed in a 100 x 15 mm petri dish filled with 2% Difco Bactoagar to maintain a high-humidity en- vironment. One 0.1 g piece of each basid- iocarp was arranged equidistant from the other species in the petri dish. The number of termites in a dish that were on each basid- iocarp was recorded at five minutes, 24 hours, and six days after set-up. Data were VOLUME 89, NUMBER 3 analyzed for each time interval with anal- ysis of variance (SAS GLM procedure, SAS Institute, Cary, North Carolina). RESULTS Termite-fungal associations.— We col- lected fruiting bodies of one ascomycete ge- nus and thirty species of basidiomycetes comprising nine families and two orders (Table 1). All of the basidiomycetes col- lected cause white rots. In several cases, dif- ferent fungal species were found in the same log, so that not all decays were directly as- sociated with termites. Coptotermes and Reticulitermes were never found together. In several cases, Coptotermes and Retic- ulitermes attacked wood decayed by the same species of fungi. Hyphoderma sp., Phellinus gilvus, Schizopora paradoxa, Ganoderma lucidum, and Stereum ostrea were associated with both Coptotermes and Reticulitermes, and Kalotermes was found with Reticulitermes in a log decayed by Phellinus gilvus. We observed Reticuli- termes feeding on basidiocarps of Hexa- gonia hydnoides, Inonotus cuticularis, and Ganoderma lucidum, indicating that fruit- ing bodies as well as decayed wood were phagostimulatory. Bioassays with fungal fruiting bodies.— The results for C. formosanus response to the fruiting bodies are shown in Table 2. At the five-minute check, significantly more termites were on the Phellinus gilvus basid- iocarp (P < .001). After 24 h, there was a significant difference in the number of ter- mites on the different basidiocarps (P < .05), with slightly more termites on P. gi/vus and Ganoderma lucidum than the others. At six days, there were also significant differences in attraction of the different species (P < .02), again with slightly more termites on basidiocarps of P. gilvus and G. lucidum. There were significant differences among colonies after 5 min (P < .001) and 24 h (P < .05), but not after six days (P > .05). Table 3 lists the results for Reticulitermes response to the basidiocarps. At five min- 419 utes, there were no significant differences among termites on the different basidio- carps (P > .05). Significantly more termites were on the G. /ucidum basidiocarp after 24 h (P < .001) and also after six days (P < .001). There were significant differences in response among colonies at the six-day check (= 7001). DISCUSSION Most fungal species associated with ter- mites in our study were white rot basidio- mycetes, demonstrating that Coptotermes and Reticulitermes do not avoid these species in nature. Whether white rots ac- tually attracted termites or were incidental to termite attack on the host wood remains to be determined. We observed no brown rot fungi in this study, possibly because brown rot fungi pre- fer softwood hosts (Gilbertson, 1981), and we sampled primarily hardwoods. In ad- dition, white rot fungi outnumber brown rot fungi (Gilbertson, 1980), so that we were less likely to encounter brown-rotted wood. Field observations of subterranean termites associated with brown rots have been lim- ited to a few accounts (Esenther et al., 1961; Williams, 1965). Kovoor (1964) reported that wood de- cayed by the white rot fungus Ganoderma applanatum (Pers.) Pat. 1s toxic to Micro- cerotermes (Termitidae). In our study, we did not encounter G. applanatum, but both Coptotermes and Reticulitermes were col- lected from wood decayed by G. /ucidum, and basidiocarps of this species were at- tractive to both genera in laboratory tests. It is clear that detailed information on ter- mite-fungal associations in nature must be gathered before we can make generaliza- tions about the effects of different decay species on termite feeding. Although subterranean termites can sur- vive on sound wood of some species (Smythe and Carter, 1970; Mannesmann, 1973), my- cophagy may be important to natural host selection. Almost all of the subterranean 420 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Fungi associated with termites in southern Louisiana. W = white rot, L = live tree, D = dead wood, R = Reticulitermes spp., C = Coptotermes fomosanus, K = Kalotermes approximatus, t = termite workers not associated with soldiers. Number of fungus-termite associations found is indicated in parentheses. Collection sites were near Baton Rouge (BR), Lake Charles (LC), Livingston Parish (LV) or New Orleans (NO). Fungal Species Rot Host Wood (L/D) Site Termite Basidiomycetes Tremellales Auriculariaceae Auricularia polytricha (Mont.) Sacc. W Celtis laevigata Willd. (L) NO (E() W Quercus shumardii Buckly (D) NO C(1) Tremellaceae Clitocybe tabescens (Scop:Fr.) Bres. W pine (D) LE R (1) W tree trunk (D) (only mycelium) IW: @G@) Aphyllophorales Auriscalpiaceae Gloeodontia discolor (Berk. et Curt.) Boid. W tree trunk (D) BR ED) Corticiaceae Bjerkandera adusta (Willd.:Fr.) Karst. W hardwood (D) BR t (1) Botryobasidium sp. W pine (D) LV R (1) Confertobasidium olivaceoalbum (Bourd. and Galz.) Jul. W wood (D) BR R (1) Hyphoderma setigerum (Fr.) Donk W wood (D) BR R (1) Hyphoderma sp. W wood (D) LEE ea) W pine (D) LV R (1) Phlebia rufa PersEr W wood (D) BR t(1) Schizopora paradoxa (Fr.) Donk W wood (D) EE @@) W wood (D) NO R (1) W wood BR t (2) W hardwood (D) LE R (1) W wood NO t (2) W hardwood (D) BR t (1) W wood (D) BR R (1) W Liquidambar styraciflua L. (D) BR R (1) Scopuloides hydnoides (Cke. and Massee in Cke.) Hjortst and Ryv. W wood (D) BR R (1) Steccherinum sp. W wood BR t(1) Trechispora farinaceae (Pers.:Fr.) Liberta W wood NO R (1) Ganodermataceae Ganoderma lucidum (W. Curt.:Fr.) Karst. W Quercus virginiana L. (L) NO (EC) W Q. virginiana L. (D) IKE R (1) W wood (D) NO R (1) W Acer rubrum L. (L) NO C(1) W A. rubrum (L) NO tah) VOLUME 89, NUMBER 3 421 Table 1. Continued. Fungal Species Rot Host Wood (L/D) Site Termite Hymenochaetaceae Tnonotus cuticularis (Bull.:Fr.) Karst. W Quercus niger L. (L) BR R (1) Phellinus gilvus (Schw.) Pat. W hardwood (D) NO K (1) W hardwood (D) IW“ R (1) W hardwood (D) LV R (1) W hardwood (D) NO R (1) WwW Quercus falcata var. pogodifolia (L) Le t (1) W wood BR t (1) Phellinus robustus (Karst.) Bourd. et Galz. W Fraxinus berlanderiana A.DC. (L) NO Ga) Phellinus sp. W hardwood (D) BR R (1) Lachnocladiaceae Asterostroma sp. W Salix sp. (D) NO R (1) Asterostroma-like W Acer rubrum L. (L) NO @a) Polyporaceae Coriolus versicolor (L.:Fr.) Murr. W tree trunk (D) NO R (2) Fomes fomentarius (L. ex Fries) Kickx W hardwood (D) NO C (1) Hexagonia hydnoides (Fr.:Sw.) M. Fidalg. W Quercus shumardii (Buckly) NO C(1) Nigroporus vinosus (Berk.) Murr. W pine (D) LE R (1) Perenniporia sp. Murr W hardwood (D) Le t(1) WwW wood (D) BR t(1) Pycnoporus sanguineus (L.:Fr.) Murr W hardwood (D) EG t (1) Trametes versicolor (L.:Fr.) Pilat W wood (D) LE C(1) Trichaptum sector (Ehrenb.:Fr.) W wood (D) BR R (1) Stereaceae Stereum hirsutum (Willd.) Fr. W hardwood (D) BR R (1) WwW wood (D) ILe (@(0) Stereum ostrea (Blume et Nees:Fr.) Fr. W hardwood (D) EE C (2) W wood (D) NO R (3) W Quercus shumardii (Buckly) (D) BR C (1) Ascomycetes Hypoxylon sp. hardwood (D) Le R (1) hardwood (D) BR R (1) hardwood (D) BR t (1) wood (D) NO t (1) pS 22 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 2. Mean (+ SD) of numbers of Coptotermes formosanus workers on fungal basidiocarps after five min, 24 h and six d. Basidiocarps were presented in a four-way choice test to 100 termite workers. Two replications of six colonies were tested. Time Interval Phellinus gilvus Ganoderma lucidum Pycnoporus sanguineus Hexagonia hydnoides 5 min SOLES os! OF6r== = NOP O10F2=70'02 053-2) OFS? 24h fase 2228 13208 O:7 =, 0:42 10722 126% 6d 2 AGEN 98 RAE tee Dale 0:20:57 OF 27240552 P Value 5 min 24h 6d Colony .0009 .0472 .0974 Basidiocarp .0008 .0486 .0149 Colony = basidiocarp .0013 0102 .4817 ' Means followed by the same letter in rows are not significantly different at P < .05, Tukey’s studentized range test. termites we have found in Louisiana were in rotted wood. Becker and Lenz (1976) found that several soft rot and brown rot fungi are attractive to Heterotermes indicola (Wasmann), Reticulitermes flavipes (Kol- lar), Coptotermes amanii (Sjostedt), and Nasutitermes nigriceps (Haldemann), and Hendee (1935) determined that the damp- wood termite, Zootermopsis angusticollis (Hagen), survives better on decayed than on sound wood. Decayed wood may be favor- able for several reasons, including enhanced nutrient content (La Fage and Nutting, 1978), decreased density (Wilcox, 1978), and detoxified wood allelochemicals (Williams, 1965; Martin, 1979). Some decays can be critical to a termite’s ability to use a host wood. Williams (1965) found that Coptotermes niger Snyder can- not feed on one of its primary hosts, Pinus caribaea Morelet, in Belize unless the heart- wood is first decayed by the brown rot fun- gus Lentinus pallidus Berk. et Curt. Simi- larly, C. formosanus cannot eat sound heartwood of baldcypress, Taxodium dis- tichum (L.) Rich (Carter et al., 1981), al- though this species is an important Copto- termes host in Louisiana. However, Coptotermes survived well on baldcypress heartwood decayed by the white rot fungus Rigidoporus sp. and an unidentified deuter- omycete, (Waller and La Fage, unpublished Table 3. Mean (+ SD) of numbers of Reticulitermes sp. workers on fungal basidiocarps after five min, 24 h and six d. Basidiocarps were presented in a four-way choice test to 100 termite workers. Two replications of five colonies, and one of a sixth colony, were tested. Time Interval Phellinus gilvus Ganoderma lucidum Pycnoporus sanguineus Hexagonia hydnoides 5 min Or4 St 105524 OPE 0233 0:07=:0,03 0.1 == 1023 24h Sree 162 8.4 + 6.63 O:3ie 105° 0.4 + 0.5° 6d Osi ORSe WAS ace 40. Onl 2.073" 0:6) 25 1822 P Value 5 min 24h 6d Colony 5057 .0624 .0003 Basidiocarp 1219 .0001 .0001 Colony x basidiocarp OF 0246 .0001 ' Means followed by the same letter in rows are not significantly different at P < .05, Tukey’s studentized range test. VOLUME 89, NUMBER 3 observations). Similarly, Perry et al. (1985) found that termite attack on heartwood of Eucalyptus and Pinus in Australia is sec- ondary to fungal infection, which in turn is always associated with fire damage. Depen- dence on fungal decay for wood use there- fore may be widespread in termites. Further work may reveal the full extent of the importance of fungal decay in termite biology. Collins (1980) suggested that rhinotermitids are limited to mesic regions of Malaysia because they require decay fun- gi in their diets. More information on ter- mite distributions and natural food prefer- ences will help us understand the roles of these important detritivores in nature. ACKNOWLEDGMENTS We thank T. Sparks for making helpful comments on the manuscript and the Lou- isiana Pest Control Association for provid- ing financial support. This paper was ap- proved for publication by the Director of the Louisiana Agricultural Experiment Sta- tion as manuscript number 86-17-0261. LITERATURE CITED Allen, T. C., R. V. Smythe, and H. C. Coppel. 1964. Response of twenty-one termite species to aqueous extracts of wood invaded by the fungus Lenzites trabea Pers. ex Fr. J. Econ. Entomol. 57: 1009- 1010. Amburgey, T. L. 1979. Review and checklist of the literature on interactions between wood-inhabit- ing fungi and subterranean termites: 1960-1978. Sociobiology 4: 279-296. Amburgey, T. L. and R. H. Beal. 1977. White rot inhibits termite attack. Sociobiology 3: 35-38. Becker, von G. and M. Lenz. 1976. Einflus von Mo- derfaulepilzen in Holz auf Frastatigkeit, Galerie- bau und Entwicklung einiger Termiten-Arten. Z. ang. Entomol. 80: 232-261. Carter, F. L., J. K. Mauldin, and N. M. Rich. 1981. Protozoan populations of Coptotermes formosa- nus Shiraki exposed to heartwood samples of 21 American species. Mat. und Org. 16: 29-38. Collins, N. M. 1980. The effect of logging on termite (Isoptera) diversity and decomposition processes in lowland dipterocarp forests. Jn J. I. Furtado, ed., Tropical Ecology and Development. Inter- national Society of Tropical Ecology, Kuala Lum- pur. 423 Esenther, G. R. and R. H. Beal. 1979. Termite con- trol: Decayed wood bait. Sociobiology 4: 215-222. Esenther, G. R., T. C. Allen, J. E. Casida, and R. D. Schenefelt. 1961. Termite attractant from fun- gus-infected wood. Science 134: 50. Gilbertson, R. L. 1980. Wood-rotting fungi of North America. Mycologia 72: 1-49. 1981. North American wood-rotting fungi that cause brown rots. Mycotaxon 2: 372-416. 1984. Relationships between insects and wood-rotting basidiomycetes Jn Q. Wheeler and M. Blackwell, eds., Fungus-Insect Relationships: Perspectives in Ecology and Evolution. Columbia Univ. Press, New York. 514 pp. Hendee, E. C. 1935. The role of fungi in the diet of the common damp-wood termite, Zootermopsis angusticollis. Hilgardia 10: 499-525. Howard, R. W. and M. I. Haverty. 1981. Seasonal variation in caste proportions of field colonies of Reticulitermes flavipes (Kollar). Environ. Ento- mol. 10: 546-549. Howard, R. W., F. Matsumura, and H. C. Coppel. 1976. Trail-following pheromones of the Rhino- termitidae: Approaches to their authentication and specificity. J. Chem. Ecol. 2: 147-166. Howard, R. W., S. C. Jones, J. K. Mauldin, and R. H. Beal. 1982. Abundance, distribution, and colony size estimates for Reticulitermes spp. (Isoptera: Rhinotermitidae) in southern Mississippi. Envi- ron. Entomol. 11: 1290-1293. Kovoor, J. 1964. Modifications chimiques provo- quees par une termitide Microcerotermes edenta- tus dans du bois de peuplier sain or partiellement degrade par des champignons. Bull. Biol. France Belgique 98: 491-509. La Fage, J. P. 1986. Practical considerations of the Formosan subterranean termite in Louisiana: A 30-year problem. A paper prepared for the Sym- posium: “Biology, Ecology and Control of the For- mosan Subterranean Termite,” Pacific Branch of the Entomological Society of America, Honolulu, Hawaii, 1985. La Fage, J. P. and W. L. Nutting. 1978. Nutrient dynamics of termites. Jn M. V. Brian, ed., Pro- duction Ecology of Ants and Termites. Cambridge Univ. Press, Cambridge, U.K. ai PAY. Me lamashing.. even atess Nema SU aK Fujii, and R. H. Ebesu. 1983. Living plants in Hawaii attacked by Coptotermes formosanus. Proc. Haw. Entomol. Soc. 24: 283-286. Mannesmann, R. 1973. Comparison of twenty-one commercial wood species from North America in relation to feeding rates of the Formosan termite, Coptotermes fomosanus Shiraki. Mater. Org. 8: 105-109. Martin, M. M. 1979. Biochemical implications of insect mycophagy. Biol. Rev. 54: 1-21. 424 Perry, D. H., M. Lenz, and J. A. L. Watson. 1985. Relationships between fire, fungal rots and termite damage in Australian forest trees. Aust. For. 48: 46-53. Sands, W. A. 1969. The association of termites and fungi, pp. 495-524. Jn K. Krishna and F. M. Wees- ner, eds., Biology of Termites. Vol. 1. Academic Press, New York. Smythe, R. V. and F. L. Carter. 1970. Feeding re- sponses to sound wood by Coptotermes formosa- nus, Reticulitermes flavipes, and R. virginicus (Isoptera: Rhinotermitidae). Ann. Entomol. Soc. Am. 63: 841-847. Su, N., J. P. La Fage, and G. R. Esenther. 1983. Ef- fects of a dye, Sudan Red 7 B, on the Formosan subterranean termite, Coptotermes formosanus Shiraki (Isoptera: Rhinotermitidae). Mat. Org. 18: 127-133. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Waller, D. A. and J. P. La Fage. 1987. Nutritional ecology of termites. Pp. 487-532. In F. Slansky, Jr. and R. Rodriguez, eds., The Nutritional Ecol- ogy of Insects, Mites and Spiders. Wiley, New York. Weesner, F. M. 1965. The termites of the United States. Handbook of the National Pest Control Association. New Jersey. 1969. External anatomy, pp. 19-47. In K. Krishna and F. M. Weesner, eds., Biology of Ter- mites. Vol. 1. Academic Press, New York. Wilcox, W. W. 1978. Review of literature on the effects of early stages of decay on wood strength. Wood Fiber 9: 252-257. Williams, R. M. C. 1965. Infestation of Pinus cari- baea by the termite Coptotermes niger Snyder. Proc. 12th Int. Congr. Entomol., Lond., pp. 675- 676. PROC. ENTOMOL. SOC. WASH. 89(3), 1987, pp. 425-439 TAXONOMIC OBSERVATIONS ON UNITED STATES TEPHRITIDAE (DIPTERA), WITH DESCRIPTIONS OF NEW SPECIES F. L. BLANC AND RICHARD H. FOOTE (FLB) 5309 Spilman Ave., Sacramento, California 95819; (RHF) Systematic Ento- mology Laboratory, BBII, Agricultural Research Service (Retired), present address: Box 166, Lake of the Woods, Locust Grove, Virginia 22508. Abstract.—Seven new species of Tephritidae from the United States are described: Eutreta modocorum Blanc, E. navajorum Blanc, Gymnocarena flava Foote, Trupanea vi- ciniformis Foote, Urophora claripennis Foote, U. setosa Foote, and Valentibulla dodsoni Foote. Gymnocarena bicolor Foote is transferred to Mylogymnocarena (n. comb.), Pa- roxyna americana Hering is a new synonym of P. genalis (Thomson), and Xanthaciura chrysura (Thomson) is further described. A review of Gymnocarena and Mylogymno- carena and taxonomic observations on all included species are presented. While identifying large numbers of fruit flies in the course of preparing a handbook of the Tephritidae of North America north of Mexico, we have encountered a number of taxonomic problems and hitherto un- described species. The present paper pre- sents our solutions to some of these prob- lems and the descriptions of seven new species which need names prior to publi- cation of the handbook. Letter abbreviations representing the var- ious private and institutional collections from which specimens were borrowed for this study, and in which type material of the newly described species is deposited, are listed in our acknowledgment section. Eutreta Loew Icaria Schiner, 1868: 276 (type species, Try- peta sparsa Wiedemann, 1830: 492; preocc. Saussure, 1853). Eutreta Loew, 1873: 276 (type species, 7ry- peta sparsa Wiedemann, 1830: 492, des. by Coquillett, 1910: 543). Stoltzfus (1977) clarified the status of this genus and its component species in his ex- cellent revision of the New World fauna. Since then, three additional species have been discovered: EF. coalita Blanc (Foote and Blanc, 1979) from California, and the fol- lowing ones from California and Arizona. Eutreta modocorum Blanc, NEw SPECIES Fig. | Diagnosis.—In profile, oral margin only slightly produced; costal margin without hyaline spots; antenna short, not attaining oral margin; wing black; pleuron pinkish amber. Female. — Head: General color pink; frons pinkish gray; lunule dark gray; antenna pink to light brown, 0.4 mm long, reaching about */; of way to oral margin; arista black, amber near base; face pink, turning more pale gray near median and oral margins, without spots; oral margin only slightly produced; parafacial with whitish pollinosity; palpus pink with 8-10 anteroventral setae ranging from white near base to dark brown near tip; lower fronto-orbitals, anterior upper fronto-orbitals, ocellars, and inner verticals brownish black but lighter near base; genal 426 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON | . VOLUME 89, NUMBER 3 bristle light brown; posterior upper fronto- orbitals white; postverticals and postoculars white but with pink near tips. Thorax: Prescutum and scutum black with light gray pollinosity, covered with short, white setae, bristles light brown blending to black at tips; scutellum mahogany brown, cinereous dorsolaterally, with 2 pairs of prominent dark brown bristles; postscutel- lum white; postnotum black, covered with silvery white pollinosity except for a pair of black spots dorsolaterally; pleuron pinkish amber; legs and halter amber yellow; wing shining coal black blending to dark brown at extreme base and in anal cell and alula; a few widely scattered, very faint gray spots in cells br, r,,;, and dm; a prominent round white spot at caudal margin of cell cua,, lying adjacent to end of vein A,CuA,, 2 small angular white spots near base of cell cua,, lying against vein CuA,; apical crescent white, extending from about midway be- tween apices of veins R,,,and R,,; to about ¥; the distance from vein M to vein CuA,, the inner arc slightly flattened into almost a straight line and its posterior tip curving distally to wing margin. Abdomen: Tergites and sternites red, the former evenly covered with small black se- tae and with slightly longer ones along pos- terior edge, 6th tergite with a pair of small black spots near lateral extremity and just caudad of 5th tergite; a row of 8-10 large dark brown spines on terminal margin of 6th tergite; oviscape dark amber, terminal margin narrowly black with 2 prominent setae. Male.— Not known. Body length.— Female 4.8 mm, including oviscape. Type specimens.— Holotype, ¢, 4 mi. E. Davis Creek, Modoc Co., Calif., 25.V1I.80, — 427 G. Steck, ex stem tip galls on low-growing Artemisia sp. (USNM). Paratype, °, same data as holotype (USNM). Discussion. —Eutreta modocorum re- sembles E. diana (Osten Sacken) and E. di- visa Stoltzfus. It can be distinguished from diana by its almost total lack of small gray- white wing spots, the prominent white spot adjacent to the terminus of vein A, +CuA,, by the pink head, pinkish amber pleuron, and by the entirely amber-yellow legs. It differs from the male of divisa by its lack of two prominent white diagonal slash marks on the wing and from the female of divisa by the lack of small gray to white spots in cells r, and r,,;, by the prominent white spot adjacent to the terminus of vein A,+CuA,, and by the characteristic color- ation of its head, pleuron, and legs. The name modocorum honors the Modoc Indians, who inhabited the area that in- cludes the type locality. Eutreta navajorum Blanc, NEw SPECIES ig. 2 Diagnosis.—In profile, oral margin strongly produced; costal margin with light areas or spots; oviscape at least as long as last 2 tergites; setae of oral margin and lower postgena largely brown or black; apical wing crescent well developed in both sexes and reaching anteriorly beyond vein R,,,; light spots in wing disk small. Female. — Head: Frons slightly wider than long, a faint, narrow pale white triangle straddling ocellar triangle posteriorly, its slender point extending anteriorly to frontal suture; face without spots, bearing antennal grooves from antennal bases to oral margin. Parafacials with silvery-white pollinosity extending up over fronto-orbital plate; or- bito-antennal spot lacking; genal margin Figs. 1-10. Right wings of Tephritidae. 1, Eutreta modocorum, female. 2, E. navajorum, female. 3, Gym- nocarena flava, female. 4, Trupanea viciniformis, female. 5, T. viciniformis, male. 6, Urophora claripennis, female. 7, U. setosa, male. 8, Valentibulla dodsoni, female. 9, V. dodsoni, male. 10. Xanthaciura chrysura, female. 428 posterior to genal bristle with a row of 4-6 medium-sized black setae, occasionally with one or more white setae in this row; 3 pairs long black lower fronto-orbitals; anterior upper fronto-orbital long and black, pos- terior white, shorter, strongly reclinate; in- ner verticals long and black; about 12 thick, white postoculars; ocellars long, black, ex- tending slightly antero-laterad. Thorax: Dark brown; notum and pleuron with scattered short, white setae; halter light brown; postscutellum greyish white; post- notum brownish black with grey pollinosity centrally; legs concolorous with general body color, tarsi lighter brown. Wing dark brown- ish black with uniformly small, distinct, greyish white spots not coalescent, rather evenly distributed except lacking in an api- cal band adjacent to apical white crescent but extending to wing base and present on anal lobe and alula; apical white crescent as narrow as, or narrower than, subapical brown band and extending anteriorly just beyond vein R,,,; and posteriorly about '3 of distance from vein M to CuA,; costa without marginal spots beyond the area sur- rounding apex of vein R,. Abdomen: Dark brown; tergites and ster- nites with evenly distributed fine dark brown setae; oviscape dark brownish black near base and apex, blending to mahogany brown in middle. Male.—All head, body, leg, and wing characters essentially as in female, without any evident sexual dimorphism except for genital structures. Type specimens.— Holotype, °, Sunny- side Cyn., Huachuca Mts., Cochise Co., Ar- izona, 9.VII.1940, D. E. Hardy (USNM). Allotype, ¢, same locality and date as ho- lotype, R. H. Beamer (USNM). Paratypes, 26, all same locality and date as holotype: DOE. Hardy § 6, 3.93 R. He Beamer 6:6; 2 oe be wenagal 3.01. Katert 26> and: L. J. Lipovsky 2 °. Depositories of para- types: 14 6, 8 2 (USNM); 1 4, 1 @ (CAS); 1 6, 1 2 (FLB). Discussion. — This species is similar to EF. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON intermedia Stoltzfus and runs to that species in Stoltzfus’ key (1977). It can be readily distinguished from intermedia by its lack of an orbito-antennal black spot. At the present time this species is not known except from the type locality. Its spe- cific name is given in honor of the Navajo Indians of the southwestern United States. Gymnocarena Hering Spilographa Loew: Aldrich (part), 1905: 604 (catalog). Euaresta Loew: Aldrich (part), 1905: 612 (catalog). Oedicarena Snow: Curran, 1934: 289 (in key), figs. 7, 30 (wing, head). Gymnocarena Hering, 1940: 4 (type species, Oedicarena diffusa Snow, 1894, by orig. des.).— Foote, 1960a: 112 (review). — Foote, 1965: 675 (catalog).— Wasbauer, 1972-17 (hosts): Studies subsequent to the junior author’s review of this genus (Foote, 1960a) showed that its species and those of Mylogymno- carena Foote were possibly congeneric. Many characters of the two genera are strik- ingly similar. In lateral view, the profile of the head is unusual in that the parafacial at the level of the antenna is quite wide, the face 1s receding, and the postgena 1s swollen, giving the head outline a slanted aspect with respect to the axis of the eye. Both genera are brown to yellow bristled without any important differences in the structure and placement of the setae except for the pres- ence of certain femoral bristles in the species of Gymnocarena (see below). The body and wing structures are also quite similar. Both genera have a hairless frons, crossvein r-m is apicad of the midpoint of the discal cell, and the dorsocentral bristles are in or very close to a line drawn between the supra- alars. The recent discovery of an undescribed species resembling tricolor has focused our attention on this situation, and our detailed examination of the species of both genera VOLUME 89, NUMBER 3 reveal several differences that distinguish two rather clear-cut groups: Gymnocarena: Background color of wing disk completely or partly yellowish; cell r, with 2 hyaline spots apicad of pterostigma: veins r-m and dm-cu nearly parallel; femora of all 3 legs somewhat enlarged, at their thickest part about 3 x diameter of respec- tive tibiae. Mylogymnocarena: Background color of wing disk uniform brown, although it may fade to completely hyaline basally; cell r, with only | hyaline area apicad of ptero- stigma; veins r-m and dm-cu lying at an angle to each other so that their posterior extensions converge at or near posterior wing margin; femora not markedly enlarged, at their greatest diameter not more than about 2x as thick as the respective tibiae. The species of the two genera exhibit yet another distinguishing characteristic: Those of Gymnocarena occur below 3000 feet east of the Continental Divide, while those of Mylogymnocarena are found at relatively high altitudes in the western mountain chain of the North American continent. KEY TO THE KNOWN SPECIES OF GYMNOCARENA 1. Background color of wing uniformly light yel- low, the hyaline markings sometimes difficult to distinguish from it; apical-most hyaline mark in cells r,,, and r,,, joined across vein R,,; 5 lou 0 ag AER RE a AA diffusa (Snow) — Apical half of wing dark brown, basal half fad- ed brown, sometimes yellowish brown; apical- most hyaline marks in cells r,,, and r,,, en- tirely separate, not at all joined across vein its < a Sn Re ed RS Bee eo 2 2. Hyaline mark at apex of pterostigma ending above vein r-m; apical 3 of discal cell and all of vein r-m included in the dark brown area anterior to them; face concave, with a rather sharp\central carina ........... tricolor (Doane) — Hyaline mark at apex of pterostigma ending distinctly distad of anterior end of vein r-m; apical 3 of discal cell and posterior half of vein r-m lying in a yellowish area similar to that in basal half of wing; lower half of face swollen, clearly visible beyond parafacial in profile .. BPRS ES Late ats Bees cae D flava Foote, new species 429 Gymnocarena diffusa (Snow) Oedicarena diffusa Snow, 1894: 161 (lec- totype female, Kansas).—Coquillett, 1899: 261 (taxonomy).—Snow, 1903: 219 (Kansas).—Curran, 1934: 290, fig. 30 (head). — Knowlton and Harmston, 1937: 145 (Utah).— Byers et al., 1962: 180 (type data). Straussia diffusa: Coquillett, 1899: 261 (taxonomy).—Essig, 1938: 602 (note). Spilographa diffusa: Aldrich, 1905: 604 (catalog).— Washburn, 1905: 118 (Min- nesota). Strauzia diffusia [error]: Cresson, 1907: 100 (New Mexico). Gymnocarena diffusa: Hering, 1940: 4 (type data).—Foote, 1960a: 113 (review).— Foote, 1962: 174 (type designation).— Foote, 1965: 676 (in catalog).—Was- bauer, 1972: 117 (hosts). A number of workers have contributed some interesting information about this species since 1970. Wasbauer (1972) and Hilgendorf and Goeden (1981) state that the species has been found associated with /e- lianthus annuus L. and possibly other He- lianthus species; Beirne (1971) and Lipp and Schulz (1970) indicate that it may be of some economic importance on these hosts; and Kamali and Schulz (1971, 1973, 1974) pre- sent additional information on_ artificial diet, biology, and ecology, and a description of immature stages. The outstanding characteristic of this species is the very pale yellow background color of the wing with which the hyaline spots contrast but little. Other important distinguishing characters are presented in the accompanying key. Gymnocarena tricolor (Doane) Euaresta tricolor Doane, 1899: 191 (lecto- type male, South Dakota). — Huber, 1927: 48 (parasite).—Foote, 1966: 125 (type designation). Tephritis tricolor. Coquillett, 1899: 264 (taxonomy). 430 Gymnocarena tricolor: Quisenberry, 1950: 10 (taxonomy).— Foote, 1960a: 113 (re- view).— Foote, 1965: 676 (in catalog). Since one of us (Foote, 1960a) published his review, no additional information con- cerning this species has come to light. The synonymy given above includes a complete account of the literature. The characters dis- tinguishing frico/or are presented in the dis- cussion of the following species. Gymnocarena flava Foote, NEw SPECIES Fig. 3 Diagnosis.—A large, completely yellow species; all body bristles yellow; dorsocen- tral bristles in or very close to a line drawn through the supra-alars; frons bare; apical anterior quarter of wing disk dark brown, remainder yellowish; hyaline spot imme- diately apicad of pterostigma terminating posteriorly at a point apicad of end of vein r-m; apical hyaline spots on cells r,,; and m not connected across vein M. Female.— Head: In lateral view, 0.8 x as wide as high; gena 0.3 x as wide as height of eye; frons bare, rather bulbous immedi- ately posterior to lunule, 0.7 as wide at vertex as long; antenna about 0.6 x as long as face, 3rd antennal segment broadly rounded apically, only about 1.2 as long as greatest width; 3 pairs lower fronto-or- bitals, 2 pairs upper fronto-orbitals, ocellars and verticals longer and stronger than other head bristles; postgena somewhat swollen, beset with numerous long, stout setae. Thorax: Dorsocentrals in or near a line through supra-alars; 2-3 pairs anepister- nals, 1 pair anepimerals, | pair katepister- nals; 2 pairs scutellars; subscutellum and postnotum yellow; forefemur with 2 rows prominent dorsal setae, | row long antero- ventral setae; hindtibia with a row of short, evenly spaced setae. Apical anterior 4 of wing disk brown, contrasting rather strongly with yellow background color on basal por- tion of wing disk; the following hyaline spots present: 2 hyaline wedges in cell r, imme- diately distad of pterostigma, both contin- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ued into cell r,,;, the proximal wedge end- ing at a point distad of vein r-m; 4 rounded spots in a line, | each in cells r,,, and r,,,; and 2 in cell m; cell m with 2 additional elongated spots; large round spot in cell r,, ; at anterior terminus of vein dm-cu; other discal spots with indistinct borders in prox- imal '2 of wing; proximal yellowish color- ation of disk invading distal 4 of discal cell, proximal 4 of cell m, and enclosing lower ’> of vein r-m. Abdomen: Entirely yellow; oviscape about 1.1 x as long as preceding 2 segments com- bined. Male. —All head, body, and leg characters essentially as in female, without any evident sexual dimorphism except for genital struc- tures. Hyaline wedges distad of stigma tend to coalesce in cell r,,,, and an additional small hyaline spot present in cell r,, 5. Type specimens.— Holotype, °, Kalsow Prairie, Manson, Iowa, 3.VII.1973, W. Bryan Stolzfus (USNM). Paratypes: 1 4, same data as holotype (ISU); 1 6, 2 mi. s. Ames, Iowa, along railroad, 7.VII.1973, W. Bryan Stoltzfus (USNM); 1 @, same data except 4.VII.1973 (ISU). Discussion.—Gymnocarena flava most closely resembles tricolor in wing pattern; in both species the basal 0.75 of the wing disk 1s yellowish in contrast to the dark brown background color of the anterior dis- tal quarter. In this respect, both these species differ from diffusa, in which the background color is an evenly distributed light yellow color throughout. Gymnocarena flava is most readily distinguished from tricolor by the characters presented in the accompa- nying key. Nothing is known about the hosts of either flava or tricolor. Mylogymnocarena Foote Mylogymnocarena Foote, 1960a: 111 (type species, Urellia apicata Thomas, 1914, by orig. des.).—Foote, 1965: 669 (in cata- log). The characters distinguishing this genus and Gymnocarena are given in the discus- VOLUME 89, NUMBER 3 sion of the latter. To date no new infor- mation has been forthcoming concerning the relatively rare species which comprise this genus. Our recent study of structural char- acters shows that Gymnocarena_ bicolor Foote more closely resembles M. apicata than G. diffusa (Snow); bicolor is here trans- ferred to Mylogymnocarena. The following key will serve to distinguish the two known species of the latter genus. KEY TO THE KNOWN SPECIES OF MYLOGYMNOCARENA — Dark area of wing confined to apical 3 of wing disk; pterostigma slightly yellowed; cell r,,, with 2 hyaline spots apicad of terminus of vein R, , ;; cell m with at least 3 hyaline spots .......... Oh LR Re a eA Pe Oe eran apicata (Thomas) — Apical *4 of wing disk, including part of ptero- stigma, with dark marking; cell r,,, completely dark apicad of terminus of vein R,,,; cell m with | large hyaline spot which sometimes con- tains a small dark central area .... bicolor (Foote) Mylogymnocarena apicata (Thomas) Urellia apicata Thomas, 1914: 428 (holo- type female, Colorado). Mylogymnocarena apicata: Foote, 1960a: 111 (review).— Foote, 1965: 669 (in cat- alog). In addition to the holotype, the only other specimen we have seen was collected at Te- palcates, 30 mi. w. Durango, Durango, Mexico, 8400 ft., 4-8.XII.1972 by Powers, Viers, and McNeill at “black and white lights.”” A very small fourth hyaline spot present in cell m is the only character we could find that distinguishes this specimen from the holotype. Mylogymnocarena bicolor (Foote), NEw COMBINATION Gymnocarena bicolor Foote, 1960a: 113 (holotype male, Indian Creek Canyon, Chiricahua Mts., Ariz.).—Foote, 1965: 676 (in catalog). Structurally, apicata and bicolor are so similar that they can be distinguished easily only by their respective wing patterns. 431 This species was described from a single male caught at an altitude of 6100 ft. in Arizona. Since then, we have seen an ad- ditional male captured 5 mi. nw. of Colonia Juarez, Chihuahua, Mexico, at 5000 ft. el- evation by G. S. Forbes 24.VHI.1979. Its wing pattern agrees in all respects with that of the holotype except for the lack of a dark spot at the center of the large hyaline area in cell m. We have also seen a Mexican male re- sembling bicolor from Hwy. 40, 6.5 mi. e. Potrerillos, Sinaloa, collected 21.VIII.1964 by E. I. Schlinger. The wing pattern differs from that of bicolor in having less well de- fined hyaline markings in cells r,,r,,;, and m; it may represent yet a third species of Mylogymnocarena. Metatephritis Foote Metatephritis Foote, 1960a: 110 (type species, fenestrata Foote, 1960a: 110, by orig. des.). To date this genus has been represented only by three Wyoming males of the sole species, fenestrata Foote (Foote, 1960a: 110). The genus 1s distinctly tephritine and in all characters but the wing pattern it re- sembles most closely the species of Neote- phritis and Euarestoides in size, habitus, and setal distribution. Gary Steck, formerly University of Tex- as, Austin, reared an additional male of fe- nestrata at Plum Valley Camp Ground, Warren Mts., Modoc Co., California, 25.VI.1980 from small stem-tip galls on a low-growing Artemisia species, the only ad- ditional record we have seen of this rare, unusual species. The wing pattern of this California specimen differs from that of the holotype illustrated by Foote (1960a: 108, fig. 2) in that a round hyaline spot is present at the center of the anal lobe contiguous with the posterior wing margin; another round hyaline spot is centered on the posterior margin of cell cua, between the termination of vein CuA, and the next proximal hyaline 432 area; and the latter is wider, occupying more of the posterior proximal half of cell cua,. Metatephritis may be accommodated in the existing key to the genera of California Tephritidae (Foote and Blanc, 1963: 6) with the following emendation: 30a(20) Wing disk almost completely brown-bor- dered, leaving large central hyaline area occupying cells r, r,,;, and discal cell .. Serer Sho Vie one eyntg Metatephritis Foote Be Se ea eer 31 31(30a) Paroxyna Hendel Paroxyna Hendel, 1927: 146 (type species, Trypeta tessellata Loew, by original des- ignation). Until the appearance of Novak’s revision of this genus (Novak, 1974), there had been no means of distinguishing among the 15 or so known species occurring 1n the United States and Canada. Nine species were added by Novak in 1974. A new synonymy in this widespread genus is recorded herewith. Paroxyna genalis (Thomson) Trypeta genalis Thomson, 1869: 585 (syn- types male, female; California). Paroxyna difficilis americana Hering, 1944: 11 (holotype male; So. Fork, Rio Grande R., Colo.). NEw SYNONYMY. As a result of studying a large number of specimens of P. genalis and P. americana collected at numerous localities in the west- ern United States, we conclude that these two forms are conspecific. The characters most often used to separate them are 1) the presence or absence of small hyaline spots in the black area of cell r, below the ptero- stigma, 2) the size of the hyaline spots in cell ‘n,:,, 3) the spresence!or absence of 2 dark bands across cells dm and cua,, and 4) whether the scutum is striped (with gray and brown) or is brown without stripes. The three wing characters have been observed in all degrees of intergradation, some individuals exhibiting the character for “americana” on PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON one wing while possessing the “genalis” character on the other. The stripeless char- acter of the scutum, which is recorded as occurring in only part of the “americana” population, also shows all degrees of inter- gradation from long prominent stripes, through faint short anterior stripes, to a to- tally brown or golden brown pronotum. Studies of genitalia by Novak (1974) show no definitive differences, and biological data are too inadequate to substantiate the species separation. Trupanea Schrank Trupanea Guettard, 1762: 171 (unavailable name, author not binominal). Trupanea Schrank, 1795: 147 (type species, Trupanea radiata Schrank, 1795: 147 by monotypy) (= stellata Fuessly). No species of Trupanea occurring in America north of Mexico have been de- scribed since the junior author revised the genus (Foote, 1960b). The distinctive species described here is of interest because of its morphological characteristics and because it was collected from two host plants never before recorded for the genus. Trupanea viciniformis Foote, NEw SPECIES Figs. 4, 5 Diagnosis.— Small species, length of wing about 2.25 mm; head quadrangular in pro- file, only slightly higher than long; thorax and abdomen gray tomentose without other distinguishing marks; oviscape shining black, about 2 x as long as terminal abdom- inal tergum; no dark ray through discal cell to posterior wing margin; apical 0.25 of cell br with broad infuscation which fades into proximal hyaline area; hyaline mark at apex of pterostigma lying at an oblique angle to horizontal wing axis. Female.— Head: Ground color brownish yellow; frons partially gray tomentose, length from vertex to lunule about equal to width between eyes at vertex; in profile quadran- VOLUME 89, NUMBER 3 gular, only slightly higher than long; gena about 0.2 x as high as eye; genal bristle in- conspicuous; oral margin projecting mark- edly forward under antenna; 3rd antennal segment about 0.3 x as long as head height, arista yellow. Thorax: Ground color of scutum and scu- tellum shining black but densely gray to- mentose, covered with thickened whitish setulae about as long as distance between their bases; ground color of pleural sclerites yellow, densely gray tomentose; postscutel- lum and metanotum more densely gray to- mentose. Wing about 2.25 mm long; pattern (Fig. 4) with usual 7rupanea configuration, a complete dark bar through pterostigma with an indistinct proximal border, fusing completely with a completely dark area oc- cupying apical 0.25 of cell br, the latter also gradually fading into the adjacent proximal hyaline area; hyaline mark at apex of ptero- stigma extending into cell r,,, but strongly constricted where it crosses vein R,,, and proximad of that point; stem of dark Y-shaped mark in apex of cell r,,; with a thickened stem; apex of discal cell with brown borders lying on veins dm-cu and M but no dark bar crossing the disk of that cell. Legs yellow, without distinguishing characteristics. Abdomen: Tergites dark gray tomentose without other distinguishing marks; ovi- scape shining black, about 2 as long as terminal tergite. Male.— Wing pattern (Fig. 5) much as in female except apical Y-shaped mark in cell r,,; with a slender stem connecting it with the large preapical dark area of wing disk; hyaline mark at apex of pterostigma not constricted at vein R,,;. Other characters as in female except for postabdomen. Type specimens.— Holotype, °, Texas A&M Ranch, 15 mi. wsw. Uvalde, Mav- erick Co., Texas, 1.1[X.1976, D-Vac, Xan- thocephalum sarothrae, Robbins and Seedle (USNM). Allotype, ¢, same data as holotype except 13.X.1976, sweeping (same _ host) 433 (USNM). Paratypes: 3 2°, Hwy. 57, 9 mi. w. LaPryor, Chaparrosa Ranch, Zavala Co., Texas, 2.1X.1976, D-Vac, Ericameria aus- trotexana Robbins and Seedle (2, USNM; 1; FLB). Discussion.—7rupanea viciniformis re- sembles vicina (Wulp) in many ways but has no dark bar running into or through the disk of the discal cell, the hyaline bar at the apex of the pterostigma is more oblique with re- spect to the longitudinal axis of the wing, and the proximal borders of the dark bar running through the pterostigma and the dark area in the apex of cell br both blend gradually into the next adjacent proximal hyaline area. This species is named to reflect its very close affinities to 7. vicina. The host data presented above unfortunately do not nec- essarily reflect the true host of this species, which is not known. Urophora Robineau-Desvoidy Euribia Meigen, 1800: 36 (type species, Musca cardui Linnaeus, 1758: 600, by des. Hendel, 1927: 41. Suppressed by ICZN 1963: 339). Urophora Robineau-Desvoidy, 1830: 769 (type species, Musca cardui Linnaeus, 1758: 600, by des. Westwood, 1840: 149). The genus Urophora comprises well over 100 species distributed mainly in the Pa- laearctic, Nearctic, and Neotropical regions. A few species have been reported from Af- rica, and one is known from the Oriental Region. The genus is unique among the Te- phritidae in that vein CuA, encloses cell cup transversely without any kind of angular ex- tension along vein A,+CuA,. Some of the American species may well be more closely related to other tephritid genera, but de- tailed studies of these possible relationships are so far lacking. Steyskal has ably reviewed the Palaearctic and New World species in his pictorial key (1979). The two species described in this 434 paper are the first to be discovered since that date. Urophora claripennis Foote, NEw SPECIES Fig. 6 Diagnosis.— Wing disk without dark markings except for a moderate brownish clouding in pterostigma in some specimens; none of the veins pigmented; pterostigma less then '2 as wide as long; legs entirely yellow; pleuron and scutellum black, dis- tinctly tomentose. Female.— Head: In profile, higher than long, ratio of height to length 1.3:1; fron- tofacial angle 110°, junction of face and frons not particularly angulate; face retreating, oral margin not projecting anteriorly; eye broad- oval; gena wide, about 0.6 x as wide as eye height; mouthparts geniculate, about 0.8 x as long as head; frons nearly parallel-sided, about |.2 x as wide at vertex as length from vertex to lunule; antenna about * as long as face, arista light at base, dark apically: occiput tomentose black, remaining por- tions of head amber colored; all head bris- tles black. Thorax: Ground color of pleuron, scu- tum, scutellum, subscutellum, and postno- tum evenly black without any distinctive markings but evenly and densely tomentose with a slight golden cast; dorsocentrals at level of supra-alars; | pair katepisternals, 1 pair anepimerals; scutal setulae black, slen- der, individually about as long as intersetal distances; all thoracic bristles slender, black; halter amber colored. All coxae brownish black basally, remainder of coxae and all legs entirely yellow; mid- and hindfemora and tibiae without any outstanding bristles, either singly or in rows. Wing disk trans- parent (Fig. 6), color pattern completely ab- sent except for a very light brown clouding in apical '2 of pterostigma in some speci- mens; all veins yellowish; pterostigma less than '2 as wide as long; vein r-m situated very slightly distad of middle of discal cell. Abdomen: Ground color of tergites black, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON subshining, with light blackish tomentum; oviscape shining black, slender, as long as or longer than abdomen, beset with slender black setulae dorsally and ventrally. Male.— Not known. Type material.—Holotype ¢, Cameron Cot, Texas, “Ssh 1928) J" Gaeshaw: (USNM). Paratypes: 2 2, same data as ho- lotype (USNM); 1 °, Brazoria Co., Texas, 100.111.1928, R. H. Beamer (UK). Discussion.—Among the Urophora oc- curring in America north of Mexico, the only species without a pattern of dark discal stripes or spots are timberlakei Blanc and Foote and claripennis. Contrasting to the new species described here, in timberlakei the femora are largely black, the pleuron and scutellum are shining black rather than tomentose, the pterostigma is marked more darkly and is shorter than half its width, some of the veins are brown, and the black Oviscape is shorter than the abdomen. Urophora setosa Foote, NEw SPECIES Fig..7/ Diagnosis. — Wing with extensive pattern of transverse stripes, crossveins lying in dark bands; scutellum entirely black; at least bas- al halves of all femora black; scutum lightly tomentose with numerous short, upstand- ing bristles in addition to acrostichals and dorsocentrals; parafacial about as wide as 3rd antennal segment; anterior '2 of frons amber, posterior '2 black, the line of de- marcation between these 2 colors sharp. Male.— Head: In profile, head squarish, about as long as high; oral margin moder- ately and bluntly projecting anteriorly; eye broad oval; gena 0.4 x as wide as eye height; parafacial as wide as 3rd antennal segment; fronto-facial angle about 100°; mouthparts geniculate, about as long as head length; frons between eyes parallel-sided, about as wide as distance from vertex to lunule; an- terior '2 of frons, first 2 antennal segments, face, parafacial, and gena amber colored, remainder of head black; antenna about % VOLUME 89, NUMBER 3 as long as face, 3rd segment brownish black, arista amber at base, black apically; all head bristles shining black. Thorax: Pleuron glistening black; hu- merus and transverse and scutoscutellar su- tures glistening black, remainder of scutum black with coarse blackish tomentum which appears somewhat silvery when viewed from behind; about 20 short, black bristles scat- tered from anterior portion of scutum to acrostichals; dorsocentrals comparatively short; scutellum, subscutellum, and post- notum shining black; 2 pairs scutellars; 2 pairs katepisternals, 1 pair anepimerals; dorsocentrals situated about halfway be- tween transverse suture and supra-alar; all thoracic bristles black; haltere amber. Each coxa with a touch of silvery tomentum; bas- al 7/3 of femora black, remainder of legs am- ber; 4th and 5th tarsomeres of all legs slight- ly darker than preceding ones. Wing disk (Fig. 7) colorless with the following brown- ish-black marks: brown spot with indistinct borders at extreme base of wing; | narrow transverse band at level of bm-cu extending from vein R, to vein A,CuA,; irregular band from costa through pterostigma and vein r-m to middle of cell cua,; dark mark in cells 15, and fr,,5 immediately distad of pterostigma; irregular transverse band from costa at apex of cell r,,, through vein dm- cu to posterior wing margin; quadrate spot filling apex of cell r,,; with a slightly lighter circular area at its center; vein r-m situated at middle of discal cell. Abdomen: Entire surface subshining black with slight black tomentum; tergites 1 and 2 apparently fused and together about as long as tergites 3 and 4 together; tergite 5 about ' as long as tergites 3 and 4 together; protandrium and remainder of visible post- abdomen black. Female.— Not known. Type material. — Holotype 4, Tar Canyon, Kings Co., California, 4.1V.1979, A. J. Gil- bert and A. Bookout (USNM). Discussion.— Two characters combine to make this very distinctive among American 435 species of Urophora: the numerous short upstanding scutal setae (for which the species is named) and the half-amber, half-black frons with a distinct line of demarcation between the two colors. In Steyskal’s (1979) key to American species, sefosa runs to ba- jae Steyskal, from which it 1s easily sepa- rated by the presence of the two characters discussed above. From the other species with an entirely black scutellum occurring in America north of Mexico it is distinguished also by details of the wing pattern. Valentibulla Foote and Blanc Valentibulla Foote and Blanc, 1959: 149 (type species, 7rypeta californica Co- quillett, 1894: 73, by orig. des.).— Foote and Blanc, 1963: 91 (review, Califor- nia).—Foote, 1965: 670 (in N.A. cata- log).—Wasbauer, 1972: 142 (hosts).— Steyskal and Foote, 1977: 154 (key, known species). — Foote and Blanc, 1979: 175 (taxonomy). This genus was originally proposed by Foote and Blanc (1959) for three quite dis- tinctive species, munda (Coquillett), cali- fornica (Coquillett), and thurmanae Foote; in the same paper, Euaresta mundula Co- quillett was synonymized with californica. Steyskal and Foote (1977) indicated that the species described as munda by Foote and Blanc (1959) was actually an undescribed species, and Foote renamed that species steyskali. Foote added C. mundulata to the genus in 1979 (Foote and Blanc, 1979), and yet another species, collected in New Mex- ico, 1s added here. Valentibulla dodsoni Foote, NEw SPECIES Figs. 8, 9 Diagnosis.—Scutum dark brownish gray tomentose, thickly set with expanded yel- lowish white setulae; hyaline incisions in cell m much wider than intervening dark areas; cell r, with only 2 hyaline costal in- cisions apicad of pterostigma, a 3rd (apical) spot entirely lacking; legs entirely yellow; 436 apical Y-shaped mark in cell r,,; with an- terior arm much shorter than posterior arm, causing the hyaline area between them to lie at an angle to vein r,s. Female.— Head: Frons, face, and gena yellow; in profile, 1.5 x as high as long; frons meeting face at about 110°; eye 0.8 x as high as head, the gena rather wide, genal bristle yellowish white; parafacial nearly as wide as 3rd antennal segment at its widest point; lower 0.3 of face barely visible in profile beyond parafacial; frons quadrangular, nearly as wide at vertex as length from ver- tex to antennal base; lunule about 0.6 x as long as wide; numerous prominent yellow- ish white setulae on apical *4 of frons; an- tenna about 0.6 x as long as face, 3rd seg- ment broadly rounded apically, basal '2 of arista yellow, apical '2 black. Thorax: Scutum heavily tomentose, dark gray to dark brown, thickly set with yellow- ish white setulae; pleural sclerites also heavily tomentose, dorsal ' of pleuron yel- low, including humerus and posterior '2 of notopleuron, katepisternum and _ ventral halves of anepimeron and anepisternum dark brownish gray; sternum yellow; scu- tellum, subscutellum and postnotum con- colorous with scutum, but apical margin of scutellum narrowly rimmed with yellow. Wing about 3.2 mm long, pattern (Fig. 8) of hyaline marks highly contrasting with a very dark brown ground color; 2 hyaline areas, | at each end of cell c; pterostigma completely dark, almost 3 x as long as great- est width; 2 hyaline spots completely cross- ing proximal 0.6 of cell r,, the proximal- most continued into cell r,,,; across vein R,,;, apex of cell r, completely dark; large, prominent bulla present in cell r,, 5; anterior arm of Y-shaped mark in apex of cell r,_; distinctly shorter than posterior arm, caus- ing the intervening hyaline area to lie at a distinct angle to vein R,,;; the 2 proximal hyaline spots in cell m partly or wholly co- alesced, in latter case appearing as a very large single spot of 2 in that cell; cell cua, and anal lobe almost entirely hyaline. Legs PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON entirely yellow, without distinguishing characteristics. Abdomen: Tergites and oviscape subshin- ing black; sternites black, lightly tomentose; tergites and apical 0.3 of oviscape sparsely set with short, slender, transparent setulae; oviscape thickened at base in lateral view, about 1.1 x as long as 2 apical tergites to- gether. Male. — Wing pattern (Fig. 9) much as in female except anterior arm of Y-shaped mark in cell R,,,; distinctly narrower than posterior arm, tapering from base and dis- appearing, or nearly so, at junction of costa and vein R,,;, the apicalmost hyaline spot in cell r,,, rarely if ever crossing vein R,,;. Other characters as in female except for those of postabdomen. Type specimens.—Holotype, °, Jemez Valley, Sandoval Co., New Mexico, on Chrysothamnus nauseosus ssp. bigelovii, 14.V.1982, G. Dodson (USNM). Allotype, 6 (USNM) and all paratypes same data (1 9, FLB; 1 6, GD; 1 6, USNM). Discussion. — Valentibulla dodsoni is a distinctive member of the genus, lacking an apical spot in cell r,, and having an oblique apical spot in cell r,,.;. The posterior third of the wing is largely hyaline. The species is named for Mr. Gary Dod- son, Department of Biology, University of New Mexico, Albuquerque, who collected the first known specimens. The key to the known species of Valen- tibulla presented by Steyskal and Foote (1977) may be amended to admit dodsoni as follows: 7(8) Legs, especially middle and hind ones, with femora and tibiae largely blackish aie Mates io i A ale Hid tale munda (Coquillett) Legs usually wholly yellowish Cell r,,; with 3 hyaline incisions apicad of pterostigma; anterior arm of Y-shaped mark in cell r,,, as long as posterior arm ae ee A ee californica (Coquillett) — Cellr,,, with only 2 hyaline incisions apicad of pterostigma; anterior arm of Y-shaped mark in apex of cell R,,; distinctly shorter than posterior arm dodsoni Foote, new species 8(7) VOLUME 89, NUMBER 3 Xanthaciura Hendel Xanthaciura Hendel, 1914a: 86 (1914b: 45). Type species, 7ryvpeta chrysura Thomson (orig. des.). A distinctive genus comprising about 16 quite similar species restricted to the New World, Xanthaciura can be recognized by the presence of a very dark, relatively nar- row wing having two very prominent hya- line triangles based on the costa just distad of the pterostigma (Fig. 11) in addition to other, rounded discal markings and rather extensive hyaline areas along the posterior wing border. A fourth species of this genus is here not- ed as occurring in North America north of Mexico. Xanthaciura chrysura (Thomson) Fig. 10 Trypeta chrysura Thomson, 1869: 580 (Guanabara, Rio de Janeiro, Brazil).— Aczél, 1950: 128 (taxonomy).—Aczél, 1952: 255 (taxonomy).— Foote, 1967: 57, 65 (in catalog). The first species of this genus to be known from North America, insecta, was described by Loew in 1862. About 60 years later, Phil- lips (1923) added a second species from Co- lumbia, Missouri (tetraspina Phillips), and later, Benjamin (1934) described connex- ionis from Florida. Still later, the junior au- thor (Foote, 1967) noted that chrysura, a South American species and type of the ge- nus, was mentioned by Aczeél (1952) as oc- curring in Florida. This name was involun- tarily omitted from the catalog of North American Diptera (1965), and the reported presence of this species has largely been ig- nored since that time. Aczél (1952) indicated he had examined a female of chrysura with the following data: ‘“Norania, Florida, Dec. 9, 1939, R. M. Gra- ham, on orange tree.’’ Recently, we have seen a series of four males and two females collected and presented to the USNMNH by Carl Stegmaier, labeled as follows: 437 ‘“‘Homestead Air Force Base, Homestead, Florida, Nov. 23, 1970, reared from flowers of Mikania sp. (hempvine).” In addition to these two Florida localities, chrysura is known to occur from Central America south to Brazil, but it has not been recognized to date from the southwestern United States, Mexico, or the Caribbean area. Among the North American species of Xanthaciura with only one pair of scutellar bristles, chrysura most closely resembles the widespread insecta (Loew), from which it may be separated by its entirely black pleu- ral sclerites and the nature of some of the hyaline markings along the posterior border of the wing (Fig. 10). In insecta, the apical- most hyaline marking in cell cua, crosses vein CuA, and extends anteriorly into the extreme posterior apical corner of the discal cell, while in chrysura, almost the entire api- cal posterior quarter of the discal cell is filled by the invasion of two entirely coalesced hyaline areas in this region. ACKNOWLEDGMENTS We hereby thank the following individ- uals and institutions who made specimens available for this study: P. H. Arnaud, Jr., California Academy of Sciences, San Fran- cisco (CAS); E. I. Schlinger, University of California, Berkeley (CIS); George Byers, University of Kansas, Lawrence (UK); W. Bryan Stoltzfus, Oskaloosa, Iowa (WBS); Gary Dodson, University of New Mexico, Albuquerque (GD); and Gary Steck, Uni- versity of Texas, Austin (GS). Some speci- mens are deposited in the National Museum of Natural History, Washington, D.C. (USNM); those belonging in the collection of the senior author are designated (FLB). LITERATURE CITED Aczél, M. L. 1950. A revision of the genus “‘Xan- thaciura” Hendel based on Argentine species. Acta Zool. Lilloana 8(1949): 111-146. . 1952. Further revision of the genus Xantha- ciura Hendel. Acta Zool. Lilloana 10: 245-280. Aldrich, J. M. 1905. A catalog of North American Diptera. Smithson. Misc. Collect. 46(1444): 1-680. 438 Beirne, B. P. 1971. Pest Insects of Annual Crop Plants in Canada, II. Diptera. Mem. Entomol. Soc. Can. No. 78: 48-70. Benjamin, F. H. 1934. Descriptions of some native trypetid flies with notes on their habits. U.S. Dep. Agric. Tech. Bull. 401, 95 pp. Byers, G. W., F. Blank, W. J. Hansen, D. F. Beneway, and R. W. Frederickson. 1962. Catalogue of the types in the Snow Entomological Museum. Part III (Diptera). Kans. Univ. Sci. Bull. 43(5): 131- 181. Coquillett, D. W. 1894. New North American Try- petidae. Can. Entomol. 26: 71-75. 1899. Notes and descriptions of Trypetidae. J. N.Y. Entomol. Soc. 7: 259-268. 1910. The type-species of the North Amer- ican genera of Diptera. Proc. U.S. Natl. Mus. 37(1719): 499-647. Cresson, E. T., Jr. 1907. Some North American Dip- tera from the southwest. Paper II. Trans. Am. Entomol. Soc. 33: 99-108. Curran, C. H. 1934. The Families and Genera of North American Diptera. C. H. Curran (privately printed). 512 pp. Doane, R. W. 1899. Notes on Trypetidae with de- scriptions of new species. J. N.Y. Entomol. Soc. 7(2): 177-193, pls. THI, IV. Essig, E.O. 1938. Insects of Western North America. Macmillan Co., N.Y. 1035 pp. Foote, R. H. 1960a. Notes on some North American Tephritidae, with descriptions of two new genera and two new species. Proc. Biol. Soc. Wash. 73: 107-117. 1960b. A revision of the genus 7rupanea in America north of Mexico. U.S. Dep. Agric. Tech. Bull. 1214, 29 pp. 1962. The types of North American Tephrit- idae in the Snow Museum, the University of Kan- sas. J. Kans. Entomol. Soc. 35: 170-179. 1965. Family Tephritidae, pp. 658-678. In Stone, A. et al., eds., A Catalog of the Diptera of America North of Mexico. U.S. Dep. Agric. Handb. 276, 1696 pp. 1966. Notes on the types of Tephritidae de- scribed by R. W. Doane (Diptera). Proc. Entomol. Soc. Wash. 68: 120-126. 1967. Family Tephritidae, No. 57, pp. 57.1- 57.91. In Vanzolini, M., ed., A Catalogue of the Diptera of the Americas South of the United States. Secr. Agric., Sao Paulo, Brazil. Foote, R. H. and F. L. Blanc. 1959. A new genus of North American fruit flies (Diptera: Tephritidae). Pan-Pac. Entomol. 35: 149-156. 1963. The fruit flies or Tephritidae of Cali- fornia. Calif. Insect Surv. Bull. 7: 1-117. 1979. New species of Tephritidae from the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON United States, Mexico, and Guatemala, with re- visionary notes. Pan-Pac. Entomol. 55: 161-179. Guettard, J.E. 1762. Observations qui peuvent servir a former quelques caractéres de coquillages. Acad. R. des Sci., Hist., avec Mém. Math. & Phys., 1756 (Mém.): 145-183. Hendel, F. 1914a. Die Gattungen der Bohrfliegen (Analytische Ubersicht aller bisher bekannten Gattungen der Tephritinae). Wien. Entomol. Ztg. 33(3/4): 73-98. . 1914b. No. 3. Die Bohrfliegen Sudamerikas. Abhandl. Ber. K6nigl. Zool. und Anthropol.-Eth- nogr. Mus. Dresden 14(1912): 1-84, pls. I-IV. 1927. 49. Trypetidae. Jn Lindner, E., ed., Die Fliegen der Palaearktischen Region, vol. 5, 1 fg. 16-19: 1-221. Hering, E.M. 1940. Neue Arten und Gattungen. Si- runa Seva |: 1-16. 1944. Neue Gattungen und Arten von Fruchtfliegen der Erde. Siruna Seva 5: 1-17. Hilgendorf, J. H. and R. D. Goeden. 1981. Phytoph- agous insects reported from cultivated and weedy varieties of the sunflower, Helianthus annuus L., in North America. Bull. Entomol. Soc. Am. 27(2): 102-108. Huber, L. L. 1927. A taxonomic and ecological re- view of the North American chalcid flies of the genus Callimome. Proc. U.S. Natl. Mus. 70(Art. 14) (No. 2663): 1-114. International Commission on Zoological Nomencla- ture. 1963. Opinion 678. The suppression under the plenary powers of the pamphlet published by Meigen, 1800. Bull. Zool. Nomencl. 20: 339-342. Kamali, K. and J. T. Schulz. 1971. Rearing sunflower maggots, Neotephritis finalis, Strauzia longipennis and Gymnocarena diffusa (Diptera: Tephritidae) on artificial diets. Proc. No. Centr. Br., Entomol. Soc. Am. 26(1/2): 85-86. 1973. Characteristics of immature stages of Gymunocarena diffusa (Diptera: Tephritidae). Ann. Entomol. Soc. Am. 66: 288-291. 1974. Biology and ecology of Gymnocarena diffusa (Diptera: Tephritidae) on sunflower in North Dakota. Ann. Entomol. Soc. Am. 67: 695- 699. Knowlton, G. F. and F. C. Harmston. 1937. Utah Diptera. Proc. Utah Acad. Sci., Arts Lett. 14: 141- 149, Linnaeus, C. 1758. Systema Naturae. Regnum Ani- male. Tomus 1, Editio Decima, Holmiae. 823 pp. Lipp, W. V. and J. T. Schulz. 1970. Characterization and quantification of damage caused by selected species of sunflower feeding insects. Proc. No. Centr. Br., Entomol. Soc. Am. 25(1): 27. Loew, H. 1873. Monographs of the Diptera of North America. Part III. Review of the North American VOLUME 89, NUMBER 3 Trypetina. Smithson. Misc. Collect. 11(256): 1- 3515 piss Xo XE Meigen, J. W. 1800. Nouvelle classification des mouches a deux ailes (Diptera L.) d’apres un plan tout nouveau. Paris. 40 pp. Novak, J. A. 1974. A taxonomic revision of Dioxyna and Paroxyna (Diptera: Tephritidae) for America north of Mexico. Melanderia 16: 1-53. Phillips, V. T. 1923. A revision of the Trypetidae of northeastern America. J. N.Y. Entomol. Soc. 31(3): 119-154, pls. 18, 19. Quisenberry, B. F. 1950. The genus Euaresta in the United States (Diptera: Tephritidae). J. N.Y. Entomol. Soc. 58: 9-38. Robineau-Desvoidy, J. B. 1830. Essai sur les My- odaires. Mém. pres. div. sav. Acad. Roy. Sci. l’Inst. France. Sci. Math. Phys., vol. 2, 813 pp. Schiner, I. R. 1868. 1. Diptera, 388 pp., 4 pls. Jn Wiillerstorf-Urbair, B. von (in charge), Reise der Osterreichen Fregatte Novara. Zool., Vol. 2, Abt. l= Sects|B: Wien: Schrank, F. 1795. Naturhistorisches und 6kono- mische Briefe uber das Donaumoor. Mannheim. 2A pp spl: Snow, W. A. 1894. Descriptions of North American Trypetidae, with notes. Kans. Univ. Quart. 2: 159- 174, pls. VI, VII. 1903. A preliminary list of the Diptera of Kansas. Kans. Univ. Sci. Bull. 2(5): 211-221. Steyskal, G.C. 1979. Taxonomic studies on fruit flies 439 of the genus Urophora (Diptera: Tephritidae). Entomol. Soc. Wash., Washington, D.C. 61 pp. Steyskal, G. C. and R. H. Foote. 1977. Revisionary notes on North American Tephritidae (Diptera), with keys and descriptions of new species. Proc. Entomol. Soc. Wash. 77: 146-155. Stoltzfus, W. B. 1977. The taxonomy and biology of Eutreta (Diptera: Tephritidae). Iowa State J. Res. 51(4): 369-438. Thomas, F.L. 1914. Three new species of Trypetidae from Colorado. Can. Entomol. 46: 425-428. Thomson, C. G. 1869. Diptera. Species Novas Des- cripsit, Insekter, pp. 443-614. Jn Konglika Sven- ska Fregatten Eugenies Resa Omkring Jorden, Pt. DreZoo leslie Wasbauer, M.S. 1972. An annotated host catalog of the fruit flies of America north of Mexico (Diptera: Tephritidae). Calif. Dep. Agric. Bur. Entomol. Occa. Pap. No. 19, 172 pp. Washburn, F. L. 1905. The Diptera of Minnesota. Minn. Agric. Exp. Stn. Bull. 93: 19-168. Westwood, J.O. 1840. Order XIII. Diptera Aristotle (Antliata Fabricius, Halteriptera Clairv.), pp. 125- 158. In his An Introduction to the Modern Clas- sification of Insects. Synopsis of the genera of Brit- ish insects. London. Wiedemann, R. W. 1830. Aussereuropaische Zwel- fliigelige Insekten, Vol. 2. Hamm, pp. xii + 684, pls. 7-10. PROC. ENTOMOL. SOC. WASH. 89(3), 1987, pp. 440-443 THE LARVAE OF TWO NORTH AMERICAN DIVING BEETLES OF THE GENUS ACILIUS (COLEOPTERA: DYTISCIDAE) JAMES F. MATTA AND DiRK EDWARD PETERSON Department of Biological Sciences, Old Dominion University, Norfolk, Virginia 23508. Abstract. —The larvae of two dytiscid beetles Acilius abbreviatus and Acilius fraternus dismalus are described and illustrated. Their life histories are briefly discussed. North American, Aci/ius adults were last reviewed by Hilsenhoff (1975) for north- eastern North America and Larson (1975) for northwestern North America. The status of the larval associations was reviewed by Wolfe (1980). Until now only one subspe- cies, Acilius fraternus fraternus (Harris), was positively associated with the adult stage. Associations of Acilius semisulcatus Aubé are in doubt because of possible misiden- tifications of the adult stage and inconsis- tencies in the larval description (Wolfe, 1980). We have reared the larva of an ad- ditional subspecies, Acilius fraternus dis- malus Matta and Michael and of one species, Acilius abbreviatus Aubé. They are de- scribed in this paper. METHODS A method for rearing small dytiscid lar- vae was presented by Matta and Peterson (1985). This method has worked equally well with larger larvae and was used in rearing the larvae described here. Measurements were made with an ocular micrometer in a Wild M8 microscope at 10 x (body length to the nearest 0.1 mm) or 50 x (other measurements to the nearest 0.02 mm). When ratios are given for segmented appendages, the basal segment is given first and the value given in the ratio is the length in millimeters. Acilius abbreviatus Aube Third instar larva.— Length 28 mm. Col- or: Yellow-brown dorsally, pale ventrally, with dark patches at lateral and posterior margins of dorsal sclerites of meso- and metathorax and abdominal sclerites | through 6. Coxae with dark markings ba- sally. Head with a dark stripe on the lateral margin and with brown markings on mid- line (Fig. 1a). Head: Elongate, sides slightly tapering posteriorly, about 1.4 times as broad ante- riorly as at cervical constriction. Labroclyp- eus slightly convex, with distinct thickening apically; apical margin with multiple rows of laminate setae. Ecdysial line branching at middle of head. Lateral margin with 12- 14 spines on a carina extending from cer- vical constriction to ocellar region. Six ocelli present in a circle laterally on the anterior third of head: 2 posterolateral, 1 ventrolat- eral, 1 anterolateral, and 2 larger dorsolat- eral ocelli. Antennae 4-segmented, appear- ing 6-segmented, 2nd and 3rd segments annulate basally; ratio 0.21:0.28:0.36:0.08. Mandible falciform, grooved on inner mar- gin, with minute teeth on inner ventral mar- gin and with a toothlike tuft of hairs on inner margin near base. Maxilla (Fig. 1b, c) with a longitudinal row of setae dorsally, no setae ventrally, 5—6 fine setae on dorsoapical edge; galea spinelike; maxillary palpus VOLUME 89, NUMBER 3 441 Fig: 1: maxilla. d, Ventral surface of labium. 3-segmented, appearing 4-segmented with the 3rd segment annulate near base; ratio 0.17:0.20:0.25. Labium (Fig. 1d), excluding ligula, about as long as wide; ligula slightly shorter than labium (0.30:0.50 mm), api- cally bifid for one third its length; labial palpus 2-segmented, ratio 0.50:0.37. Thorax: Pronotum elongate, about twice as long as other thoracic segments, which are subequal in length. Spiracles present on meso- and metathorax. Legs with segment ratios as follows: foreleg, 1.42:0.63:1.56: 1.06:0.48; midleg, 1.82:0.46:1.88:1.32:0.74; hindleg, 1.86:0.42:1.87:1.34:0.61. Femora, tibiae, and tarsi fringed with a line of fine swimming hairs on anterior and posterior margins and with small setae scattered over dorsal surface. Abdomen: Segments 7-8 completely sclerotized, ring-like; ratio 2.5:3.0 mm; cer- ci short and thin, ratio to eighth abdominal Acilius abbreviatus third instar larva. a, Head. b, Dorsal surface of maxilla. c, Ventral surface of segment 0.88:3.0 mm. Spiracles located lat- erally on segments | through 7. All segments with a lateral fringe of hairs, these fine and sparse on segments 3 to 6, long and dense on segments 7 and 8. Material examined.— Described from two specimens, one reared, collected from a spring-fed pool near the margin of Swann Lake, Glacier Co., Montana, July 13, 1983. The pool was about one meter deep, con- tained a large amount of rooted aquatic vegetation, especially along its margins, and appeared fish free. The larvae were collected near the water surface in open water near the aquatic vegetation. Acilius fraternus dismalus Matta and Michael Third instar larva.—Length 24 mm. Col- or: Brown to black dorsally, pale ventrally, darker at lateral and posterior margins of 442 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 2. of maxilla. d, Ventral surface of labium. dorsal sclerites of meso- and metathorax and abdominal sclerites | through 6. Legs yel- low-brown to dark brown. Coxae with black markings basally. Head with a dark stripe on the lateral margin and with dark mark- ings on midline (Fig. 2a). Head: Elongate, sides slightly tapering posteriorly, about 1.6 times as broad ante- riorly as at cervical constriction. Labroclyp- eus slightly convex with distinct thickening apically; apical margin with multiple rows of laminate setae. Ecdysal line branching at middle of head. Lateral margin with 14 spines on a carina extending from a cervical constriction to ocellar region. Six ocelli present in a circle laterally on the anterior third of head: 2 posterolateral, 1 ventrolat- eral, 1 anterolateral, and 2 larger dorsolat- eral ocelli. Antennae 4-segmented, appear- ing 6-segmented, 2nd and 3rd segments annulate basally; ratio 0.28:0.33:0.33:0.06. Acilius fraternus dismalus third instar larva. a, Head. b, Dorsal surface of maxilla. c, Ventral surface Mandible falciform, grooved on inner mar- gin, with minute teeth on inner ventral mar- gin and with a toothlike tuft of hairs on inner margin near base. Maxilla (Fig. 2b, c) with a longitudinal row of setae dorsally, no setae ventrally, no setae on dorsoapical edge; galea spinelike; maxillary palpus 3-seg- mented, appearing 4-segmented with the 3rd segment annulate near base; ratio 0.16:0.22: 0.26. Labium (Fig. 2d), excluding ligula, about as long as wide, ligula slightly shorter than labium (0.30:0.50 mm), apically bifid for three fourths its length; labial palpus 2-segmented, ratio 0.62:0.40. Thorax: Pronotum elongate, about twice as long as other thoracic segments, which are subequal in length. Spiracles present on meso- and metathorax. Legs with segment ratios as follows: foreleg 1.20:0.28:0.90:1.04: 0.92; midleg 1.60:0.40:1.45:1.30:0.95; hindleg 1.50:0.43:1.38:1.38:1.05. Femora* VOLUME 89, NUMBER 3 tibiae, and tarsi fringed with a line of fine swimming hairs on anterior and posterior margins; with small setae scattered over dorsal surface. Abdomen: Segments 7-8 completely sclerotized, ring-like; ratio 2.5:3.6 mm; cer- ci long and tapering, ratio to eighth abdom- inal segment 1.70:3.6 mm. Spiracles located laterally on segments | through 7. All seg- ments with a lateral fringe of hairs, these fine and sparse on segments 3 to 6, long and dense on segments 7 and 8. Variation.— Body color varies from yel- low brown to black, which seems to corre- late well with the habitat. Lighter specimens are found in open, sunlit, temporary pools while darker specimens are found in deeply shaded pools with a leaf litter bottom. Material examined.— Described from 53 specimens, 4 reared, collected from the Dis- mal Swamp, Suffolk, Virginia. Collection dates ranged from April 13 to April 29, 1981. Larvae are found at the surface in open water areas of temporary pools (generally fish free) and in flooded woodlands. They are usually associated with debris of some kind, either decaying leaves or, more usually, the dead blades of flooded tufts of grass that are fre- 443 quently found in the temporary pools. On the southeastern coastal plain these pools usually flood in late fall and winter and are dry by mid summer. A, fraternus may be separated from A. abbreviatus by the more deeply bifid ligula, longer cerci, longer labial palpus, and dif- fering leg segment ratios, particularly those for the femora and tarsi. 4. f fraternus may be separated from A. f. dismalus by the pres- ence of dorsoapical setae on the maxillae in A. f. fraternus. LITERATURE CITED Hilsenhoff, W. L. 1975. Notes on Nearctic Acilius (Dytiscidae) with the description of a new species. Ann. Entomol. Soc. Am. 68: 271-274. Larson, D. J. 1975. The predaceous water beetles (Coleoptera: Dytiscidae) of Alberta: systematics, natural history and distribution. Quaest. Entomol. 11: 245-498. Matta, J. F. and Peterson, D. E. 1985. The larvae of six nearctic Hydroporus of the subgenus Neoporus (Coleoptera: Dytiscidae). Proc. Acad. Nat. Sci. Phila. 137: 53-60. Wolfe, G. W. 1980. The larva and pupa of Acilius fraternus fraternus (Coleoptera: Dytiscidae) from the Great Smoky Mountains, Tennessee. Coleopt. Bull. 34: 121-126. PROC. ENTOMOL. SOC. WASH. 89(3), 1987, pp. 444-447 A CLADISTIC ANALYSIS OF THE GENERA OF THE LESTONOCORINI (HEMIPTERA: PENTATOMIDAE: PENTATOMINAE) CARL W. SCHAEFER AND IMTIAZ AHMAD Department of Ecology and Evolutionary Biology, University of Connecticut, U-43 Storrs, Connecticut 06268; (IA) Permanent address: Department of Zoology, Karachi University, Karachi-32, Pakistan. Abstract. —The four genera of the Lestonocorini (Gynenica, Lestonocoris, Neogynenica, Umgababa) are analyzed cladistically. The tribe is known from Africa and the Indian subcontinent. Their foodplants, all Acanthaceae, are listed. The genus Gynenica Dallas has for some time avoided tribal placement in the Pen- tatominae. Leston (1953) noted its differ- ences from other genera, and concluded “‘it probably deserves tribal status within the Pentatominae” (p. 182). In 1980, Ahmad and Mohammad created the pentatomine tribe Lestonocorini for Lestonocoris n. gen., and placed Gynenica therein; the former ge- nus was so named in order that the tribe might honor Dennis Leston, who first sug- gested the taxon. Included in Lestonocorini with these two genera were Umgababa Les- ton (raised from subgeneric rank within Gy- nenica) and Neogynenica Y ang (raised from synonymy with Gynenica). Four years later, Shafee and Azim (1984) created the tribe Gynenicini to include Gynenica; they did not mention the Lestonocorini, although a year later Azim and Shafee (1985) com- mented that “the tribe Gynenicini” is “closely related to Lestonocorini Ahmad & Mohammad, but differs from it in the shape of the female genitalia and last abdominal tergum”’ (p. 9). Shafee and Azim (1984) did not examine other genera placed by Ahmad and Mohammad (1980) in their Lestono- corini, and placed Gynenica alone in Gy- nenicini. The two tribes are the same, as is suggested by the inclusion of the same genus in each, and 1s evident from an examination of specimens and the literature. We syn- onymize Gynenicini Shafee and Azim 1984 with Lestonocorini Ahmad and Moham- mad 1980. METHODS Specimens of the following species were examined: Gynenica marginella Dallas (type species), G. affinis Distant, G. basilewskyi Leston, G. carayoni Leston, G. funerea Hor- vath, G. malaisi Leston, G. rustica Distant, G. qadrii Ahmad and Mohammad (type specimen); Lestonocoris karachiensis Ah- mad and Mohammad (type specimen); Um- gababa capeneri (Leston) (paratype speci- men), and U. tellinii (Schouteden). We have been forced to rely on illustrations and de- scriptions for several other species: The type and apparently only specimen of Neogy- nenica izzardi Yang appears to have been mislaid (Museum National d’Histoire Na- turelle; pers. comm., J. Carayon, Jan. 9, 1987). Several requests to the Zoological Survey of India, and to Azim and Shafee, remain unanswered, and so we have not seen Gynenica ghaurii Mathew and G. ala- mi Shafee and Azim. VOLUME 89, NUMBER 3 Table 1. apomorphic states indicated by 1, 1’, etc. (see Fig. 1). 445 Characters and their states in the genera of Lestonocorini. Plesiomorphic states indicated by 0, a d Clypeus and b c Paraclypei Pronotol Humeral Angles Scutellum Female Genitalia Gynenica subequal 0 prolonged, often spined 2 longer than broad acuminate l Neogynenica paraclypel prolonged 2 longer than broad (acuminate?) | longer 1 Lestonocoris subequal 0 not spined 0 not longer than broad subacute 0 Umgababa subequal 0 somewhat spined 1 not longer than broad acuminate l Polarity was determined by outgroup comparison with the family Pentatomidae and the superfamily Pentatomoidea. DISCUSSION OF CHARACTERS Tables 1, 2 Clypeus and paraclypei (character a).— Relative lengths of the clypeus and para- clypei vary so greatly in the Pentatomoidea, we can only suggest that the commonest relationship (subequality) is plesiomorphic here. However, we note that where else- where in the superfamily the paraclypei are longer, they often converge in front of the clypeus; they do not in Neogynenica. Also, the paraclypei in Neogynenica are some- what pointed, an unusual feature which is probably correlated with their being pro- longed. Humeral angles (character b).—Spined humeral angles occur elsewhere in the Pen- tatomidae, but sufficiently infrequently and sporadically that their presence here can be considered apomorphic. That is, the infre- quency of spined angles elsewhere in the family makes it unlikely that they have aris- en more than once within the Lestonocor- ini. The angles are produced but not sharply so in Umgababa, which represents a con- dition intermediate between that of Lesto- nocoris and Gynenica-Neogynenica. Scutellum (character c.)—We could not determine the polarity of the length-breadth relationship of the scutellum (character c); but see the Discussion of Cladogram. Female genitalia (character d).— The acu- minate female genitalia, a particularly char- acteristic feature of this tribe, was what first struck Leston. We believe that the some- what more rounded, less pronounced gen- italia of Lestonocoris is plesiomorphic. Neo- gynenica 1s known only from a single male Table 1. Continued. h é Dorsal Conjunctival Spermathecal bulb Ventral Rim of Genital Capsule Paramere Appendage Gynenica round 0 deeply incised medially 1 with dorsal lobe 1 short l slightly incised medially and sublaterally 1’ Neogynenica ? ? ? ? Lestonocoris round 0 slightly incised without dorsal lobe 0 well developed 0 medially and sublaterally 1’ Umgababa oval 1 slightly incised without dorsal lobe 0 _ short l sublaterally 4 446 Lestonocoris Umgababa ( Pakistan, | sp.) (Africa, 2 spp.) Lestonocorini Fig. 1. (now unavailable), but the specimen (as fig- ured by Yang [1935]) 1s so close to Gynenica we believe the female genitalia will, like Gy- nenica’s, be acuminate. Spermathecal bulb (character e).—The lack of projections on the bulb (as occur, e.g., in the Podopinae [Schaefer, 1983]) is doubtless plesiomorphic, being character- istic of most nonpentatomid pentatomoids. Because round bulbs occur in the primitive pentatomoids, we believe the oval shape in Umgababa is apomorphic. Genital capsule (character f).—The sev- eral types of emargination in the ventral rm differ in each genus. These differences may be generic autapomorphies, and we postu- late each has arisen from an original non- emarginate condition. Paramere and conjunctiva (characters g and h).—A dorsal lobe on the paramere is sufficiently unusual in the Pentatomidae that its presence in Gynenica is apomorphic. A long dorsal conjunctival lobe is common in Cladogram of Lestonocorini genera. Polarities: PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Neogynenica Gynenica (Africa, 8 spp. Indo- Pakistan, 4 spp. ) ( India, ! sp.) ff! g fi We. OS bi bz. Ot (see Table 1). N\ 1p! the Pentatomidae, and the short condition is apomorphic. DISCUSSION OF CLADOGRAM Fig. | Based upon the seven characters for which polarities could be reasonably deduced (characters a, b, d—h), we constructed Fig. 1. Only one homoplasy (f’) had to be in- voked. Unfortunately, important genitalic characters of Neogynenica remain un- known. Assuming the hypothesis in Fig. | is correct, we can suggest that a scutellum longer than broad (character c), whose po- larity we could not determine a priori, is apomorphic; it occurs in the Gynenica- Neo- gynenica clade. A test of the hypothesis will occur when specimens of Neogynenica become avail- able. The cladogram predicts that this genus will have acuminate female genitalia, per- haps a round spermathecal bulb, a genital capsule probably with some emargination, VOLUME 89, NUMBER 3 a short dorsal conjunctival appendage; and its paramere may have a dorsal lobe. FOODPLANTS The records we have indicate that these bugs feed on Acanthaceae (Scrophulariales): Lestonocoris karachiensis Ahmad and Mo- hammad has been raised through several generations on Barleria (Ahmad and Mo- hammad, 1982), Gynenica alami and G. af- finis feed on Crossandra (Shafee and Azim, 1984), and Mathew (1980) records Barleria as the hostplant of G. ghaurii. These bugs and plants are found generally in dry areas. ACKNOWLEDGMENTS We are very grateful to M. J. Spring for the cladogram and R. Govern for her pa- tience and typing. LITERATURE CITED Ahmad, I. and F. A. Mohammad. 1980. A new tribe, a new genus and two new species of the subfamily 447 Pentatominae Amyot et Serville (Hemiptera: Pen- tatomidae) from Pakistan and their relationships. Trans. Shikoku Entomol. Soc. 15: 11-25. Ahmad, I. and F. A. Mohammad. 1982. An esti- mation of immature survivorship for Lestenocoris [sic] karachiensis Anmad and Ali Muhammad [sic] (Pentatomoidea: Pentatominae) in Pakistan. Che- mosphere 11: 1011-1014. Azim, M. N. and S. A. Shafee. 1985. Historical re- view and family-group names of the family Pen- tatomidae (Heteroptera). Indian J. Syst. Entomol. 2: 1-24. Leston, D. 1953. A review of Gynenica Dallas 1851. Rev. Zool. Bot. Afr. 48: 179-195. Mathew, K. 1980. A new species of Gynenica from south India (Heteroptera: Pentatomidae). Orient. Insects 14: 379-382. Schaefer, C. W. 1983. Host plants and morphology of the Piesmatidae and Podopinae (Hemiptera: Heteroptera): further notes. Ann. Entomol. Soc. Am. 76: 134-137. Shafee, S. A. and M. N. Azim. 1984. A new tribe of Pentatominae (Heteroptera: Pentatomidae). In- dian J. Syst. Entomol. 1: 1-S. Yang, W. 1935. Descriptions of a new family and three new genera of heteropterous insects. Ann. Mag. Nat. Hist. (10)16: 476-482. PROC. ENTOMOL. SOC. WASH. 89(3), 1987, pp. 448-457 SPATIAL AND TEMPORAL DISTRIBUTION OF SHORE FLIES IN A FRESHWATER MARSH (DIPTERA: EPHYDRIDAE) J. L. Topp AND B. A. FOOTE Department of Biological Sciences, Kent State University, Kent, Ohio 44262; JLT present address: Department of Entomology, Ohio Agricultural Research and Develop- ment Center, The Ohio State University, Wooster, Ohio 44691. Abstract. —Information is given on the diversity, and spatial and temporal distribution of 25 species of Ephydridae occurring in a freshwater marsh located near Kent in north- eastern Ohio (Portage County). Eight different vegetation zones were recognized in the marsh, with the greatest number of ephydrid species (21) occurring in the vegetation type dominated by yellow water lily, Nuphar lutea (L.) Sibth. & Smith. It is suggested that the ephydrid community organization in Nuphar is influenced by an abundance of high quality food and presence of structural refuges that provide oviposition sites and protection from inclement weather conditions and predators. The family Ephydridae (shore flies) 1s one of the largest entities within the acalyptrate Diptera, consisting of over 1400 described species distributed in all the major faunal regions of the world (Rohdendorf, 1974). More than 400 Nearctic species are known (Deonier, 1979) with most species occurring in aquatic and semi-aquatic habitats. Most studies confirm the association of ephydrids with wetlands, although a few workers have reported the presence of ephydrids in more xeric habitats (Bahrmann, 1978; Steinly, 1984). Several species exist in stressful hab- itats such as crude oil pools (Thorpe, 1930), highly alkaline waters (Wirth, 1971) and thermal springs (Collins, 1975), but most occur in less unusual habitats, including mud shores along drainage ditches (Thier and Foote, 1980), sedge meadows (Scheiring and Foote, 1973), and inland marshes (Deonier, 1965; Todd, 1985). Information available on the trophic hab- its of ephydrids indicates that adults and larvae of many species ingest algae (Dahl, 1959; Deonier, 1972; Foote, 1979; Blair and Foote, 1984), whereas other species are sa- prophagous, feeding on decaying animal tis- sue (Bohart and Gressitt, 1951; Steinly and Runyan, 1979), or plant detritus (Eastin and Foote, 1971; Busacca and Foote, 1978). Larvae of some ephydrid species are leaf- miners in wetland macrophytes (Grigarick, 1959; Deonier, 1971), and a few are pred- ators (Sturtevant and Wheeler, 1953; Simp- son, 1975). The present study gives information on the spatial and temporal distribution of adult Ephydridae occurring in a northeastern Ohio freshwater marsh. Data are also given on species richness, diversity, evenness, and similarity for ephydrid assemblages occur- ring in the 8 different vegetation types found in the marsh. This information is then used to propose factors that may influence ephydrid community organization. MATERIALS AND METHODS The freshwater marsh utilized in this study was located near Kent, Ohio, 0.8 km east of the Kent State University campus. The VOLUME 89, NUMBER 3 Key to vegetation types Carex stricta (CS) Carex lacustris (CL) Eleocharis smallii (ES) Nuphar lutea (NL) Open Areas Phalaris arundinacea (PA) Sparganium eurycarpum (SE) Typha latifolia (TL) Typha-Phalaris (TP) It @ Quadrat Fig. 1. marsh measured approximately 0.5 h and contained a spatial mosaic of 8 vegetation types, most of which existed in nearly monoculture condition (Fig. 1): Phalaris arundinacea L. (canary reed grass), Typha- Phalaris (equally divided between cattail and grass), Carex lacustris Willd. (sedge), Carex stricta Lam. (sedge), Eleocharis smallii Britt. (spike rush), Typha latifolia L. (broad-leaved cattail), Nuphar lutea (L.) Sibth. & Smith (yellow water lily), and Sparganium eury- carpum Engelm. (burreed). Open areas, rep- resented by standing water or bare sub- strate, were located within some of the vegetation types. For collecting purposes, a5 x 10m quad- rat was established within each vegetation type. Sampling for adult ephydrids was con- ducted between June and October, 1984, by using a simplified version of the pan trap described by Grigarick (1959). A 38.5 x 14.0 x 4.5 cm yellow plastic container was filled with a detergent-water solution to a depth of approximately 2 cm. Yellow was 449 Map of freshwater marsh showing distribution of 8 vegetation types. chosen because Disney et al. (1982) found it to be most effective in collecting Diptera from grass-like areas. One pan was placed in the center of each quadrat biweekly, and left in place for 24 hours. Trapped insects were removed and stored in 70% ethyl al- cohol. Additional specimens were obtained by sweeping vegetation with a standard 30.5 cm aerial insect net. Sweeping was con- ducted biweekly before placement of the pan traps and consisted of 10 back and forth sweeps along the 10 m center line of each quadrat. All sweep sampling was done in the early afternoon hours. Adult flies obtained from sweeps and pan traps were combined to attain species lists for Ephydridae occurring in each vegetation type. Relative abundance and percent pres- ence values were calculated for all species collected from each vegetation type. Rela- tive abundance values were further cate- gorized according to the method developed by Scheiring and Foote (1973) and were de- fined as follows: 1-2%, rare (r); 3-8%, oc- 450 No. of species PA T™ CL PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON cS ES We NL SE Vegetation Type Rigs2: casional (occ); 9-14%, common (c); 15-25%, abundant (a); and 26-—100%, very abundant (va). As a measure of diversity, the Shannon- Weaver Index (H’) (Pielou, 1966) was used because it incorporates both species rich- ness and evenness. Separate measures of di- versity (H’), richness (s), and evenness (J’) were calculated for each vegetation type, each collection date, and each month (June— October) within each vegetation type. The degree of ephydrid similarity among the vegetation types was calculated accord- ing to Morisita’s Index (Morisita, 1959). Morisita’s Index ranges from 0 (no similar- ity) to | (complete similarity), and refers to the probability that individuals randomly drawn from each of two vegetation types will belong to the same species, relative to the probability of randomly selecting a pair Number of species of Ephydridae collected in 8 vegetation types of the freshwater marsh. of individuals of the same species from one of the vegetation types. RESULTS A total of 1304 ephydrids of 25 species and 11 genera was collected from the 8 quadrats established within the marsh (Ta- ble 1). Nuphar lutea contained the highest species richness (Fig. 2) and number of in- dividuals (Fig. 3). Species richness in the marsh increased fairly steadily from the on- set of the study in early June (12 spp.) to a peak in mid-July (16 spp.) (Fig. 4). From August to mid-September, species richness remained fairly constant (X = 13) but dropped drastically at the end of September (3 spp.). A secondary peak occurred in early October (9 spp.), followed by another de- crease at the end of the study in late October (4 spp.). Most individuals of Ephydridae VOLUME 89, NUMBER 3 451 70 65 60 3 65 50 x ®@ 465 3) 8 40 3 3 < oe + 0 2 ew 2 0) c 20 15 10 4 0 rn ee PA TP jo & cs i=) TL NL cs 1 18 20 22 78 148 666 131 Vegetation Type Fig. 3. were collected in late June, mid-July and late August (Fig. 5). Values for s, H’, and J’ were obtained for each vegetation type in the marsh (Table 2). Species richness was highest in N. /utea (21), followed by E. smallii and T. latifolia, both with 14 species. High diversity values were found in T. Jatifolia (0.92), E. smallii (0.92), and S§. eurycarpum (0.85). The J’ values in- dicate that individuals were most evenly distributed among the component ephydrid species in 7ypha-Phalaris (0.86), followed by E. smallii (0.80) and T. latifolia (0.80). Similarly, values for s, H’, and J’ were obtained for each sampling date (Table 3). Species richness was highest in mid-July (16), whereas the highest diversity value was obtained in early July (1.03). The J’ values indicate that individuals were most evenly Relative abundance of species of Ephydridae collected in 8 vegetation types of the freshwater marsh. distributed among the component species in early July (0.87) and early August (0.82). Values for s, H’, and J’ were also obtained for each vegetation quadrat during each of the 5 months of the study (Table 4). Values obtained for quadrats containing 10% or more of the total number of ephydrids col- lected from the marsh are discussed here. Those quadrats are: N. /utea (68%), T. la- tifolia (11%), and S. eurycarpum (10%). Ty- pha latifolia had the most species (10) in June and July; N. /utea (15), in August; and S. eurycarpum (10), in July. Typha latifolia (0.82), and S. eurycarpum (0.81) had their highest diversities in June, and N. /utea (0.81) had its highest diversity in July. Peak evenness was found in August for 7. /ati- folia (0.96), in October for N. lutea (0.73), and in September for S. eurycarpum (0.97). No. of species Relative Abundance % 52 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON nin 6-1 66 621 7-9 7-19 86 8-17 8-30 913 930 10-11 10-27 Time Fig. 4. Number of species of Ephydridae occurring at 12 time intervals in the freshwater marsh. an aa oa ey ey 8-31 215 lis Hi Fa “Time Fig. 5. Relative abundance of species of Ephydridae occurring at 12 time intervals in the freshwater marsh. VOLUME 89, NUMBER 3 Table 1. Species of Ephydridae collected in a fresh- water marsh. Brachydeutera argentata (Walker) Dichaeta caudata (Fallén) Discocerina obscurella (Fallén) Hydrellia cruralis Coquillett Hydrellia discursa Deonier Hydrellia griseola (Fallén) Hydrellia harti Cresson Hydrellia spp. (females) Lytogaster excavata (Sturtevant and Wheeler) Notiphila adjusta Loew Notiphila avia Loew Notiphila bella Loew Notiphila deonieri Mathis Notiphila eleomyia Mathis Notiphila nudipes Cresson Notiphila olivacea Cresson Notiphila pauroura Mathis Notiphila scalaris Loew Notiphila solita Walker Notiphila taenia Mathis Notiphila n. sp. Notiphila spp. (females) Ochthera anatolikos Clausen Paracoenia fumosalis Cresson Scatella stagnalis (Fallén) Setocera atrovirens (Loew) Typopsilopa atra Loew All vegetation quadrats were compared for similarity of ephydrid species (Table 5). The ephydrid community in Nuphar showed very low similarity (0.22-0.31) to those found in any other vegetative type. In con- trast, the ephydrid community in Phalaris was similar to that in C. lJacustris (0.96); Typha-Phalaris was very similar to C. stric- ta (0.99) and quite similar to Sparganium (0.92); C. lacustris was very similar to C. stricta (0.97); C. stricta was quite similar to Eleocharis (0.92); and Typha was quite sim- ilar to Sparganium (0.91). Field collections indicate that N. /utea produced 84% of all ephydrid species and 68% of all ephydrid individuals taken dur- ing the study (Figs. 2, 3). A total of 886 individuals was collected in this vegetation type, comprising seven genera and 21 spe- cies (Table 6). Notiphila spp. dominated 453 Table 2. Species richness (s), diversity (H’), and evenness (J’) of Ephydridae in 8 marsh vegetation types. Vegetation S H’ ad PA l 0.00 0.00 EP 6 0.67 0.86 OU 5 039 0.56 CS 7 0.62 0.73 ES 14 0.92 0.80 ele 14 0:92 0.80 NL 21 0.81 0.61 SE 13 0.85 0.76 the Nuphar association, comprising 76% of the total ephydrids collected. Three species were collected exclusively from water lily vegetation. Notiphila eleomyia Mathis was very abundant (42%), Notiphila bella Loew was abundant (22%), and an undescribed species of Notiphila was occasional (3%). D. caudata (Fallen) was also common in Nu- phar (14%). Notiphila bella was mainly col- lected between early June and early August, with peak emergence in early June (Table 7). This species occurred in 50% of the sam- ples taken during the study (Table 6). No- tiphila eleomyia was collected between early June and mid-July, and between mid-Au- gust and mid-October. Peak abundances were reached in late June and late August (Table 7). This species was a fairly constant member of the Nuphar ephydrid fauna, as Table 3. Temporal species richness, diversity, and evenness in marsh Ephydridae. Date s H’ J 6-1 0 0.00 0.00 6-6 12 0.58 0.53 6-21 14 0.68 0.59 7-9 15 1.03 0.87 7-19 16 0.87 On72 8-6 14 0.95 0.82 8-17 14 0.91 0.79 8-30 1] 0.66 0.66 9-13 13 0.91 0.81 9-30 3 0.26 0.54 10-11 9 0.59 0.61 10-27 4 0.38 0.62 454 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 4. Temporal values for ephydrid species richness (s), diversity (H’), and evenness (J’) in 8 marsh vegetation types. Vegmiliype: 93s H' ui S H’ J’ s PA a = OU = _ 0 TP 0:00 4 052, 0:86 2 CE 1 0.00 — 5) OS 3a 079, 4 CS 0 — = 2) 028 +0.92 6 ES Ue = 6 0.70 0.90 10 SUL, 10° 0:82 O582> 105) 0:81 05831 6 NL O 20:4- {Ora9s 4s OLS O71 0tme 15 SE 2 -O:31.~ WONS5 a0 0/5580 78 7 it occurred in 75% of the samples taken (Ta- ble 6). The undescribed Notiphila sp. was less patterned in occurrence, attaining slight peaks in abundance during late June and late August (Table 7). This species was ob- tained in 25% of the samples taken (Table 6). Dichaeta caudata was collected between early June and late August, and between late September and late October with peak abundance in late August (Table 7). This species occurred in >80% of the samples taken (Table 6). DISCUSSION Wetland habitats have long been recog- nized as supporting a relatively rich ephyd- rid fauna (Dahl, 1959; Deonier, 1965; Scheiring and Foote, 1973; Busacca and Foote, 1978). Comparison of the quantita- tive variables s, H’, and J’ in this study in- dicates that differences exist among the ephydrid communities associated with var- August September October H’ J s H’ J s H i — = 1 = a= 0 = — 0.28 0.92 0 0.00 ~ 0 _ — 0.60 1.00 1 = = 1 0.00 _ 0.66 0.85 1 0.00 — 1 0.00 — O95) F095 5 0540) (058 45 105 2a R0rSG OF75— 70196 1 0.00 -- 1 0.00 a 0.80 0.68 10 064 064 7 0.62 0.73 O87 7091 ») 0:68 0:97 “4950/57 530395 ious marsh vegetation types. Nuphar lutea, T. latifolia, and S. eurycarpum contained relatively large and diverse ephydrid assem- blages, with species of Notiphila being among the more common ephydrids encountered. In northeastern Ohio, Busacca and Foote (1978) repeatedly collected 4 species of No- tiphila from a marsh containing cattails, sedges, spike rushes, and true rushes. The present study indicates that N. /utea also represents a particularly suitable habitat for many Notiphila spp., with their abundance probably the result of several interacting factors. The water lilies grew in approxi- mately 25-30 cm of water throughout the study period. Whether the presence of standing water had a direct effect on Diptera in the marsh was not determined, but other authors have examined its importance. Thier and Foote (1980) reported that per- iodic inundation had a significant positive effect on the population growth of several Table 5. Ephydrid community similarity in 8 marsh vegetation types. PA TP Gl ES ES TL NL SE PA — 0.64 0.96 0.83 0.64 0.35 0.23 0.68 ue 0.82 0.99 0.90 0.85 0.29 0.92 CE — 0.97 0.80 0.26 0.28 0.57 CS = 0.92 0.70 0.29 0.79 ES _ 0.68 0.31 0.78 IE — 0.22 0.91 NL — 0.24 SE VOLUME 89, NUMBER 3 ephydrid species associated with a mud shore, although Cameron (1976) noted that adult insects were not affected by inundating tides in a California salt marsh. Larvae of Notiphila spp. are not affected by standing water and survive submerged while feeding on anaerobic sediments because they obtain oxygen through sharp spiracular spines that are inserted periodically into intercellular spaces in the roots of aquatic plants (Busac- ca and Foote, 1978). During October and November, 1985, over 30 third-instar lar- vae of Notiphila spp. were obtained from mud surrounding Nuphar roots, suggesting that the life cycle of species of this genus can be completed in the Nuphar association. This method of obtaining oxygen while liv- ing in anaerobic sediments may explain, in part, the abundance of Notiphila in Nuphar. The occurrence of certain ephydrids in N. lutea may also be influenced by the avail- ability of potential food resources, such as broad leaves, thick stems, and bulbous flow- ers. However, there is no evidence that adult Notipthila are capable of utilizing water lily vegetation itself as a food source. According to Waitzbauer (1976), Notiphila adults usu- ally feed on fluids of decaying plant mate- rial. Utilization of Nuphar pollen and nectar by adult Notiphila is also known (van der Velde and Brock, 1980). However, water lily vegetation may indirectly enhance food availability for larvae of Notiphila spp. Water lilies in the marsh were continually in various stages of decomposition, reach- ing a maximum in mid-August. Decaying Nuphar leaves have been shown to harbor a rich microflora (Wallace and O’Hop, 1984), and larvae of many Notiphila spp. ingest microorganisms associated with de- caying vegetation (Busacca and Foote, 1978). Similarly, both larvae and adults of D. cau- data are scavengers on bacteria and yeasts associated with plant detritus (Scheiring and Foote, 1973). The continual decomposition of water lilies may thus provide a high qual- ity food resource. Although the availability of food re- 455 Table 6. Ephydridae collected from the yellow water lily vegetation type. No. of Relative Percent Species Inds. Abundance Presence B. argentata 12 35 25.00 D. caudata 127. 14.33 83.33 FH. cruralis 2 0:23 8.33 H. discursa 22. 2.48 25.00 H. griseola 18 2.03 41.67 H. harti l 0.11 8.33 Hydrellia spp. 6 0.68 33.33 N. bella 194 21.90 50.00 N. deonieri 6 0.68 8.33 N. eleomyia 369 41.65 75.00 N. nudipes 17 1.92 33.53 N. olivacea 5 0.56 25.00 N. pauroura 13 1.47 25.00 N. scalaris 3 0.34 16.67 N. solita 5 0.56 16.67 N. taenia 4 0.45 16.67 Notiphila n. sp. 24 PTA 25.00 Notiphila spp. 36 4.06 58.33 O. anatolikos l 0.11 8.33 S. stagnalis 19 2.14 41.67 S. atrovirens 2 0.23 16.67 Total species = 21 Total inds. = 886 sources for adults and larvae is likely to be the most important factor contributing to the large and diverse ephydrid fauna in N. lutea, these flies require much more from this plant than just a food source; they also require hiding places from predators, pro- tection from adverse weather conditions, and oviposition sites. In all these respects, the structural complexity of N. lutea may render it more suitable for ephydrid utili- zation than the other marsh plants. The broad, flat leaves of Nuphar formed a dense covering over the surface of the water for most of the study, and were held above the substrate at various heights by thick stems. Ephydrid adults may have been able to es- cape predation by hiding among the various layers of vegetation much more effectively in Nuphar than in the other marsh plants, which were structurally much simpler, con- sisting of straight, slender leaf blades or un- branched stalks. Similarily, Nuphar leaves 456 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 7. Temporal distribution of Ephydridae in the yellow water lily vegetation type. June July Species l 6 21 9 19 B. argentata - — - — D. caudata — 1 2 2 4 H. cruralis _ = — = — H. discursa — _~ — 6 H. griseola —- = _ _ l H. harti — _ _ _ — Hydrellia spp. — _ _ - — N. bella — 93 34 16 36 N. deonieri _ — . eleomyia — 7 . nudipes — — . olivacea = _ . pauroura _ — . scalaris — — solita — _ taenia ~ — Notiphila n. sp. ~ Notiphila spp. - O. anatolikos _ — S. stagnalis _ - S. atrovirens — — to I Ss Next 10 SS eS eS || | | rll eoresy Il TRIS wey | may have aided in protecting adult Ephyd- ridae from adverse weather conditions. For example, the varying heights of water lily leaves could offer ephydrids shelter from intense sunlight, thereby guarding against desiccation, or provide protection from heavy rainfall and strong winds. Oviposi- tion sites available in N. /utea include thick stems, large flowers, and broad leaves. Ob- servations made during this study revealed that Nuphar flowers served as an important Oviposition site for Notiphila spp. During early June, 1985, many Notiphila eggs were discovered inside Nuphar flowers at the base of the petals, often in small clusters of 5-10 eggs. In summary, collection data indicate that the Ephydridae form a dominant dipteran family in a northeastern Ohio freshwater marsh, with species of Notiphila being es- pecially abundant. Utilization of different vegetation types appeared to be important in structuring the ephydrid community, with the majority of species and individuals col- | are be Se | | | | lacey |) ell = Se ise | lected from the yellow water lily, N. /utea. The flight period for most species was be- tween late June and late August, with peak emergence in mid-July. The availability of food resources 1s suggested as the most im- portant factor influencing the abundance of shore flies in Nuphar. The ability of certain ephydrid larvae to survive submerged in water, and the structural complexity of Nu- phar vegetation are also suggested as influ- encing ephydrid community organization in that particular vegetation type. ACKNOWLEDGMENTS We are indebted to B. K. Andreas, Cuy- ahoga Community College in Cleveland, Ohio, for aid in determining species of plants found in the study site and to A. D. Huryn, Department of Entomology at the Univer- sity of Georgia, for determining species of Notiphila. This research was supported by a grant (DEB-79 12242) from the National Science Foundation. VOLUME 89, NUMBER 3 LITERATURE CITED Bahrmann, R. 1978. Okofaunistische Untersuchun- gen an Ephydriden verschiedener Rasenbiotope in Thuringen (DDR) (Diptera). Dtsch. Entomol. Z. 25: 337-348. Blair, J. M. and B. A. Foote. 1984. Resource parti- tioning in five sympatric species of Scate/la (Dip- tera: Ephydridae). Environ. Entomol. 13: 1336- 1339. Bohart, G. E.and J. E. Gresitt. 1951. Filth-inhabiting flies of Guam. Bull. Bishop Mus. 204, 152 pp. Busacca, J. D. and B. A. Foote. 1978. Biology and immature stages of two species of Notiphila with notes on other shore flies occurring in cattail marshes (Diptera: Ephydridae). Ann. Entomol. Soc. Am. 71: 457-466. Cameron, G. N. 1976. Do tides affect coastal insect communities? Am. Midl. Nat. 95: 279-287. Collins, N.C. 1975. Population biology of a brine fly (Diptera: Ephydridae) in the presence of abundant algal food. Ecology 56: 1139-1148. Dahl, R. G. 1959. Studies on Scandanavian Ephyd- ridae (Diptera Brachycere). Opusc. Entomol. Suppl. 15: 1-224. Deonier, D. L. 1965. Ecological observations on lowa shore flies (Diptera, Ephydridae). Proc. Iowa Acad. Sci. 71: 496-510. 1971. A systematic and ecological study of Nearctic Hydrellia (Diptera: Ephydridae). Smiths. Contr. Zool. 68: 1-147. 1972. Observations on mating, oviposition- ing, and food habits of certain shore flies (Diptera: Ephydridae). Ohio J. Sci. 72: 22-29. 1979. Introduction—a prospectus on re- search in Ephydridae, pp. 1-19. 7m Deonier, D. L., ed., First Symposium on the Systematics and Ecol- ogy of Ephydridae (Diptera). N. Am. Benthol. Soc. Disney, R. H. L., Y. Z. Erzinclioglu, D. J. de C. Hen- shaw, D. Howse, D. M. Unwin, P. Withers, and A. Woods. 1982. Collecting methods and the adequacy of attempted fauna surveys, with refer- ence to Diptera. Field Studies 5: 607-621. Eastin, W. C. and B. A. Foote. 1971. Biology and immature stages of Dichaeta caudata (Diptera: Ephydridae). Ann. Entomol. Soc. Am. 64: 271- 279. Foote, B. A. 1979. Utilization of algae by larvae of shore flies (Diptera: Ephydridae), pp. 61-71. Jn Deonier, D. L., ed., First Symposium on the Sys- tematics and Ecology of Ephydridae (Diptera). N. Am. Benthol. Soc. Grigarick, A. A. 1959. Bionomics of the leaf miner, 457 Hydrellia griseola (Fallen), in California (Diptera: Ephydridae). Hilgardia 29: 1-80. Morisita, M. 1959. Measuring the dispersion of in- dividuals and analysis of the distributional pat- terns. Mem. Fac. Sci. Kyushu Univ. Ser. E. (Biol.). 2: 215=235. Pielou, E.C. 1966. Species-diversity and pattern di- versity in different types of biological collections. J. Theor. Biol. 13: 131-144. Rohdendorf, B. 1974. The Historical Development of Diptera. Univ. Alberta Press. xv + 360 pp. Scheiring, J. F. and B. A. Foote. 1973. Habitat dis- tribution of the shore flies of northeastern Ohio (Diptera: Ephydridae). Ohio J. Sci. 73: 152-166. Simpson, K. W. 1975. Biology and immature stages of three species of Nearctic Ochthera (Diptera: Ephydridae). Proc. Entomol. Soc. Wash. 77: 129- 162. Steinly, B. A. 1984. Shore fly (Diptera: Ephydridae) community structure in a xeric grass habitat. Proc. Entomol. Soc. Wash. 86: 749-759. Steinly, B. A.and J.T. Runyan. 1979. The life history of Leptopsilopa atrimana (Diptera: Ephydridae), pp. 139-147. In Deonier, D. L., ed., First Sym- posium on the Systematics and Ecology of Ephyd- ridae (Diptera). N. Am. Benthol. Soc. Sturtevant, A. H.and M. R. Wheeler. 1953. Synopsis of Nearctic Ephydridae (Diptera). Trans. Am. Entomol. Soc. 79: 151-261. Thier, R. W. and B. A. Foote. 1980. Biology of mud- shore Ephydridae (Diptera). Proc. Entomol. Soc. Wash. 84: 517-535. Thorpe, W. H. 1930. The biology of the petroleum fly (Psilopa petrolii Coq.). Trans. Entomol. Soc. Lond. 78: 331-343. Todd, J. L. 1985. The Community Organization of Acalyptrate Diptera in a Freshwater Marsh. M.S. Thesis, Kent State University, 135 pp. van der Velde, G. and Th. C. M. Brock. 1980. The life history and habits of Notiphila brunnipes Ro- bineau-Desvoidy (Diptera, Ephydridae), an aut- ecological study on a fly associated with nym- phaeid vegetations. Tijdschr. Entomol. 123: 105- 127e Waitzbauer, W. 1976. Die Insettenfauna mannlicher Blutenstande von Typha angustifolia. Zool. Anz. 196: 9-15. Wallace, J. B.and J. O’Hop. 1984. Life on a fast pad: Waterlily leaf beetle impact on water lilies. Ecol- ogy 66: 1534-1544. Wirth, W. W. 1971. The brine flies of the genus Ephy- drain North America (Diptera: Ephydridae). Ann. Entomol. Soc. Am. 62: 357-377. PROC. ENTOMOL. SOC. WASH. 89(3), 1987, pp. 458-467 NEW SPECIES AND RECORDS OF NEW CALEDONIA ALLUAUDOMYIA (DIPTERA: CERATOPOGONIDAE) FRANCIS E. GILES AND WILLIS W. WIRTH (FEG) Loyola College in Maryland, Baltimore, Maryland 21210; (WWW) Systematic Entomology Laboratory, BBII, Agricultural Research Service, USDA, and Florida State Collection of Arthropods, Florida Department of Agriculture and Consumer Services, P.O. Box 1269, Gainesville, Florida 32602. Abstract.—Three new species of Alluaudomyia from New Caledonia are described: epsteini, mouensis and poguei. The hitherto unknown male of 4. me/anesiae Clastrier is described and figured, and new distribution records are given for this species and for A. tillierorum Clastrier. A key is presented for identification of females of the nine known species of Al/uaudomyia Kieffer known to occur in New Caledonia. The predaceous midges of the genus 4/- luaudomyia Kieffer are worldwide in dis- tribution. The species of the southern hemi- sphere appear to be numerous but are as yet poorly known. Spinelli and Wirth (1984) list 23 Palearctic, nine Nearctic, and 31 Ori- ental species, compared with 17 Neotropi- cal, 39 Afrotropical, and 27 Australasian species. Clastrier (1986) recently described six new species of Al/uaudomyia from a small collection (15 specimens from six lo- calities ranging from 30 to 850 m elevation) from New Caledonia. This is an example of the rich fauna awaiting study in the Pacific area. We recently received a large collection of ceratopogonids taken in New Caledonia in 1984 by Michael G. Pogue and Marc Ep- stein of the University of Minnesota in St. Paul. In the genus A//uaudomyia we studied 128 specimens from four localities ranging from 100 to 1200 m elevation. We segre- gated these into five species. Two had been described by Clastrier, and three are de- scribed here for the first time. The Alluaudomyia species of the South Pacific are rather easily identified, due to comprehensive revisions previously pub- lished by Tokunaga and Murachi (1959) for Micronesia, Wirth and Delfinado (1964) for the Oriental Region, Tokunaga (1963) for New Guinea, and Debenham (1971) for Australia and New Guinea. Wirth and Del- finado (1964) divided the Oriental species into five rather well-defined groups, but De- benham (1971) found these groups unwork- able for the Australasian fauna. After a comprehensive review of all the described species of A//uaudomyia world- wide, Clastrier settled on mathematical combinations of three female characters to arrange the species in eight artificial sec- tions. These characters are: presence or ab- sence of interfacetal pubescence on the eyes, number of spermathecae (one or two), and presence or absence of an appendix or di- verticulum on the spermatheca near the base of the neck. Using this system Clastrier found that all the Palearctic and Neotrop- ical species fell in his section 8 (annulata, bella, maculipennis and xanthocoma groups); the Afrotropical and Oriental species fell in four sections: 2, 5, 6, 8 (an- VOLUME 89, NUMBER 3 nulata, bella, maculipennis, marmorata and parva groups); and the Australasian species were the most diverse, falling in six sections: ipa? 3555, 6, 8. No Species have yet been found that would fall in Clastrier’s sections 4 and 7. The New Caledonia species are all closely related, having a wing pattern with only two or three small black spots, all found on the radial veins, and only some vague dark streaks distally along the posterior veins in some species. All nine known New Cal- edonian species except one have diverticula on the spermathecae (Clastrier’s sections 1, 2 and 3) and all except one have two sper- mathecae (Clastrier’s sections 1, 2 and 6, the species with one spermatheca falling in Clastrier’s section 3). For explanation of terms, ratios, and measurements see Wirth and Delfinado (1964); morphological terminology follows Downes and Wirth (1981). All specimens listed were collected in new Caledonia at UV light by M. Pogue and M. Epstein, and were mounted on slides in phenol-balsam by the technique of Wirth and Marston (1968). Holotypes and allotypes of new species will be deposited in the National Museum of Natural History, Smithsonian Institution, Washington, D.C.; paratypes, as available, will be deposited in the follow- ing museums: Muséum National d’ Histoire Naturelle, Paris; Australian National Insect Collection, CSIRO, Canberra; Bishop Mu- seum (Honolulu); British Museum (Natural History), London; and California Academy of Sciences, San Francisco. KEY TO FEMALES OF NEW CALEDONIAN SPECIES OF ALLUAUDOMYIA |. | Spermathecae without diverticulum ...... Oe nr pacifica Clastrier Spermatheca(e) with diverticulum 2. One spermatheca present; eyes with inter- facetal airs. 74.7 shee eh. boucheti Clastrier - Two spermathecae present; eyes with or without interfacetal hairs ............... 3 3h. EVESIDATC ER ee eee ee ete ee ese ene 4 = Ey esupubescente amma oe sere iene ae i 4. Yellowish species, thorax yellow on dorsal 459 half; spermathecae hyaline .............. Oe Sis Me eas cee epsteini Giles and Wirth - Brownish species, thoracic dorsum brown- ish, at least in part; spermathecae pigmented 5. | Spermathecae large, diameter 0.065-0.070 mm; scutum yellowish with prominent brown patches on anterior half .......... Se orn De pease ene poguei Giles and Wirth - Spermathecae small, diameter 0.035-0.037 mm; scutum brownish, pale mottlings not prominent if present 6. Fore and mid tibiae with narrow brown band at midlength; wing with distinct linear dark spot over proximal portion of vein R1 ete Ra oR ee ne ee tillierorum Clastrier ~ Fore and mid tibiae with broad brown band at midlength; wing without dark spot over proximal portion of vein RI .. exiguwa Clastrier Fore and hind tibiae with broadly dark mid- portion and narrow pale bands on each side; scutellum brown except in midportion ... Fhe eee oe ee mouensis Giles and Wirth - Fore and hind tibiae with narrow dark band at midlength and broad pale bands on each side;scutellumientinely pale: ...<.5... 5... 8 8 Scutellum with two pairs of setae in mid- portion, none laterally; scutum with one ob- scure pale area on each side, antennal ratio 0.83; palpal segments four and five with lengths in proportion of 20:27 ........... deine Seva wR eT A Ae melanesiae Clastrier - Scutellum with two pairs of lateral setae, none in midportion; scutum with two pale areas on each side; antennal ratio 0.91; palpal seg- ments four and five with lengths in propor- tion of 25:26 neocaledoniensis Clastrier 7(3). Alluaudomyia epsteini Giles and Wirth, New SPECIES Fig--1 A large pale yellowish species with con- trasting dark brown lower half of pleuron and broad contrasting brown bands on legs; wing creamy yellowish with two small blackish anterior spots; eyes bare; sperma- thecae two, hyaline, with short diverticula; male gonostylus slender and bent hooklike on distal half, parameres with slender, ball- like tips. Holotype female. — Wing length 1.52 mm; breadth 0.66 mm. Head: Eyes contiguous, bare. Antenna 460 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig: 1 gonostylus; f, aedeagus and parameres; g, genitalia, aedeagus and right gonostylus omitted; h, femur and tibia of (left to right) fore, mid, and hind legs. (Fig. la) pale brown, narrow bases of fla- gellomeres 2-8 pale; flagellomeres with lengths in proportion of 20-17-16-16-16- 17-18-19-21-23-23-22-22: antennal ratio (9-13/1-8) 0.80; sensilla basiconica present on flagellomeres 9-12. Palpus (Fig. 1c) brown; lengths of segments in proportion of 6-8-14-11-13; palpal ratio 2.3. Mandible with 11-12 teeth. Thorax: Uniformly yellowish on dorsal Alluaudomyia epsteini, a—d, h, female; e-g, male: a, antenna; b, wing; c, palpus; d, spermathecae; e, half, pleural and sternal areas dark brown. Legs (Fig. 1h) yellowish; coxae, trochanters and bases of femora dark brown; brown on proximal halves of fore and mid femora, proximal 0.6 on hind femur, mid femur with narrow apex blackish; tibiae with bases broadly pale except narrow blackish ring at base of mid tibia; apices of tibiae brown, distal 0.4 on fore tibia, 0.3 on mid tibia, and 0.25 on hind tibia; hind tibia with small VOLUME 89, NUMBER 3 infuscation on extensor side at midlength. Tarsi pale, hind basitarsus dark; hind tarsal ratio 2.92. Claws long, slender and unequal on all legs. Wing (Fig. 1b): Creamy white, veins only slightly infuscated; two prominent small black spots, first dark spot covering apex of costa and vein R4+5 and extending 3 way caudad across cell r4+5; second spot prox- imad of r-m crossvein extending from vein R to vein M1 +2. Costal ratio 0.61; macro- trichia numerous on distal portion of wing. Halter pale, knob slightly infuscated. Abdomen: Pale brownish with ventral pigmented pattern on segments 3-7, last three segments yellowish, cerci whitish. Spermathecae (Fig. 1d) two, hyaline, with diverticula subequal in length to the slender, well-developed necks; slightly unequal, measuring 0.065 x 0.045 mm and 0.058 x 0.041 mm including necks, diverticula 0.007 mm long. Male allotype.—Wing length 1.27 mm, breadth 0.42 mm; costal ratio 0.60. Similar to female with usual sexual differences. An- tenna with flagellomeres 2-8 fused, plume brownish; lengths of flagellomeres in pro- portion of 39-18-19-17-16-17-15-15-15-23- 48-48-42; antennal ratio (10—-13/1-9) 0.94. Genitalia (Fig. 1g): Yellow, proximomesal and distolateral margins of gonocoxite brown; gonostylus brown on distal half. Ninth sternite with broad, shallow, caudo- median excavation, ventral membrane spiculate; ninth tergite hyaline, with short, bluntly pointed, divergent apicolateral pro- cesses. Gonocoxite strongly arcuate, bases stout and nearly meeting mesad, narrowed on distal half; gonostylus (Fig. le) slender, somewhat sickle-shaped but abruptly bent at midlength, with sharp-pointed tip, bear- ing 5—7 long slender setae along flexor mar- gin. Aedeagus (fig.1 f) with rounded basal arch extending to half of total length, basal arms dark brown; distal portion tapering to moderately slender, short, rounded tip abruptly bent ventrad. Parameres (Fig. 1 f) separate, moderately stout and parallel on 461 proximal 0.6; narrowly constricted on distal third, the slender distal portion sinuately bent mesad, ventrad, and caudad and end- ing in swollen capitate tip. Distribution. — New Caledonia. Types. — Holotype 8, allotype 6, Mt. Dzu- mac, 760 m, 27.11.1984. Paratypes, 19 °, 7 6, same data. Discussion.—The species is named for Marc Epstein of the University of Minne- sota in recognition of his interest and help in collecting New Caledonia ceratopogo- nids. Differs from related species by the characters given in the key. Alluaudomyia poguei Giles and Wirth, New SPECIES Fig; 2 A relatively large yellow and brown mot- tled species; wing with two prominent an- terior dark spots; scutum yellowish on pos- terior half, brownish anteriorly with prominent mottlings; legs with prominent pale and dark bands; eyes bare; spermathe- cae two, with short diverticula; male gono- stylus sharp-pointed distally but not curv- ing, parameres straight distally with slightly bilobed tips. Holotype female.— Wing length 1.18 mm; breadth 0.50 mm. Head: Brown; eyes narrowly separated to contiguous, bare. Antenna (Fig. 2a) pale brown, flagellomeres 2—8 narrowly pale ba- sally; lengths of flagellomeres in proportion of 18-14-14-14-15-15-15-14-18-18-18-19- 20; antennal ratio 0.78; sensilla basiconica present on flagellomeres 9-12. Palpus (Fig. 2b) brown, segment five darkest; lengths of segments in proportion of 5-8-12-10-11; segment three with small shallow pit bear- ing three long sensilla; palpal ratio 2.4. Mandible with 10-11 coarse teeth. Thorax: Yellowish brown with dark brown mottling dorsad; anterior half of scu- tum dark brown with small yellowish patches; pleuron dark brown on lower half; scutellum yellowish, pale brown mesad; postscutellum brownish. Legs (Fig. 2e) yel- 462 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig..2: lowish; coxae and trochanters dark brown; femora with narrowly blackish apices; fore leg brownish with narrow pale rings sub- apically on femur and sub-basally on tibia; mid femur brownish with broad subapical pale band; mid tibia brownish with broad sub-basal and narrow subapical pale bands, a faint infuscation usually present at prox- imal third of length; hind femur brownish Alluaudomyia poguei, a—e, female; fh, male: a, antenna; b, palpus; c, wing; d, spermathecae; e, femur and tibia of (left to right) hind, mid, and fore legs; f, parameres; g, aedeagus; h, genitalia, aedeagus and right gonostylus omitted. with proximal half pale and narrow sub- apical pale ring; hind tibia pale with dark bands at midlength and at apex; tarsi pale, hind basitarsus dark. Hind tarsal ratio 2.36; tarsal claws long, slender, and unequal on all legs. Wing (Fig. 2c): Two prominent small dark spots; first covering apex of costa and vein R4+5 and extending lightly in a caudolat- VOLUME 89, NUMBER 3 eral direction; second dark spot extends from apex of vein M1+2 just before r-m cross- vein across apex of basal cell to include vein R; strong pigmentation along the lengths of medial and mediocubital veins and their branches and on anal vein. Costal ratio 0.55; six strong setae on vein R4+5 and eight on radial sector; macrotrichia numerous on distal half of wing. Halter pale, knob slightly infuscated. Abdomen: Pale brown with dark mottling; ventral pigment pattern on segments 3-9; cerci pale. Spermathecae (Fig. 2d) two, heavily sclerotized, ovoid with diverticula subequal in length to the well-developed slender necks; slightly unequal, measuring 0.084 x 0.068 mm and 0.073 x 0.061 mm including necks. Allotype male.—Wing length 1.16 mm, breadth 0.36 mm; costal ratio 0.51. Similar to the female with usual sexual differences. Antenna with lengths of flagellomeres in proportion of 26-10-10-10-10-10-10-10-10- 15-26-28-25, antennal ratio (9—13/1-8) 0.76; flagellomeres 2-10 fused, plume yellowish brown. Hind tarsal ratio 2.40. Genitalia (Fig. 2h) brown, tergum nine hyaline distally. Sternum nine with shallow caudomedian excavation, ventral membrane with minute spicules; tergum nine tapering to rather broad, truncated tip with long slender par- allel apicolateral processes. Aedeagus (Fig. 2g) with rounded basal arch extending to a fourth of total length; main body broad and tapering distally to bluntly pointed tip (as seen in allotype). In some specimens, ap- parently due to compression of cover-slip, the aedeagus appears broader and more quadrate, with two sublateral slender scler- otized blades separating out and flanking the slender, tapering, longer median pro- cess. Parameres (Fig. 2f) appearing as a pair of moderately stout straight sclerites, prox- imal portion of each spindle-shaped, taper- ing distally to a slightly expanded, twisted tip with ventrolateral lobe slightly longer than the low dorsomesal expansion. Distribution. — New Caledonia. 463 Types. — Holotype 2, Mt. Dzumac, 760 m, 27-28.11.1984. Allotype 4, same data but 16— 17.1.1984. Paratypes, 44 2, 13 4, same data, both dates; 2 2, Ponandou River, 5 km S Touho, 22.11.1984. Discussion.—This species, the most abundant in our collections, 1s dedicated to Michael G. Pogue of the University of Min- nesota in appreciation of his interest and cooperation in making available to us this superb ceratopogonid collection. Charac- ters to separate 4. poguei from related species are given in the key. Alluaudomyia mouensis Giles and Wirth, New SPECIES Fig. 3a-e A moderately large dark brown species with two-spotted wing, slightly infuscated along veins and membrane slightly dusky; eyes hairy; spermathecae two, small and subequal, each with moderately short di- verticulum. Male unknown. Female holotype. — Wing length 1.15 mm; breadth 0.50 mm. Head: Dark brown. Eyes nearly contig- uous, hairy. Antenna (Fig. 3a) dark brown; lengths of flagellomeres in proportion of 15- 10-9-10-11-11-12-12-17-18-19-19-19, an- tennal ratio 0.98. Palpus (Fig. 3b) dark brown, short; lengths of segments in pro- portion of 5-10-13-9-12; third segment slender, palpal ratio 2.0, with five long sen- silla borne in broad, irregular, very shallow pit. Mandible with 9-10 teeth. Thorax: Dark brown, scutum somewhat darker punctate, scutellum lighter brown. Legs (Fig. 3e) dark brown, knee spots black- ish; femora with narrow pale subapical rings; fore and mid tibiae with narrow sub-basal and subapical pale rings; hind tibia with broad sub-basal and subapical pale rings; tarsi pale, basitarsus dark; hind tarsal ratio 2.74. Claws slender and unequal on all legs, shortest on hind leg; lengths in proportion (fore, mid, hind) of 28:11, 28:11, and 20:7. Wing (Fig. 3c): Slight infuscation on membrane and darker infuscated lines along 464 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VOSTRO csi Ce a A ir Fig: 3: e, femur and tibia of (left to right) fore, mid, and hind legs; f, genitalia, parameres and right gonostylus omitted; g, parameres. veins; costa and radial veins darker toward bases; two prominent small blackish spots, first dark spot covering apex of costa and vein R4+5, second spot proximad of r-m crossvein extending from vein R to vein M1+2. Costal ratio 0.59: macrotrichia nu- Alluaudomyia mouensis female, a—e; A. melanesiae, f-g: a, antenna; b, palpus; c, wing; d, spermathecae; merous on distal portion of wing. Halter pale. Abdomen: Dark brown, cerci pale. Sper- mathecae (Fig. 3d) two, globular with short slender necks; each with moderately long diverticulum; slightly unequal, measuring VOLUME 89, NUMBER 3 0.043 x 0.038 mm and 0.039 x 0.033 mm including necks, diverticula 0.017 mm long. Male.— Unknown. Distribution. — New Caledonia. Types.— Holotype 2°, | 2 paratype, Mt. Mou, 1200 m, 13.11.1984. Discussion.—The species takes its name from its habitat at high elevation on Mt. Mou. It is closely related to 4. melanesiae Clastrier and 4. neocaledoniensis Clastrier, but can be distinguished from these species by its larger size and darker color with more restricted pale leg markings, features that follow the tendency for animals in cooler habitats and higher elevations to be larger and darker than those in hotter, drier cli- mates. Alluaudomyia melanesiae Clastrier Fig. 3f-g Alluaudomyia melanesiae Clastrier, 1986: 190 (@; New Caledonia; fig. genital seg- ments, spermathecae, scutellum). Female diagnosis.— Wing length 0.75 mm, breadth 0.39 mm; costal ratio 0.53. A small dark brown species with whitish scu- tellum and halteres. Wing pale with slight cloudiness, especially along veins; two prominent small blackish spots in usual lo- cation at end of costa and proximad of r-m crossvein. Legs dark brown with narrow pale rings, subapically on femora and at base of hind femur, sub-basally and subapically on fore and mid tibiae, broad base and a broad subapical area on hind tibia pale; hind tarsal ratio 2.74. Female claws slender and un- equal, smallest on hind legs, measuring on fore, mid and hind legs 0.070 and 0.032 mm, 0.066 and 0.034 mm, and 0.048 and 0.020 mm respectively. Antenna and palpus short, dark brown; antennal ratio 0.83. Mandible with 9-10 teeth. Spermathecae two, unequal, globose with distinct slender necks, each with moderately long divertic- ulum; measuring 0.046 x 0.034 mm and 0.030 x 0.030 mm, not including necks. Male description. — Wing length 0.99 mm, breadth 0.36 mm; costal ratio 0.46. Similar to the female with the usual sexual differ- 465 ences. Genitalia (Fig. 3f): Ninth sternite with broad shallow caudomedian excavation, ventral membrane with coarse spicules; ninth tergum tapering to short, tapering, hyaline apicolateral processes. Gonocoxite broad basally, the anteromesal corners nearly meeting mesad, curving and tapering to nearly straight, more slender distal por- tion; gonostylus abruptly curved at base, ta- pering and evenly curved distally to sickle- shaped, slender, pointed distal portion, 4— 5 fine setae borne midway on inner margin. Aedeagus with basal arch rounded and ex- tending to nearly half of total length, main body with slightly convex lateral margins, tapering to slender buttonlike tip bent ven- trad. Parameres (Fig. 3g) nearly straight and moderately swollen on proximal half, nar- rowed distally and evenly curved caudolat- erad, ending in slightly broadened, bladelike distal portion. Distribution. — New Caledonia; original- ly described from a unique ° from Mine Galliéni, 166°20'55”E, 21°54'33”S, 800 m, ‘““maquis haut, sur peridotite.”’ Specimens examined.— Mt. Dzumac, 760 m, 27.11.1984, 10 9, 6 6. Five km E Grand Lac, Plaine des Lacs, 21-—25.1.1984, 3 6. Discussion.—The previously unknown male is described from a specimen from Mt. Dzumac. Clastrier (1986) in his key sepa- rates A. melanesiae from A. neocaledonien- sis Clastrier by two minor characters: scu- tellum with two pairs of setae located near midline and the genital sclerotization of the eighth sternite broad and massive in me- lanesiae, and scutellum with two pairs of setae located at lateral ends and genital sclerotization short and linear in neocale- doniensis. We have identified our series as melanesiae on the basis of the scutellar se- tae. Alluaudomyia tillierorum Clastrier Alluaudomyia tillierorum Clastrier, 1986: 191 (male, female; New Caledonia; figs.). Female diagnosis. — Wing length 0.93 mm, breadth 0.41 mm; costal ratio 0.53. A moderately large, brownish mottled species 466 with three prominent small blackish spots on wing; one just before r-m crossvein, sec- ond at end of costa and vein R4+5, and third, a linear spot along vein RI. Legs brown with prominent pale bands, narrow and subapical on femora, and broad on tib- iae; tibiae yellowish except narrow apices blackish, fore tibia with narrow sub-basal brown ring and broad extensor infuscation at midlength; mid tibia with narrow sub- basal brown ring in midportion; hind tarsal ratio 2.64. Antenna and palpus moderately long, brownish, antennal flagellomeres 2-8 narrowly pale at bases; antennal ratio 0.86; lengths of palpal segments in proportion of 16-30-34-26-33. Eyes bare; mandible with ten teeth. Spermathecae two, globose with distinct slender necks, each with moderately long diverticulum; slightly unequal, mea- suring 0.050 x 0.037 mm and 0.043 x 0.033 mm. Male diagnosis. — Wing length 0.79 mm, breadth 0.30 mm; costal ratio 0.47. Similar to female with usual sexual differences. Genitalia: Ninth sternum with caudal mar- gin convex, with coarse spicules on mem- brane subapically; ninth tergum short and tapering, caudal margin weakly bilobed, without distinct apicolateral processes. Gonocoxite arcuate, broad at base, ventral apodemes nearly touching mesad; gono- stylus long and curved, sickle-shaped, end- ing in slender point, 5-5 slender setae scat- tered along inner margin. Aedeagus with basal arch extending to about a third of total length, main body with convex lateral mar- gins, tapering abruptly on distal third to bluntly pointed tip with angular point turned ventrad. Parameres straight on proximal third, gradually broadened toward midpor- tion, then abruptly narrowed and continued as apically swollen, fingerlike process di- rected slightly ventrolaterad. Distribution. — New Caledonia. Original- ly described from 1 ¢ (holotype) and 7 ° from Vallée de la Ouinné, 730 m, humid Araucaria forest; upper trail of the Riviére PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Bleue, 230 m in humid forest; and Mine Galliéni 800 m, high scrub on peridotite. Specimens examined.— Mt. Dzumac, 760 m, 14-17-1984) 2°36, 5 o ‘sames 27= 28.11.1984, 4 6, 5 2. Five km E Grand Lac, Paline des Lacs, 300 m, forest, 22= 25.11.1984, 1 9. Ponandou River, 5 km §S touho, 100 m, 22.11.1984, 1 2. ACKNOWLEDGMENTS We are grateful to the collectors, Michael G. Pogue and Marc Epstein, for paying spe- cial attention to the collection of cerato- pogonids at their UV light while in new Cal- eonida, and to Donald W. Davis for calling to their attention our interest in New Cal- edonia biting midges. We also thank Edwin F. Cook of the University of Minnesota for permission to retain and study the New Cal- edonia collections, made while Michael Po- gue was on a travel grant from the Univer- sity of Minnesota. William L. Peters of Florida Agricultural and Mechanical Uni- versity at Tallahassee kindly provided maps of New Caledonia and much helpful liter- ature and information on its insect fauna. LITERATURE CITED Clastrier, J. 1986. Ceratopogonidae de la Nouvelle- Calédonie IV. Genre A//uaudomyia (Diptera, Nematocera). Description de six espéces nou- velles; simplification de Videntification des fe- melles. Cah. ORSTOM Ser. Entomol. Med. Par- asitol. 23: 187-201. Debenham, M. L. 1971. Australasian Ceratopogon- idae (Diptera: Nematocera). Part XV: The genus Alluaudomyia Kieffer in Australia and New Guinea. Proc. Linn. Soc. N.S.W. 96: 128-174. Downes, J. A. and W. W. Wirth. 1981. Ceratopo- gonidae [chapter] 27, pp. 393-421. Jn McAlpine, J. F. et al., eds., Manual of Nearctic Diptera. Vol. 1. Res. Branch Agric. Can. Ottawa. Monograph 27: 674 pp. Spinelli, G. R. and W. W. Wirth. 1984. The Neo- tropical predaceous midges of the genus 4//uau- domyia (Diptera: Ceratopogonidae). Proc. Ento- mol. Soc. Wash. 86: 673-702. Tokunaga, M. 1963. New Guinea biting midges (Dip- tera: Ceratopogonidae), 3. Pac. Insects 5: 211-279. Tokunaga, M. and E. K. Murachi. 1959. Insects of VOLUME 89, NUMBER 3 467 Micronesia (Diptera: Ceratopogonidae). Insects Wirth, W. W. and N. Marston. 1968. A method for Micronesia 12: 103-434. mounting small insects on microscope slides in Wirth, W. W. and M. D. Delfinado. 1964. Revision Canada balsam. Ann. Entomol. Soc. Am. 61: 783- of the Oriental species of A//uaudomyia Kieffer 784. (Diptera, Ceratopogonidae). Pac. Insects 6: 599- 648. PROC. ENTOMOL. SOC. WASH. 89(3), 1987, pp. 468-477 A NEW ARBOREAL CARABODES FROM EASTERN NORTH AMERICA (ACARI: ORIBATIDA: CARABODIDAE) R. MARCEL REEVES Department of Entomology, University of New Hampshire, Durham, New Hampshire 03824. Abstract.—Carabodes dendroetus n. sp. is described. It is the first North American member of the genus found primarily in arboreal habitats that shows a preference for the bark and branches of conifers or hardwoods rather than rotten logs and leaf litter on the forest floor. It is distributed from Nova Scotia and New Brunswick south through New England and Pennsylvania to western North Carolina. Oribatid mites reach their highest diver- sity and abundance in forest litter and soil where densities have been estimated at 55,000-—425,000/m * (Wallwork, 1983). The epigeal oribatid fauna, while not as diverse and abundant, was considered by Wallwork to offer considerably more promise in eco- logical studies because of the more isolated and simplified microhabitats available for study. Trave (1963) compared the edaphic with the epigeal fauna and found three of six species of Carabodes were present in nearly equal numbers in these two habitats. The thick, hard exoskeleton of Carabodes was suggested as the most important factor in their survival in epigeal zones. Luxton (1972) considered Carabodes to be panphy- tophages and thus better able to exploit a variety of habitats because of their wide food preferences. During an examination of organisms ex- tracted from fir and spruce branches using the hot lye process of Miller and McDougall (1968) for removal of spruce budworm lar- vae, I found the following seven species of Carabodes: granulatus Banks, labrynthicus (Michael), niger Banks, wil/manni Bernini and three undescribed species. One of these undescribed species I had previously col- lected in large numbers from sugar maple, oak and white pine bark but rarely from forest litter, and then usually as single spec- imens. Thus, I believe this new species to be primarily arboreal, hence the name ““dendroetus” from the Greek meaning tree dweller. Terminology and abbreviations are those developed by F. Grandjean, as summarized by Balogh and Mahunka (1983). All mea- surements are in micromillimeters and made from unmounted specimens. Line drawings were made from dissected specimens and may be a composite of more than a single specimen. Collection/extraction methods varied from using Berlese funnels to extract mites from litter or from sifted litter, ex- traction from branches using a hot lye so- lution and collecting mites directly from bark and foliage using a microscope. Names of the following collectors are abbreviated: D.S.C.—D. S. Chandler, A.G.—A. God- frey, R.H.H.—R. H. Hutchins, and R.M.R.—R. M. Reeves. Carabodes dendroetus Reeves, NEw SPECIES Adult. — Measurements: Mean length (measured from tip of rostrum to posterior VOLUME 89, NUMBER 3 igs: 1 2) edge of notogaster): female (n = 23) 475 (range 439-513); males (7 = 26) 419 (range 390-454). Mean notogastral width (mea- sured at widest point): females (n = 22), 262 (range 245-291); males (m = 26), 227 (range 194-252). Mean height (measured from genital-anal area dorsally to highest point of notogaster): females (n = 21) 216 (range 184-247): males (n = 24) 188 (range 168- 209). Integument: Brown. Body and antiax- ial surfaces of femora I-IV and trochanters III and IV covered with pits. Prodorsum: 469 Carabodes dendroetus, adult. 1, Dorsal view. 2, Ventral view. Prodorsal length: females (7 = 23) 141 (range 124-161); males (n = 26) 132 (range 112- 146). Prodorsum nearly completely covered with pits, pits most uniform in size and shape between rostral and lamellar setae (Fig. 15). Lamellae with very weakly developed pits medially, lateral margin smooth. Dorsal se- jugal groove narrow, pits becoming very ir- regular near groove (Fig 13). Rostral setae (ro) very minutely barbed, slightly arched mesad, 9-12 long. Lamellar setae (/e) longer (12-16), more strongly barbed and arched 470 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 3-5. pd, and pd, 4, Palpus, antiaxial view, trochanter removed. 5, Chelicera, paraxial view. mesad than ro, inserted near anterior mar- gin of lamellae (Figs. 1, 3, 15). Interlamellar setae (in) similar in shape to /e but longer (20-25), inserted on prodorsal surface above internal medial edge of lamellae. Ventral wall of bothridium with a notch (Fig. 16). Sensillus (ss) short, capitate, head spinose on distal two thirds (Fig. 16). Notogaster: Notogaster with moderately sized, round pits (2-8 diameter) separated from adjacent pits by at least (usually more than) the pit diameter (Fig. 14). Laterally and posteriorly notogastral pits become less uniform in shape, often coalescing (Figs. 11, 12, 19). All notogastral setae similar: long, thin, nearly uniform in diameter throughout, barbed and slightly enlarged distally (mi- nute barbs below tip visible only at high Carabodes dendroetus, adult. 3, Lateral view with legs removed, integumental pits not shown on magnification and in SEM Fig. 10). Setal lengths diminish posteriorly with setae p,— Pp; approximately three fourths length of ta. Setal lengths (n = 4) of ta: 24-28, te 18-25, ti: 20-24, ms: 18-24, r,: 20-24, r,: 18- 22, r;: 18-20, p,;: 16-20, p,: 16-20, p;: 16- 20. Length of ta and ti one half to two thirds distance between insertions of ta-ti and fi- ms setae respectively. Seta fa positioned an- teromediad of ¢i so that fa, ti, ms, and r, form a nearly straight longitudinal row, with te and r, forming a parallel row (Figs. 1, 11). Gnathosoma: Diarthic. Mentum with pits. Setal positions as in Figs. 2, 17. Palpal for- mula: 0-2-1-3-9 (+1 solenidion) (Fig. 4). Chelicera chelate (Fig. 5). Tragardh’s organ (Trg) present. Ventral surface: Pits on ven- tral surface similar to those on prodorsum VOLUME 89, NUMBER 3 471 Figs. 6-9. Carabodes dendroetus, adult. 6, Leg I. 7, Leg II. 8, Leg III. 9, Leg IV. Antiaxial views, trochanters removed. Carabodes dendroetus, adult. 10, Seta r, (2000 x). 11, Dorsal aspect (220 x). 12, Lateral aspect Figs. 10-12. (240 x). 472 VOLUME 89, NUMBER 3 Figs. 13-16. and notogaster; least modified in shape near center of plates (Figs. 17, 20). Epimeral plates (ep]—ep4, Fig. 2) divided by distinct epimeral furrows (Fig. 17). Epimeral setal formula 3- 1-3-3; seta /blongest (8—9), others 3-6 long. Four pairs of genital and | pair of aggenital setae (ag) present (Figs. 2, 18, 20); short (3-5), minutely barbed. Two pairs of anal (an) and 3 pairs of adanal (ad) setae present (Figs. 2, 20); an,, an>, and ad; sim- ilar (6-8 long), minutely barbed; ad, (12- 473 Carabodes dendroetus, adult. 13, Dorsal sejugal groove detail (830). 14, Pits on notogaster between setae ¢/ and ms (625 x). 15, Prodorsum, anterior view (490 x). 16, Sensillus (3530 x). 15) and ad, (10-13) similar in shape to dor- sal notogastral setae, but shorter. Lateral surface: Most of the lateral surface below the lamella, bothridium and antero-lateral edge of the notogaster and above acetabula I-IV with small tubercles (Fig. 3). Tutorium narrow s-shaped ridge, becoming more dis- tant from lamellar edge anteriorly. Carina extending from near middle of tutorium ventrally and anteriorly to below insertion of ro (Fig. 3). Pedotectum I (pd,) covering 474 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 17-20. Carabodes dendroetus, adult. 17, Ventral view of gnathosoma and epimera (590 x). 18, Genital plates (1330). 19, Posterior view of notogaster (445 x). 20, Ventral view of genital anal area (770). base of acetabulum I, widest ventrally, ta- pering dorsally to near bothridial base. Exo- bothridial seta (ex) absent (tubercles in area where ex should be, make presence of al- veolar remnant of ex difficult to determine). Pedotectum II (pd,) a short, blunt tooth par- tially covering acetabulum II. A short, blunt discidium (d/) is present between acetabula III and IV. Legs: Pits present on the an- tiaxial surfaces of femora I and II and tro- chanters and femora III and IV (Figs. 6-9, 12). Porose areas on femora I-IV and tro- chanters III and IV on paraxial surface. Se- tation (I to IV, solenidia in brackets), tro- chanters 1-1-2-1, femora 4-4-3-2, genua 3(1)-3(1)-1(1)-2, tibiae 4(2)-3(1)-2(1)-2(1), tarsi 15(2)-15(2)-15-12. One dissected spec- imen lacking ft” on one tarsus III. Ventro- distal edge of femora III and IV with well developed spur (Figs. 8, 9). Tarsi mono- dactyl. Immatures. — Unknown. VOLUME 89, NUMBER 3 Fig. 21. Known distribution of Carabodes dendroetus. 475 476 Material examined.— Holotype: adult 4, USA, NEW HAMPSHIRE, Carroll Co., 2.5 mi NW Wonalancet, The Bowl, VII-30-85, D.S.C., sifted rotten beech logs; deposited in Canadian National Collection (Type no. 19463). Paratypes: | °, same data as holo- type except VI-21-85, sifted fir/hemlock leaf litter; 1 9, 1 mi NW Wonalancet, Spring Brook, VII-10-85, D.S.C, sifted rotten hem- lock/fir logs; 1 3, same data except IX-5-85; 1 4, same data except V-15-85, sifted litter along stream; | 6, Five Fingers Pt., Squam Lake, Sandwich, X-12-64, R.M.R., humus at base of white pine; | ?, Belknap Co., S. Barnstead, VII-2-65, R.M.R., Quercus ru- bra foliage; 1 6, Coos Co., Hellgate, Dead Diamond R., VII-13-74, R.M.R., moss on ground in spruce-fir stand; 6 6, 8 2, 1 sex undetermined, 3 mi NE Errol (Rt. 16), IX- 3/4-83, A.G., extracted from spruce branch- es; 6 4, 8 2, 6 sex undetermined, same data except extracted from fir branches; 1 4, 17 mi N Crystal, [X-9/11-83, A.G., extracted from spruce branches; 2 sex undetermined, same data except extracted from fir branch- es; 1 2, 1 sex undetermined, 6 mi SW Errol (Rt. 16), [X-3/4-83, A.G., extracted from fir branches; | °, 2 sex undetermined, 5 mi NE Errol (Rt. 16), [X-3/4-83, A.G., extract- ed from spruce branches; 3 @, 1 sex unde- termined, 13 mi. NW Errol, Dartmouth Grant, IX-9/11-83, A.G., extracted from fir branches; | 3, | sex undetermined, 8 mi NW Errol (Rt. 16), [X-9/11-83, A.G., extracted from fir branches; | 4, 1 2, same data except IX-3/4-83; 1 6, 10 mi N Milan (Rt. 16) IX- 3/4-83, A.G., extracted from spruce branch- es; 2 6, 1 2, Grafton Co., Campton, I-6-72, R.H.H., white cedar; | sex undetermined, Ellsworth, VIII-1-66, R.H.H., sugar maple, bark; 1 8, 1 2, same data except VI-15-66; 4 6,42, Plymouth, IV-30-70, R.H.H., sugar maple bark; 1 9°, Strafford Co., Somers- worth, IV-13-77, R.M.R., pine-oak litter; 2 6,2 sex undetermined, Durham, II-17-69, R.M.R., dead bark from top of oak; 24 4, 11 2, 7 sex undetermined, Durham, IX-14- 70, R. M. R., oak bark, cordwood; 6 4, 5 8, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 10 sex undetermined, Durham, V-6-71, R.M.R., white pine bark; | 6, NORTH CAROLINA, Great Smoky Mts. Nat. Pk., 4000’, IlI-27-75, D. R. Blais, litter at base of tree; 1 6, Coweeta Hyd. Sta., Shope Fork Rd., 3000’, V-30-83, D.S.C., sifted oak and ash litter; 1 6, same data except 2450’, V-29-83, sifted oak and cherry litter; 1 9, PENNSYLVANIA, Somerset Co., 3 mi E Berlin, VII-10-84, D.S.C., sifted rotten wood: | 4, | ¢, Warren Co., Hearts Content Nat. Area, Allegheny Nat. For., V-29-85, R.M.R., white pine litter; 1 6, Huntington Co., Alan Seeger Nat. Area, V-30-85, R.M.R., bark of chestnut oak; 2 sex unde- termined, CANADA, NEW BRUNS- WICK, 8 mi W Lake George, VII-18-68, E. E. Lindquist, moss and lichens on white pine trunk; | sex undetermined, same data except under bark white pine infested with Ips pini; 2 sex undetermined, NOVA SCO- TIA, Cape Breton Highlands Nat. Pk., Lone Shieling, VIII-26-83, M. Sharkey, soil, litter and moss. Paratypes are deposited in the United States National Museum, Washing- ton, D.C., Museum of Comparative Zool- ogy, Cambridge, Massachusetts, Canadian National Collection, Biosystematics Re- search Centre, Ottawa, and author’s collec- tion. This species is found in eastern North America from New Brunswick and Nova Scotia south to North Carolina (Fig. 21). Remarks. — Carabodes dendroetus is most similar to those species of the genus that have ta, ti, ms and r, in a longitudinal row, all notogastral setae similar in shape and lack a deep cervical cavity. The long rod- like, distally barbed notogastral setae, size and spacing of pitting and the short, barbed capitate sensillus will distinguish C. den- droetus from other Carabodes species in North America. The leg setal formula of C. dendroetus was compared to that of C. niger Banks (Norton, 1978), C. minusculus Berlese (Bernini, 1976) and C. will/manni Bernini (Bernini, 1975) and found to be similar. The positions of (pv) on tarsus I and (pv) and (ff) VOLUME 89, NUMBER 3 on tarsus II in C. dendroetus are more sim- ilar to C. niger than to C. minusculus. Also, the size and shape of (v) on tibia I does not vary in C. dendroetus and C. niger as much as in C. minusculus. The ventrodistal spur of femur III and IV is more developed in C. dendroetus than in either C. niger or C. minusculus. ACKNOWLEDGMENTS Scientific Contribution Number 1466 from the New Hampshire Agricultural Ex- periment Station. The Central University Research Fund, University of New Hampshire, provided fi- nancial support for the scanning electron micrographs used in this description. I thank the many collectors for providing material for examination, in particular, April God- frey and Donald S. Chandler. I especially acknowledge Valerie Behan-Pelletier, Bio- systematics Research Centre, Ottawa, Can- ada, for encouragement and help in the preparation and review of the manuscript. I thank also Donald S. Chandler and John 477 F. Burger, University of New Hampshire, for review of the manuscript. LITERATURE CITED Balogh, J. and S. Mahunka. 1983. The Soil Mites of the World, Vol. 1: Primitive Oribatids of the Pa- laearctic Region. Elsevier: New York. 372 pp. Bernini, F. 1975. Notulae oribatologicae XII. Una nuova specie di Carabodes affine a C. minusculus Berlese 1923 (Acarida, Oribatei). Redia 56: 455- 471 and pls. 1-4. . 1976. Notulae oribatologicae XIV. Revisione di Carabodes minusculus Berlese 1923 (Acarida, Oribatei). Redia 59: 1-49 and pls. 1-9. Luxton, M. 1972. Studies on the oribatid mites of a Danish beechwood soil. I. Nutritional biology. Pe- dobiologia 12: 434-463. Miller, C. A. and G. A. McDougall. 1968. A new sampling technique for spruce budworm larvae. Can. Dep. For. Rur. Dev., Ottawa, Ont. Bi-Mon. Res. Notes 24: 30-31. Norton, R. A. 1978. Notes on Nathan Banks species of the mite genus Carabodes (Acari: Oribatel). Proc. Entomol. Soc. Wash. 80: 611-615. Travé, J. 1963. Ecologie et biologie des Oribates (Acariens) saxicole et arboricole. Vie Milieu. 14: 1-267. Wallwork, J. A. 1983. Oribatids in forest ecosystems. Ann. Rev. Entomol. 28: 109-130. PROC. ENTOMOL. SOC. WASH. 89(3), 1987, pp. 478-482 POPULATIONS OF TRAP-NESTING WASPS NEAR A MAJOR SOURCE OF FLUORIDE EMISSIONS IN WESTERN TENNESSEE W. NELSON BEYER, GARY W. MILLER, AND W. JAMES FLEMING Patuxent Wildlife Research Center, U.S. Fish and Wildlife Service, Laurel, Maryland 20708. Abstract. —Trap-nesting wasps were collected from eight sites at distances of from 1.2- 33.0 km from an aluminum reduction plant in western Tennessee. The sites had similar topographies, soils, and vegetation, but differed in their exposure to fluoride, which was emitted in large quantities from the plant. It was postulated that if fluoride emissions had greatly changed the insect community then relative densities of their predators would have varied accordingly. However, the degree of fluoride pollution was unrelated to the relative densities of the wasps and to the number of cells provisioned with prey. Monobia quadridens, Trypargilum clavatum, and T. lactitarse were found to have two complete generations in western Tennessee. 7rypargilum collinum rubrocinctum has at least two generations, and Euodynerus megaera probably has three generations. Six other wasp species and a megachilid bee were also collected. The effect of an environmental contam- inant, such as fluoride, on insect commu- nities is difficult to evaluate because of the diversity of insect species and the natural fluctuations in their populations. Levin (1982) and Moriarty (1983) suggested a need for toxicological studies on the functioning of ecosystems, rather than just investiga- tions on the effects of chemicals on indi- viduals or on populations of particular species. Our approach was to study “‘trap- nesting” wasps, described by Krombein (1967), at several sites having a gradient of exposure to fluoride. If the composition of an insect community were greatly changed because of environmental contamination, it would be reasonable to expect changes in the populations of these predators. Wood- well (1970) suggested that “obligate carni- vores high in the trophic structure, are at a disadvantage because the food chain con- centrates the toxin and, what is even more important, because the entire structure be- neath them becomes unstable.” Although this statement may be an oversimplifica- tion, since many ecological and physiolog- ical factors influence the movement of en- vironmental contaminants in food chains, the choice of a group of predators for study- ing the effects of fluoride seems reasonable. The life histories and prey of the species studied here are known (Krombein, 1967). They belong to several families, but have similar nesting habits. All seek out holes in wood, where they construct cells, parti- tioned with soil or debris. Cells are provi- sioned with prey, an egg laid, and the cells and hole sealed. Adults emerge either a few weeks after the egg is laid or the following year, after the larvae have overwintered. These wasps can be readily studied because they can be attracted to wooden blocks that have holes drilled in the ends, called “‘trap- nests.”” In addition to providing informa- tion on species distributions, life histories, prey, nesting habits, and parasites, the trap- VOLUME 89, NUMBER 3 nesting technique seems to have the poten- tial for comparing relative densities of wasps at different sites. The technique may prove suitable for studying community indices, such as species diversity. Our intent was to determine whether pop- ulations of trap-nesting wasps were reduced at sites greatly polluted with fluoride. The study was conducted concurrently with studies by the Patuxent Wildlife Research Center on the effects of fluoride on song- birds and on the decomposition of the litter horizon. We also hoped to gain information on the life histories of these wasps in Ten- nessee. METHODS Eight sites were selected at various dis- tances from the Consolidated Aluminum Company’s (CONALCO) aluminum reduc- tion plant on Kentucky Lake, in Humphreys County, Tennessee. The plant emits more than 3000 tonnes of fluoride per year when operating at peak capacity; from 1980 to 1984 the plant operated at about 25-50% of capacity and the fluoride emissions were reduced proportionally (Tennessee Valley Authority, unpublished data). Compared to emissions at other industrial plants in the United States these emissions are very high. Sites 1, 2, and 5 were in Humphreys Co.; 3, 4, and 6 in Benton Co.; and 7 and 8 in Henry Co. Sites 1-8 were located at 1.2, 1.4, 3.6, 4.4, 4.5, 8.8, 32, and 33 km from the plant (Fig. 1). From the study of the effects of fluoride on the decomposition of litter we know that the acid-extractable fluoride con- centrations in the litter decreased progres- sively with the distance of the sites from the plant (Sites 1-8; 695, 440, 107, 102, 50, 46, 16, 12 mg/kg F). Sites 1 and 2 should be considered very contaminated. All sites were on upland ridges or slopes, had similar soils, and were dominated by a mixture of red and white oaks (Quercus spp.) and shagbark hickory (Carya ovata [Mill.] K. Koch). The sites were selected so that the only apparent ecological difference was the exposure to the 479 y 10 km N CONALCO Plant 2 i Ay Si Evac IS o%* y ; 4 Ba \ we Fig. 1. The 8 sampling sites located at distances of from 1.2 to 33 km from the CONALCO plant in north- western Tennessee. emissions from the aluminum reduction plant. In April 1984 traps were set out in a Strat- ified random manner. At each site we laid down a 100-m transect with six predesig- nated randomly located points, one in each of six equal lengths of the transect. Each randomly selected point that fell within 5 m of another point was replaced by another randomly selected point. We selected the tree closest to each point and mounted a platform to hold each set of traps about 1.5 m high. The traps were cut from clear spruce Q.76% 2:7 ¥. 16:0.cm) Pompilidae SS) Dipogon sayi Banks Sphecidae Podium rufipes Fabricius 0 Isodontia auripes (Fernald) 0) Trypargilum clavatum (Say) 5 Trypargilum lactitarse (Saussure)* 5 Trypargilum collinum rubrocinctum (Packard) } Megachilidae Osmia pumila Cresson 0 Unidentified | Total occurrences i) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Occurrence of wasps and bees in wooden trap-nests at eight sites located at different distances from No. Traps Used by Species at Sites 1-8 2 3 4 5 6 7 8 Total 0 0 l 0 0 0 0 1 0 0 0 0 0 0 2 2 2 0 0 0 0 0 0 2 Z 0 3 3 l 0 2 1] 3 0 2 0 5 5 l 16 0 l 0 0 0 0 0 l 0 0 0 0 0 0 3 3 0 0 0 3 0 0 4 di 2 8 Z 3 1 3 l 25 0 0 1 Th 2D 5 0 18 6 l | 2 0 - 1 17 0 2 0 0 0 0 0 2 0 l 3 l 3 2D 0 11 15 13 13 Ss 12 17 14 116 * Cited as T. striatum (Provancher) (Krombein, 1967), a junior synonym. gether and attached to the platform with rubber bands. About once every three weeks, from May 13 to September 13, traps with nests were collected and replaced by empty ones. The open end of the collected traps was wrapped in a nylon screen and secured with a rubber band to prevent the emerged wasps from escaping. Some of the larger wasps chewed through the nylon, so wire screens were added to some of the traps. To break the diapause of larvae that did not emerge in 1984 we moved the traps with closed cells to an unheated greenhouse on October 5, and brought them back inside on March 3. Traps with cells that were still closed by August 1985 were opened. Emerging adults were keyed to genus or to species (when keys were available), and then compared with specimens in the National Museum of Nat- ural History, Washington, D.C. Nomencla- ture (Table 1) follows Krombein et al. (1979). For some species the architecture of the nest (Isodontia auripes) or the shape of the co- coon (7rypargilum) was distinctive enough to permit identification. Wasps from about 10% of the traps could not be identified: these individuals had been parasitized, had died for other reasons, or had escaped from the traps. Spearman rank correlation coefficients (Conover, 1980) were used to determine whether the distances of the sites from the plant were related to either the number of traps used or the number of cells provi- sioned per site. Additional analyses were planned only if the correlation coefficients were significantly different from zero. RESULTS AND DISCUSSION Fluoride emissions seemed to have no discernible effect on the populations of trap- nesting wasps. The number of trap nests used by wasps was about the same at each VOLUME 89, NUMBER 3 site, varying only from 12 to 19 (Table 1). An average of 14 traps was used at sites | and 2, which 1s close to the average of 14.7 for the six sites farther from the plant. The number of traps used was not significantly correlated with the distance of the sites from the plant (Spearman rank correlation coef- ficient = 0.00, P > 0.05). Sites 1 and 2 were exposed to extremely high concentrations of fluoride, but we collected 6 of the total of 11 species of wasps at one or the other of these sites. We cannot rule out the pos- sibility that populations of particular species of wasps have been reduced or eliminated close to the plant. However, when all trap- nesting wasps are considered as a unit we conclude that the populations are just as high close to the plant as far from it. The numbers of provisioned cells per site provide a second way of evaluating the ef- fects of fluoride on trap-nesting wasps. They were as follows: site 1-43, site 2-39, site 3- 61, site 4-41, site 5-81, site 6-37, site 7- 69, site 8-44. These numbers were not sig- nificantly correlated with distances of the sites from the plant (Spearman rank cor- relation coefficient = 0.26, P > 0.05). Many invertebrate prey were required to support the wasp populations. For example, wasps of the three species of 7rypargilum we col- lected provision their cells with means rang- ing from | 1 to 16 spiders (Krombein, 1967). These species provisioned an average of 3.6 cells per trap in Tennessee. Each spider re- quires many prey, sO we estimate that the 60 Trypargilum collected in the study were dependent on many thousands of prey. Di- pogon Ssayi also preys on spiders. All of the eumenids prey on caterpillars, Podium ru- fipes preys on cockroaches, and [sodontia auripes preys on Orthoptera (Krombein, 1967). Field studies relating fluoride to popula- tions of other kinds of insects have been reviewed by Alstad et al. (1982). Popula- tions of some herbivores, such as the pine bud moth (Exoteleia dodecella), increased in areas polluted with fluoride, presumably 481 either because the host plants were weak- ened or because the numbers of parasites or predators were suppressed. Populations of silkworms, honeybees, some bark beetles (Pityokteines) and the European pine shoot moth (Rhyacionia buoliana) were shown to be lower than normal in areas polluted with fluoride. In a study of invertebrates collect- ed near an aluminum reduction plant, scav- engers tended to have the highest concen- trations of fluoride, followed by predators, omnivores, and herbivores (Buse, 1986). There are conflicting opinions on whether fluoride is accumulated in food chains (Al- stad et al., 1982). Hymenoptera as a group tend to accumulate relatively high concen- trations of fluoride, compared to other kinds of insects (Dewey, 1973). However, the main route of exposure (respiration, food, groom- ing) of fluoride to Hymenoptera has not been identified (Alstad et al., 1982), and the trap- nesting wasps may have a much lower ex- posure to flouride than species such as hon- eybees, which forage for pollen. The species we obtained had all been col- lected previously in trap-nests (Krombein, 1967). Five of the species nested frequently enough to permit generalization about their life histories in Tennessee. Monobia quad- ridens, Trypargilum clavatum, and T. lac- titarse have two complete generations, the first emerging mainly in August, and the second the following spring. /. quadridens and 7. lJactitarse selected the 12.7 mm bor- ings almost exclusively, and 7. clavatum showed a preference for the 6.4 mm borings but selected some of the 4.8 mm borings. These species probably have a single gen- eration in northern states, but two genera- tions in Florida and North Carolina (Krom- bein, 1967). The seasonal changes in the populations of 7rypargilum collinum ru- brocinctum were less clear, and all that we could conclude was that there were at least two generations per year. This species se- lected the 4.8 mm borings almost exclu- sively. Euodynerus megaera is bivoltine in North Carolina (Krombein, 1967). Several 482 traps collected from Tennessee in June had wasps emerging in July, and several traps collected in July and August had wasps emerging in August or September. None of the wasps emerged in the spring, which sug- gests that there was a third generation in the fall that we did not obtain. This species showed a strong preference for the 6.4 mm borings. About 13% of the traps were parasitized by Diptera, Encyrtidae, or other parasites. The mutillid wasp, Sphaeropthalma pen- sylvanica (Lepeletier), parasitized nests of T. clavatum and T. lactitarse, as previously reported (Krombein, 1967), but it also par- asitized a nest of /sodontia auripes. In evaluating the trap-nesting technique for studying the effect of contaminants on insect populations we find two drawbacks. Combining all of the wasps yields large sam- ple sizes, but we note that the species feed on prey belonging to different food chains. Generalizing requires disregarding many of the important differences between species, if we are to examine the functioning of the whole insect community. Also, although we assume that the numbers of wasps collected in the traps vary with relative densities, there may be other ecological variables, such as the number of suitable natural holes pres- ent, that would make the traps more at- tractive to wasps at some sites than at oth- ers. The advantage of the trap-nesting tech- nique is that the results are relevant to the community. Populations of forest insects are frequently variable, making it difficult to relate a pollutant to changes in a population (Alstad et al., 1982). Unless all of the pop- ulations of individual species vary in syn- chrony with each other, the combined pop- ulations of several species should be more stable and easier to evaluate than the pop- ulations of individual species. If the species PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON have overlapping niches then the function- ing of the community may remain similar even if the populations of the species change relative to each other. The technique also enables sampling of populations over their entire season. If the populations of trap- nesting wasps were reduced at particular sites then it would be reasonable to examine community variables, such as species di- versity. ACKNOWLEDGMENTS We thank Donald R. Clark and Matthew C. Perry for reviewing the manuscript, Ter- ry Griswold for identifying Osmia pumila, and Arnold Menke for permitting us to ex- amine the wasp collection at the National Museum of Natural History. LITERATURE CITED Alstad, D. N., G. F. Edmunds, Jr., and L. H. Weinstein. 1982. Effects of air pollutants on insect popula- tions. Ann. Rev. Entomol. 27: 369-384. Buse, A. 1986. Fluoride accumulation in inverte- brates near an aluminum reduction plant in Wales. Environ. Pollut. Ser. A Ecol. Biol. 41: 199-217. Conover, W. J. 1980. Practical Nonparametric Sta- tistics. John Wiley & Sons, New York, 493 pp. Dewey, J. E. 1973. Accumulation of fluorides by in- sects near an emission source in western Montana. Environ. Entomol. 2: 179-182. Krombein, K. V. 1967. Trap-Nesting Wasps and Bees: Life Histories, Nests, and Associates. Smithsonian Press, Washington, D.C. 1i-—vi & 570 pp. Krombein, K. V., P. D. Hurd, Jr., D. R. Smith, and B. D. Burks. 1979. Catalog of Hymenoptera in America North of Mexico. Vol 2, Apocrita (Acu- leata). Smithsonian Institution Press, Washington, D.C. 2209 pp. Levin, S. A. (ed.) 1982. New Perspectives in Ecotox- icology. Ecosystems Research Center, Cornell University, Ithaca, N.Y., ERC Report No. 14. Moriarty, F. 1983. Ecotoxicology, the Study of Pol- lutants in Ecosystems. Academic Press, New York, 233 pp. Woodwell, G. M. 1970. Effects of pollution on the structure and physiology of ecosystems. Science 168: 429-433. PROC. ENTOMOL. SOC. WASH. 89(3), 1987, pp. 483-488 TWO NEW SPECIES OF BOTOCUDO FROM VERTICAL ROCK FACES IN INDONESIA (HEMIPTERA: LYGAEIDAE) JAMES A. SLATER AND DAN A. POLHEMUS (JAS) Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs, Connecticut 06268; (DAP) University of Colorado Museum, 3115 South York Street, Englewood, Colorado 80110. Abstract. — Botocudo hebrodes and Botocudo polhemusi are described as new species from Bacan Island, Indonesia. Both species were taken living together on vertical rock faces adjacent to a flowing river near the coastal plain. The habitat was that usually occupied by species of Hebridae. A discussion of possible feeding habits and of the occurrence of other lygaeids that are known to live on vertical rock faces is included. Figures include a dorsal view and details of the abdomen and of the male genitalia. Detricolous lygaeids in the wet tropics often must adopt unusual strategies for sur- vival because the mature seeds upon which they feed usually either germinate quickly or are attacked by fungi once they have reached the ground. Thus we find species, in genera whose members are usually ground litter bugs, feeding on seeds in bird drop- pings on leaves, on seeds in bat guano in caves, and especially upon plants that retain mature seeds on the plants for a consider- able period of time. Recently the junior author and his father discovered the lygaeids described in this pa- per living in a most unusual habitat on the island of Bacan in Indonesia (Bacan is the current Indonesian name for the island that in most past biographic and systematic pa- pers has been called Batchian). It is just west of the large island of Halmahera. One of the most effective methods of collecting many species of Hebridae and Veliidae is to splash water upon vertical rock faces to wash the insects into a net held below. On September 23, 1985, the junior author and his father were using this technique on Bacan Island in the hope of obtaining specimens of He- bridae. The site was at less than 100 meters elevation just at the point where the narrow coastal plain with its coconut plantations meets the first forested outliers of the Gun- ung Sabela massif. Here a swift shallow riv- er flows through a narrow gap between large rock outcrops. The vertical outcrops are of a tan sedimentary rock (probably sand- stone) and are partially shaded by disturbed primary rain forest. From these rocks the collecting party washed down a series of tiny insects that they thought were unusual look- ing hebrids or veliids since they occupied a typical hebrid/veliid habitat on the rocks just above the water line. It 1s of great in- terest that not only did the insects prove to be antillocorine lygaeids of the genus Bo- tocudo, but that two closely related species were present and that hebrids were absent. The absence of Hebridae (small Veliidae were present) is in itself unusual since they did occur on the mountain at 350 meters. The presence of the lygaeids in such an unusual habitat and the absence of Hebri- dae raised several interesting questions. What were the lygaeids feeding on? Were they competitively excluding the Hebridae? 484 What was their relationship to the veliids and were the two lygaeid species in direct competition with each other? While no def- inite answers can be given, some probabil- ities can. First, the lygaeid species show no morphological modifications that suggest an adaptation to the unusual habitat, nor to any modification in their feeding habits. All members of the tribe to which these lygaeids belong (including several congeneric species) feed on mature seeds of angiosperms. While the collecting party did not see any signs of vegetation on the rock face it is highly prob- able that the tiny insects were feeding either on minute seeds falling from the vegetation above or upon seeds continually being washed against the sides of the stream. The presence of numerous adults but no nymphs suggests the exploitation of a temporary food source. While Lygaeidae living on vertical rock faces is most unusual it is not unprece- dented. Baranowski and Slater (in press) de- scribe a new genus and species of antillo- corine from Trinidad that feeds upon the seeds of Pilea microphylla (L.) Liebm. that grows on the vertical rock or concrete sur- faces of culverts, bridges and in rocky areas near streams and waterfalls. These authors collected this species together with a mem- ber of the Ozophora pallescens complex and a species of Botocudo by placing a net below the Pi/ea plants on the vertical faces and brushing the insects into the net held below. The senior author of the present paper also took a long series of a species of Bo- tocudo running over rocks in a dry stream bed at El Valle, Panama, during the dry sea- son. These small antillocorine lygaeids, species of which also inhabit caves (Slater, 1984), thus appear to be, at least in part, opportunists that rapidly move in to exploit a seed crop. We conclude that the Bacan Island species are probably not predatory and thus would not be in competition with Hebridae. On the other hand, they might PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON well be prey items of the associated Veli- idae. That two very closely related species of Botocudo were taken also suggests the ex- ploitation of a temporary seed source. It is difficult to believe that if both species were primarily adapted to living on these vertical rock faces that they would not be in direct competition with one another. Both species are similar in size, shape and color and have mouth parts of approximately the same length. It is not unusual for closely related seed feeding lygaeids to be found together (even as breeding assemblages) but when this occurs it always appears to involve an abundant and temporary food source. Many species of Sti/bocoris in Africa and Ozo- phora in the Neotropics are striking ex- amples. RELATIONSHIP OF THE LYGAEID SPECIES As noted above neither of the Botocudo species described below shows any unusual morphological adaptations. The abdominal spiracle and trichobothrial patterns are found in many other species of the genus. The sperm reservoir (Fig. 2) 1s generalized. The genus Botocudo may not be a mono- phyletic taxon as presently delimited but these two species are certainly part of a monophyletic group of species which is found in the Orient, Australia, and the Neo- tropics. All measurements are given in mm. CL numbers following locality data refer to codes used by the junior author to reference ecological notes. Botocudo hebrodes Slater and D. Polhemus, New SPECIES Body elliptical (Fig.1). Head dull red. Pronotum and scutellum with ground color chocolate brown. Pronotum marked with yellow as follows: a transverse stripe along anterior margin from meson laterad to inner edge of compound eye; an irregular macula near center of pronotum; extreme humeral 485 neo VOLUME 89, NUMBER 3 ) Se Botocudo hebrodes, dorsal view. rig. I. 486 angles. A large diffuse reddish brown mac- ula present behind dark calli. Extreme pos- terior margin of pronotum pale. Posterior one-third of scutellum white. Hemelytra white or very pale yellow with strongly con- trasting dark brown punctures. Corium with three dark brown maculae, one along lateral margin at level of apex of scutellum, a small one along apical corial margin at place of maximum concavity of margin and a large subapical macula covering entire distal por- tion of corlum. Membrane uniformly hya- line. Lateral and ventral surfaces dark choc- olate brown. Legs (including femora) and antennae pale yellow. Proximal one-third to one-half of third antennal segment reddish brown. Head subshining, pronotum and scutel- lum somewhat pruinose, evenly and con- spicuously punctate. Dorsal surface thickly clothed with short conspicuous upright sil- very hairs. Head moderately declivent anteriorly. Tylus extending to middle of first antennal segment. Vertex convex, eyes large and ses- sile. Length head 0.46, width 1.24, inter- ocular space 0.34. Pronotum lacking a well defined anterior collar and without a transverse impression. Anterior pronotal lobe not strongly convex. Lateral margins of pronotum calloused but not sharply carinate, strongly and almost evenly narrowing from humeral angles to anterior margin. Length pronotum 0.60, width 1.20. Scutellum lacking a median carina. Length scutellum 0.74, width 0.74. Clavus with three rows of punctures. Length claval com- missure 0.20. Lateral corial margins evenly narrowing posteriorly, not sinuate. Midline distance apex clavus-apex corium 0.62. Midline distance apex corium-apex abdo- men 0.48. Metathoracic scent gland auricle strongly bent posteriorly. Evaporative area small, forming an arc around auricle, with outer margin rounded not covering posterior lobe of metapleuron. Fore femur with four very small spines PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ventrally on distal half (appearing mutic in most views) plus a row of close set spinules running entire length on inner ventral sur- face of femur. Labium reaching between mesocoxae. Length labial segments I 0.28, II 0.42, 1110.24, IV 0.22. Gular trough elon- gate, extending entire ventral length of head. Antennae slender, terete, fourth segment not thickened nor markedly fusiform. Length antennal segments I 0.32, II 0.62, III 0.52, IV 0.56. Total body length 2.92. Abdomen with well-developed scent gland scars between terga 3-4, 4-5, and 5-6, that between 5 and 6 the largest. Inner latero- tergites present on segments 4, 5, and 6 (Fig. 7). Spiracles 2 and 4 located on sternal shelf, those on segments 3 and 5 below shelf (Fig. 3). Trichobothria of sterna 4 and 5 arranged in a linear row, two posterior trichobothria both anterior to spiracle 5 (Fig. 3). Paramere (Fig. 6) very small, trianguloid, and with a short distally rounded thumb-like posterior projection. Sperm reservoir as in polhemusi. Opening of genital capsule as in Fig. 4. Holotype: 6. INDONESIA: Bacan Is.: Maluku Province, Kab. Maluku Utara., swift shallow river 3 Km. S. of Labuha nr police station. 50-100 M. 23.1X.11985 (CL2134) (J. T. and D. A. Polhemus). In National Museum of Natural History (USNM). Type No. 100060. Paratypes: 7 6, 14 2, same data as holo- type. In J. A. Slater and J. Polhemus col- lections. This species is readily separable from all known species in the Eastern Hemisphere. China (1930) notes that Botocudo swezeyi from Samoa is readily distinguished from all other species by the uniformly red-brown pronotum and scutellum. This is not strictly true if the entire Oriental and Australian fauna is considered. It is true that many of the Botocudo species do have a bicolored pronotum (/ongicornis Barber; flavicornis Signoret; yasumatsul, japonicus, and _for- mosanus Hidaka; assimulans Bergroth; pa- tricius, signandus, and fraternus Distant). From the remaining described species he-— brodes differs as follows; swezeyi China has - VOLUME 89, NUMBER 3 487 Figs. 2-7. view. 4, B. hebrodes, genital capsule, dorsal view. 5, B. polhemusi, paramere. 6, B. hebrodes, paramere. 7, B. hebrodes, abdomen, dorsal view. a dark fuscous brown fourth antennal seg- ment; marianensis Usinger has a pale v-shaped mark on the scutellum, or (var. scutellatus) the entire posterior one-half of the scutellum is yellow; rennellensis Scud- der also has the distal one-half of the scu- tellum pale yellow and has the entire distal one-half of the corium and the end of the clavus dark brown; validulus Bergroth has the first antennal segment almost as long as segment three and apparently has only the apex of the corlum with a dark macula; pronotalis has the entire pronotum yellow; scudderi has a similar corial coloration as hebrodes but a shining rather than hirsute and more sub-quadrate shaped pronotum and dark brown femora. Botocudo polhemusi Slater and D. Polhemus, NEw SPECIES Very similar to Botocudo hebrodes in size and general coloration. Pronotum entirely dark, anterior collar narrowly reddish brown. Apical white macula occupying en- ure distal one-third of scutellum (with ex- ception of marginal punctures) and sharply truncate across anterior end. Dark hemel- Botocudo spp. 2, B. polhemusi, sperm reservoir, dorsal view. 3, B. hebrodes, abdomen, lateral ytral maculae midway along costal margin and adjacent to concave region of apical corial margin larger than in hebrodes. All femora reddish brown becoming paler dis- tally. Tibiae, tarsi, antennae, and labium pale yellow. Dorsal body surface nearly gla- brous, pronotum lacking a dense covering of distinct upstanding hairs. Tylus attaining middle of first antennal segment. Vertex convex. Length head 0.34, width 0.58, interocular space 0.30. Anterior pronotal lobe appreciably swollen above posterior lobe and evenly so across middle. Anterior collar area narrow and well defined posteriorly by a line of coarse punctures. Lateral pronotal margins sinuate, obtusely calloused; posterior margin concave. Length pronotum 0.58, width 1.04. Length scutel- lum 0.60, width 0.64. Hemelytral shape as in hebrodes. Length claval commissure 0.16. Midline distance apex clavus-apex corlum 0.50. Midline distance apex corium-apex membrane 0.42. Metathoracic scent gland auricle, fore femora and evaporative area as in hebrodes. Labium barely attaining ante- rior margin of mesocoxae. Length labial seg- ments 10:22" I0!30: Il 0:18, TV-0.18. 488 Antennae relatively short and stout, seg- ment II enlarged at distal end, segments III and IV fusiform. Length antennal segments I 0.30, II 0.40, III 0.34, IV 0.44. Total body length 2.52. Sperm reservoir with conventionally shaped elliptical cup and large wings that curve posteriorly to end in distal enlarge- ment (Fig. 2). No evident holding sclerites. Vesica with three prominent coils and a well developed helicoid process. Paramere very small, blade straight and tapered to a sharp point; posterior projection elongately pro- truding and truncate at distal end (Fig. 5). Abdominal segmentation, placement of trichobothria, spiracles and shape of open- ing of genital capsule as in hebrodes. Holotype: 6. INDONESIA: Bacan I. Ma- luku Province, Kab. Maluku Utara., swift shallow river 3 Km S. of Labuha nr. police station. 50-100 M. 23.1X.1985 (CL2134) (J. T. and D. A. Polhemus). In National Museum of Natural History (USNM). Type No. 100061. Paratypes: 3 4, 1 2. Same data as holotype. In J. A. Slater and J. Polhemus collections. We are pleased to name this species for John Polhemus in recognition of his many contributions to Hemipterology. It is remarkable that two such similar and apparently closely related species should be taken together in the same unusual habitat. Botocudo polhemusi may be distinguished from hebrodes by the much more swollen anterior pronotal lobe; by the lack ofa dense clothing of upright hairs on the surface of the pronotum and scutellum and by the rel- atively much shorter antennal segments. This can be expressed by the ratio of the second antennal segment to the interocular space. In hebrodes the second antennal seg- ment is always considerably more than one and one-half times the interocular distance, mean 1.73 (n = 5), whereas in polhemusi the mean is 1.40 (n = 4) and there is no overlap in the ratios. A glance at the mea- surements of the holotypes will clearly show that the third and fourth antennal segments PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON are also much shorter in po/hemusi than they are in hebrodes. The paramere of pol- hemusi has a longer and more truncated posterior projection (Fig. 5) than does that of hebrodes (Fig. 6). In addition to these structural differences there are color differ- ences that will separate the two species. B. hebrodes (Fig. 1) has considerable pale col- oration on the pronotum, the white apical scutellar macula has an irregular anterior margin and the femora are uniformly pale yellow. In polhemusi the pronotum is uni- formly dark, the anterior margin of the white scutellar macula is evenly truncate and the femora are for the most part reddish brown. Botocudo scudderi Slater (described from Ceylon) is apparently a member of this com- plex and agrees with po/hemusi in having brown femora and a non-hirsute pronotum and scutellum. However, scudderi may readily be separated by its uniformly pol- ished and shining dorsal body surface, dark brown antennae, small non-truncated white scutellar spot, less elevated anterior pro- notal lobe and evenly convex, non-sinuate lateral pronotal margins. ACKNOWLEDGMENTS We thank Mary Jane Spring and Eliza- beth Slater (University of Connecticut) for preparation of the illustrations and assis- tance with the manuscript respectively. This work was sponsored in part by a grant from the National Geographic Soci- ety, Washington, D.C. and by the National Science Foundation. LITERATURE CITED Baranowski, R. and J. A. Slater. New genera and species of Antillocorini from Trinidad and Brazil (He- miptera: Lygaeidae). Fla. Entomol. (In press.) China, W.E. 1930. Insects of Samoa & other Samoan Terrestrial Arthropoda Pt. II. Hemiptera. Facsc. 3: 81-162. London. British Museum (Natural His- tory). Slater, J. A. 1984. On the biology of cave inhabiting Antillocorini with the description of a new species from New Guinea (Hemiptera: Lygaeidae). J. N.Y. Entomol. Soc. 91: 424-430. PROC. ENTOMOL. SOC. WASH. 89(3), 1987, pp. 489-499 ON THE IMMATURE STAGES OF PSALYDOLYTTA FUSCA (COLEOPTERA: MELOIDAE) RICHARD B. SELANDER AND ALIDA A. LAURENSE (RBS) Department of Genetics and Development, University of Illinois at Urbana- Champaign, Urbana, Illinois 61801, U.S.A.; (AAL) Integrated Pest Management Project, Crop Protection Service Department, Ministry of Agriculture, Yundum, Western Divi- sion, The Gambia. Abstract. —Egg masses laid by 27 females of Psalydolytta fusca (Olivier) from The Gambia contained a mean of 125.2 eggs. Mean incubation time at 27°C was 22.3 days in 19 masses that produced larvae. Within individual egg masses the hatching period extended over a period of 2—9 days (mean 5.1). Anatomical characteristics of the triungulin (first instar) larva, described in detail, confirm that Psalydolytta is an epicautine. The larva is the largest known in Meloidae and has unusually abundant setae on the dorsum of the head and body. Triungulin larvae of P. fusca and first grub larvae presumed to represent this species were found in egg pods of the grasshopper Cataloipus fuscocoeruleipes (Sjéstedt) at Tumani Tenda, Western Division, The Gambia. This is the first record of a meloid preying on an acridid of the subfamily Epyrepocnemidinae. The genus Psa/ydolytta Péringuey 1s rep- resented in Africa by 42 species and in India by an additional 10 (Kaszab, 1954; Saha, 1979; Selander, 1986). In both regions adults feed primarily or exclusively on grasses, in- cluding wild species of Andropogon and Cymbopogon as well as cultivated species of Eleucine, Oryza, Panicum, Pennisetum, Setaria, Sorghum, and Zea. Feeding, which generally occurs at night, destroys flowers and developing grains. When adults are nu- merous they may seriously reduce crop yields or, at worst, destroy an entire harvest. In addition, adults attracted to lights at night in public places are sometimes a nuisance because, when crushed against the skin, they cause blistering and other irritation (Gig- lioli, 1965; Zethner et al., 1985; and other references in Selander, 1986). In the taxonomic literature of the Meloi- dae Psalydolytta has been associated con- sistently with the genus Epicauta Dejean, which is assigned, together with its close relatives, to a separate tribe or subtribe of Meloinae (e.g., Kaszab, 1954, 1959, 1969; Selander, 1955; MacSwain, 1956; Saha, 1979). Indeed, there is little to distinguish Psalydolytta as a genus apart from Epicauta except the peculiar conformation of the mandibles of the adult, and even this is par- alleled to a certain extent in some Nearctic and Ethiopian species that undoubtedly be- long in Epicauta. The mandibles in Psaly- dolytta are enlarged, strongly bent backward (vaulted), and do not overlap distally as they do in most Meloidae. It has been conjec- tured that these modifications are in some way adaptive in feeding on the inflores- cences of wild grasses (Selander, 1986). Ad- ditional evidence for associating Psalydo- lytta with Epicauta is found in Fletcher’s (1914) report that young larvae of the In- dian P. rouxi (Castelnau) fed freely on the eggs of the pyrgomorphid Colemania 490 sphenarioides Bolivar in the laboratory and that a pupa was found associated with the eggs of this grasshopper in nature. Fletcher did not describe the larva of P. rouxi, and there are no other references to the imma- tures of Psalydolytta in the literature. Recently, interest in the biology of Psa- lydolytta has been stimulated by economic losses resulting from attacks on pearl millet (Pennisetum americanum) in West Africa by several species, the most important of which are P. vestita (Dufour), in Mauritania and Mali, and P. fusca (Olivier), in southern Senegal, The Gambia, and northeastern Guiné-Bissao. Partial results of a three-year study of the bionomics, economic impact, and control of P. fusca in The Gambia are given in the present paper. Specifically, the egg stage is treated briefly, the triungulin (first instar) larva is described in detail, and evidence concerning the mode of larval de- velopment 1s reported. The principal results of the study will be published elsewhere (Zethner and Laurense, 1987). EGG STAGE Between 18 September and 20 October 1985, 50 gravid females of Psalydolytta fus- ca collected in pearl millet fields, at a light trap in Yundum, and at lights at the Yun- dum airport were confined individually in cotton-stoppered glass vials 10 cm in height and 2.5 cm in internal diameter. Each vial contained a strip of paper that, by limiting the movement of the female, prevented eggs from being trampled after they had been laid. No food was provided. This procedure was adopted after trials in which adults were maintained in groups of 2—1 1 in plastic cages with earheads of pearl millet as food re- sulted in short survival (median 10 days) and little oviposition. (In retrospect it ap- pears that the poor results were caused by overcrowding.) Over a period of 1-5 days of confinement (mean = 2.0 days), 27 of the females ovi- posited, producing from 36 to 225 eggs each PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (mean = 125.2, SE = 9.08). Comparable figures for P. rouxi reported by Fletcher (1914) were 50 and 125, respectively. The egg masses of both species are numerically smaller than average for Meloidae but well within the normal range for species of Epi- cauta, which characteristically produce rel- atively large eggs in masses of at most a few hundred (Adams and Selander, 1979; Se- lander, 1981). Individually, the eggs of P. fusca are cream colored, weakly tapered, about 3.5 mm in length, and roughly '4 as wide as long. Shortly after deposition, eggs were re- moved from the glass vials and, using a fine brush, transferred in groups of five to plastic vials 7 cm in height and | cm in internal diameter in which they were incubated in darkness at 100% R.H. and 30°C. Hatching was recorded in 15 of the egg masses from individually confined females and in four additional masses from females kept in plastic cages with food material. Among these 19 masses the percentage of hatching ranged from 0.5 to 89.9, with a mean of only 33.8 (SE = 6.48). The low mean value is attributable, we believe, partly to failure of females, under starvation, to fertilize eggs properly and partly to damage to eggs as a result of handling. Except for four cases in which only one or two eggs hatched, hatch- ing of the eggs in a mass occurred over a period of 2—9 days (mean = 5.1, SD (pooled estimate) = 1.42, n = 15). Mean days to hatching in the 19 masses ranged from 20.1 to 25.0 days, with an unweighted mean of means of 22.3 days (SE = 0.31). According to Fletcher (1914), eggs of P. rouxi, at un- specified temperature, hatched in about 15 days. DESCRIPTION OF TRIUNGULIN LARVA Figs. 1-3, 4b Anterior, median and posterior rows of setae on a sclerite are denoted AR, MR, and PR, respectively. Within a row, a seta is referenced by its ordinal position from the VOLUME 89, NUMBER 3 midline of the body. Lengths of segments of the antenna and maxillary and labial pal- pl were measured on the ventral midline. Color medium brown; pronotum dark brown in posterior 7 and along lateral mar- gins in anterior 3. Sclerotized cuticle with weak, transverse reticulations, becoming obsolete medianly on dorsum and venter of head and medianly on pronotum; weak ten- dency for reticulations on abdominal pleu- rites and posterior abdominal tergites to be drawn into short spines; membranous cu- ticle, except that of clypeus, smooth. Setae in general long, conspicuous; those on dor- sum of head, thorax, and abdomen unusu- ally numerous, for most part bifid (very rarely trifid) at apex; ventral setae normal in number, very rarely divided at apex. Head (excluding labrum) as long as pro- and mesothorax combined, barely wider than long, widest across stemmata; sides constricted, sinuate behind middle, reduc- ing basal width to /,, maximum width. Stem of epicranial suture about *% as long as head; lateral arms attaining antennal foramina. Stemma unusually large and prominent, perfectly spherical; diameter greater than 1-'2x width of antennal segment II, greater than width of maxillary palpal segment II. Epicranium with 114-118 setae (12-14 on front, 10-14 lateral, 10 ventral); dorsal setae posteriad of level of stemmata clustered as in Fig. 1, not arranged in recognizable rows; major ocular seta inserted just mesad of stemma, not bifid, longer than antenna, at least 2 x as long as most setae outside ocular area; seta directly posteriad of stemma lon- ger than most other setae, inserted very near stemma; four minor setae on each side of epicranium at base, arranged longitudinally in nearly straight line; sensory cone mesad of posteriormost minor seta and another just anteriad of major ocular seta. Four or (as shown in Fig. 1) five setae on each side of frontal area, their insertions inset from branch of epicranial suture but roughly par- alleling it; setae subequal in length. Row of 491 4 setae on anterior margin of front; AR, much longer than other frontal setae; sen- sory cone and pit between AR, and AR.,. Clypeus semimembranous, microstriate. Labrum with anterior '2 of dorsal surface membranous in median 2; anterior margin rounded, very long seta ('2 as long as man- dible) on anterior corner, shorter one sub- marginally between corner and midline, and short, heavy marginal seta between these two setae; median transverse of 6 setae (MR, nearly at lateral margin); 4 setae on each lateral margin. Venter of labrum densely set with spines that project anteriad, anterior- most spines projecting well beyond anterior labral margin; each side with seta near lat- eral margin and another near midline at level of clypeolabral suture. Gula sharply tapered posteriad, reducing minimum width to /,, maximum width; length of gula '2 greatest width of head; gular setae exceeding anterior margin of gula, length of setae about % gular width. Antenna about 4 as long as head; segment I 2x as wide as long; II sinuate, nearly 3x as long as I, nearly 2 x as long as wide; III % as long as II, 2 x as long as wide; sensory cone of II perfectly conical, % as wide as long, as long as, and nearly 2 as wide as, III; short, setiform organ in mem- brane beside cone; 3 apical setae on seg- ments II and III (1 dorsal, 2 lateral), seta on posterior (outer lateral) margin of each seg- ment longer than other two; terminal seta of III short, 14x as long as II. Mandible robust; length (ventral condyle to apex) ¥, head length, 2 x basal mandibular width; 6 coarse, rounded teeth on mesal margin; proximal seta slightly longer than distal seta, as long as gular seta. Maxilla with cardo slender; stipes each with 3 long setae and 2 sensory pits ventrally near base, middle seta longer than others, longer than maxillary palpal segment III, mesal margin of stipes with several seta; mala with 2 long and 2 short, rather stubby setae. Maxillary palpus ¥, as long as mandible; segments I and II equal in length, 2'2x as wide as long, seta 492 on mesal margin of II as long as longest malar seta; III 2’ as long as II, nearly 2 x as long as wide, widest at basal *%, lateral margin evenly curved; sensory area of III extending about % length of segment, not overlapping onto ventral surface, papillae rodlike, separated by average distance equal to their length, not obscuring one another in dorsal view, 2-segmented sensory appen- dix *3as long as IJ. Labium with prementum II transverse, emarginate anteriorly, with pair of short setae and sensory pits; pre- mentum I deeply emarginate anteriorly, with pair of short and pair of long setae; pair of long setae on anterior labral margin between palpi; dorsal surface of prementum I dense- ly spinous, with anteriormost spines visible along anterior margin in ventral view (not shown in Fig. 1). Labial palpus '2 as long as maxillary palpus, slender, cylindrical; seg- ment I 2'4x as wide as long, with short ventral seta; II weakly tapered, 4*4 x as long as I, 24x as long as wide, mesal margin nearly straight, lateral margin slightly curved, with | dorsal seta, inserted at basal 3, attaining sclerotized distal margin, 2-seg- mented sensory appendix slightly narrower than that of maxillary palpus. Thorax with line of dehiscence complete on pro- and mesonotum, weak, incomplete on metanotum. Pronotum widest at base, about 1! as wide as long, as wide as head and 7, as long; sides nearly straight; about 134 setae, not arranged in recognizable rows. Mesonotum a little more than % as long as pronotum, nearly as wide; sides rounded, convergent anteriad; about 42 long setae. Metanotum similar to mesonotum; about 46 long setae. Thoracic venter not sclero- tized; each sternum with 4 setae in two rows; setae in AR of prosternum very short, widely separated from each other; setae of meso- and metasternum much longer than longest setae on prosternum. Abdomen moderately tapered; tergite V slightly shorter than metanotum, '3 as long as wide. All long setae on tergites I-VIII divided at apex. Long setae on tergites I- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VIII arranged in median and posterior (marginal) rows (MR actually well posteriad of middle); MR setae shorter than PR setae; MR with 16 or 18 setae; PR with 32 setae on I-V, 28 on VI and VII, 20 on VIII. Spine- like evaginations at bases of PR setae strongly developed on tergites I-IV, weak on V, absent on VI-IX; no evaginations at bases of MR setae; tergite V with PR setae ¥, as long as tergite. One pair of caudal setae on segment IX, a little shorter than seg- ments VIJ-IX combined. Seta just laterad of caudal seta thicker than other tergal setae, ’; as long as caudal seta. Setae on pleurites and abdominal venter not divided at apex, generally longer than tergal setae. Pleurites large, distinctly separated from tergites, wider than long; pleurite I with 2 setae, II- VIII each usually with 3 (sometimes with only 2) long setae. Sternites of segments I- VII reduced to pair of small, narrow scler- ites; sternites VIII and X well developed, undivided; sternum I with 2 setae of equal length on each sternite; H-IX each with PR of 6 setae, PR1 longer than rest on II-VII, inserted at posterior end of sternite; H-VIII with MR of 2 setae, these shorter than PR,, inserted at anterior end of sternite on I- VII. Spiracles round. Mesothoracic spiracle ventral; diameter *% that of stemma; scle- rous ring produced anteriad to support an erect seta. Abdominal spiracles set in lateral ’; of pleurite, equidistant from anterior and posterior margins; spiracle I /, diameter of mesothoracic spiracle; H-VIII progressive- ly slightly smaller; VIII about “% diameter of I. Legs with coxae 2 x as long as wide. Fem- ora I-III progressively slightly longer, tibiae I-III markedly so. All lanceolate setae heavy. Coxae each with 4 long setae, one of which is lanceolate on I and II. Trochantins each with 4 setae, 3 of which are lanceolate on I and IJ. Femur I with 6 anterior and 5 pos- terior lanceolate setae; I] and HI each with 5 anterior and 4 posterior lanceolate setae; femur III 4 as deep as long. Tibiae with VOLUME 89, NUMBER 3 493 ! Fig. 1. Triunguli L1G ee? | ST: ie EC . SSS / Bhi \ x 16 Se cae ees PEE. \ i> n larva of Psalydolytta fusca, dorsal (left) and ventral (right) views of head and thorax. 494 belt ttl Fig. 2. Triungulin larva of Psalydolytta fusca, dor- sal (left) and ventral (right) views of abdomen. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON heavy curved seta dorsally at apex; tibia III with 11 setae in anteroventral rows. Tar- sungulus III nearly % as long as tibia; an- terior seta 2 as long as tarsungulus; poste- rior seta % as long as anterior seta. Body length 5.2 (5.1—5.3) mm; caudal seta length 1.0 mm. Material studied.—12 larvae, from eggs laid by a female taken at light at the Yun- dum airport, Western Division, The Gam- bia, 17 September 1985, by A. A. Laurense; eggs 18 September 1985, larva ca. 7 October 1985. Remarks.—On the basis of the anatomy of the triungulin larva P. fusca the genus Psalydolytta clearly belongs in the Epicau- tina. The larva agrees with MacSwain’s (1956) diagnoses of the Epicautina and Epi- cauta except for the large number of setae on the dorsal surface of the head and body and runs to Epicauta in his key to genera of Meloidae. [MacSwain’s specification that the long ventral (“‘tactile”’) seta of the femur is inserted near the middle of that leg seg- ment in the Epicautina 1s erroneous.] With- in the Epicautina distinctive features of the larva of P. fusca are its (1) large body size, (2) the size and shape of its stemmata, and (3) numerous details of chaetotaxy resulting from a proliferation of setae. As far as we can determine, the triungulin of P. fusca is the largest yet described in the Meloidae. The only larvae of comparable size are those of the South American species Epicauta leopardina (Haag-Rutenberg) (4.9 mm long) (Agafite1 and Selander, 1980), the North American species Megetra cancellata (Brandt and Erichson) (4.0-—4.5 mm) (Se- lander, 1965), and the Asian species My- labris quadripunctata (Linnaeus) (4.0-4.5 mm) (Priamikova and Iukhnevitch, 1958). Otherwise, meloid triungulins do not ex- ceed 4 mm in length, and most are consid- erably smaller. The larval stemmata are larger and more prominent in P. fusca than in other epicautines and are, among Meloi- dae, unusual in being perfectly spherical in shape. VOLUME 89, NUMBER 3 3 Fig. 3. Triungulin larva of Psalydolytta fusca, fore- (a), mid- (b), and hindleg (c), anterior views. The most obvious diagnostic feature of the triungulin of P. fusca is the large number of setae on the head, body, and (to a lesser extent) legs. MacSwain (1956) specified a maximum of 60 pronotal setae in the Epi- cautina, but P. fusca has nearly twice that number. In most Meloidae the triungulin has six setae in MR and 10 in PR on ab- dominal tergites I-VIII. The number in PR is increased to 12 or 14 in several New World Epicautina (Agafitei and Selander, 1980; Se- lander and Agafitei, 1982) and reaches a maximum of 16 in E. funesta (Chevrolat) (Selander and Agafitei, 1982) and FE. dubia (Fabricius) (Zakhvatkin, 1954) (attained also in the mylabrine genus Coryna Billberg, ac- cording to MacSwain, 1956). In contrast, Psalydolytta fusca has 16-20 setae in MR and 28-32 in PR. The tergal MR setae in P. fusca are also unusual for Epicautina in being more than '2 as long as the PR setae. Ven- trally the pattern of setation is more nearly normal in P. fusca, although here again there are more setae than usual (six rather than four in PR of abdominal sterna II-VIII and three rather than two on the pleurites). The number of lanceolate setae on the femora (11) is greater than usual for Epicautina but is matched in four species of the North American Vittata Group of Epicauta, where it varies intraspecifically from nine to 11 (Agafitei and Selander, 1980). One might expect a positive correlation between num- ber of setae and body size in meloid triun- gulins, but except in the case of the lanceo- late femoral setae there is no indication of such a relationship in the available data. Apical division or branching of setae on the head, thorax, and abdomen of the triun- gulin larva has been recorded heretofore only 496 in the genus Epicauta, where it takes a va- riety of forms (Fig. 4). In the Old World the setae are simply bifid in the Palearctic species E. ruficeps (iliger), E. dubia, E. erythro- cephala (Pallas), and E. rufidorsum (Goeze) [= EF. verticalis (Illiger)] and the Ethiopian species EL. albovittata (Gestro) (Cros, 1938; Zakhvatkin, 1929, 1954) and undivided in Palearctic species E. gorhami Marseul (Chu and Wang, 1956; Nagatomi and Iwata, 1958) and E. megalocephala (Gebler) (Zakhvat- kin, 1954). Since the four species having bifid setae represent three of the 16 species groups of Old World Epicauta recognized by Kaszab (1952, 1953), we may presume that the characteristic 1s of widespread oc- currence among Old World Epicauta. In the New World, where triungulin larvae of many species have been described, branched setae are known in only three species. In the Vit- tata Group, as studied by Agafitei and Se- lander (1980), E. monachica (Berg) has bifid setae similar to those of P. fusca.' In the same group the setae of EF. temexa (Adams and Selander) are basically bifid, but there is a strong tendency for one or both of the branches of the seta to be further subdivided and the incidence of purely trifid setae is much higher than in P. fusca. Finally, in E. niveolineata (Haag-Rutenberg), represent- ing the subgenus Macrobasis LeConte, there are, typically, multiple branches, some of which arise from the sides of the seta, well before the apex (Selander and Agafitei, 1982). The functional significance of setal branching is unknown, although one might suppose that adaptation to burrowing in a particular type of substrate, such as sand, is involved. In view of the limited nature of the de- scriptive material available for larvae of the ' MacSwain (1956), Agafitei and Selander (1980), and Selander and Agafitei (1982) erred in attributing branched setae in Old World species of Epicauta only to E. albovittata. Agafitei and Selander (1980) also erred in characterizing the setae of E. monachica as divided several times apically, rather than as bifid. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Old World species of Epicauta, it is not pos- sible to determine which Old World species group is phenetically closest to P. fusca or, for that matter, whether the Psalydolytta is phenetically closer to one or more of the New World groups of the genus than to the Old World groups. Kaszab (1952) described his Group I of Epicauta, which includes E. cognata (Haag-Rutenberg) and relatives in the Oriental Region and E. canescens (Klug) and relatives in the Ethiopian Region, as forming a link between Epicauta and Psa- /ydolytta on the basis of similarities in the conformation of the adult mandibles. Un- fortunately, the triungulin larva of this group is unknown. Among the New World groups, the Vittata Group is perhaps the most sim- ilar to Psalydolytta. However, judging from Cros’ (1938) description of the triungulin larva of the Ethiopian E. albovittata, there is at least an equally strong phenetic rela- tionship between Psa/lydolytta and that species. THE MODE OF LARVAL LIFE Adults of Psalydolytta fusca attacking pearl millet spend their adult lives in the millet fields, feeding at night and hiding in the bases of the plants during the day. Cap- tive females have been observed laying eggs in cavities in the soil, and it is probable that soil in and near millet fields is commonly used for oviposition. Although we were not aware in 1985 of Fletcher’s (1914) associ- ation of P. rouxi with the eggs Colemania sphenarioides, the systematic position of Psalydolytta suggested that the larva of P. fusca would prove to be a predator of grass- hopper eggs. Further, knowledge of the adult behavior suggested that larvae would be most readily found near infested millet fields. After several unsuccessful attempts to find grasshopper eggs by scraping off the top lay- er of soil in square-meter quadrats at several localities, one of us (AAL) finally discovered a substantial number of egg pods of acridid Cataloipus fuscocoeruleipes (Sj6stedt) in soil at Tumani Tenda, Western Division, in an VOLUME 89, NUMBER 3 Ml il 497 | e Fig. 4. Apically branched PR setae of abdominal tergite III of the triungulin larvae of Epicauta albovittata (after Cros, 1938) (a), Psalydolytta fusca (b), E. monachica (c), E. temexa (d), and E. niveolineata (e). area of weeds and shrubs between swamp- rice fields and a pearl millet field that was heavily infested with adult Psalydolytta fus- ca.” Meloid larvae were found in three Ca- taloipus egg pods on 25 October 1985 and in four pods on 7 November 1985. All of * In the classification of Dirsh (1965) Cataloipus Bo- livar is placed in the Eyprepocnemidinae, a subfamily of Acrididae with which larval Meloidae have not been associated previously. the pods containing larvae showed evidence of having been punctured. All were subse- quently kept in darkness at 100% R.H. and 29-30°C in an attempt to rear the meloid larvae. An inventory of the contents of the pods follows, using the nomenclature of lar- val phases of Selander and Mathieu (1964). Except as noted, all larvae died in the instar in which they were found. Pod 1: One triungulin (T,) larva, which escaped. 498 Pod 2: A \ast-instar first grub larva (FG,;?) that ecdysed to the coarctate (C) phase two weeks after collection. Pod 3: An FG larva of undetermined in- star. Pod 4: A T, larva positively identified as Psalydolytta fusca by comparison with reared triungulins and an FG, larva that reached FG, in captivity. The left side of the head of the T, possessed a large black scar, possibly the result of combat with the other larva. Pod 5: An FG, larva and two FG larvae of undetermined instar. The left midleg of the FG, larva was missing except for the coxa, but there was no indication of injury to the cuticle. Pod 6: A l\ast-instar FG larva (FG;?) that ecdysed to the C phase two weeks after col- lection. Pod 7: An FG, larva, not fully fed, and the exuvia of FG, (of which only the head capsule was preserved). This larva may have been injured during handling; when exam- ined after death it had a large bubble on the left side of the thorax, as though hemo- lymph had extruded and coagulated. Comparison of the FG larvae and exuvia with respect to size, setation, and numerous characters of the antennae, mouthparts, and legs leads us to believe that all of the grubs represent a single species. The instar of the exuvia identified as FG, can be specified confidently on the basis of its resemblance to a T larva in several characters, particu- larly those of the legs. The larva that pro- duced this exuvia is, then, an FG,, and since the exuvial head capsule associated with the larva in pod 7 is larger than that of the FG,, it must represent FG, and its associated lar- va FG;. Very likely the two larvae that reached the C phase were, when found, in instar FG, also, since this is commonly the last instar of the FG phase in Meloidae. The assignment of instar numbers, as well as our contention that the larvae are conspecific, is supported by the fact that a plot of the logarithm of the width of the frontal sclerite of the head against known or inferred instar PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON number is quite linear (n = 6, r = .990, P= .00015). The estimated regression equation for the relationships 1s Y = —.2640 + .1274x. As for the identification of the FG larvae, the exuvia of the FG, larva is so large as virtually to rule out its representing any species but P. fusca. In particular, the size of the FG, would seem definitely to exclude the possibility of its representing Mylabris holosericea (Klug), a relatively small mel- oid, as an adult, often occurring in millet fields with P. fusca, although generally less abundantly. Additional evidence bearing on the identity of the larvae associated with Cataloipus is the fact that the spiracles of the C larvae obtained by rearing occupy mound-like protuberances of the cuticle that appear to sag posteriad, tipping the spirac- ular cones in that direction. Until now this feature has been regarded as diagnostic of the genus Epicauta, but its occurrence in Psalydolytta would not be surprising, given the close relationship of the two genera. At the very least it would seem to establish that the larvae are epicautine. In summary, while it is clear that T, lar- vae of Psalydolytta fusca are capable of en- tering the egg pods of Cata/loipus, our data provide only circumstantial evidence that P. fusca larvae eat the eggs of this grass- hopper, since the T, larvae found in the field had not fed to an appreciable extent and the FG larvae cannot be identified with cer- tainty as P. fusca at this time. Nevertheless, we are reasonably confident that P. fusca is, in fact, a larval predator of Cataloipus.* ACKNOWLEDGMENTS The collaboration leading to this paper began at a conference on the bionomics of Meloidae in Dakar, Senegal, in August 3 After submission of this paper for publication, one of us (AAL), by releasing groups of T, larvae on moist sand containing egg pods of Cataloipus fuscocoeru- leipes, successfully reared two larvae of P. fusca to the C larval phase. The C larvae from this rearing are similar to those obtained earlier from FG larvae col- lected in the field. VOLUME 89, NUMBER 3 1985. This conference was sponsored and supported financially by the Integrated Pest Management Project, Permanent Interstate Committee for Drought Control in the Sa- hel (CILSS). Participation in it by one of us (RBS) was supported by the Food and Ag- riculture Organization, United Nations. We are pleased to acknowledge the valuable contribution of O. Zethner to the planning and execution of the study. LITERATURE CITED Adams, C. L. and R. B. Selander. 1979. The biology of blister beetles of the Vittata Group of the genus Epicauta (Coleoptera, Meloidae). Bull. Am. Mus. Nat. Hist. 162(4): 137-266. Agafitei, N. J. and R. B. Selander. 1980. First instar larvae of the Vittata Group of the genus Epicauta (Coleoptera: Meloidae). J. Kans. Entomol. Soc. 53: 1-26. Chu, H.-F. and L.-Y. Wang. 1956. On the life-history of the legume blister beetle, Epicauta gorhami Marseul, with a discussion of hypermetamor- phosis. [In Chinese, with English summary.] Acta Entomol. Sinica 6: 61-73. Cros, A. 1938. Considerations générales sur le genre Epicauta Redtenbacher. Etude biologique sur Epi- cauta albovittata Gestro. Mem. Soc. Entomol. Ital. 16: 129-144, pl. 3. Dirsh, V.M. 1965. The African genera of Acridoidea. Cambridge University Press, Lond, x1i + 579 pp. Fletcher, T. B. 1914. Some South Indian insects. Su- perintendent, Government Press (Madras, India), vil + 565 pp. Giglioli, M. E.C. 1965. Some observations on blister beetles, family Meloidae, in Gambia, West Africa. Trans. R. Soc. Trop. Med. Hyg. 59: 657-663. Kaszab, Z. 1952. Die palaarktischen und orienta- lischen Arten der Meloiden-Gattung Epicauta Redtb. Acta Biol. Acad. Sci. Hung. 3: 573-599. 1953. Revision der aethiopischen Arten der Meloiden-Gattung Epicauta Redtb. Acta Biol. Acad. Sci. Hung. 4: 481-513. 1954. Die Arten der Meloiden Gattung Psa- lydolytta Per. (Coleoptera, Meloidae). Acta Zool. Acad. Sci. Hung. 1: 69-103. 1959. Phylogenetische Beziehungen des fli- gelgedders der Meloiden (Coleoptera), nebst Be- schreibung neuer Gattungen und Arten. Acta Zool. Acad. Sci. Hung. 5: 67-114. 1969. The system of the Meloidae (Coleop- tera). Mem. Soc. Entomol. Ital. 48: 241-248. 499 MacSwain, J. W. 1956. A classification of the first instar larvae of the Meloidae (Coleoptera). Univ. Calif. Publ. Entomol. 12, iv + 182 pp. Nagatomi, A. and K. Iwata. 1958. Biology of a Jap- anese blister beetle, Epicauta gorhami Marseul (Coleoptera, Meloidae). Mushi 31: 29-46, pl. 4. Priamikova, M. A. and L. A. Iukhnevich. 1958. Key to the triungulins of blister-beetles (Coleoptera, Meloidae) of the tribe Mylabrini in the fauna of the USSR. [In Russian.] Entomol. Obozr. 37: 176- 182. [English translation in Entomol. Rev. 37: 139- 145.] Saha, G. N. 1979. Revision of Indian blister beetles (Coleoptera: Meloidae: Meloinae). Records Zool. Surv. India 74(1): 1-146. Selander, R. B. 1955. The blister beetle genus Lins- leya (Coleoptera, Meloidae). Am. Mus. Novit. 1730, 30 pp. . 1965. A taxonomic revision of the genus Me- getra (Coleoptera: Meloidae) with ecological and behavioral notes. Can. Entomol. 97: 561-580. . 1981. Evidence for a third type of larval prey in blister beetles (Coleoptera: Meloidae). J. Kans. Entomol. Soc. 54: 757-793. . 1986. An annotated catalog and summary of bionomics of blister beetles of the genus Psaly- dolytta (Coleoptera, Meloidae). Insecta Mundi. [In press. | Selander, R. B. and N. J. Agafitei. 1982. First-instar larvae of the Uniforma Group of the genus Epi- cauta (Coleoptera, Meloidae). Proc. Entomol. Soc. Wash. 84: 138-148. Selander, R. B. and J. M. Mathieu. 1964. The on- togeny of blister beetles (Coleoptera, Meloidae) I. A study of three species of the genus Pyrofa. Ann. Entomol. Soc. Am. 57: 711-732. Zakhvatkin, A. A. 1929. The biology and morphol- ogy of the parasites of the egg-pods of Acrididae in Central Asia. 1. Description of the triungulin of Epicauta erythrocephala Pall. (Col., Meloidae). {In Russian.] Sredneaziatsk. Inst. Zashch. Rast. (Tashkent) 15 (1928), 7 pp. 1954. Parasites of Acrididae of the Angara Region. [In Russian.] Trudy Vsesoyuz. Entomol. Obshch. 44: 240-300. Zethner, O. and A. A. Laurense. 1987. Investigations on the economic importance and control of the adult blister beetle Psalydolytta fusca (Olivier) (Coleoptera: Meloidae), a serious pest of pearl mil- let in The Gambia. Trop. Pest Manag. [In press.] Zethner, O., S. B. Sagnia, A. A. Laurense, and S. Bruce- Oliver. 1985. CILSS Integrated Pest Manage- ment Project Annual Report 1985. Entomology. Yundum, The Gambia, 110 pp. PROC. ENTOMOL. SOC. WASH. 89(3), 1987, pp. 500-501 SEX ATTRACTANT FOR SCOPARIA BIPLAGIALIS (LEPIDOPTERA: PYRALIDAE) H. G. Davis, L. M. MCDONOUGH, AND D. C. FERGUSON (HGD, LMM) Yakima Agricultural Research Laboratory, Agricultural Research Ser- vice, USDA, 3706 West Nob Hill Boulevard, Yakima, Washington 98902; (DCF) Sys- tematic Entomology Laboratory, Agricultural Research Service, BBII, USDA, % National Museum of Natural History, NHB 168, Washington, D.C. 20560. Abstract. —(Z)-11-Hexadecen-1l-ol acetate was discovered to be a sex attractant for Scoparia biplagialis. This is the first example of a sex attractant for Scopariinae. During the last 15 years, a multitude of sex pheromones of insects have been iden- tified (Inscoe, 1982). Sex pheromones are used in crop protection programs as attrac- tants in traps which indicate the presence and change in population level of a target economic species. Also, sex pheromones are useful for studying taxonomic relationships (Roelofs and Brown, 1982). In 1980, while conducting other research, we noted many specimens of a small species of moth in one of our traps, and we sub- sequently identified the species as Scoparia biplagialis Walker (Pyralidae: Scopiari- inae). In 1981 and 1982 we conducted fur- ther tests to confirm our observations and to determine the response of S. biplagialis to different amounts of the attractive com- pound. Here we report the results of the tests conducted in 1981 and 1982. MATERIALS AND METHODS Lures were prepared by adding 100 ul of a dichloromethane solution of (Z)-11-hex- adecen-1-ol acetate to No. | red rubber sep- ta (West Co., Phoenixville, PA). The amount of acetate used was 0.0 and 1.0 mg per sep- tum in 1981 and 0.0, 0.1, 1.0, and 10.0 mg per septum in 1982. Lures were placed in the bottom of wing traps (Howell, 1972) which had been coated with polyisobutyl- ene to provide a sticky surface (Pherocon 1C® traps, Zoecon Corp., Palo Alto, CA). Traps were hung at a height of 2 m in filbert trees. In 1981, one control and one test trap each were placed in two different abandoned orchards (Sherwood in Washington County and Dundee in Yamhill County) in the Wil- lamette Valley, Oregon from | July to 27 August. In 1982 five replicates of the four treatments were arranged randomly along a row of filbert trees in the Sherwood orchard from 6 July to 24 August. In 1981, trap catches were recorded weekly and in 1982 on 22 July and 24 August. The 1982 data were analyzed by Duncan’s New Multiple Range Test (P = 0.05). Insects were removed from the sticky traps Table 1. Number of S. biplagialis captured in traps containing the lure (Z)-1 1-hexadecen-1-ol acetate, dur- ing tests conducted from 6 July to 24 August 1982 in the Willamette Valley, Oregon. Each dosage was tested with 5 replicates. Means followed by the same letter are not significantly different, ANOVA and DMRT (P = .05). Lure Dosage/Trap . (mg) Catch/Trap and Significance 0) Oa 0.1 16.6 b 1.0 20.4 b 10.0 2.8 a VOLUME 89, NUMBER 3 for taxonomic identification as follows. When the insects were to be examined from the dorsal view only, the cardboard surface to which they adhered was cut from the rest of the trap and the undisturbed insect was positioned on an insect pin. Insects selected for ventral and lateral views were freed from the sticky surface by washing several times with hexane. After soaking in hexane for 24 hours, the insects were placed on filter pa- pers and transferred to a relaxing jar. When the specimens became soft, they were at- tached to minuten nadeln. RESULTS AND DISCUSSION In 1981 in both orchards the traps which contained (Z)-11-hexadecen-1-ol acetate captured S. biplagialis (172 at the Sherwood orchard and 41 at the Dundee orchard). Neither control trap captured any S. bipla- gialis. Over half of the captured S. bipla- gialis were examined and all were found to be males. At Sherwood 163 (95%) and at Dundee 38 (93%) were caught by 6 Aug. In 1982, the dosage tests (Table 1) showed that the traps containing 0.1 and 1.0 mg of (Z)-11-hexadecen-1l-ol acetate per septum captured significantly more moths than the traps containing 10.0 mg or no lure. Of the 199 total, 121 (61%) were captured by the 22 July count. Based on the trap catches during 1981 and 1982, S. biplagialis ap- pears to have a major flight period during July and early August. Munroe (1972) lists the immature forms of North American Scoparia as undescribed but notes that certain European species are known to feed underground on the roots of ragwort (Senecio spp.) and others are be- lieved to tunnel in mosses. The noxious weed Tansy ragwort, Senecio jacobaea L., grows abundantly in western Oregon and Wash- ington (Hepworth and Guelette, 1979) and is prevalent in both the Sherwood and Dun- dee abandoned orchards, and thus is a pos- sible host. According to Munroe (1972), S. biplagi- 501 alis is found widespread along the northerly parts of the continent that extends from Newfoundland to the Carolinas and west- ward across Canada and the northern one- half of the United States to the Pacific Coast and northward to the Aleutian Islands. He recognized five noticeably different popu- lations and treated them as subspecies, based on the material from the following places: Natashquan, Quebec; Maine; Kaslo, British Columbia; Victoria, British Columbia; and Afognak, Alaska. The population in the Willamette Valley, Oregon, probably should be referred to S. biplagialis pacificalis Dyar, originally described from Victoria, British Columbia, although Oregon could also have subspecies fernaldalis, or populations that are intermediate. Further taxonomic inves- tigation is needed, as Munroe (1972: 41) indicated in his remark to the effect that some of the subspecies might really be species with poor morphological differen- tiation. This is the first example of a sex attractant for a species of the subfamily Sco- pariinae. In view of the great variation of size, coloration, etc., within S. biplagialis and the extensive distribution of this species in northern North America, Z1 1-16:Ac may be helpful in solving taxonomic problems. LITERATURE CITED Hepworth, H. H. and L. O. Guelette. 1979. Tansy ragwort. Pacific Northwest Ext. Publ. No. 175. Coop. Ext. Service, Oreg. State Univ. 4 pp. Howell, J. F. 1972. An improved sex attractant trap for codling moths. J. Econ. Entomol. 65; 609-611. Inscoe, M. N. 1982. Insect attractants, attractant pheromones, and related compounds, pp. 201- 295. In Kydonieus, A. F. and Beroza, M., Insect Suppression with Controlled Release Pheromone Systems. CRC Press, Inc., Boca Raton, Florida. Munroe, E. G. 1972. Pyraloidea (in part). 7» Dom- inick, R. B. et al., The Moths of American North of Mexico, Fascicle 13.1. E. W. Classey, Ltd. and The Wedge Entomological Research Foundation, London. 304 + xx pp., 13 col. pls. Roelofs, W. L. and R. L. Brown. 1982. Pheromones and evolutionary relationships of Tortricidae. Ann. Rev. Ecol. Syst. 143: 395-422. PROC. ENTOMOL. SOC. WASH. 89(3), 1987, pp. 502-511 A REVIEW OF THE HOLARCTIC GENUS TERRILIMOSINA (DIPTERA: SPHAEROCERIDAE), WITH DESCRIPTIONS OF NEW SPECIES FROM NEPAL AND JAPAN S. A. MARSHALL Department of Environmental Biology, University of Guelph, Guelph, Ontario N1G 2W1, Canada. Abstract. —Terrilimosina unio from Nepal and Java, T. smetanai from Nepal and Pa- kistan, 7. deemingi from Nepal, 7. brevipexa from Japan, and T. longipexa from Japan and Nepal are described as new. A cladogram and a key to the nine world species of Terrilimosina are provided and new distributional data are given. The genus Jerrilimosina Rohaéek, wide- spread in the Holarctic and Oriental Re- gions, constitutes a distinctive monophy- letic group within the sphaerocerid subfamily Limosininae. As the generic name suggests, species of this genus are largely terricolous and are often associated with small deposits of wet, decaying vegetation in forests or peatlands. One species, 7. ra- covitzal Bezzi, 1s troglophilic and polysa- prophagous. The major synapomorphies for Terrilimosina are a posteriorly rounded dis- cal cell, a comb-like row of bristles on the surstylus, and a shortened, frame-like dis- tphallus. Members of this genus are further characterized by a telescoping female ab- domen, weakly sclerotized distiphallus, wing venation with R,,,; and R,,; sinuate, and the costa extending beyond the tip of R,.;. A full generic description can be found in Rohaéek (1983) and a key to the four pre- viously known species can be found in Mar- shall (1985). No eastern Palaearctic species were previously known, although Deeming (1969) mentioned the existence of unde- scribed species similar to 7. schmitzi (Duda) in Nepal, and Richards (1961) recorded T. racovitzai, the type species of Terrilimosina, from Afghanistan. The identity of the spec- imens from Afghanistan 1s confirmed, and the specimens previously examined by Deeming are included in the type series of new species described below. Collection abbreviations. — Biosystemat- ics Research Institute, Ottawa, Canada (CNC); University of Guelph Collection (GUE); the Silesian Museum, Opava, Czechoslovakia (OPAVA): the Geneva Mu- seum (GM); British Museum of Natural History (BMNH); Universitetets Zoolo- giske Museet, Copenhagen, Denmark (UZMC). KEY TO THE SPECIES OF 7ERRILIMOSINA Terrilimosina species are superficially very similar, but each has distinctive, species- specific genitalic characteristics which should be examined to confirm identifica- tions. 1. Eye height less than 1.5 times genal height. Mid tibia ventrally with only an apical bristle. Eu- rope, Afghanistan and eastern North America. Seep S hh Nitierre pormaedi le YAO Se 2 racovitzai (Bezzi) — Eye height more than 2.0 times genal height. Mid tibia with a mid ventral bristle. 2. Wing shorter than body, even in dried speci- mens. Epiproct without bristles. Europe. .... Dye Te SERED CIN Pee sudetica (Rohaéek) VOLUME 89, NUMBER 3 — Wing longer than body. Epiproct with 2 bris- tles. 3. Forecoxa yellow, contrasting with dark pleu- non Ne palris scene cree deemingi n. sp. — Forecoxa brown (sometimes very pale, ap- peaninedintyavellow)ime4 eee sae 4 4. Sternite 5 of male concave and simple pos- teromedially. Epiproct twice as long as wide. Folanctichmeme. joc eee foe schmitzi (Duda) — Sternite 5 of male with a posteromedial lobe covered with modified setulae. Epiproct at most leSitimesyasslongeasiwides.. 4... ne os 5 5. Syntergite 1+2 at least 1.5 times as wide and as long as tergite 3 and at least partly darker than tergite 3. Surstylus with 2-5 very short, blunt bristles apically on posteroventral lobe (Figs. 2, 32). Hypoproct concave anteriorly, setulose on less than posterior half (Figs. 6, 34). so of Ce RCE ER GR Sei cate a ee em 6 — Syntergite 1+2 not markedly darker or wider, and no more than 1.5 times as long as tergite 3. Surstylus with a comb-like row of bristles on inner ventral surface. Hypoproct convex an- teriorly, setulose on more than posterior half. OES WARLS 5A RA te ADI Basta CeCe Oat IISA 7 6. Syntergite 1 +2 entirely darkly pigmented, twice as long as tergite 3. Surstylus with about 5 very short, blunt bristles apically on posteroventral lobe (Figs. 1, 2). Anterior margin of epiproct EXCISedi(hg.16)sapal., aks. woe. brevipexa Nn. sp. — Syntergite | +2 with an anteromedial pale area, 1.5 times as long as tergite 3. Surstylus with 2 small blunt bristles apically on posteroventral lobe (Fig. 32). Anterior margin of epiproct en- tire (Fig: 36)! Nepal; Java: .......... unio N. sp. 7. Surstylus elongate, with a comb-like row of short bristles (Figs. 23, 24). Sternite 5 of male long, posteromedial lobe covered with single bristles. Tergite 8 of female entirely dark; ex- tending lateroventrally as broad, shining lobes (Fig. 28). Nepal, Pakistan. ...... smetanal n. sp. — Surstylus subquadrate in lateral view, with a comb-like row of long bristles (Figs. 15, 16). Sternite 5 of male short, posteromedial lobe covered with tufts of setulae. Tergite 8 of fe- male with tripartate pigmentation and not ex- tending to ventral surface (Fig. 22). 8. Sternite 5 of male with a prominent, dark, ba- sally narrowed posterior lobe covered with tufts of long, flattened setulae (Fig. 19). Spermathe- cae cup-shaped; invagination at about 140 de- grees from base (Fig. 21). Japan, Nepal. SE A ne. pee ns RE AN longipexa n. sp. — Sternite 5 of male with an inconspicuous, broadly based, posteromedial lobe covered with patches of short, flattened setae. Spermathecae peanut-shaped; invagination apical. Nearctic. ET ee ere RT epee eee ahes pexa Marshall 503 All of the species described below resem- ble the most common species, 7. schmitzi, in external features such as mesotibial chae- totaxy as well as general features of internal genitalia such as the short hypandrium and the anteriorly setulose paramere. Terrilimosina brevipexa Marshall, New SPECIES Description.— Body length 1.9-2.1 mm. Weakly punctate, pruinose, dark brown to black; pleural sutures, tarsi, and tips of fem- ora light brown. Interfrontal plate 1.5 times as high as wide, bordered by 4 subequal interfrontal bristles. Eye height 2.5 times genal height at point of maximum eye height; genal shining area restricted to a narrow strip below anterior half of eye. Dorsocentral bristles 1n 2 pairs, prescutellar pair 2.5 times as long as anterior pair, equal to or slightly longer than intrapostalar bristles. Wing slightly infuscated; second costal sector shorter than third; halter luteous. Syntergite 1+2 uniformly black, wider than and twice as long as tergite 3; tergite 3 light brown. Male terminalia.—Sternite 5 convex and densely setulose posteromedially; these set- ulae enlarged and flattened at apex of pos- teromedial lobe (Fig. 5). Surstylus laterally setulose at base; ventrally with anterior and posterior lobes, posterior lobe with an in- ternal comb of short, stout bristles at apex (Figs. 1, 2). Paramere broad, strongly bent; apex broadly bifid (Fig. 3). Basiphallus nar- row, not extending ventrally beyond point of articulation with distiphallus. Distiphal- lus very long, narrow; with relatively dark proximal dorsal and ventral sclerites and pale distal dorsal and ventral sclerites. Female terminalia.—Tergite 8 with tri- partite pigmentation, wide median area pale and shining, lateral areas pruinose, ventro- lateral lobes shining brown; epiproct slight- ly longer than wide, emarginate anteriorly, setulose and with 2 bristles on posterior third (Fig. 8). Sternite 8 simple, darker and more than twice as large as hypoproct. Hypoproct deeply emarginate anteriorly, setulose on 504 Zs RATAN Ss, Ce PTT REN Figs. 1-8. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Terrilimosina brevipexa. 1, Male terminalia, left lateral. 2, Surstylus, ventral. 3, Aedeagus and associated parts. 4, Male terminalia, posterior. 5, Male sternite 5. 6, Female terminalia, ventral. 7, Spermathecae. 8, Female terminalia, dorsal. posterior third (Fig. 6). Spectacles-shaped sclerite visible only as 2 lightly sclerotized rings. Spermathecae cylindrical, bent and widened apically, with a small apical in- vagination (Fig. 7). Types.— Holotype ¢ (CNC) and 9 2 para- types (GUE, CNC). JAPAN. Shikoku, Isi- zuchi Mt. National Park, Tsuchigoya, 1400 m, 11-18.viii.1980, S. Peck. Etymology.—The specific epithet refers to the short comb-like row of short bristles on the surstylus. Comments.—This species is most easily recognised on the basis of distinctive char- acters of the abdomen, including the large, dark syntergite 1+2, the very distinctive distiphallus, the bent paramere, the short comb of bristles on the surstylus, and the deeply emarginate hypoproct. Terrilimosina deemingi Marshall, New SPECIES Figs. 9-14 Description. — Body length 1.8-2.0 mm. Body pruinose brown, legs light brown, coxa yellow. Interfrontal plate subequal in height and width, bordered by 4 subequal inter- frontal bristles. Eye height 2.5—2.8 times ge- nal height at point of maximum eye height; genal shining area restricted to a narrow strip below anterior half of eye. Dorsocentral bristles in 2 pairs, prescutellar pair 3 times as long as anterior pair, 1.5 times as long as intrapostalar bristles. Wing clear; second costal sector shorter than third; halter lu- teous. Abdominal syntergite | +2 uniformly brown, slightly wider and 1.4 times as long as tergite 3; tergite 3 similarly pigmented. VOLUME 89, NUMBER 3 505 Figs. 9-14. Male terminalia.—Sternite 5 convex and uniformly setulose posteromedially (Fig. 11). Surstylus basally subquadrate and laterally setulose; ventrally with a long, tapered an- terior lobe and a posteroventral lobe with a loose comb of 7 stout bristles (Fig. 10). Par- amere bent anteriorly and apically pointed (Fig. 9). Basiphallus narrow, not extending ventrally beyond point of articulation with distiphallus. Distiphallus long, narrow; with a dark, flat proximal dorsal sclerite, large lateroventral sclerite, and pale, thin distal dorsal and ventral sclerites surrounded by finely spinulose membrane. Female terminalia.—Tergite 8 entirely pruinose, with tripartite pigmentation, me- dian part pale brown; epiproct slightly lon- ger than wide, posterior third setulose and with 2 bristles (Fig. 14). Sternite 8 large, shield-shaped, very darkly pigmented, set- ulose and setose (Fig. 12). Hypoproct small, transverse, setulose on posterior third. Spectacles-shaped sclerite not visible except as large, weakly sclerotized rings. Sperma- thecae small, bent apically, with invagina- tion at about 120 degrees from duct (Fig. 14). Types. — Holotype 6 (CNC) and 172 para- types (47 6, 125 9, GUE, CNC, OPAVA). NEPAL. Between Ghopte and Thare Pati, Terrilimosina deemingi. 9, Aedeagus and associated parts. 10, Male terminalia, left lateral. 11, Male sternite 5. 12, Female terminalia, ventral. 13, Spermathecae. 14, Female terminalia, dorsal. 3200 m, 23-26.i1v.1985, Flight Intercept Trap, A. Smetana. Other paratypes: NE- PAL. Below Thare Pati, 3300 m, 9- 13.iv.1981, pitfall trap on a clearing in a mixed Abies, Acer, Rhododendron forest, A. Smetana (3 2, GUE); Taplejung Distr., San- gu, c. 6200’, mixed vegetation by stream in gully,” x0. 1961=1-1962, sR. L. Coe’ (1. *6, BMNH). The latter specimen was listed as Leptocera (Limosina) sp. (2) near parami- nima (Duda) by Deeming (1969). Etymology. — This species ia named after J. Deeming, in recognition of his 1969 paper on Sphaeroceridae from Nepal. Comments.— Terrilimosina deemingi 1s easily recognized by its pale legs, especially the yellow coxae, and is further character- ized by unique male terminalia and dis- tinctive features of the female terminalia such as the dark, shield-shaped sternite 8. Terrilimosina longipexa Marshall, NEw SPECIES Figs. 15-22 Description. — Body length 1.6-1.9 mm. Body pruinose brown; legs light brown. In- terfrontal plate 1.5 times as high as wide, bordered by 4 short interfrontal bristles. Eye 1.9-2.1 times as high as gena at point of maximum eye height; anterior part of gena 506 Kocthnn Etat Wafimeranines Shun oudgigghs nH HR EY 1 9 Figs. 15-22. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Terrilimosina longipexa. 15, Male terminalia, left lateral. 16, Surstylus, ventral. 17, Aedeagus and associated parts. 18, Male terminalia, posterior. 19, Male sternite 5. 20, Female terminalia, ventral. 21, Spermathecae. 22, Female terminalia, dorsal. with a triangular shining area. Dorsocentral bristles in 2 pairs, prescutellar pair twice as long as anterior pair, 1.5 times as long as intrapostalar bristles. Mid tibia with antero- apical bristle over half as long as ventroap- ical bristle (much smaller in all congeners). Wing clear, second and third costal sectors subequal, halter luteous. Abdominal syn- tergite 1+2 with a large anteromedial pale area, as wide and 1.5 times as long as tergite 3; pigmentation of tergite 3 similar to pale part of syntergite. Male terminalia.—Sternite 5 transverse, shorter than sternite 4, darkly pigmented; posterior margin long setose; posteromedial lobe constricted basally, distally with sev- eral short, stout marginal bristles and cov- ered with tufts of small, flat setulae (Fig. 19). Surstylus subquadrate in lateral view, with 6 stout bristles anteroventrally, a comb-row of long bristles on inner ventral surface, and a distally expanded, setose lobe on inner dorsal surface (Figs. 15, 16, 18). Paramere simple, broad basally, strongly narrowed apically (Fig. 17). Basiphallus very narrow, frame-like, extending ventrally beyond point of articulation with distiphallus. Distiphal- lus broad, with lightly pigmented dorsal and ventral sclerites subequal in length. Female terminalia.—Tergite 8 with tri- partite pigmentation, narrow median part pale and shining, lateral parts pruinose, ven- trolateral lobes shining brown; epiproct broader than long, posterior third setulose and with 2 bristles (Fig. 22). Sternite 8 sim- ple, entirely setulose, twice as large as hy- poproct; hypoproct setulose except for con- vex anterior process (Fig. 20). Internal VOLUME 89, NUMBER 3 genital sclerotization distinct, spectacles- shaped sclerite with small rings and a broad, bifid median piece. Spermathecae large, cup- shaped, invagination deep, about 140 de- grees from duct (Fig. 21). Types.— Holotype ¢ (CNC) and 19 para- types (10 4, 9 29, CNC, GUE, OPAVA). JA- PAN. Shikoku, Ishizuchi Mt. National Park, Omogo Valley, 700 m, 18—25.vi11.1980, dung trap, S. Peck. Other paratypes: JA- PAN. Shikoku, Ishizuchi Mt. National Park, Omogo Valley, 1400 m, 1 1-18.vii1.1980, S. Peck (1 6, 2 2, GUE). NEPAL. Katmandu, Godavan, 5000’) 1.v.1967: ~ 5400’, 2.vili.1967; 6000’, 12.viii.1967, Canadian Nepal Expedition (3 6, CNC); Taplejung Distr., Dobhan, c. 3500’, spray-splashed rocks in River Maewa, 25.1.1962, R. L. Coe (1 6, BMNH). The latter specimen was listed in Deeming (1969) along with 2 specimens of P. unio as Leptocera (Limosina) sp. (1) near paraminima (Duda). Etymology.—The specific epithet refers to the long, comb-like row of long bristles on the surstylus. Comments.—This species is most easily recognized by its complex surstylus, dis- tinctive male sternite 5, large cup-shaped spermathecae, and the unique internal scler- otization in the female abdomen. It is closely related to the Nearctic 7. pexa, as indicated by the similar surstylus and pattern of or- namentation on the male fifth sternite. The deeply invaginated, cup-shaped spermathe- cae are more similar to those of 7. racovitzai and 7. sudetica than to the long-cylindrical spermathecae of 7. pexa, but this is inter- preted as homoplasous similarity resulting from superficially similar modifications of the plesiomorphic long-cylindrical type of spermatheca. Terrilimosina smetanai Marshall, NEw SPECIES Figs. 23-30 Description. — Body length 2.2—2.6 mm. Body dark brown to black, pruinose, mi- nutely punctate; legs brown. Interfrontal 507 plate 1.5 times as high as broad, bordered by 4 subequal interfrontal bristles. Eye height 2.5 times genal height at point of maximum eye height; genal shining area broad but only extending along anterior half of lower mar- gin of eye. Dorsocentral bristles in 2 pairs, prescutellar pair 2.5 times as long as ante- rior pair, 1.5 times as long as intrapostalar bristles. Wing slightly infuscated, second costal sector subequal to third; halter lu- teous. Abdominal syntergite 1 +2 uniformly dark brown; tergite 3 similarly dark brown, slightly narrower and about 0.9 times as long as syntergite at middle. Male terminalia. —Sternite 5 twice as long as sternite 4, very darkly pigmented; pos- teromedially with a setose, convex lobe; posteromedial lobe pale distally with dark setae around base of pale part (Fig. 27). Sur- stylus elongate; posterior part laterally se- tose; anterior part dorsally with pale, blunt serrations, ventrally with an internal comb- like row of short bristles (Fig. 24). Paramere distally broad, apex broadly bifid. Basi- phallus short, broad relative to congeners, not extending ventrally (Fig. 25). Disti- phallus very narrow, elongate; with proxi- mal and distal ventral and dorsal sclerites; distal dorsal sclerite long and apically curved. Female terminalia.—Tergite 8 complete- ly pigmented, shining and bare medially, pruinose laterally and with very dark, bare lateroventral lobes. Epiproct subequal in length and width, weakly emarginate ante- riorly, bare except for 2 posterior bristles (Fig. 30). Sternite 8 simple, entirely setu- lose, subequal in size to hypoproct; hy- poproct convex anteriorly, setulose on posteromedial half only (Fig. 28). Specta- cles-shaped sclerite visible as lightly scler- otized rings only. Spermathecae long-cylin- drical, slightly widened and with a small evagination apically (Fig. 29). Types. — Holotype 6 (CNC) and 19 para- types (13 6, 6 29, CNC, GUE, OPAVA). NE- PAL. Below Thare Pati 3300 m, 13.i1v.1981, pitfall trap on a clearing in a mixed Abies, 508 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 23-30. Terrilimosina smetanai. 23, Male terminalia, left lateral. 24, Surstylus, ventral. 25, Aedeagus and associated parts. 26, Male terminalia, posterior. 27, Male sternite 5. 28, Female terminalia, ventral. 29, Spermathecae. 30, Female terminalia, dorsal. Acer, Rhododendron forest, A. Smetana. Other paratypes. NEPAL. Between Ghopte and Thare Pati, 3200 m, 23—26.iv.1985, FIT, A. Smetana (100 4, 100 2, labeled as paratypes out ofa much longer series, GUE, CNC); North slope above Syabru, intercept trap on wet meadow in subalpine forma- tion, 3800 m, 17-19.1v.1985, A. Smetana (2 6, 2 9, GUE); Taplejung Distr., Sangu, c. 6200', mixed vegetation by stream in gully, x1.1961, R. L. Coe (1 6, BMNH, listed by Deeming (1969) as Leptocera (Limosina) sp. (1) near schmitzi (Duda)); Kuwapani, 2250 m, 6.iv.1984, I. Lobl, no. 6 (1 2, GM). PA- KISTAN. Marghuzar, 1300 m, 8.v.1983, C. Besuchet—I. Lobl, No. 2b (1 6, 2 9, GM); Malam Jabba, 2500-2600 m, 18.v.1983, C. Besuchet—I. Lobl, No. 17a (1 6, GM); Mur- ree, 2100 m, 5.vi.1983, C. Besuchet—I. Lobl, No. 40 (1 6, GM). The latter 5 spec- imens have not been examined by the au- thor and are included as paratypes at the request of Dr. L. Papp, Hungarian Natural History Museum, who has compared them to the above description and illustrations. Etymology.—Terrilimosina smetanai 1s named after A. Smetana, the collector of much of the type series of this species and of many other important series of sphae- rocerids from Nepal. Comments. — This species 1s unique in the genus in having a completely sclerotized female abdominal tergite 8. This dark, lateroventrally shining sclerite is even dis- tinctive on dried, pinned specimens. Ter- rilimosina smetanai 1s also easily recog- nized by its distinctive elongate distiphallus, short basiphallus, and elongate surstylus with a short internal comb-like row of bris- tles. Terrilimosina unio Marshall, New SPECIES Figs. 31-36 Description. — Body length 1.4 mm. Body dark brown, pruinose; legs light brown. In- VOLUME 89, NUMBER 3 509 Figs. 31-36. Terrilimosina unio. 31, Aedeagus and associated parts. 32, Male terminalia, left lateral. 33, Male sternite 5. 34, Female terminalia, ventral. 35, Spermathecae. 36, Female terminalia, dorsal. terfrontal plate 1.5 times as high as wide, bordered by 4 short interfrontal bristles. Eye 2.5 times as high as gena at point of max- imum eye height; anterior part of gena with a small triangular shining area. Dorsocen- tral bristles in 2 pairs, prescutellar pair twice as long as anterior pair; 1.5 times as long as intrapostalar bristles. Wing clear to faint- ly infuscated anteriorly, second costal sector 0.7 times as long as third; halter dark brown. Syntergite 1+2 with an anteromedial pale area and dark anterolateral patches; 1.5 times as wide and as long as tergite 3. Ter- gite 3 pale brown, very pale around its edges. Male terminalia.—Sternite 5 pale pos- teromedially, with 2 spinulose lobes (Fig. 33). Surstylus laterally setulose at base; ven- trally with a small anterior lobe and an elon- gate posteroventral lobe terminating in two short, stout bristles (Fig. 32). Paramere sim- ple, broadest distally (Fig. 31). Basiphallus narrow, not extending ventrally beyond point of articulation with distiphallus. Dis- tiphallus long, narrow; with a large, dark proximal dorsal sclerite and paler proximal lateroventral sclerites; distally with a finely spinulose membrane. Female terminalia.—Tergite 8 with tri- partite pigmentation, wide median area pale and shining, lateral areas pruinose, ventro- lateral lobes shining brown; epiproct slight- ly longer than wide, anterior edge entire, bare except for 2 bristles (Fig. 36). Sternite 8 simple, darker and more than twice as large as hypoproct (Fig. 34). Hypoproct deeply emarginate anteriorly, setulose on posterior third. Spectacles-shaped sclerite visible as 2 large, lightly sclerotized rings. Spermathecae short cylindrical, bent and widened apically, with a distinct apical in- vagination (Fig. 35). Types.— Holotype ¢ (BMNH). NEPAL. Taplejung Distr., river banks below Tam- rang Bridge, c. 5500’, x—x1.1961, R. L. Coe. Paratypes: NEPAL. Taplejung Distr., be- tween Sangu and Tamrang, c. 5200’, mixed shrubs in deep gorge, x—x1.1961, R. L. Coe (1 4, abdomen missing, BMNH); mixed plants by damp cliff in deep river gorge, i- 11.1962, R. L. Coe (1 4, 2 2, BMNH); JAVA. ‘“Kandang Badok. v. Tjibodas, 18.vili.1922, Dr. Th. M. /1/6,1922” (Chibodas, near Mt. Gede, 3000 m) (1 4, 3 2, UZMC). Comments.—The holotype and one male paratype were among the specimens listed as Leptocera (Limosina) sp. (1) near para- minima (Duda) by Deeming (1969). Spec- imens of 7. /ongipexa were also included in that series. Another paratype male and one female were included by Deeming (1969) as Leptocera (Limosina) sp. 2 near schmitzi (Duda). One type female was listed by Deeming (1969) under Leptocera (Limosi- na) sp. (2) near paraminima (Duda) with some uncertainty as to its placement. The distinctive pigmentation and size of both PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ASIA ‘~ 5 i Y os Y ©” So S ¥ & @) Pe ®& £& .& & g ~ 3 y ¢ » & Y S " § & o < 2 S Pi £ RY e Oo) < OF e & Q Sy 6 & 5 6 v ND O 11+ 10 +++ 18 ++ lowes 15 ++ S++ P14 +451 744 7+ 134445 16++ 6+ 5++ 124+ 37 Att 3++ 2+ | +o Fig. 37. Phylogenetic hypothesis for the genus Terrilimosina. Characters marked (+) are thought to be especially subject to misinterpreted polarity or homoplasy. Those marked (+ +) or (+++) are thought to have a higher probability of being true synapomorphies. The numbers refer to the following putative synapomorphies: 1, surstylus with a row of bristles. 2, discal cell rounded. 3, basiphallus short, frame-like. 4, distiphallus largely membranous. 5, row of bristles on surstylus comb-like with bristles long and appressed. 6, distiphallus shortened and distally expanded. 7, paramere basally broadened. 8, posteromedial setulae on male sternite 5 flattened. 9, basiphallus extending posteriorly beyond posterior junction between distiphallus and basiphallus. 10, flattened bristles of male sternite 5 in small, scale-like tufts. 11, surstylus short, with an internal comb of long bristles. 12, sternite 5 of male densely setose posteromedially. 13, distiphallus elongate, with additional distal sclerites. 14, subanal plate broadened. 15, hypoproct reduced, emarginate. 16, paramere distally enlarged. 17, comb-row of surstylus on ventral lobe. 18, greatly enlarged syntergite 1 +2. male and female syntergites 1+2 was the major character used to associate the sexes, an association confirmed by chaetotaxy, ve- nation, eye:gena ratios and the congruent cladistic affinities of both male and female specimens listed above with male and fe- male specimens of the closely related 7. brevipexa. Etymology.—The name wnio, Latin for “one,” was chosen because the holotype was first recorded by Deeming as Leptocera (Li- mosina) sp. (1) near paraminima (Duda). Terrilimosina pexa Marshall 1985 New distributional records.—UNITED STATES. Alaska, Elliot Highway, mi. 27.8, White Mountain Trail, 16-18.vui.1985, dung traps, S. Marshall; Chena Ridge Road, 5 m1. W Fairbanks, 27.vii-12.viii.1984, malaise trap in birch and spruce, S.&J. Peck; Chena Hot Springs, mi. 50 Hot Springs Road, 28.vii-12.viii.1984, FIT, spruce moss taiga, S.&J. Peck; 12 mi. E Summit, 86 mi. ENE Fairbanks, 54 mi. SW Circle Rt. 6, 1105 m, VOLUME 89, NUMBER 3 6-13.vii. 1984, carrion trap in tundra, S.&.J. Peck. New Hampshire, Carr Co., 2.5 mi. NW Wonalancet, 2-17.x.1985, FIT, D. Chandler (1 4). Because this species was previously known only from eastern North America (New Brunswick, Ontario, Arkansas, Oklahoma), its discovery in Alaska is significant. PHYLOGENY AND ZOOGEOGRAPHY A phylogenetic hypothesis is presented in Fig. 37. One interesting aspect of this hy- pothesis is the suggestion that 7. /ongipexa, common to Nepal and Japan, shares a more recent common ancestor with the Nearctic species 7. pexa with any other species. These species in turn form the sister group of a European species. The other major species group of Terrilimosina is entirely Asian or Oriental in distribution. ACKNOWLEDGMENTS I am indebted to A. Smetana of the Bio- systematics Research Institute, Ottawa and Si S. Peck of Carleton University, Ottawa for providing me with the large numbers of Asian insect trap samples from which most of the specimens examined during this study were obtained. B. Pitkin of the British Mu- seum kindly sent me the Deeming type ma- terial from Nepal. J. Rohacéek, A. Norrbom, and L. Papp read and commented on the manuscript. LITERATURE CITED Deeming, J. C. 1969. Diptera from Nepal. Sphae- roceridae. Bull. Br. Mus. Nat. Hist., Entomol. 23: 53-74. Marshall, S. A. 1985. The genera Yenolimosina and Terrilimosina in North America. Proc. Entomol. Soc. Wash. 87: 759-769. Richards, O. W. 1961. Contribution 4 l’étude de la faune d’Afghanistan. 42. Diptera, Sphaeroceridae. Entomol. Mon. Mag. 97: 177-179. Rohaéek, J. 1983. A monograph and reclassification of the previous genus Limosina Macquart (Dip- tera, Sphaeroceridae) of Europe. Part 11. Beitr. Entomol. 33: 3-195. PROC. ENTOMOL. SOC. WASH. 89(3), 1987, pp. 512-526 SCANNING ELECTRON MICROSCOPIC STUDY OF NORTH AMERICAN POGONOMYRMEX (HYMENOPTERA: FORMICIDAE)' STEPHEN W. TABER, JAMES C. COKENDOLPHER, AND OSCAR F. FRANCKE (SWT, OFF) Department of Biological Sciences, Texas Tech University, Lubbock, Texas 79409; present addresses: (SWT) Division of Biological Sciences, University of Texas, Austin, Texas 78712; (OFF) Crown Cork de Mexico, S. A., 134 Poniente No. 583, Col. Industrial Vallejo, Mexico 16, D. F. (JCC) Department of Entomology, Texas Tech Uni- versity, Lubbock, Texas 79409.? Abstract.—A SEM study of 16 Pogonomyrmex taxa from the U.S.A. is presented. Taxa representing both groups previously referred to as subgenera and all four complexes of the nominate subgenus are examined. Some previously used surface structure terminology is corrected for the genus and the head of a bilateral gynandromorph of P. occidentalis is illustrated. The problematic nature of P. huachucanus is discussed. The ant genus Pogonomyrmex Mayr 1s confined to the New World and is repre- sented in North America by 29 species (Cole, 1968; MacKay, 1980; Snelling, 1981la; MacKay et al., 1985). These ants are com- monly known as “harvesters” because they store seeds within their nests, possibly as a food source. Most of these ants are xerophi- lous and only one species occurs east of the Mississippi River in the U.S.A. The greatest concentration of species in the U.S.A. is in the desert Southwest. Pogonomyrmex are the dominant ant throughout much of their range. Damage to cultivated crops through seed harvesting, grass cutting, and construction of large, du- rable mounds adversely affects both the farmer and urban resident. These insects often build nests near the road shoulder where extensive tunneling causes the road ' A portion of the senior author’s M.S. thesis, De- partment of Biological Sciences, Texas Tech Univer- sity, Lubbock, Texas 79409. ? Address reprint request to J.C.C., Department of Entomology, Texas Tech University. surface to crumble and collapse, forming potholes. Medically, harvester ants are 1m- portant because of their highly algogenic stings which cause anaphylactic shock in hy- persensitive victims. These ants are also of taxonomic interest because of the poorly understood relationships between conge- ners. Wheeler (1902) divided Pogonomyrmex into two subgenera, Ephebomyrmex Wheel- er and the nominate subgenus, based on the heavier sculpturing, smaller size, and re- duced psammophore of Ephebomyrmex. The subgenera created by Wheeler were retained by Creighton (1950) in his key to the ants of North America. Creighton con- sidered that the criteria chosen for the rec- ognition of Ephebomyrmex were inappro- priate and that the reproductive forms required further study. He noted that the thoraces of females of P. (E.) imberbiculus lack the elevated scutellum of Pogonomyr- mex sens. str. (Creighton, 1956). Angular metasternal flanges and an irregular or ab- sent row of gular hairs (ammochaetae) that form the psammophore distinguish Ephe- VOLUME 89, NUMBER 3 bomyrmex workers from those of the other subgenus in his key (Creighton, 1950). Gallardo (1932) claimed that certain South American species of Pogonomyrmex had incomplete psammophores and that this condition was found in both subgenera. Al- though these species seemed to be clear ex- amples of intermediate forms, Wheeler specified the absence of a psammophore as a distinguishing character between the two groups. Gallardo recommended that the subgeneric distinction be abandoned. At the other extreme, an attempt to raise Ephebomyrmex to full generic status was made by Kusnezov. According to Kusnezov (1959), Wheeler distinguished the subgenus Ephebomyrmex on the basis of the follow- ing characteristics: (1) the absence of a psammophore, (2) a small head, (3) highly curved mandibles, and (4) heavy sculptur- ing on the head and thorax. Kusnezov (1949) agreed with the importance of the psam- mophore but argued that the sculpture, head size, and mandible differences were not ac- ceptable characters for discrimination. He described Ephebomyrmex as the most primitive taxon within the genus, yet Po- gonomyrmex sens. str. could not have evolved from these ants because there are secondary derivations in the wing venation of Ephebomyrmex males and females. In addition, maxillary palp conformation and the presence or absence of a psammophore were considered definitive generic differ- ences (Kusnezov, 1959). Consequently, Kusnezov (1959) treated Ephebomyrmex and Pogonomyrmex as distinct genera in his key to the ants of Patagonia. Snelling and George (1979) and Snelling (1981b) regarded Ephebomyrmex as a sep- arate genus because its members possess poorly developed psammophores, matinal/ crepuscular foraging habits, and are omniv- orous. In the key to the desert ants of Cal- ifornia (Snelling and George, 1979), P. im- berbiculus is separated from the nominate subgenus by virtue of its distinctly off-center eye and a prominent ridge connecting the 513 weakly developed epinotal spines. MacKay and MacKay (1984) and Wheeler and Wheeler (1985) recently concurred with the treatment of Ephebomyrmex as a separate genus. The North American species of Pogono- myrmex were revised by Cole (1968), and this work is currently the standard refer- ence. Cole divided the North American members of the genus into two subgenera, Ephebomyrmex Wheeler and the nominate subgenus. The latter group contained four complexes and 24 species, whereas Ephe- bomyrmex contained three species and was not subdivided into complexes. Pogonomyrmex huachucanus Wheeler is presently considered a member of Ephe- bomyrmex and was described on the basis of workers only (Wheeler, 1914). Wheeler noted that the gular area enclosed by the psammophore and the sculpture of P. huachucanus suggest that this ant is “‘transi- tional between the species of Pogonomyrmex sens. str. and the subgenus Ephebomyrmex Wheeler.”’ Nevertheless, Wheeler placed that species in the nominate subgenus. Creigh- ton (1952) eventually described the repro- ductive forms of P. huachucanus. Cole agreed with Wheeler’s subgeneric concept and considered that the subgeneric distinction was upheld by certain “‘defini- tive characters” on the reproductive female thorax. Creighton (1956) first discovered these differences: The outline of the thorax was supposed to be smooth in Ephebomyr- mex and disrupted in Pogonomyrmex sens. str. Although the illustration by Creighton suggests that P. huachucanus is allied with the nominate subgenus, the drawing by Cole (1968) indicates a closer affinity to Ephe- bomyrmex. One of the illustrations might be inaccurate or the character might be vari- able. A single reproductive female exam- ined during the current study closely resem- bles Cole’s figure. The revision by Cole (1968) of Pogono- myrmex and all previous taxonomic de- scriptions were based upon external char- 514 acters only. Certain taxa were distinguished primarily on differences in surface detail, and scanning electron microscopy 1s ideally suited to reveal these differences. In an effort to better define some of these characters, an SEM study was undertaken. MATERIALS AND METHODS Sixteen taxa, representing both subgenera of Pogonomyrmex and all four complexes of the nominate subgenus were available for study. The collection data, museum and fig- ure numbers for these samples follow (TTU# = catalogue number of voucher se- ries in the entomological collection of The Museum, Texas Tech University): Subgenus Pogonomyrmex Mayr badius complex Pogonomyrmex badius (Latreille). Figs. 2— 6; Florida: Leon Co., Tallahassee. 25 Sep- tember 1985, TTU# 6957. barbatus complex Pogonomyrmex apache Wheeler. Figs. 11- 12; Arizona: Yavapai Co., Clarksdale. | July 1985, TTU# 6831. Pogonomyrmex barbatus (Smith). Figs. 13- 16; Texas: Brazos Co., College Station. 2 June 1985, TTU# 6964. Pogonomyrmex desertorum Wheeler. Figs. 7-10; Arizona: Cochise Co., 4.8 km E Portal. 20 June 1985, TTU# 6754. Pogonomyrmex rugosus Emery. Figs. 17- 20; Texas: Garza Co., 4.8 km E South- land. 27 August 1985, TTU# 6949. maricopa complex Pogonomyrmex californicus (Buckley). Figs. 41-45; California: Kern Co., Bakersfield. 28 June 1985, TTU# 6925. Pogonomyrmex californicus [estebanius sensu Pergande]. Figs. 46—47; California: San Bernardino Co., 4.8 km E Apple Val- ley. 29 June 1985, TT UF 6802. Pogonomyrmex comanche Wheeler. Figs. 1, 31-37; Texas: Houston Co., Grapeland. 24 September 1985, TTU# 6953. Pogonomyrmex magnacanthus Cole. Figs. 48-49; California: Riverside Co., 1.6 km W Indio. 30 June 1985, TTU# 6825. Pogonomyrmex maricopa Wheeler. Figs. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 38-40; Texas: El Paso Co., 30.4 km E El Paso. 19 June 1985, TTU# 6746. occidentalis complex Pogonomyrmex brevispinosus Cole. Figs. 25-27; California: Kern Co., 17.6 km E Shafter. 28 June 1985, TTU# 6782. Pogonomyrmex montanus MacKay. Figs. 28-30; California: San Bernardino Co., 9.6 km E Fawnskin. 29 June 1985, TTU# 6927. Pogonomyrmex occidentalis (Cresson). Figs. 21-22; New Mexico: Grant Co., 17.6 km W Jct. US 180 & NM 78. 2 July 1985, TTU# 6834. Pogonomyrmex subnitidus Emery. Figs. 23- 24; California: Los Angeles Co., 8 km S Palmdale. 27 June 1985, TTU# 6763. Subgenus Ephebomyrmex Wheeler Pogonomyrmex huachucanus Wheeler. Figs. 55-58; Arizona: Cochise Co., Huachuca Mts., Miller Canyon. 21 June 1985, TTU# 6696. Pogonomyrmex imberbiculus Wheeler. Figs. 50-54; Arizona: Cochise Co., 4.8 km E Portal. 20 June 1985, TTU# 6745. The insects were preserved in 80% eth- anol and were thoroughly dried before mounting. Representatives of the worker castes were fixed to aluminum stubs, coated with gold/palladium in a sputtering device, and placed in the vacuum chamber of the microscope. A Jeol JSM-25S scanning elec- tron microscope and a Polaroid camera us- ing Polaplan 4x5 Land Type 52 Film were used to photograph the ants. The head and thorax of each species were photographed, and additional distinctive or illustrative fea- tures were selected from certain taxa. The characters and general morphology (Fig. 1) discussed are those considered important in the revision of the genus (Cole, 1968). Ter- minology generally follows that used by Cole, but occasional use was made of works by Torre-Bueno (1962) and Harris (1979). RESULTS AND DISCUSSION The sculpturing of the head and thorax as well as the appearance of the spaces be- tween the rugae covering these structures VOLUME 89, NUMBER 3 clypeus mandible occiput scape frontal lobe frontal triangle epinotum mesonotum pronotum ‘ \ CAS PS Fig. 1. have been considered important specific characters, and these spaces are usually de- scribed as “smooth” or “punctate.” During this study many specimens were identified according to this system using a dissecting microscope, but data obtained from elec- tron micrographs proved this terminology to be inappropriate in every case encoun- tered. For example, the interrugal spaces of P. montanus appear strongly punctate under the light microscope which suggests that the surface is covered with punctures or pits. Electron microscopy (450 x) revealed tiny rugules that are interconnected in a manner that is best described as ‘‘areolate”’ (Harris 1979). Accordingly, this term is used in place of “punctate” in the following discussions. epinotal spine petiole C3 postpetiole psammophore (beard) gaster sting Gross morphology of Pogonomyrmex comanche worker. Subgenus Pogonomyrmex A. The badius complex.— This complex is composed of only one species, P. badius. It is the only polymorphic species in North America, the only species to occur east of the Mississippi River, and it is largely al- lopatric with other congeners over its entire range. The workers are divided into minor, medium, and major size classes, whereas all other species (except for Pogonomyrmex coarctatus Mayr of South America) main- tain only one worker size class. The large head of the major appears partially bilobed (Fig. 2), and there are no spines on the epi- notum (Fig. 3). The scutellum is developed; in all other North American species this 516 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 2-7. Morphology of Pogonomyrmex, scale line = 1.0 mm. 2-6, P. badius: 2, Major; head, frontal view. 3, Major; thorax, lateral view. 4, Minor; head, frontal view. 5, Minor; thorax, lateral view. 6, Minor; head, lateral view. 7, P. desertorum: head, frontal view. structure 1s present only in the reproductive castes. The minor worker is distinguished by the shape of its head (Fig. 4), which is narrowed behind the eyes. The thorax is less devel- oped (Fig. 5) than that of the major worker, and a lateral view of the head (Fig. 6) reveals the beard for which the genus was named. B. The barbatus complex.—As illustrated by P. desertorum, these taxa are distin- guished from all other members of the sub- genus by a broad head, extended clypeal lobes, straight and parallel cephalic rugae (Fig. 7), and the presence of long, erect hairs on the venter of the petiolar peduncle (Fig. 8). This species is distinguished by the com- bination of very fine cephalic rugae (Fig. 9), tapering epinotal spines, and a prominent VOLUME 89, NUMBER 3 Syh7/ Figs. 8-13. Morphology of Pogonomyrmex, scale line for Fig. 9 = 0.1 mm, all others = 1.0 mm. 8-10, P. desertorum: 8, Thorax, lateral view. 9, Cephalic rugae, frons. 10, Thorax, dorsolateral view. 11-12, P. apache: 11, Head, frontal view. 12, Thorax, lateral view. 13, P. barbatus: head, frontal view. ventral lobe on the petiolar peduncle (Fig. 10). Pogonomyrmex apache is readily identi- fied by its deeply excised clypeus (Fig. 11). Like P. desertorum, this species possesses fine cephalic rugae, but the thorax bears no dorsal spines (Fig. 12). Pogonomyrmex barbatus is characterized by coarser cephalic rugae (Fig. 13) than the two previous taxa. The head of this species, like other members of the group, is broad. Unlike P. apache, this species possesses well- developed epinotal spines (Figs. 14 and 15). A close-up view of the sculpturing and the epinotal spiracle is presented (Fig. 16). The head of P. rugosus bears the coarsest cephalic rugae (termed costulate by Harris, 1979) in the complex (Figs. 17 and 18). 518 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 14-19. Morphology of Pogonomyrmex, scale line for Figs. 15, 16 and 18 = 0.1 mm, all others = 1.0 mm. 14-16, P. barbatus: 14, Thorax, lateral view. 15, Epinotal spines. 16, Epinotal spiracle. 17-19, P. rugosus: 17, Head, frontal view. 18, Head, lateral view. 19, Thorax, lateral view. Widely spaced, wavy rugae and epinotal spines are evident on the pronotum and epi- notum, respectively (Fig. 19). The rugae are often connected with coarse reticulations (termed porcate by Harris, 1979) (Fig. 20). One of the taxonomic problems presented by the genus is the occurrence of forms that are apparently intermediate between P. bar- batus and P. rugosus. Harris (1979) pub- lished photomicrographs of a head and tho- racic spiracle region of what he identified as ‘**Pogonomyrmex barbatus rugosus.” C. The occidentalis complex.— Members of this complex are characterized by subquadrate heads and more divergent ce- phalic rugae than those of the barbatus com- VOLUME 89, NUMBER 3 Figs. 20-25. 19 " Qn. LP ig TS TIS Morphology of Pogonomyrmex, scale line for Fig. 20 = 0.1 mm, all others 1.0 mm. 20, P. rugosus: epinotal spiracle. 21-22, P. occidentalis: 21, Head, frontal view. 22, Thorax, lateral view. 23-24, P. subnitidus: 23, Head, frontal view. 24, Thorax, lateral view. 25, P. brevispinous: head, frontal view. plex (Fig. 21). The offset basal mandibular tooth which is diagnostic for P. occidentalis is also present (Fig. 21). Epinotal spines are always present in this group and the thoracic dorsum is slightly arched (Fig. 22). The cephalic interrugal spaces of P. sub- nitidus are not areolate as are those of P. occidentalis, but instead are glabrous (Fig. 23). Most of the thoracic interrugal spaces present the same smooth appearance and the spines are well developed (Fig. 24). Members of the occidentalis complex generally possess wide antennal scape bases (Fig. 25). The cephalic interrugal spaces of P. brevispinosus are areolate unlike those of P. subnitidus, and the epinotal spines are short (Fig. 26). Areolate thoracic rugae sur- round the metathoracic spiracle (Fig. 27). 520 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 26-31. Morphology of Pogonomyrmex, scale line for Figs. 27, 29, and 31 = 0.1 mm, all others = 1.0 mm. 26-27, P. brevispinosus: 26, Thorax, lateral view. 27, Metathoracic spiracle. 28-30, P. montanus: 28, Head, frontal view. 29, Cephalic interrugal spaces. 30, Thorax, lateral view. 31, P. comanche: scape base. MacKay (1980) described a species of harvester ant which he associated with the occidentalis complex of Cole. The cephalic interrugal spaces of P. montanus are strong- ly areolate (Figs. 28 and 29). The thorax is highly reticulate and the basal face of the epinotum is elevated (Fig. 30). D. The maricopa complex.—Represen- tatives of all four members of this group were available for study. The workers may be distinguished from those of the occiden- talis complex by the weakly enlarged anten- nal scape base as shown by that of P. co- manche (Figs. | and 31). The cephalic rugae VOLUME 89, NUMBER 3 ra Figs. 32-37. 321 yee ae +a il apie (ib ; pt. Morphology of Pogonomyrmex, scale line for Figs. 33, 34, and 37 = 0.1 mm, all others = 1.0 mm. 32-37, P. comanche: 32, Head, frontal view. 33, Oculomandibular area. 34, Cephalic interrugal spaces. 35, Thorax, lateral view. 36, Thorax, dorsolateral view. 37, Mesothoracic spiracle. are often coarse (Fig. 32) and may form con- centric whorls around the eyes. Cephalic re- ticulation may be present (Fig. 33), but much of the interrugal surface is smooth (Fig. 34). All members of the maricopa complex pos- sess a strongly arched thoracic dorsum, and epinotal spines may be present (Figs. 1 and 35). Both spines (Fig. 36) are present in dor- solateral view, and extremely fine hairs sur- round the mesothoracic spiracle (Fig. 37). The head of P. maricopa is presented in frontal view (Fig. 38). A higher magnifica- 522 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ey eee hy Figs. 38-43. Morphology of Pogonomyrmex, scale line for Fig. 42 = 0.1 mm, all others = 1.0 mm. 38-40, P. maricopa: 38, Head, frontal view. 39, Psammophore. 40, Thorax, lateral view. 41-43, P. californicus: 41, Head, frontal view. 42, Cephalic interrugal spaces. 43, Thorax, lateral view. tion reveals the psammophore (Fig. 39). The thoracic dorsum is convex (Fig. 40). The specimen illustrated is from a population known as “‘maricopa Variant No. 3” (Cole, 1968). Workers of this variety bear epinotal armature, although the spines may be short. A species often confused with P. mari- copa is P. californicus. However, the ce- phalic rugae are coarser and reticulation is sparse or absent in the latter species (Figs. 41 and 42). The epinotum is never armed and the epipleura are smooth and costate (Fig. 43). A micrograph of the thorax is also provided by Harris (1979). The mesopleu- ron bears groups of long, barbed hairs and short, simple hairs (Fig. 44). The shallow frontal triangle is characteristic of both the occidentalis and macricopa complexes (Fig. VOLUME 89, NUMBER 3 323 Sane Figs. 44-49. Morphology of Pogonomyrmex, scale line for Figs. 44 and 45 = 0.1 mm, all others = 1.0 mm. 44-45, P. californicus: 44, Mesopleuron. 45, Frontal triangle. 46-47, P. californicus [estebanius sensu Pergande]: 46, Head, frontal view. 47, Thorax, lateral view. 48-49, P. magnacanthus: 48, Head, frontal view. 49, Thorax, lateral view. 45). Photomicrographs of the head of this species have been published by Scharf (1977: 48) under the name “California Harvester Ant.” Pogonomyrmex californicus estebanius Pergande was synonymized under P. cali- fornicus by Cole (1968). This subspecific name was given to those taxa presenting a bicolored appearance (the gaster is brown or black, whereas the anterior portion of the ant is red). Cole (1968) noted that eastern populations of P. californicus were charac- terized by finer cephalic rugae and increased “‘punctation”’ on the pronotum. A compar- ison of the previous “typical” P. californicus micrographs with those of a P. californicus 524 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 50-55. Morphology of Pogonomyrmex, scale line for Figs. 53 and 54 = 0.1 mm, all others = 1.0 mm. 50-54, P. imberbiculus: 50, Head, lateral view. 51, Head, frontal view. 52, Thorax, lateral view. 53, Pedicel, lateral view. 54, Epinotal spine. 55, P. huachucanus: head, frontal view. [estebanius sensu Pergande] specimen col- lected 197 km southeast of the former (Figs. 46 and 47) supports the contention of Cole (1968), but further study of intermediate samples is needed before the status of the bicolored populations can be accurately de- termined. Pogonomyrmex magnacanthus is a small ant which may be readily identified by a pair of exceptionally large compound eyes (Fig. 48). The thorax bears no epinotal spines (Fig. 49). Subgenus Ephebomyrmex This group is represented in the U.S.A. by three species, one of which was not avail- able for study. Pogonomyrmex imberbiculus (Fig. 50) has a relatively undeveloped psam- VOLUME 89, NUMBER 3 525 Figs. 56-59. Morphology of Pogonomyrmex, scale line for Figs. 57 and 58 = 0.1 mm, all others = 1.0 mm. 56-58, P. huachucanus: 56, Thorax, lateral view. 57, Epinotal spines. 58, Pedicel, lateral view. 59, P. occidentalis (gynandromorph): Head, frontal view. mophore, one of the characters chosen by Wheeler (1902) to justify the creation of the subgenus. The coarse rugosity and heavy interrugal reticulation shared by all mem- bers of the group 1s also apparent. The com- pound eyes of these species are placed below the center of the head (Fig. 51). The prom- inent clypeal angles are diagnostic for P. im- berbiculus (Fig. 51). Extensive reticulation and coarse rugosity cover the thorax (Fig. 52). A relatively massive postpetiole is pres- ent (Fig. 53), and numerous barbed hairs arise from the epinotal spines (Fig. 54). One of the most problematic species in the genus 1s P. huachucanus. It 1s often de- scribed as intermediate between Pogono- myrmex sens. str. and the other two North American Ephebomyrmex species. The most obvious such character is the relative size of the ants. Pogonomyrmex imberbiculus is the smallest North American species in the genus, whereas P. huachucanus 1s larger, but smaller than almost all members of the nominate subgenus. Rugosity and com- pound eye placement of P. huachucanus ap- pear “intermediate” (Fig. 55). The thorax is not as heavily sculptured (Fig. 56), and the epinotal spines are more slender than those of P. imberbiculus (Fig. 57). The less abundant hairs found on these spines are relatively simple, but the postpetiole is pro- portionately large, as is that of P. imberbi- culus (Fig. 58). Sufficient differences between species of Pogonomyrmex and Ephebomyrmex exist to recognize two genera, as currently done by several other authors. These differences will be further documented in a forthcoming paper on the chromosomes of these species. Some question on the placement of “EF.” huachucanus still remains. The present study and the unpublished data on chromosomes 526 suggest that this species represents a third group, but we refrain from describing such a group. For the present, it appears best to retain this species in Ephebomyrmex. Stud- ies on the third “subgenus” of Pogonomyr- mex and other species complexes from South America and the West Indies should be in- structive, and may aid in an accurate generic placement of “EF.” huachucanus. The only known bilateral gynandro- morph of the genus was described without the use of micrographs by Taber and Francke (1986). The head of that specimen of P. oc- cidentalis has a male right side and a pre- dominantly female left side (Fig. 59). ACKNOWLEDGMENTS Thanks are extended to Scott Stockwell for providing valuable technical advice and assistance. Sanford Porter kindly provided the Florida specimens of P. badius. M. Kent Rylander, Robert Sites, and Harlan Thor- vilson reviewed the manuscript. Willard A. Taber and Ruth Ann Taber (parents of S.W.T.) are thanked for their encourage- ment and their advice as professional bi- ologists. This study was supported in part by the Texas Department of Agriculture Inter- agency Agreement IAC (86-87)-0800, and is Contribution No. T-10-179, College of Agricultural Sciences, Texas Tech Univer- sity. LITERATURE CITED Cole, A. C., Jr. 1968. Pogonomyrmex Harvester Ants: A Study of the Genus in North America. Univ. of Tennessee Press, Knoxville. 222 pp. Creighton, W. S. 1950. The ants of North America. Bull. Mus. Comp. Zool. 104: 1-585 + 57pls. . 1952. Studies on Arizona ants (3). The habits of Pogonomyrmex huachucanus Wheeler and a description of the sexual castes. Psyche 59: 71-81. 1956. Studies on the North American rep- resentatives of Ephebomyrmex (Hymenoptera: Formicidae). Psyche 63: 54-66. Gallardo, A. 1932. Las hormigas de la Republica Argentina. Subfamilia Mirmicinas segunda sec- cion Eumyrmicinae Tribu Myrmicini (F. Smith) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Género Pogonomyrmex Mayr. An. Mus. Nac. Hist. Nat. 37: 89-169. Harris, R. A. 1979. A glossary of surface sculpturing. California Dept. Agric., Occas. Papers Entomol., No. 28, 31 pp. Kusnezov, N. 1949. Pogonomyrmex del grupo Ephe- bomyrmex en la fauna de la patagonia (Hyme- noptera, Formicidae). Acta Zool. Lilloana 8: 29 1- 307. 1959. La fauna de hormigas en el oeste de la Patagonia y Tierra del Fuego. Acta Zool. Lilloana 17: 321-401. MacKay, W. P. 1980. A new harvester ant from the mountains of southern California (Hymenoptera: Formicidae). Southwest. Nat. 25(2): 151-156. MacKay, E. E. and W. P. MacKay. 1984. Apoyo a la actual division generica de hormigas usando etologia comparativa (Hymenoptera, Formici- dae). Folia Entomol. Mex. (61): 179-188. MacKay, W. P., E. E. MacKay, J. F. Perez Dominguez, L. J. Valdez Sanchez, and P. Vielma Orozco. 1985. Las hormigas del Estado de Chihuahua Mexico: El genero Pogonomyrmex (Hymenoptera: For- micidae). Sociobiology 11: 39-54. Scharf, D. 1977. Magnifications. Photography with the Scanning Electron Microscope. Schocken Books, New York. 119 pp. Snelling, R. R. 198la. The taxonomy and distribu- tion of some North American Pogonomyrmex and descriptions of two new species (Hymenoptera: Formicidae). Bull. South. Calif. Acad. Sci. 80(3): 97-112. 1981b. Systematics of social Hymenoptera, pp. 369-435. Jn H. Hermann, ed., The Social In- sects. Vol. 2. Academic Press, New York. Snelling, R. R. and C. D. George. 1979. The Tax- onomy, Distribution, and Ecology of California Desert Ants (Hymenoptera: Formicidae). U.S. Dept. Interior, Bur. Land Management, California Desert Plan Program. 334 pp. + 331 figs. Taber, S. W. and O. F. Francke. 1986. A bilateral gynandromorph of the western harvester ant, Po- gonomyrmex occidentalis (Hymenoptera: Formi- cidae). Southwest. Nat. 31: 274-276. Torre-Bueno, J. R. de la. 1962. A Glossary of Ento- mology. New York Entomol. Soc., New York. 336 pp. + 9 pls. Wheeler, G. C. and J. Wheeler. 1985. A checklist of Texas ants. Prairie Nat. 17(2): 49-64. Wheeler, W. M. 1902. New agricultural ants from Texas. Psyche 9: 387-393. 1914. New and little known harvesting ants of the genus Pogonomyrmex. Psyche 21: 149-157. Note: The negatives for Figures 2-59 were provided by the authors and were not photographed by Allen Press, Inc. PROC. ENTOMOL. SOC. WASH. 89(3), 1987, pp. 527-531 A NEW KATHROPERLA SPECIES FROM WESTERN NORTH AMERICA (PLECOPTERA: CHLOROPERLIDAE) BILL P. STARK AND REBECCA F. SURDICK (BPS) Dept. of Biology, Mississippi College, Clinton, Mississippi 39058; (RFS) Ento- mological Laboratory, Rt. 2, Box 1072, Front Royal, Virginia 22630. Abstract. — Kathroperla takhoma, new species, is described from both sexes collected in California and Washington. Eggs of the species are compared with those of K. perdita (Banks) from scanning electron photomicrographs. Diagnoses are given for male and female Kathroperla, but nymphs of the two species are presently indistinguishable. Banks (1920) proposed the genus Kath- roperla for a unique species, K. perdita, known at that time from a single female from British Columbia. Needham and Claassen (1925) added the male description and Neave (1934) figured the nymph. Since these early studies, Kathroperla has been ac- cepted as a distinctive, monotypic genus ranging from the Sierras and Cascades to the northern Rocky Mountains (Jewett, 1959; Baumann et al., 1977). However, some females from Washington and Cali- fornia and an associated male from Mt. Rainier are unlike other Kathroper/a in the shape of the female subgenital plate and length of cercal segments, and eggs obtained from these females differ significantly from Kathroperla eggs described by Knight et al. (1965). Therefore, we are describing a new species on the basis of these specimens. Kathroperla takhoma Stark and Surdick, New SPECIES Male.— Forewing length 15 mm, general coloration and elongate head similar to K. perdita; head dorsum yellow except for dark ocellar triangle and posterolateral dark stripes (Fig. 8). Tergum 10 with a pair of small posterior patches of sensilla basicon- ica located lateral to mesal sclerotized stripe; basal cercal segments about three times as long as wide (Fig. 1). Lateral margins of vesicle strongly convergent basally giving stalked appearance (Fig. 2); vesicle surface covered with small sensilla basiconica. Ae- deagus bulbous with multiple membranous lobes and a curved pair of lateral sclerites; fine setae located in a ventrolateral band apically and more sparsely on lateral scler- ites (Fig. 3). Female.— Forewing length 18 mm. Pig- mentation and head shape similar to male; basal cercal segments three times as long as wide. Subgenital plate notched apically; lat- eral margins strongly convergent at apical third forming a distinctly offset apical tab (Fig. 4); apex of plate reaches posterior mar- gin of sternum 9. Egg.— Elongate, oval, length .3 mm, width .2 mm; collar absent (Fig. 10). Chorionic surface covered with scattered tuberculate projections; some projections arranged in irregular, poorly organized rows (Fig. 11); surface granular around projections. Micro- pyles set on projections in apical third of egg (Fig. 12). Nymph.—Presently indistinguishable from K. perdita. Etymology.—The species name, tak- homa, a noun in apposition, is derived from the Yakima Indian name for Mt. Rainier. Types.—Holotype 2 and allotype 3: 528 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON RES Figs. 1-4. Kathroperla takhoma genitalic structures. 1, Male terminalia, dorsal. 2, Male sternum 9 with vesicle. 3, Aedeagus, lateral (dorsum directed to right). 4, Female terminalia with subgenital plate. VOLUME 89, NUMBER 3 ai a TK ad ee ae \\ Ve Ve Wr \\ . tel / \\ WBS \ \\ athe wh ae Gime ON \ OW ite AGN gia EON aoe | Shi [Vs AN /\ | ee aly 2 VN Ay Nos | a | hile 4 PUA | yee ate \ \ ~ \ « | | ( / A A tS x Ss yl J ( Lia | , A ale eo pal i ! on 9 RFS 5 \ 8 | Na a \ ‘ i SEGA I NNN XN : / } / ett sf Cet) j SR il Ga / t v / 7; ash a ey Vi [jl ; |__AY NY \ f' KI Nae 7 \ | V/ Spf i f J Hf vi / 4 \g AI \ 6 2 \ va 4 ' iy 4 \ x x Yi w bs a Ve PA y aay =) leper [:! Figs. 5-9. Kathroperla genitalic structures and color patterns. 5, K. perdita female subgenital plate. 6, K. perdita left side of male terminalia showing basal cercal segments. 7, K. perdita vesicle. 8, K. takhoma head. 9 K. perdita head. WASHINGTON, Mt. Rainier National park, Falls Creek above junction Ohana- pecosh River, Hwy. 706, 15-VI-69, R. W. Baumann. Holotype and allotype deposited in the United States National Museum of Natural History. Paratype: CALIFORNIA: Mendocino Co., Barnwell Creek, 15-IV-85, R. L. Bottorff, | 2 reared (Stark collection). Diagnosis.—The female of K. takhoma may be distinguished from that of K. perdita by the length and outline of the subgenital plate as well as by egg morphology. The subgenital plate of K. perdita does not reach the posterior margin of sternum 9 and the lateral margins are not abruptly convergent near the apex (Fig. 5) as they are in K. tak- homa (Fig. 4). The two species are easily differentiated by their distinctive eggs. Eggs of K. perdita have a wide collar, and the tuberculate projections are organized into striations (Fig. 13), but eggs of K. takhoma lack a collar, and the tuberculations are scat- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 530 we © ce ¢ ore 665*. Micropyle.) 11, K. takhoma, anterior pole, Kathroperla eggs. 10, K. takhoma, lateral aspect, 266 x. 12, K. takhoma micropyle and chorionic detail, 2660 x. Figs. 10-13. 13, K. perdita, lateral aspect, 274 =. (M VOLUME 89, NUMBER 3 tered (Figs. 10, 11). Males are most easily distinguished by the cerci and by the shape of the vesicle. In K. perdita, the basal cercal segments are about twice as long as wide (Fig. 6) and the vesicle base, although ta- pered to about half the vesicle apex, is not stalk-like (Fig. 7). In K. takhoma males, the basal cercal segments are about three times long as wide and the vesicle is strongly con- vergent basally (Figs. 1, 2). Adults are also distinguished by head and pronotal color patterns. In K. takhoma, the occiput is bright yellow posterior to the oc- cipital suture except for dark lateral stripes (Fig. 8) and the lateral pronotal margins are yellow. In K. perdita, the occiput is uni- formly dark (Fig. 9) and the lateral pronotal margins are dark. Needham and Claassen (1925) noted the distinctive color pattern typical of K. takhoma in a California female that they included in the description of K. perdita. Specimens of K. perdita from the follow- ing localities were examined for this study: CALIFORNIA: El Dorado Co., North Co- sumnes Riv. at Grizzly Flats; Tehama Co., no locality. MONTANA: Broadwater Co., Deep Creek; Flathead Co., Essex Creek, Au- tumn Creek; Lake Co., Yellow Bay Creek; Missoula Co., Rattlesnake Creek. ORE- GON: Douglas Co., Muir Creek. WASH- INGTON: Mt. Rainier National Park, Ste- vens Creek below Lake Louise. ACKNOWLEDGMENTS We gratefully acknowledge the loan of Kathroperla specimens by R. Bottorff (Univ. Soi of California-Davis), R. W. Baumann (Monte L. Bean Museum, Brigham Young Univ.), M. A. Ivie (Montana State Univ.), and K. W. Stewart (North Texas State Univ.). N. D. Penny (California Academy of Sciences) graciously arranged for one of us (BPS) to dissect eggs from museum spec- imens and S. W. Szczytko (Univ. of Wis- consin-Stevens Point), D. L. Lentz (Univ. of Mississippi School of Dentistry) and A. B. Hogue (Mississippi College) assisted in the preparation of SEM micrographs. This study was supported by NSF grant #BSR 840755. LITERATURE CITED Banks, N. 1920. Perlidae. Jn New Neuropteroid In- sects. Bull. Mus. Comp. Zool. 64: 314-325. Baumann, R. W., A. R. Gaufin, and R. F. Surdick. 1977. The stoneflies (Plecoptera) of the Rocky Mountains. Mem. Am. Entomol. Soc. 31: 1-208. Jewett, S.G. 1959. The stoneflies (Plecoptera) of the Pacific Northwest. Oreg. State Monogr. Stud. Entomol. 3: 1-95. Knight, A. W., A. V. Nebeker, and A. R. Gaufin. 1965. Description of the eggs of common Plecoptera of western United States. Entomol. News 76: 105- Halil Neave, F. 1934. Stoneflies from the Purcell Range, B. C. Can. Entomol. 66: 1-6. Needham, J. G. and P. W. Claassen. 1925. A mono- graph of the Plecoptera or stoneflies of America north of Mexico. Thomas Say Foundation Ento- mol. Soc. Am. 2: 1-397. PROC. ENTOMOL. SOC. WASH. 89(3), 1987, pp. 532-547 CHIROSIA BETULETI (RINGDAHL) (DIPTERA: ANTHOMYIIDAE) A GALL-FORMER ON THE OSTRICH FERN, MATTEUCCIA STRUTHIOPTERIS, WITH NOTES ON OTHER INSECT-FERN ASSOCIATES P. VON ADERKAS AND B. V. PETERSON (PvA) Atlantic Research Laboratory, National Research Council, 1411 Oxford Street, Halifax, Nova Scotia B3H 3Z1, Canada; present address: Canadian Forestry Service— Maritimes, Fredericton, New Brunswick, E3B 5P7, Canada; (BVP) Systematic Entomology Laboratory, BBII, Agricultural Research Service, USDA, % National Museum of Natural History, NHB-168, Washington, D.C. 20560, U.S.A. Abstract. — Larvae of Chirosia betuleti cause the formation of cover galls in the vegetative leaves of the ostrich fern, Matteuccia struthiopteris. This is the first record of this antho- myiid fly in North America. Females lay their eggs on the unfurling fronds. Newly emerged fly larvae feed on glandular hairs causing the pinnules to curl around the midrib, and partially or completely loop around the feeding larvae. Fly eggs and larvae have been found contaminating the fiddleheads of this edible fern. Larvae were parasitized by bra- conid wasps of the genus Aphaereta and eulophid wasps of the genera Dimmockia and Elachertus. Larvae of the chloropid fly, E/achiptera costata, were secondary invaders of the vacated galls. Larvae of Chirosia betuleti and Elachiptera costata are described in detail for the first time. Effects of the gall on leaf growth are outlined, and the distribution of the galls in a sampled population is described. A cursory description of a gall caused by the tortricid moth, Olethreutes auricapitana, is given, and a list of the primary insects collected and identified from the fern is also provided. Species of the anthomyiid genus Chirosia Rondani have been reared from several fern species. Most descriptions of the life cycles of these flies and their effect on the host plant are from the bracken fern, Pteridium aquilinum (L.) Kuhn (Meijere, 1911; Cam- eron, 1930; Meikle, 1937; Brown and McGavin, 1982). Chirosia betuleti (Ring- dahl) is known from the fern genera Preridi- um and Athyrium in Europe (Meijere, 1911; Seidel, 1957) but has not been recorded pre- viously from Matteuccia nor from North America. Matteuccia struthiopteris (L.) Todaro, the ostrich fern, is found along flood plains in the Northern Hemisphere, where it devel- ops large clonal populations. Chirosia be- tuleti 1s closely associated with this fern species in eastern Canada, and it is the na- ture of this association, as well as the effect it has on the fern plant and population, which we describe in this paper. We also describe the unique immature stages of this fly species. During the course of this inves- tigation, a number of other insects were col- lected from Matteuccia struthiopteris or in association with vacated galls. Principal among these was the chloropid fly, Ela- chiptera costata (Loew), whose larva also is described below. A list of the other primary insects collected from the fern during this study is also provided. VOLUME 89, NUMBER 3 MATERIAL AND METHODS Field collections. —Galls from the ostrich fern were collected initially in June, 1981, from plants growing along Five Mile River, near South Maitland, Hants County, Nova Scotia. In July of the same year, galls were taken into the laboratory and placed in per- forated plastic bags. The flies that emerged from these galls were mounted and identi- fied. In the following years, galls were again collected from the same site, as well as from four other locations: Lattie’s Brook, River Herbert, and Meander River, all in Hants County, as well as at Cape Split, Kings County, Nova Scotia. Crowns of the fern were collected in late summer and in early autumn of 1983 and taken to the laboratory for dissection. In 1984, collections were made weekly on the Five Mile River site, beginning May 29, until the first frost in mid-September. In addition, plants were assessed 1n surveyed transects for a number of leaf characteristics and gall information. Plants were measured in a stand of ferns divided into transects 20 m in length. Each week a transect was cho- sen and plants selected at | m intervals. Leaf dry weight, leaf type, extension and number were measured, and the presence or absence of galls noted. Details on leaf sampling are recorded elsewhere (Aderkas and Green, 1986). In 1985, a larval collection was made June 16, from the Five Mile River site. The lar- vae were kept in Sterilite® jars on filter pa- per moistened with 0.5% Javex® in distilled deionized water. Pupae, which developed by July 16, were maintained in the same way as the larvae and kept at 4°C until No- vember, after which time, at monthly in- tervals, samples were removed and held at 20°C. In this way three males and a female were recovered. In addition, information on the occur- rence and effect of galls on leaf development was collected from plants removed from a tagged population on the Five Mile River 333 site. These excavated rhizomes and leaves were frozen until examined. The details on leaf characteristics, most of which are not relevant to this study, will be presented else- where. During the course of pulling apart these ferns and looking at various morpho- logical features, two puparia were found be- tween leaf bases of different rhizomes. Preparation of specimens.—All imma- ture stages of the flies were preserved in 70% ethanol for further processing. Specimens for scanning electron microscope study were removed from the ethanol preservative, freeze dried, and glued to the observation stub. Plant specimens were hand sectioned with a straight razor and stained with 0.5% Toluidine Blue 0 in 1% borate (aqueous). Sections of affected and unaffected tissue were viewed under a Leitz Dialux 20 mi- croscope. RESULTS Insect phenology and effects on leaf de- velopment.—Eggs of Chirosia betuleti were laid in early May on the inside of leaflets, on the unfurling portion of the coiled veg- etative leaf. They adhered to the sticky hairs, present at this time, on the midrib. Emerg- ing larvae fed on the numerous glandular trichomes in the upper quarter of the groove of the frond rachis. Feeding caused a local- ized reaction in the expanding frond such that the young leaflets in the immediate vi- cinity remained stunted, and curled over the midrib (Fig. 28). In most fronds there were further de- velopments. Along that portion of the mid- rib on which the larva(e) fed, there was a subsequent partial or, less commonly, a complete loop of the rachis. Often, only a three-quarter twist was completed such that the resulting gall was a sack-like or U-shaped alteration of the normal course of the mid- rib. In all galls, the larvae were covered by the pinnules (Fig. 27). The portion of the leaf anterior to the feeding larvae continued to develop normally (Fig. 28). At maturity, 534 the leaf was normal in overall shape and size, except for the affected portion which appeared as a looped whorl of pinnules. In some plants, leaves were found in which the upper pinnules completely failed to un- furl but were kept rolled together over a feeding larva (Fig. 28, O). These pinnules died and fell off, leaving a truncated frond. This condition was found to be caused by the larva of the lepidopteran Olethreutes au- ricapitana (Walsingham) (Tortricidae). Only one gall was formed per leaf and only rarely was the gall occupied by more than a single larva of Chirosia betuleti. Lar- vae that had eaten most of the trichomes of the rachis continued along the covering pin- nules, as well as feeding on epidermal and collenchymatous hypodermal tissues along the adaxial side of the rachis. There was no expansion or other deformation of the mid- rib. Midribs were never mined since feeding of the larva was restricted to the adaxial surface of the rachis. With continued feed- ing these areas became black from the ac- cumulation of frass and the enzymatic and bacterial breakdown of the epidermal and hypodermal tissues. Matteuccia struthiopteris sporophytes ini- tially produced vegetative leaves, followed six weeks after by fertile fronds and cata- phylls. In terms of leaf types, galls were found on vegetative fronds, the only leaf type pres- ent at the time the eggs of Chirosia betuleti were being laid. Three larval instars were present through May and June. First instar larvae were col- lected from the middle of May until the first week in June, second instar larvae were present from early June until the beginning of July, and mature larvae were found from mid-June until mid-July. Puparia, which developed from as early as mid-June until mid-July, dropped out of the galls to the soil surface or, occasionally, between leaf bases of the fern rhizome, where they re- mained over autumn and winter. In the lab- oratory, larvae were maintained for rearing to pupae, but they did not undergo any fur- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ther development over the summer. About 20% of the larvae of C. betuleti were heavily parasitized by an undescribed braconid wasp closely related to Aphaereta pallipes (Say) which is a common, widespread, polypha- gous species parasitizing a number of an- thomyiid and other muscoid flies. The new wasp will be described by Robert A. Whar- ton, Texas A&M University, College Sta- tion, Texas. These wasps were found in the larvae only after mid-June and prevented the pupae from completing their develop- ment to the adult stages. The wasps did not require a cold period prior to emergence, though they might occasionally overwinter within the host as determined from pupae studied in the laboratory. A single female of a species of the eulophid wasp, Dim- mockia Ashmead, was found in a container with parasitized and non-parasitized pupae being reared to adults, and four males and six females of Elachertus loh Schauff (de- scribed as new with the holotype female from our material) were also found. Schauff (1985) cited the host of the latter species simply as a leafroller and stated that the specimens from Nova Scotia were reared on ferns. Of 320 fern plants sampled in the weekly transects, 114 plants had insect galls on their leaves. Plants with galls also had signifi- cantly (P < 0.01: t. test) more vegetative leaves (X + SE = 6.5 + 1.0) than plants without galls (5.2 + 0.8). Fewer plants than expected had one or two galls and many more plants than expected had three or more galls. Test of fitness (G test) showed that the number of galls per plant did not follow a Poisson distribution, but indicated a clump- ing of galls. Among plants with galls, there was an average of 2.4 + 1.3 galls per plant. Occasionally, plants were observed with all leaves inhabited; in one instance, eight galls per eight leaves were found. The highest number of galls found was nine in a plant that possessed 13 leaves. Plants with galls did not necessarily have shorter average leaf lengths (galled leaves—79.0 + 22.0 mm; unaffected leaves—81.0 + 14 mm). The VOLUME 89, NUMBER 3 presence of the fly larvae distorted the ex- pansion of the pinnae about the gall, but not the overall height expansion of the leaf. At the Cape Split site, galls were observed on two plants of Dryopteris carthusiana (Villars) Fuchs, but the absence of larvae at the time of observation (late June) preclud- ed positive identification of the gall former. However, the peculiar damage indicates that a species of Chirosia probably was respon- sible for these galls as well. Infestation differed from one year to the next. In 1984, 36% of the fern plants were affected, but in 1985, only 30% were af- fected. Of the originally affected plants, 60% were infected two years in a row. In the latter part of June, pupae of Chi- rosia betuleti were falling to the ground when females of the chloropid fly, Elachiptera costata (Loew) laid their eggs 1n the vacant cover galls. Generally, only one chloropid larva inhabited each gall, but sometimes two were found, and as many as ten were count- ed jointly inhabiting one gall. Larvae fed on superficial plant tissue within the gall and pupated by mid-July; adults emerged at the end of July and the beginning of August. No parasites were found associated with the im- matures of this species. In the course of one year, F. costata com- pleted two generations. Second generation larvae inhabited late developing leaves (cataphylls) nearest the crown, destroying the young leaf tissue, and overwintered in the debris. Eggs were laid on or under the moist papery scales that covered the emer- gent cataphylls, and larvae burrowed among these scales until they reached the pinnules of the crozier. From observations on feeding larvae, it was apparent that trichomes were the food of choice, and when most of these had been consumed, the cataphyllar leaf tis- sue was eaten. The damage to the young fronds was extensive, always resulting in death of the organ. On an affected crown, which produces an average of two cataphylls per year, all cataphylls may be affected. The larvae of E. costata overwintered under the 535 papery scales, pupated with the onset of warmer weather, and emerged as adults by early June. DESCRIPTIONS OF IMMATURE STAGES Chirosia betuleti (Ringdahl) Egg.— Pale white; elongate cylindrical, ta- pering to a rounded apex at anterior end, broadly truncate at posterior end (Figs. 13- 15); length 1.44-1.69 mm. Chorionic mem- brane with a markedly reticulate sculptur- ing. First instar larva (Figs. 1-4, 16-18).— Pale yellowish white; slender, subcylindrical, ta- pering anteriorly, rounded posteriorly; 0.90- 1.61 (av 1.18) mm long on hatching, grow- ing to about 2.82 mm before moulting to second instar. Integument without any ap- parent pattern or ornamentation except for ventral surface and anterolateral margins of abdominal segments I-7, that are some- what wrinkled; creeping welts and anterior margins of these segments bearing narrow rows of minute spinules, these spinules more numerous on creeping welts than on dorsum of abdominal segments; a few spinules also present ventrally on anterior margins of thoracic segments 2 and 3, and on terminal abdominal segment where they are more numerous. Terminal segment with eight short tubercles, four dorsal and four ventral to posterior spiracles. Perianal pad relative- ly large, well-defined, each lateral half sub- divided into a smaller anterior lobe and a slightly larger posterior lobe. Anterior spi- racles absent. Posterior spiracles situated on short, cylindrical and subparallel stigmato- phores, each spiracular plate bearing two small, weakly defined but distinctly sepa- rated openings that have faintly sclerotized but incomplete margins, and a somewhat W-shaped or cordate sclerite with a heavily sclerotized, incomplete margin (Fig. 2). Cephalopharyngeal skeleton (Fig. 3) slen- der, seemingly delicate even though rather heavily sclerotized. Mandible relatively long, slender, pointed, with a single tiny accessory 536 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON d proc md .1 mm acc tth t -1 mm p NM Figs. 1-6. Chirosia betuleti, larva. 1, Habitus, first instar. 2, Posterior spiracular plate, first instar. 3, Left lateral view of cephalopharyngeal skeleton, first instar. 4, Anterior (end on) view of head segment showing mouth hooks and their dorsal sclerotized processes, first instar. 5, Left lateral view of cephalopharyngeal skeleton, second instar. 6, Anterior (end on) view of head segment showing mouth hooks, second instar. Abbreviations: acc tth, accessory tooth; ant, antenna; d brg, dorsal bridge; d corn, dorsal cornu; d proc md, dorsal process of mandible; hyphar brg, ventral bridge of hypopharyngeal sclerite; hyphar scl, hypopharyngeal sclerite; lab scl, labial sclerite; md, mandible; mx plp, maxillary palpus; pastm b, parastomal bar; phr, pharynx; tnt phgm, tentorial phragma; tntphr scl, tentoropharyngeal sclerite; v corn, ventral cornu. VOLUME 89, NUMBER 3 tooth on ventral margin, and two (occa- sionally three) prominent teeth on outside margin; also bearing a slender, heavily scler- otized, antler-like process that extends dor- sally along medial margin of antennomax- illary lobe and curves laterally and expands plate-like dorsally over antenna and maxilla (Fig. 4); this process intimately connected with integument and not free from or ex- ternal to it. Hypopharyngeal sclerite rela- tively long, moderately stout. Dental scler- ite absent. Labial sclerite small, slender, but well-developed and heavily sclerotized. Parastomal bar long, moderately stout and rather heavily sclerotized. Tentoropharyn- geal sclerite with tentorial phragma short, dorsal cornu nearly horizontal, distinctly longer and more slender than ventral cornu but becoming weaker and pointed poste- riorly; ventral cornu relatively stout and rather bluntly rounded posteriorly, with a slender clear band posteriorly below dorsal margin; ventral cornu occasionally strongly angled posteroventrally in relation to dorsal cornu (this angle seemingly decreases as the larva increases in size approaching moult to second instar). Dorsal bridge prominent, el- evated slightly above dorsal cornu, only moderately sclerotized. Mature larva (Figs. 7-9, 19-26).—Pale yellowish white; subcylindrical, tapering anteriorly, broadly rounded posteriorly; length 4.08-8.76 (av 7.25) mm; greatest width 0.85-1.92 (av 1.60) mm. Integument mostly smooth except along anterior mar- gins of segments both dorsally and ventral- ly. Creeping welts small, covered with a moderately dense concentration of minute, slightly raised, spiniform integumental pro- cesses that often are difficult to see; these processes extending dorsally along antero- lateral margin of each abdominal segment forming a continuous narrow band that ex- pands dorsally so that each segment has a more or less distinct collar or girdle ante- riorly. Antennomaxillary lobe of cephalic segment with a small, single segmented, 537 somewhat dome-shaped antenna; maxilla distinct, with about five sensory areas; and with about 15-17 oral ridges. Terminal seg- ment with integument slightly granulate, and bearing a series of 16 small but conspicuous tubercles, each with a central pore, arranged as follows: six dorsal tubercles situated in front of posterior spiracles in a nearly trans- verse row that is broken medially (two sub- median, two sublateral, two lateral); six tu- bercles with a similar arrangement behind posterior spiracles; two slightly more ven- trally placed submedian tubercles; and two submedian ventral tubercles dorsal to anus (Fig. 25); area between these two ventral tubercles and anus with a patch of 10-30 dark spinules. Perianal pad moderately large, oval, well-defined, anus situated longitudi- nally. Anterior spiracle (Figs. 7, 23, 24) somewhat flattened and fan-like with about 21 minute papillae arranged in a slightly irregular row. Posterior spiracles (Figs. 9, 25) situated on two slightly elevated cylin- drical stigmataphores; each spiracular plate bearing three oval openings set at about a 20° angle to nearest opening, each opening surrounded by a narrow partially sclerotized ring; with four spiracular hairs that are small but visible with a dissecting microscope, palmate with 4-10 (rarely more) branches that are mostly bifurcate near tips but with some simple branches; peritreme weakly sclerotized, incomplete. Ecdysial scar not evident. Cephalopharyngeal skeleton rather ro- bust, heavily sclerotized but tentoropharyn- geal sclerite mottled by paler and less dense- ly sclerotized areas (Fig. 8). Mandible short, stout, with one or two variably stout acces- sory teeth on ventral margin. Hypopha- ryngeal sclerite relatively large, stout; hy- popharyngeal bridge bowed ventrally so that in lateral view, it projects conspicuously be- low lateral arms of this sclerite. Dorsal and ventral labial sclerites U-shaped, rather weakly sclerotized, often somewhat nebu- lous and irregular in structure. Parastomal 538 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1.0 mm .3 mm 11 i 4 M. Figs. 7-12. 7-9. Chirosia betuleti, third instar larva. 7, Habitus. 8, Left lateral view of cephalopharyngeal skeleton. 9, Posterior spiracular plate. 10-12. E/achiptera costata, third instar larva. 10, Habitus. 11, Left lateral view of cephalopharyngeal skeleton. 12, Posterior spiracular plate. Abbreviation: esph, esophagus. bar apparently absent. Tentoropharyngeal at least posterior ' of dorsal margin of ven- sclerite stout; tentorial phragmata relatively tral cornu more weakly sclerotized and paler long, it and ventral cornu dark, heavily brownish; dorsal cornu broad, weakly scler- sclerotized except entire ventral marginand otized except heavily and narrowly sclero- VOLUME 89, NUMBER 3 539 Figs. 13-18. Scanning electronmicrographs of Chirosia betuleti. 13, Egg. 14, Anterior portion of egg. 15, Posterior portion of egg. 16, Head, first instar larva. 17, Ventral aspect of head of first instar larva showing labial palpi. 18, Dorsal integumental armature of segmental collar of first instar larva. 540 tized along ventral margin, posterior 3 to *, of dorsal cornu with a clear, unsclerotized ‘window.’ Dorsal bridge broad, prominent, but rather weakly sclerotized, mottled with clear, roundish spots. Puparium.—Dark yellowish to reddish brown; length 4.38-6.37 (av 5.27) mm, width 1.26-2.21 (av 1.72) mm. Somewhat barrel-shaped and showing gross external features of mature larva. First two thoracic segments somewhat flattened, metathoracic segment slightly smaller than abdominal segments; anterior segment subtruncate, weakly emarginate medially, and bearing anterior spiracles at each anterolateral cor- ner. Comments.—The specific identity of this anthomyiid fly seems reasonably certain de- spite the paucity of good adult specimens. F. C. Thompson, Systematic Entomology Laboratory, Washington, D.C., compared the male terminalia of our specimens with those of a European specimen of C. betuleti housed in the U.S. National Museum col- lection, and found them to match very closely. This identification was subsequent- ly confirmed by V. Michelsen, Zoological Museum, Copenhagen, Denmark, who also kindly compared our one good specimen with European material. In his treatment of the Palearctic anthomyiids, Hennig (1966) indicated that the life history and larval de- scriptions given under the name Chorto- phila signata Brischke by Meijere (1911) really applied to Chirosia betuleti. An ex- amination of Meiere’s figures (plate 6, fig- ures 16-19) shows that they are similar to our specimens. However, the anterior spi- racle and the mandible of Meijere’s figures show just enough differences from our ma- terial to support the possibility that there really are two species. On the other hand, it is possible that Meiere’s figures are not as accurate as they could be and thus may be of little comparative value in helping re- solve our species identification problem. In any event, we are calling our species Chi- rosia betuleti until additional comparative PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON material of all stages, from both Europe and North America, is available to enable us to revise this identification if necessary. Our specimens represent the first record of this species in North America. Huckett (1965) recorded only two species of Chirosia from North America, viz. C. hirtipes Stein, a Hol- arctic species, and C. idahensis Stein, a western U.S. species. In 1974, he listed four additional species from North America: C. delicata (Huckett), C. pusillans (Huckett), and C. stratifrons (Huckett) all from eastern U.S., and C. hystrix (Brischke) (= C. hys- tricina Rondani) a Holarctic species. From the many larvae of C. betu/eti that pupated, only five females and four males emerged. Of these, only three females and one male were good specimens; the other adults were damaged during emergence. Most of the larvae that pupated were heavi- ly parasitized by the undescribed braconid wasp mentioned earlier, which prevented further development of the flies. The above descriptions were based upon the examination of 42 eggs, 62 first instar larvae, 28 second instar larvae, 122 third instar larvae, and 21 puparia of which nine were associated with the reared adults men- tioned above. The first instar larva closely resembles the second and third instar larvae in general appearance. However, apart from size, some interesting changes in certain morphologi- cal features are dramatically evident be- tween the three larval stages. Externally, the terminal segment of the first instar larva seems to bear only eight tiny posterior lobes, four dorsal and four ventral to the posterior spiracles. In the third instar larva 16 lobes are evident although these are not all the same size. Possibly all 16 lobes are present in the first instar larva but some may be so small they are extremely difficult to discern. Also, the posterior spiracle of the first instar larva has only two incompletely defined spi- racular openings and a well-defined sclerite that has a heavily sclerotized but incom- plete cordate margin. Each spiracle of the VOLUME 89, NUMBER 3 mature larva has three openings and the ec- dysial scar 1s not evident. As in at least some of the first instar larvae of the Muscoidea, the anterior spiracle is absent. In the second instar larva the anterior spiracle is present but small and sometimes difficult to see, especially in recently moulted specimens. This spiracle in the third instar larva 1s well- developed and prominent. Anteriorly, the most notable and visible difference in the first instar larva is the presence of a scler- otized antler-like process extending dorsally from the top of each mandible. This process is darkly pigmented and extends above and over the antenna where it expands plate- like laterally. The origin and function of these structures is not known. They might be remnants of some ancestral structures, and possibly might be used in breaking out of the egg. In the second instar larva only a short remnant of this process remains; how- ever, visible traces of the position it occu- pied in the first instar larva remain in the cuticle of the second instar larva (Figs. 5, 6). All traces of this antler-like process are absent 1n the third instar larva. The man- dibles of the first and second instar larvae are long and slender whereas the mandible of the third instar larva is short and stout. In the first two instars there is a roughly quadrate sclerite, bearing a tiny but distinct aperture, that is directly behind and clearly separated from the mandible. In the mature larva this sclerite is fused with the mandible proper. It is interesting that the cephalo- pharyngeal skeletons of the first two instar larvae are quite similar except for the dif- ference in size. Normally, the cephalopha- ryngeal skeleton of the second instar larva more closely resembles that of the third in- star larva than it does the cephalopharyn- geal skeleton of the first instar larva. The hypopharyngeal sclerite in the first instar larva is stout and apparently fused with the tentopharyngeal sclerite, and the parasto- mal bar is moderately broad and sclero- tized. In the second instar larva there is a distinct line of demarcation between the hy- 541 popharyngeal and tentoropharyngeal scler- ites, yet they remain heavily sclerotized, while the parastomal bar is somewhat more slender and less sclerotized. In the third in- star larva the hypopharyngeal sclerite 1s shortened but still strongly sclerotized, the anteroventral marginal process of the ten- toropharyngeal sclerite 1s much shorter, the tentoropharyngeal phragma is much long- er, the dorsal and ventral cornua are stouter and the dorsal cornu has a distinct window- like area posteriorly. Also, the parastomal bar apparently has disappeared in the ma- ture larva. Other progressive changes can be seen in Figures 2-9. Without having rep- resentatives of associated larvae of each in- star it would be easy to suspect the three morphological forms of the larvae of this species as representing three distinct species. The only other anthomylid first instar lar- va presently known to the authors that pos- sesses similar antler-like processes extend- ing dorsally from the mandibles is that of Macateeia protuberans Malloch. However, there are some marked differences between the processes, as well as between the entire cephalopharyngeal skeletons of these two species. Karl Valley, Pennsylvania Depart- ment of Agriculture, Harrisburg, Pennsy]l- vania, who brought the first instar larva of this species to our attention, plans to de- scribe it in detail. In M. protuberans this antler-like process is shorter, broader, less sclerotized and brownish over most of its length, and the plate-like expansion over the antenna is larger, more weakly sclerotized and less discrete than in Chirosia betuleti. Again, the function of this structure is not known. This process may be more com- monly present among the first instar larvae of phytophagous anthomyiid flies than is currently known. Elachiptera costata (Loew) Mature larva (Figs. 10-12).—Creamy white; subcylindrical, tapering anteriorly, broadly rounded posteriorly. Length 3.18- 542 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 19-24. Scanning electronmicrographs of Chirosia betuleti, third instar larva. 19, Anterior view of head (partly withdrawn) and anterior thoracic segment showing armature. 20, Close-up of thoracic armature and pore. 21, Dorsal integumental armature of segmental collar of abdomen. 22, Close-up of integumental armature of collar and pore of posterior abdominal segment. 23, Anterior spiracle, dorsal view. 24, Anterior spiracle, lateral view. 3.64 (av 3.35) mm; width 0.46-0.65 (av row are largest and those of second ante- 0.58) mm. Integument smooth except for riormost row are next to largest; spinules on small creeping welts that are beset with S— abdominal segments 1-7 extending dorsally 7 irregular rows of spinules; these spinules about 4 to ’2 height of segments. Anten- are minute but those of third posteriormost nomaxillary lobe of cephalic segment with VOLUME 89, NUMBER 3 a small but prominent 2-segmented antenna having a pair of basiconic sensillae at junc- tion of the two segments, an inconspicuous maxilla, and about 15 oral ridges. Terminal segment smooth, without accessory tuber- cles, but ventrally with a small, well-devel- oped, transversely oval perianal pad that is divided into four subequal lobes; and dor- sally bearing posterior spiracles on two short cylindrical stigmatophores (Fig. 10). Each spiracular plate (Fig. 12) with three small, round openings, the lateral openings set at about 30° to 40° from central opening; each opening with a slightly thickened and lightly sclerotized rim; all three openings encircled by a lightly sclerotized, somewhat poorly defined peritreme; with four conspicuous multibranched spiracular hairs; ecdysial scar not apparent. Anterior spiracle small, fan- like, with six minute papillae. Cephalopharyngeal skeleton (Fig. 1 1) well- developed and moderately robust; mandi- ble and hypopharyngeal sclerite heavily sclerotized, tentoropharyngeal sclerite rath- er lightly sclerotized except heavily sclero- tized along anterior rim of dorsal bridge and along ventral margin of dorsal cornu and continuing ventrally along hind margin of tentorial phragma. Mandible rather long, slender, decurved, and with a prominent, ventral, pointed process. Hypopharyngeal sclerite stout, with a broad hypopharyngeal bridge. Dental sclerite small, somewhat crescent to triangular in shape; labial sclerite slender, U- to V-shaped with the two sides not fused or weakly joined anteromedially; parastomal bar long, slender and weakly to moderately sclerotized. Dorsal bridge prominent; dorsal cornu shorter than ven- tral cornu, slender and pointed posteriorly; ventral cornu broad, rounded posteriorly and lightly sclerotized. First and second instar larvae externally similar to mature larva, but first instar larva ranging from 0.37—0.60 mm in length, and the second instar larva ranging from 1.05- 2.8 mm in length. Puparium.—Brownish yellow; length 2.28-3.22 (av 2.57) mm; width 0.60-0.93 543 (av 0.72) mm. External features essentially as in mature larva except anterior two (tho- racic) segments broadly flattened; anterior margin truncate, with a shallow median notch, and bearing anterior spiracles at each anterolateral corner. Comments.—The above descriptions of E. costata were based on the examination of three mature larvae, and 24 puparia (sev- en puparia were associated with reared adults). Also available were 11 first instar larvae, two second instar larvae, and 71 reared adults without associated puparia. The above larval and pupal descriptions ap- parently are the first for this species. DISCUSSION Chirosia is a genus intimately associated with ferns. Of the 14 Palearctic species (Wiezorek, 1973), the nine species for which any hosts are known occur exclusively on ferns. These anthomyiids are either leaf roll- ers, leaf miners, or gall formers. S1x species (C. albifrons Tiensuu, C. albitarsis Zetter- stedt, C. betuleti, C. crassiseta Stein, C. hys- tericina Rondani, and C. parvicornis Zet- terstedt) have had their life cycles described (Wiezorek, 1973; Brown and McGavin, 1982), but the life cycles of the other species are poorly known or undescribed. The life cycle of Chirosia betuleti has been described by Meijere (1911) and Seidel (1957), and our observations broadly concur with their descriptions. Chirosia betuleti is not re- stricted to one species of fern. Meijere (1911) records it from Pteridium aquilinum as well as on Athyrium. Our report of this fly on the genus Matteuccia means that it is oli- gophagous, restricting itself to a few fern genera. Most gall formers are monophagous (Gerson, 1979). Several species of Chirosia (e.g. C. albitarsis, C. albifrons, C. crassiseta) have been reported only from bracken (Cameron, 1930; Meikle, 1937; Brown and McGavin, 1982), although Lawton (1976) suspects that some species of Chirosia found on bracken likely have another major host. Studies of other fern genera are scanty, since most of the efforts on fern-arthropod inter- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 25-28. 25-26. Scanning electronmicrographs of Chirosia betuleti, third instar larva. 25, Posterior end of abdomen. 26, Posterior spiracles. 27-28. Matteuccia struthiopteris. 27, Close-up view of three galls caused by Chirosia betuleti. The pinnae are curled about the rachis of the frond (0.8). 28, Fronds showing two types of galls: (C) gall caused by Chirosia betuleti characteristically placed in upper portion of the frond; (O) gall caused by Olethreutes auricapitana at the tip of the frond, which subsequently aborts (0.12). actions have concentrated on bracken for economic reasons. Monophagy does not ap- pear to be the case for a number of Chirosia species since C. parvicornis is known from ferns of the genera Asplenium, Dryopteris and Pteridium, and Chirosia hystricina is known from ferns of the genera Asplenium, Athyrium, Blechnum, Cystopteris, Dryop- teris, Matteuccia, Osmunda, Polypodium, and Polystichum (Wiezorek, 1973). The ex- clusive feeding on ferns by species of the genus Chirosia is an exception to the ma- jority of fern feeding insects in which there is usually only one species of a genus that VOLUME 89, NUMBER 3 utilizes ferns as a food source (Hendrix, 1980). Indeed, if the conditions Hendrix (1980) poses for evidence of adaptive ra- diation, such as monophagous (or oligopha- gous) feeding by a closely related group of insects are correct, then the genus Chirosia ideally fits the mode. Biological studies of the remaining species are certainly neces- sary to support this speculation. It is also interesting that most of the Chirosia-fern associations are, with one exception (C. hys- tricina on Osmunda), found on modern polypodiaceous s.]. ferns. The cover gall that Chirosia betuleti caus- es on either bracken or the ostrich fern dif- fers with each host. In bracken, the gall is at the tip of the frond (Mejere, 1911), but in Matteuccia the gall is sub-apical. Unlike other types of Chirosia galls found on the unfurling frond tips, such as those caused by C. albifrons on bracken, or C. parvicornis on a variety of genera, the gall does not result in much damage to the tip of the frond as our results clearly indicate. The portions of the frond apical to the gall develop nor- mally, with no significant decrease in height expansion of the leaf. The damage in the immediate vicinity of the gall remains lo- calized and is not enough to disrupt trans- port systems within the affected fronds, which live as long as uninfested ones. Chirosia betuleti was very specific in its feeding site. It seems from a count of eggs in newly emerged fronds that, initially, many more leaves had eggs than eventually de- veloped galls. The larvae had to travel up the cleft of the adaxial side of the curled frond to arrive at the appropriate site, and failure to find the groove of the rachis or early disorientation probably proved fatal to the insects. Although we did not carry out a detailed study concerning all predators on Matteuc- cia over the course of the season, it was readily apparent that very few were present at the beginning of the season. The only others we noted at that time were Olethreu- tes auricapitana, a lepidopteran that forms 545 a gall within the last ten or so pinnae of the unfurling frond (this gall subsequently re- sults in abortion of the leaf tip), and Spar- ganothis reticulatana Clemens, that caused localized disruption of the pinnae. Both of these species are polyphagous feeders and are not restricted to ferns. Olethreutes has been recorded from ferns by Ottosson and Anderson (1983) who found O. lacunana (Freeman) on pinnae of the bracken. Our records of Sparganothis apparently repre- sent the first of this genus on fern. Once the galls are vacated by Chirosia betuleti, they are taken over by the chloro- pid Elachiptera costata. This is the first re- port of EF. costata and the first report of any chloropid known to occur on ferns (Hen- drix, 1980). Previous available evidence suggested that the larvae of Elachiptera probably fed on decaying organic matter, such as old leaf sheaths, or decaying plant tissue and grass following damage by other insects (Sabrosky, 1948). Elachiptera cos- tata was known previously to occur in rot- ting spathes of the aroid Symplocarpus foe- tidus (L.) Nutt. (Brown, 1956; Grimaldi and Jaenike, 1983). The distributions of the fern and aroid host plants do not overlap in Nova Scotia (Roland and Smith, 1969; Aderkas and Bird, 1983). In many other provinces and states the two do occur together in floodplains. Grimaldi and Jaenike (1983) suggest that most Diptera raised in their study of Symplocarpus foetidus probably also breed in decaying matter of other plant species, an observation that has been sub- stantiated in the case of Elachiptera costata. The appearance of eggs and larvae of Chi- rosia betuleti on unfurling fronds of the os- trich fern can be considered as possible food contaminants since the ostrich fern has long been utilized as a food plant in northeastern North America (Aderkas, 1983). We have observed eggs and larvae in both frozen and freshly picked fiddleheads sold in Nova Sco- tia. The frozen fiddleheads originated from New Brunswick, where these galls also have been seen in the field. 546 Table 1. List of Some Insects Collected in Asso- ciation with the Ostrich Fern, Matteuccia struthiop- teris. Chirosia betuleti (Ringdahl) (Diptera: Anthomyii- dae)—gall former on M. struthiopteris. Elachiptera costata (Loew) (Diptera: Chloropidae)— secondary invader of galls. Tricimba melancholica (Becker) (Diptera: Chloropi- dae)— probably secondary invader of galls. Clinodiplosis sp. (Diptera: Cecidomyiidae)— secondary invader of galls. Aphaereta n. sp. near pallipes (Say) (Hymenoptera: Bra- conidae)— Larval parasite of Chirosia betuleti (new species to be described by R. A. Wharton). Elachertus loh Schauff (Hymenoptera: Eulophidae)— Larval parasite of Chirosia betuleti. Dimmockia n. sp. (Hymenoptera: Eulophidae) —par- asite, emerged from pupa of Chirosia betuleti (new species to be described by M. E. Schauff). Olethreutes auricapitana (Walsingham) (Lepidoptera: Tortricidae)—gall former on M. struthiopteris. Sparganothis reticulatana Clemens (Lepidoptera: Tor- tricidae)— feeding on M. struthiopteris. Malacosoma disstria Hubner (Lepidoptera: Lasiocam- pidae)—pupating on fertile fronds of M. struthiop- teris. Lygus rubroclarus Knight (Heteroptera: Miridae)— commonly resting on M. struthiopteris. Table | presents a list of the primary in- sects that were collected in association with the ostrich fern during our study. A number of various kinds of insects are casual visitors to this fern, but are not relevant to this study. ACKNOWLEDGMENTS We are grateful to F. C. Thompson, C. W. Sabrosky, R. J. Gagne, P. M. Marsh, M. E. Schauff, D. C. Ferguson, and T. J. Henry, Systematic Entomology Laboratory, BBII, Agricultural Research Service, USDA, Washington, D.C.; V. Michelsen, Zoologi- cal Museum, Copenhagen, Denmark; R. A. Wharton, Department of Entomology, Tex- as A&M University, College Station, Texas; and P. T. Dang, Biosystematics Research Center, Agriculture Canada, Ottawa, On- tario, Canada, for identification or confir- mation of various insects collected during PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON this project. We are also grateful to H. J. Teskey, Biosystematics Research Center, Agriculture Canada, Ottawa, Ontario, Can- ada for reading the descriptions of the im- mature stages; and Karl Valley, Bureau of Plant Industry, Pennsylvania Department of Agriculture, Harrisburg, Pennsylvania, for reading the manuscript, for helpful and stimulating discussions during the prepa- ration of the manuscript, and for providing the first instar larvae of Macateeia protu- berans. We are also grateful to Mary Lou Cooley, staff artist, Systematic Entomology Laboratory, for her careful preparation of the fine larval illustrations, and P. Malikul, technician, Systematic Entomology Labo- ratory, for taking the SEM photomicro- graphs that accompany this paper. We gratefully acknowledge the field help of E. Butterworth, New Brunswick Research and Productivity Council, Fredericton, New Brunswick; J. Moore and P. Borwein, Dal- housie University, Halifax, Nova Scotia; as well as S. Johnson, N. Lewis and G. Wil- liams, Atlantic Research Laboratory, Hal- ifax, Nova Scotia. Help with statistical anal- ysis by P. Green, Dalhousie University, Halifax, Nova Scotia, is acknowledged. Gratefully acknowledged is the support for a portion of this project that came through a Research Associateship awarded to the senior author at the laboratory of J. Mc- Lachlan, Atlantic Research Laboratory, Halifax, Nova Scotia. This paper represents NRCC publication no. 26322. LITERATURE CITED Aderkas, P., von. 1984. Economic history of Mat- teuccia struthiopteris (L.) Todaro, the edible fid- dlehead. Econ. Bot. 38: 14-23. Aderkas, P., von and C. J. Bird. 1983. The habitat of the ostrich fern (Matteuccia struthiopteris) in Nova Scotia and Prince Edward Island. Proc. N.S. Inst. Sci. 33: 131-135. Aderkas, P., von and P. E. G. Green. 1986. Leaf development of the ostrich fern, Matteuccia stru- thiopteris (L.) Todaro. Bot. J. Linn. Soc. 93: 307- 321" Brown, V. K.and G. C. McGavin. 1982. The biology VOLUME 89, NUMBER 3 ofsome mine and gall forming Diptera on bracken, Pteridium aquilinum (L.) Kuhn. J. Nat. Hist. 16: 511-518. Brown, W. L. 1956. Drosophilid and chloropid flies bred from skunk cabbage. Psyche 63: 13. Cameron, A. E. 1930. Two species of anthomyiid diptera attacking bracken and their hymenopter- ous parasites. Scott. Nat. 182: 137-141. Gerson, U. 1979. The associations between pterido- phytes and arthropods. Fern Gaz. 12: 29-45. Grimaldi, D. and J. Jaenike. 1983. The Diptera breeding on skunk cabbage, Symplocarpus foetidus (Araceae). J. N.Y. Entomol. Soc. 91: 83-89. Hendrix, S. D. 1980. An evolutionary and ecological perspective of the insect fauna of ferns. Am. Nat. 115: 171-196. Hennig, W. 1966. 63a. Anthomyiidae. Lfg. 268, Bd. VII/I, pp. 49-96. Jn E. Lindner, ed., Die Fliegen der palaearktischen Region. Schweizerbart, Stutt- gart. Huckett, H.C. 1965. Subfamily Anthomyiinae, pp. 843-869. In A. Stone, C. W. Sabrosky, W. W. Wirth, R. H. Foote, and J. R. Coulson, eds., A Catalog of the Diptera of America North of Mex- ico. U.S. Dep. Agric., Agric. Res. Serv., Agric. Handb. 276. Huckett, H. C. 1974. The Anthomyidae and Mus- cidae of the Great Smoky Mountains and Mt. Mitchell, North Carolina (Diptera). J. N.Y. Ento- mol. Soc. 82: 150-162. 547 Lawton, J. H. 1976. The structure of the arthropod community on bracken (Prteridium aquilinum (L.) Kuhn). Bot. J. Linn. Soc. 73: 187-216. Meikle, A. A. 1937. The insects associated with bracken. Agric. Progress 14: 58-60. Meijere, J. C. H., de. 1911. Uber in farnen parasi- tierende Hymenopteren- und Dipteren-Larven. Tijdschr. Ent. 54: 80-127, Pls. 5-7. Ottosson, J. G. and J. M. Anderson. 1983. Number, seasonality and feeding habits of insects attacking ferns in Britain: An ecological consideration. J. Anim. Ecol. 52: 385-406. Roland, A. E. and E. C. Smith. 1969. The flora of Nova Scotia. 2nd ed. Nova Scotia Mus., Halifax, 746 pp. Sabrosky, C. W. 1948. A synopsis of the Nearctic species of Elachiptera and related genera (Diptera, Chloropidae). J. Wash. Acad. Sci. 88: 365-382. Schauff, M.E. 1985. Taxonomic study of the Nearctic species of Elachertus Spinola (Hymenoptera: Eu- lophidae). Proc. Entomol. Soc. Wash. 87: 843- 858. Seidel, J. 1957. Pflanzengallen und Blattminen aus dem Gebiete des Schneebergers. Prirod. Sb. Os- travsk. Kraje 18: 248-270. Wiezorek, H. 1973. Zur Kenntnis der Adlerfarnin- sekten. Ein Beitrag zum Problem der biologischen Bekaémpfung von Preridium aquilinum (L.) Kuhn in Mitteleuropa. Z. Angew. Entomol. 72: 337-358. PROC. ENTOMOL. SOC. WASH. 89(3), 1987, pp. 548-551 DISCOVERY OF THE T. W. HARRIS COLLECTION AT HARVARD UNIVERSITY AND DESIGNATION OF A LECTOTYPE FOR PODOSESIA SYRINGAE HARRIS (LEPIDOPTERA: SESITDAE)'! FOSTER FORBES PURRINGTON AND DAvID G. NIELSEN Department of Entomology, Ohio Agricultural Research and Development Center, The Ohio State University, Wooster, Ohio 44691. Abstract. —T. W. Harris’ insect collection, the oldest synoptic collection in North Amer- ica, has been located in the Museum of Comparative Zoology at Harvard University. It contains the types of more than 200 species of Coleoptera and Lepidoptera described by Harris, Thomas Say, and others. Among the clearwing moths (Sesiidae) is the syntypic series of Podosesia syringae (Harris) for which we designate here a lectotype. Thaddeus William Harris’ insect collec- tion was moved across the Charles River in 1941 from Boston’s New England Museum of Natural History to Harvard University’s Museum of Comparative Zoology (MCZ) in Cambridge. The arrival at the MCZ of this important synoptic collection was noted by Philip J. Darlington, Jr. (1941), Fall Curator of Coleoptera at Harvard at that time. It is the oldest synoptic collection of insects in North America, containing the types of more than 200 species described by Harris, Thomas Say, and others. The New England Museum of Natural History changed its name in 1949 to the Museum of Science. Although it transferred the Harris collection to Harvard, the Mu- seum retained 20 volumes of Harris’ manu- scripts dealing with the insect collection, in- sects of Massachusetts, indexes, abstracts, lectures, and other relevant writings, all of which are presently housed at the Museum of Science library (C. Fruci, pers. comm.). ' Support for this research was provided by State and Federal funds appropriated to the Ohio Agricultural Research and Development Center, The Ohio State University. Journal Article No. 1-87. In the years following the transfer, the Harris collection slipped from the sight of at least some specialists who sought it out, and eventually it came to be regarded as lost or destroyed (Duckworth and Eichlin, 1973, 1977, 1978). In February 1986, one of us (F.F.P.) visited Harvard University to ex- amine the clearwing moth holdings and was promptly shown the Harris collection by MCZ Collections Manager Scott R. Shaw. Our interest in the Harris collection cen- ters on the type material of 11 clearwing moth species that Harris described from collections he made in eastern United States, primarily New England (Table 1). These specimens bear red MCZ type labels and other labels, some probably in Harris’ hand. Some series of syntypes include pupal exu- viae and even cocoons; many specimens are in poor condition. Ten of these species were originally described in Harris’ Descriptive Catalogue (1839): Aegeria polistiformis was described elsewhere (Harris, 1854). Because most of these clearwings are economically important and have been studied exten- sively, application of most names has not been in doubt. Nevertheless, since we de- scribed Podosesia aureocincta Purrington and Nielsen as a cryptic sibling species of VOLUME 89, NUMBER 3 Table 1. Harris MCZ Original Harris Name Number Aegeria caudata 87 26354 Trochilium tibiale 387 26355 Trochilium marginatum 388 26356 Aegeria tricincta 322 26357 Aegeria polistiformis 791 26358 Trochilium denudatum Silul 26359 Aegeria syringae 464 26360 Aegeria fulvipes 17 26361 Aegeria scitula 333 26362 Aegeria pyri 702 26363 Aegeria cucurbitae 249 33258 Podosesia syringae (Harris) (Purrington and Nielsen, 1977), the serendipitous discovery of the Harris types has given us an oppor- tunity to verify that aureocincta does not refer to syringae: that it is not a junior syn- onym of syringae. Until now, without ref- erence to the types, it has seemed possible that Harris had specimens of aureocincta before him, instead of syringae. This is not an unwarranted concern, be- cause Engelhardt’s monograph on the Se- sudae (1946) illustrates for P. syringae (Harris) a Podosesia saccus tip of the male genital armature clearly assignable to au- reocincta, rather than to syringae (Purring- ton and Nielsen, 1979). Difficulties of rec- ognition of this sort can obviously attend separation of sibling species and no doubt are enhanced by the operation of selection pressures that maintain mimicry in the Se- siudae. The kinds of phenological, phero- monal, and behavioral nuances that main- tain genetic distinctness between pairs of clearwing cryptic siblings are not reflected by characters that are preserved in museum specimens (Heppner and Duckworth, 1981). Neither our separation of P. aureocincta from syringae nor the subsequent separa- tion of Paranthrene pellucida Greenfield and Karandinos from Paranthrene simulans (Grote) could be made using standard char- acters of the male genital armature (Green- field and Karandinos, 1979), although with Number 549 List of Sesiidae types described by T. W. Harris held at the MCZ in the Harris collection. Present Name Alcathoe caudata (Harris) Sesia tibialis (Harris) Pennisetia marginata (Harris) Paranthrene tabaniformis (Rottemburg) Vitacea polistiformis (Harris) Sesia asilipennis (Boisduval) Podosesia syringae (Harris) Synanthedon fulvipes (Harris) Synanthedon scitula (Harris) Synanthedon pyri (Harris) Melittia cucurbitae (Harris) intensive study such features were eventu- ally located in the case of Podosesia aureo- cincta (Purrington and Nielsen, 1979). Nomenclatural stability in Podosesia has also been jeopardized by Naumann (1971) who, although he did not claim the types were missing, only that he did not examine them, nevertheless established Brooklyn, New York as type locality for syringae. He noted a female type was in the ““Boston So- ciety of Natural History” (sic), possibly to agree with Engelhardt (1946), who made the same inaccurate contentions. Apparently neither author was aware of the collection’s transfer to Harvard in 1941. It is unclear to us why Naumann felt com- pelled to establish a type locality for syr- ingae, unless he was in fact convinced the types were missing. He published original drawings of probable syringae genitalia tak- en from specimens obtained from two Eu- ropean museums, including material col- lected in Brooklyn, New York. Neither Duckworth and Eichlin (1977) nor Hodges and Miller (in litt.) follow Naumann in his Brooklyn type locality designation for syr- ingae, they indicate Massachusetts is the likely collection site (Harris gave no locality data for his syntypic Podosesia series). To verify the Harris Podosesia types as syringae auctorum, we borrowed the syn- typic series of two males and one female from the MCZ. Genital slides of both males 550 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Genitalia of Podosesia males (aedeagi removed). Top, syringae lectotype. Bottom, aureocincta, Woos- ter, Ohio, 12 September 1976, col. F. F. Purrington. VOLUME 89, NUMBER 3 were made, and in accordance with stan- dard practice we have selected, labeled, and designated as follows one of these males as lectotype. The pinned lectotype male imago has 4 labels: 1) 464 6, 2) MCZ No. 26360, 3) Aegeria syringae 4 type, genitalia slide #A, MCZ No. 26360, E. H. Metzler, 1 May 1986, 4) lectotype, F. F. Purrington, 14 Oc- tober 1986. The lectotype genitalia slide is a standard 1” by 3” glass microscope slide bearing 2 labels: 1) Aegeria syringae é type, 464 6, MCZ type No. 26360, slide #A, E. H. Metzler, 1 May 1986, 2) lectotype, F. F. Purrington, 14 October 1986. We have designated the second male syn- type as paralectotype. The pinned paralec- totype male imago bears the same label data as the pinned lectotype except the notation that its genitalia are on slide #B. That gen- italia slide bears 2 labels: 1) Aegeria syringae 6 type, 464 6, MCZ type No. 26360, slide #B, E. H. Metzler, 1 May 1986, 2) paralec- totype, F. F. Purrington, 14 October 1986. The single female is also designated as paralectotype, with label data as follows: 464 2, MCZ No. 26360, paralectotype, F. F. Pur- rington, 14 October 1986. All three types and the genitalia slides prepared by us are in the MCZ, Harvard University. Both lectotype and paralectotype Podo- sesia males clearly show the genital char- acter state that we have determined is uniquely associated with syringae (Harris). The tip of the saccus of syringae is truncate or somewhat knobbed (Fig. 1-top), whereas in its cryptic sibling aureocincta the tip of the saccus (Fig. 1-bottom) is strongly bifur- cated, the corners elaborated into prongs. ACKNOWLEDGMENTS We thank Scott R. Shaw (MCZ, Harvard) for his hospitality, for providing the Harris Podosesia types, and for fruitful discussion. The genital preparations were made with consummate skill by Eric H. Metzler (De- partment of Natural Resources, Ohio), whom we thank for his craftsmanship, ad- vice and comments. W. Donald Duckworth 551 (Bishop Museum, Honolulu) and Charles A. Triplehorn (The Ohio State University, Co- lumbus) offered encouragement and re- viewed the manuscript. We appreciate the opportunity afforded us by Scott E. Miller (Bishop Museum, Honolulu) to preview a draft catalogue of MCZ (Harvard) micro- lepidoptera type material. LITERATURE CITED Darlington, P. J., Jr. (1941). The insect collection of Thaddeus W. Harris (1795-1856). Entomol. News 22S: Duckworth, W. D.and T. D. Eichlin. 1973. The type- material of North American clearwing moths (Lepidoptera: Sesiidae). Smithson. Contrib. Zool. 148, 34 pp. 1977. A classification of the Sesiidae of America north of Mexico (Lepidoptera: Sesioi- dea). Occasional Papers in Entomol. No. 26, Calif. Dep. Food Agric., Sacramento, 54 pp. 1978. The clearwing moths of California (Lepidoptera: Sesiidae). No. 27, Calif. Dep. Food Agric., Sacramento, 82 pp., 7 pl. Engelhardt, G. P. 1946. The North American clear- wing moths of the family Aegeriidae. U.S. Nat. Mus. Bull. 190, 222 pp. Greenfield, M. D. and M. G. Karandinos. 1979. A new species of Paranthrene (Lepidoptera: Sesi- idae). Proc. Entomol. Soc. Wash. 81: 499-504. Harris, T. W. 1839. Descriptive catalogue of the North American insects belonging to the Linnaean genus Sphinx in the cabinet of Thaddeus William Harris, M.D., librarian of Harvard University. Am. J. Sci. Arts. 36: 282-320. 1854. Report on some of the diseases and insects affecting trees and vines. Proc. Am. Pomo- logical Soc. Cong., pp. 197-217. Heppner, J. B. and W. D. Duckworth. 1981. Clas- sification of the superfamily Sesioidea (Lepidop- tera: Ditrysia). Smithson. Contrib. Zool. 314, 144 pp. Naumann, C. M. 1971. Untersuchungen zur Syste- matik und Phylogenese der holarktischen Sesiiden (Insecta: Lepidoptera). Bonner Zoologische Mon- ographien (Bonn), 1: 1-190. [English Translation: 1977. Studies on the systematics and phylogeny of holarctic Sesiidae (Insecta: Lepidoptera). 208 pp., Washington: Smithson. Institution.] Purrington, F. F. and D. G. Nielsen. 1977. Biology of Podosesia (Lepidoptera: Sesiudae) with descrip- tion of a new species from North America. Ann. Entomol. Soc. Am. 70: 906-910. 1979. Genitalic difference between males of Podosesia aureocincta and P. syringae (Lepidop- tera: Sesiidae). Ann. Entomol. Soc. Am. 72: 552- 55): PROC. ENTOMOL. SOC. WASH. 89(3), 1987, pp. 552-558 LIFE HISTORY OF NEOTEPARITIS FINALIS (LOEW) ON NATIVE ASTERACEAE IN SOUTHERN CALIFORNIA (DIPTERA: TEPHRITIDAE) R. D. GoEDEN, T. D. CADATAL, AND G. A. CAVENDER Department of Entomology, University of California, Riverside, California 92521. Abstract. — Neotephritis inornata (Coquillett) is synonymized with N. finalis largely based on examination of reared specimens. New host-plant records for N. finalis are presented, and selected published records confirmed or questioned. An affinity for hosts in the subtribe Verbesininae of the tribe Heliantheae (Asteraceae) is noted. The biology of this multivoltine tephritid in flower heads of Encelia farinosa Gray is described, one of a succession of host-plant species in which it reproduces throughout the year in various parts of southern California. The primary, hymenopterous, solitary, larval or larval-pupal parasites, Eurytoma vernonia Bugbee (Eurytomidae) and Preromalus sp. (Pteromalidae) are reported from N. finalis. Neotephritis finalis (Loew) commonly in- fests the flower heads of many Asteraceae in southern California. Herein we record our collective findings on this heretofore lit- tle-known fly studied irregularly by us since 1980. TAXONOMY Foote (1960) reviewed the three species of Neotephritis known from North America. Only N. finalis and Neotephritis inornata (Coquillett) are known from California (Foote and Blanc, 1963), the latter species only from two solitary, swept or trapped male specimens. Swept and reared speci- mens of Neotephritis collected by RDG in California appeared to vary in the key char- acters that have been used to separate in- ornata and finalis. This prompted a detailed examination of specimens of Neotephritis reared from extensive samplings of mature flower heads of native Asteraceae from cen- tral and southern California to determine if the two species are distinct. The key provided by Foote and Blanc (1963) separated finalis from inornata by two characters: specimens with two light spots in cell r,,; immediately anterior of vein M and the ovipositor sheath darkened apically were considered /finalis; specimens with three small hyaline spots in r,,; and a basally darkened ovipositor sheath were called inornata. Steyskal (1972) provided a key to 11 species of Neotephritis in which he separated finalis from inornata by two additional characters: in finalis, cell dm (discal cell) with a distinct hyaline spot close to its postero-apical corner and the scutel- lum largely rufous medially; in ‘nornata, cell dm without a distinct hyaline spot close to its postero-apical corner and the scutellum rufous along its hind margin. Two hundred and forty-five mounted specimens of Neotephritis reared from five genera and 12 species of Asteraceae from 24 different locations during 1980-86 by RDG were examined. All of these speci- mens would key to either finalis or inornata in Steyskal (1972) and Foote (1960). These specimens, which were largely randomly se- VOLUME 89, NUMBER 3 lected and mounted, comprised 126 (51.4%) 6 and 119 (48.6%) 2 and represented sub- samples from more than twice this number of reared flies. Among the males examined, there were 12 (9.5%) from eight locations that lacked an apical spot in the postero- apical corner of cell dm on both wings, 7 (5.6%) that lacked a spot on one wing, but with a faint or small spot, or in two in- stances, a prominent spot on the other wing, and eighty (63.5%) 6 with a large spot on both wings, and 27 (21.4%) with a large spot on one wing and a tiny or faint spot on the other. Similarly, among females, 9 (7.6%) from seven locations lacked the spot on both wings, and 4 (3.4%) lacked a spot on one wing, but possessed a faint, small, or, in one case, a prominent spot on the other. Eighty- seven (73.1%) 6 had prominent spots in both wings, and 19 (16.0%) showed greatly re- duced or faint spots in one or both wings. Clearly, this wing character used by Steyskal (1972) varies too continuously among both sexes of specimens from southern Califor- nia to segregate them into different species. None of the above 245 reared specimens examined had a partially or wholly rufous scutellum. Instead, both sexes possessed a yellowish-white central longitudinal stripe on the scutellum that varied in length from a narrow posterior band to a prominent api- cal spot to nearly bisecting this segment, and was bounded by dark lateral stripes and an overlay of dark to light grey microtomen- tum. Examination of the male syntype by A. L. Norrbom (in /itt. 1987) as well as one of the two female syntypes of inornata from the type locality, 1.e. Las Vegas Hot Springs, New Mexico, on loan to RDG from the Na- tional Museum of Natural History (USNM) collection, confirmed that the scutellum of both was rufous brown with the lateral and posterior third yellowish. Rufous or not, scutellum markings varied too much among reared specimens in southern California to use this character to segregate them. Similarly, ovipositor sheath color was ex- amined in 102 of the females reared by RDG 33 that did not have this structure discolored by dried feces or obscured by overlapped wings. In all but 2 (2.0%) 2, a dark apical band was present, which varied from nar- row to broad. More importantly, 75 (73.5%) showed at least some basal darkening of the ovipositor sheath. Only a single specimen had a unicolored, undarkened, rufous-yel- low ovipositor sheath. Obviously, this char- acter also showed too much continuous variation in reared specimens to support its use in distinguishing /nornata. This leaves the key character mentioned by Coquillett (1902), and used by Foote (1960), Foote and Blanc (1963), and Steys- kal (1972) to separate inornata from finalis, i.e. three versus two spots in cell r,,; 1m- mediately anterior of vein M. Eight (0.5%) of 201 6 reared by RDG during 1980-86 had the three spots diagnostic of inornata on one wing; 2 (0.1%) had the three spots on both wings (in one case, four spots on one wing). Similarly, 7 (3.7%) of 190 2 had the three spots on one wing; | (0.5%) 2, the three spots on both wings. Possession of these spots was the one basis for retaining, mounting, and separately storing specimens reared during this period. These specimens varied in the ovipositor sheath and scutel- lum characters described above similar to specimens with only two spots in r,,; in both wings. No correlation in any of three characters was detected. The specimens with three spots were reared from four genera and seven species of Asteraceae, and from eight widely separated locations in southern California along with specimens with two spots in all cases. The central or proximal spots usually were the smaller of the three, but all three spots were equally large in one wing of three specimens, 2 4 and | 9, and on both wings of | 2. Had this latter female been trapped or swept, like all reported specimens of /nornata, not reared among a large series of specimens with only two spots in r,,5, it, too, would probably have been identified as inornata. Comparison of this individual with three specimens of /nornata 554 on loan from the USNM, each identified by different tephritid taxonomists, showed no obvious distinctions unaddressed above. Short series of specimens from the same locations in Arizona, Oregon, and Utah se- lected from the USNM collection by A. L. Norrbom and loaned to RDG also showed variation in the number of spots in cell r,, ; and other characters addressed above, which demonstrated that this variation was not confined to southern California. Therefore, the senior author (RDG), independent of his coauthors, recommends that N. inornata (Coquillett, 1902) be synonymized with N. finalis (Loew, 1862), because the name 1s based on a character, which is rare but con- tinuously variable among specimens that otherwise appear conspecific. DISTRIBUTION AND Host PLANTS Foote (1960, p. 149) termed N. finalis ‘one of the most commonly encountered tephri- tids in North America.” Foote and Blanc (1963, p. 35) described its geographic range as “Southern Canada; northern Mexico; the Continental United States except Alaska and New England.” N. finalis is widely distrib- uted throughout California (Foote and Blanc, 1963, Map 35), including coastal central California and the Mojave Desert, areas unmarked by collection points on its state distribution map in Foote and Blanc (1963) (Goeden, unpublished data). New host-plant genus or species records determined by comparison with Wasbauer (1972) are reported for N. finalis from southern California. These represent rear- ings from samples of mature flower heads collected and processed by RDG, as de- scribed elsewhere (Goeden, 1985). Host- plant records that are not reported in Was- bauer (1972) are labeled with a double or single asterisk for genera and species, re- spectively. Among multiple samples of a particular plant species, only the sample from which the most individuals was re- covered is reported. The plant nomencla- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ture used follows Munz (1974). Rearing rec- ords are listed alphabetically. * Balsamorhiza deltoidea Nuttall, 16 4 and 9 2, Burnt Peak, Angeles Nat. Forest, NW Los Angeles Co., 8 vi 1983; *Encelia fru- tescens Gray, 8 6 and 6 2, Zzyxx offramp on Interstate Hwy. I-15, SW of Baker, San Ber- nardino Co., 29 iv 1981; *Encelia virginen- sis A. Nelson, 8 6 and 2 2, Bautista Canyon, Riverside Co., 29 iv 1980; **Geraea canes- cens Torrey and Gray, 41 éand 32 2, Hidden Springs (NW of Indio), Riverside Co., 14 iv 1981; *Helianthus gracilentus Gray, 2 6 and 2 2, Moreno, Riverside Co., 19 v 1981: *H. niveus (Bentham) Brandegee ssp. fe- phrodes (Gray) Heiser, 34 6 and 40 2, sand hills near Glamis, Imperial Co., 28 1 1982; **Viguiera deltoidea Gray var. parishii (Greene) Vasey and Rose, | 2, Chino Can- yon, Riverside Co., 3 iv 1980; *Wyethia ovata Torrey and Gray, 2 6 and 3 9, Filaree Flat, Laguna Mountain, Cleveland Nat. Forest, San Diego Co., 14 vi 1984. In addition, the following rearing records confirmed host records for N. finalis in Was- bauer (1972): Encelia californica Nuttall, 17 6and 13 9, Seacliff, Ventura Co., 1 iv 1981; Encelia farinosa Gray, 18 é6and 12 2, above Parker Dam, SE San Bernardino Co., 30 ii 1984: Helianthus annuus L., 1 6 and 2 9, Canebrake, Kern Co., 8 ix 1986; Wyethia mollis Gray, 14 6 and 13 2, Bald Eagle Peak, Piute Mountain, Sequoia Nat. Forest, NE Kern 'Co., 2/vi1 1982: Additional hosts of N. finalis reportedly include Aster spinosus Bentham, Balsamor- hiza sagittata (Pursh) Nuttall, Dahlia sp., Eriophyllum lanatum (Pursh) Forbes, Gail- lardia aristata Pursh, Heliathella uniflora (Nuttall) Torrey and Gray, Helianthus nut- tallii (Torrey and Gray, Verbesina (as Actinomeris) sp., Wyethia helenioides (deCandolle) Nuttall, and Xanthium stru- marium L. (as enceliodes) (Wasbauer, 1972). Balsamorhiza, Encelia, Helianthus, He- lianthella, Geraea, Verbesina, Viguiera, and Wyethia all are members of the subtribe VOLUME 89, NUMBER 3 Table 1. native Asteraceae in southern California. 555 Examples of synphagy among Neotephritis finalis and Trupanea sp. in flower head samples from Number (%) of Tephritidae Reared Host Plant Replicate N. finalis Trupanea spp. Encelia californica 1 30 (19.0) T. wheeleri 128 (81.0) 2 12 (857) T. wheeleri 52 (81.3) 3} 20 (9.1) T. wheeleri 199 (90.5) T. nigricornis 1(OES) E. farinosa 1 1 (3.2) T. nigricornis 30 (96.8) D2 30 (46.9) T. nigricornis 34 (53.1) E. frutescens 1 14 (45.2) T. nigricornis 17 (54.8) D, 14 (87.5) T. nigricornis Di(12=5) E. virginensis ] 10 (66.7) T. wheeleri 4 (26.6) T. nigricornis 1 (6.7) 1e@eS) T. nigricornis 64 (98.5) 3 1 (0.6) T. nigricornis 166 (99.4) Geraea canescens 73 (92.4) T. jonesi 4 (5.1) T. bisetosa 225) 22 7 (46.7) T. bisetosa 7 (46.7) T. jonesi 1 (6.6) 3 23 (95.8) T. jonesi 1 (4.2) 4 Ill (7323) T. bisetosa 4 (26.7) Verbesininae of the tribe Heliantheae, and thus form a natural assemblage of host plants (Munz and Keck, 1959). Unconfirmed host records from the genera Dahlia and Xan- thium, which belong to other subtribes of Heliantheae, are somewhat less suspect than records from Aster in the tribe Astereae or Eriophyllum and Gaillardia in the tribe He- lenieae (Munz and Keck, 1959; Bailey, 1975). However, Hilgendorf and Goeden (1983) never reared N. finalis from flower heads or otherwise collected this fly from X. spinosum L. or X. strumarium during extensive faunistic surveys of these weeds in southern California. Synphagy.—Cavender (1981) reported that N. finalis was the third most common species of Tephritidae in the flower heads of cultivated and wild sunflower, H. annuus, in southern California after Paracantha cul- taris (Coquillett) and Trupanea_ bisetosa (Coquillett). During 1980-86, RDG consis- tently reared N. finalis together with one or two species of 7rupanea from samples of flower heads of four species of Encelia and Geraea canescens (Table 1). These samples were collected from different locations, usu- ally in different years. Trupanea bisetosa, T. Jonesi Curran, T. nigricornis (Coquillett), and 7. wheeleri Curran variously were as- sociated with N. finalis in these flower heads (Goeden, 1985). The same two or three species usually emerged from flower heads of each plant species; however, none of the four 7rupanea species was recovered from all samples (Table 1), nor was N. finalis al- ways recovered (Goeden, unpublished data). The proportions of these synphagous te- phritid species varied among samples and hosts, with N. finalis acting either as the dominant, middle level, or subordinate species, sometimes in different samples from the same species of host plant (Table 1). This interpretation assumes that the num- bers of flies of each species reared from each sample reflect the results of interspecific competition for their common flower-head resources. Synphagy is a common form of 556 resource sharing among flower head-infest- ing Tephritidae in southern California (Goeden, unpublished data). Indeed, syn- phagy apparently is a characteristic life his- tory strategy for certain genera, e.g. Nearctic Urophora (Goeden, 1987). BIOLOGY The biology of N. finalis was studied in the field and insectary by TDC largely in and near the Botanic Garden of the Uni- versity of California, Riverside, during 1982 and 1983 on one of its most common and widespread native hosts in southern Califor- nia, Encelia farinosa (Table 1). As the flow- er heads of this host, as well as cultivated and wild sunflowers, commonly contained Trupanea nigricornis and Melanagromyza viridis (Frost) (Diptera: Agromyzidae), the studies of Cavender (1981), published in part as Cavender and Goeden (1982, 1983), helped TDC distinguish N. finalis during the present study. Immature flower heads pro- tected from oviposition by small plastic bags in the field for up to 4 days were uncovered and offered to caged females in the insectary of the Division of Biological Control, during studies of fecundity, longevity, and behav- ior during courtship, mating and grooming. Insectary conditions were 27 + 2°C, 40- 60% RH, and 16-h photoperiod. Cages used were described by Gilstrap and Goeden (1974). Egg.—The egg is smooth, translucent white, elongate ellipsoidal, blunt posterior- ly, but tapered anteriorly. Fifty eggs mea- sured 1.15 + 0.10 (* + SEM) (range, 1.09- 1.24) mm long by 0.27 + 0.02 (range, 0.23- 0.31) mm wide. The egg was laid singly be- tween the florets with its posterior end touching the pappus above its juncture with the achene. In the insectary, 116 (98.3%) of 118 eggs hatched in 2 days; the remaining two eggs, in 3 days. Larva.— Before hatching, the embryo took about 10 min. to reverse its position inside the egg, then fed for about an hour on a gel- like substance inside the posterior end be- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON fore exiting through a posterior slit-like break in the chorion. Upon hatching, the larva immediately entered the immature achene and began to feed. After eating through the achene, the larva moved laterally into an adjacent achene and continued to tunnel in this fashion through a succession of disc achenes throughout its growth and devel- opment. Dissection of field-collected flower heads of E. farinosa indicated that first in- stars perforated 3 to 6 achenes, second in- stars tunneled through an additional 3 to 9 achenes, and third instars attacked another 7 to 11 achenes, of which the contents of 3 to 5 were completely consumed. When lar- vae fed on sunflower achenes, the first stadi- um lasted about 2 days (range, 2-4; N = 118); the second stadium, about 3 days (range, 2-6; N = 114); and the third stadi- um, about 7 days (range, 4-14; N = 82). Pupa.—Pupariation occurred at the ter- minus of the larval mine in a cell made of frass, slightly larger than the puparium. The puparium is oblong, slightly tapering ante- riorly, translucent light brown to medium brown, with very distinctive, dark brown remnants of the anterior and posterior sp1- racles. Thirty 6 puparia averaged 3.5 + 0.2 (range, 3.1-3.9) mm in length by 1.5 + 0.1 (range, 1.2-1.6) mm in greatest width. Twenty 2? puparia averaged 3.7 + 0.1 (range, 3.4-3.9) mm long by 1.6 + 0.1 (range, 1.4—- 1.6) mm wide. The puparium rested atop the receptacle, usually with its anterior end from which the adult emerges facing out- ward, separated from the outside by a thin layer of frass. Fifty-three adults emerged from their puparia an average of 9 (range, 6-14) days after pupariation under insectary conditions. Adult. — Mating was not observed in na- ture. In insectary cagings, mating behavior was initiated as a male approached a female while flexing his wings alternately. One wing was thrust forward perpendicular to the long axis of the body, while the other wing was held outward at about 15°. The wings then alternated in these positions at about | s VOLUME 89, NUMBER 3 intervals, while the male moved forward at about 2 mm per s. Meanwhile, the female mainly stood still, but also alternately flexed her wings in response. When the pair faced each other about 7 mm apart, the male brought both wings backward at 15°, as the female reversed her position and faced away. The male then flexed both wings forward together and quickly mounted her poste- riorly. Coupling followed in about 4 s, after the female lifted her ovipositor, into which the male inserted his aedeagus. Coupling lasted until the female disengaged herself from the generally passive male. Nine pairs were observed in copula an average of 6 (range, 2-11) times, for an average of nearly 3 h (range, 36 min.—7.5 h) for 30 individual matings timed from start to finish in the insectary. The nonreceptive or just-mated female dislodged the male by brushing him off with her hindlegs, by moving her ovi- positor downward and away from the male, or by folding her wings over her abdomen as a barrier to copulation. Fighting was observed in the field and insectary. This involved males that faced each other about 7 mm apart and alternately flexed their wings forward. The defender then suddenly rushed forward and butted the intruder with its head. Females similarly employed forward thrusts of their bodies, sometimes accompanied by synchronized forward thrusts of their wings, to confront and ward off intruders, including ants that usually ignored these actions. Generally, however, individual flies exhibited nonag- gressive behavior and spent considerable time resting or grooming themselves. In the insectary, 45- to 60-day-old females were continuously quiescent for 3 to 5 h, except for the normal pumping motion of their mouthparts. In the field and the insectary, males were motionless for as long as 2 h. Both sexes groomed their wings with their hindlegs; the head, eyes, and antennae were groomed with the front legs. The charac- teristic rubbing of both front or hindlegs after grooming apparently removed debris. Sa) Adults occasionally exhibited a swaying movement in the field and insectary, where- upon the body was rocked from side to side. This behavior usually was exhibited during the approach of a moving object, and pos- sibly helped the fly to distinguish an in- truder or potential mate. During prepara- tion for flight, both sexes assumed a characteristic posture with the head lifted, the long axis of the bodies slanted upward, and the front and middle legs extended out- ward. Oviposition and ovipositor probing was observed in the field from 7 AM to 5 PM on spring days; however, these activities peaked during mid-day. These were the most common activities of fecund females on flower heads in the field. Instances of males attempting to mate with ovipositing females were Observed, but these nonreceptive fe- males fended off the males with thrusts of their heads, flew away, or behaved as de- scribed above to prevent mating. The ovipositing female walked upon a flower head while examining it, occasionally stopping to probe the florets with her ovi- positor. Upon locating a suitable site, she bent her ovipositor downward and inserted it between the florets, the tip reaching the precise location previously described. Dep- osition of each egg took about 20 to 60 s, after which searching behavior resumed. Sometimes a female would interrupt her search to walk down the pedicel for a short distance then return to the head and con- tinue her examination. Dissections of 20 E. farinosa flowers heads in which females had recently oviposited yielded an average of 3 (range, 1-5) eggs. Twenty pairs (each | ¢ and 1 2) produced an average of 37 (range, 11 to 141) eggs in insectary cagings. Individual females laid an average of 19 (range, 1-36) eggs per day on days when eggs were laid. The mean ovi- positional period was 21 (range, 16-30) days, the mean age at last oviposition was 41 (range, 26-56) days, and the mean ovipo- sitional period was 16 (range, 1-54) days. 558 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Seasonal history.—Neotephritis finalis is multivoltine in southern California. Adults have been collected throughout the year, with reproduction occurring in low desert areas in winter, e.g. on Helianthus niveus, and continuing 1n a succession of low and high desert, interior valley, and coastal, host- plant species in blossom in spring along a gradient of increasing altitude and rainfall, e.g. on Geraea and Encelia. In summer, this reproduction by N. finalis continues in hosts in mountain areas, e.g. Balsamorhiza and Wyethia. In the fall, this reproductive ac- tivity and, probably, adult migration, again moves downward and continues in hosts such as fall-blooming species of Helianthus. Natural enemies.—Eurytoma vernonia Bugbee (Erytomidae) and Preromalus sp. (Pteromalidae), which are primary, solitary, larval or larval-pupal parasites, were reared from flower heads of Encelia farinosa. Both species were confirmed as parasites of N. finalis and probably also parasitize Trupa- nea nigricornis and, perhaps, Me/anagro- myza viridis in these heads. ACKNOWLEDGMENTS We thank A. L. Norrbom, Systematic Entomology Laboratory, Agricultural Re- search Service, for arranging loans of spec- imens from the U.S. National Museum of Natural History to RDG and for helpful comments on the manuscript. F. L. Blanc (retired), California Dept. of Food & Agric., Sacramento, kindly also loaned RDG se- lected specimens from his private collection and reviewed the manuscript. The parasites and agromyzid associate of N. finalis were identified by E. E. Grissell and G. C. Stey- skal (retired), respectively, at the Systematic Entomology Laboratory. We thank also G. Gordh and J. A. McMurtry for their reviews of early drafts of this paper. LITERATURE CITED Bailey, L. H. 1951. Manual of Cultivated Plants. MacMillan Pub. Co., Inc., New York, 1116 pp. Cavender, G. L. 1981. Life Histories of Trupanea bisetosa (Coquillett) and Paracantha cultaris (Co- quillett) on Helianthus annuus L. (Compositae) in Southern California, with Notes on Trupanea ni- gricornis (Coquillett) (Diptera: Tephritidae). M.S. Thesis, Univ. Calif., Riverside, 187 pp. Cavender, G. L.andR.D.Goeden. 1982. Life history of Trupanea bisetosa (Diptera: Tephritidae) on wild sunflower in southern California. Ann. Entomol. Soc. Am. 75: 400-406. 1983. On distinguishing 7rupanea bisetosa (Coquillett) from 7. nigricornis (Coquillett) (Dip- tera: Tephritidae). Proc. Entomol. Soc. Wash. 85: 275-281. Coquillett, D. W. 1902. New acalypterate Diptera from North America. J. N.Y. Entomol. Soc. 10: 177-191. Foote, R. H. 1960. The species of the genus Neote- phritis Hendel in America north of Mexico (Dip- tera, Tephritidae). J. N.Y. Entomol. Soc. 68: 145- 151. Foote, R. H. and F. L. Blanc. 1963. The fruit flies or Tephritidae of California. Bull. Calif. Insect Surv. Te IS) foro Gilstrap, F. E. and R. D. Goeden. 1974. Biology of Tarachidia candefacta, a Nearctic noctuid intro- duced into the U.S.S.R. for ragweed control. Ann. Entomol. Soc. Am. 67: 265-270. Goeden, R. D. 1985. Host-plant relations of Tru- panea spp. (Diptera: Tephritidae) in southern Cal- ifornia. Proc. Entomol. Soc. Wash. 87: 564-571. 1987. Host-plant relations of native Uro- phora spp. (Diptera: Tephritidae) in southern Cal- ifornia. Proc. Entomol. Soc. Wash. 89: 269-274. Hilgendorf, J. H. and R. D. Goeden. 1983. Phytoph- agous insect faunas of spiny clotbur, Xanthium spinosum, and cocklebur, Xanthium strumarium, in southern California. Environ. Entomol. 12: 404— 411. Loew, H. 1862. Diptera Americae septentrionalis in- digena. Centuria secunda. Berlin. Entomol. Ztschr. 6: 185-232. Munz, P. A. 1974. A Flora of Southern California. Univ. Calif. Press, Berkeley and Los Angeles, 1086 pp. Munz, P. A. and D. D. Keck. 1959. A California Flora. Univ. Calif. Press, Berkeley and Los An- geles, 1681 pp. Steyskal,G.C. 1972. A preliminary key to the species of Neotephritis Hendel (Diptera: Tephritidae). Proc. Entomol. Soc. Wash. 74: 414-416. Wasbauer, M. W. 1972. An annotated host catalog of the fruit flies of America north of Mexico (Dip- tera: Tephritidae). Calif. Dep. Agric. Bur. Ento- mol. Occas. Pap. 19, 172 pp. PROC. ENTOMOL. SOC. WASH. 89(3), 1987, pp. 559-561 A NEW SPECIES OF IRENANGELUS FROM COSTA RICA (HYMENOPTERA: POMPILIDAE: CEROPALINAE) HOWARD E. Evans Department of Entomology, Colorado State University, Fort Collins, Colorado 80523. Abstract.—Irenangelus eberhardi n. sp. is described from Costa Rica. This is the tenth known Neotropical species of these cleptoparasitic Pompilidae and the first to have been reared from a host (Auplopus semialatus Dreisbach). The pompilid genus /renangelus has been known from nine Neotropical species and several from the Oriental and Australian regions (Evans, 1969). Although the genus has several unique features, common struc- tural characters with Ceropales suggest that, like members of that genus, these wasps are cleptoparasites of other Pompilidae. Wil- liams (1919) discovered this with respect to a Philippine species, J. /uwzonensis (Rohwer). He found a small larva attached to an egg of Tachypompilus analis (Fabricius) which had been laid ventrally near the base of the metasoma of the spider prey; he later reared several Jrenangelus from Tachypompilus cocoons. He also reared an Jrenangelus from a cocoon of Auplopus nyemitawa (Rohwer). Nothing further has been added in the more than half a century since Williams’ report. Thus I welcome the opportunity to describe a new species from Costa Rica which has supplied further information on the biology of these rare and unusual wasps (Wcislo et al., in press). Irenangelus eberhardi Evans, New SPECIES Fig. 1 Holotype female.— Length 7.5 mm; fore- wing 6.8 mm. Ground color yellow, marked with black as follows: greater part of man- dibles; extreme sides of clypeus; front with a pair of streaks from just above antennal sockets, nearly connecting with a band be- tween tops of eyes, also with a spot in front of median ocellus and a streak behind ocelli connecting to black on occiput; pronotum with a transverse band anteriorly; mesoscu- tum with three broad longitudinal bands; scutellum black anteriorly and posteriorly, otherwise yellow; postnotum black as well as two longitudinal stripes on propodeum; extreme base of first tergite black as well as broad apical bands on tergites 1—5; small streaks on mesopleura and along suture be- tween meso- and metapleura; mid and hind coxae streaked with black as well as inner surface of mid and hind femora and tibiae and outer surface of hind femora; tarsi yel- lowish brown. Scape yellow, flagellum black except apical 4.5 segments light reddish brown. Wings faintly tinged with yellow; ex- treme tip of forewing faintly clouded; stigma yellowish brown. Labrum emarginate, 2.0 times as wide as its median height; clypeus 1.56 times as wide as its median height. Scape cylindrical; fla- gellum filiform, flagellomere | 1.7 times as long as thick. Genae deeply grooved behind each eye, posterior margin of groove round- ed below, sharp above. Tubercle above an- tennal sockets well developed, bisected by a groove that extends to median ocellus; laterad of the groove the front is deeply pit- 560 Big. 1: ted all the way to eye margins and ocelli; vertex broadly depressed laterad of lateral ocelli. Mesoscutum with notauli distinct, linear, scutum without a median ridge; post- notum 0.7 times as long as metanotum, de- pressed and slightly broadened medially but not notably extended backward into pro- podeum, latter with a shallow median impression. All claws dentate. Third sub- marginal cell broader than second both above and below, receiving second recur- rent vein very slightly beyond middle. Allotype male.—Length 7.3 mm; fore- wing 6.8 mm. Color as in female except apical five segments of antennae light yel- lowish brown. Structure of head and thorax also as described for female; abdomen less PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Trenangelus eberhardi Evans, new species. Male genitalia, ventral aspect. tapered and compressed apically; last tergite shallowly emarginate; subgenital plate quadrate, wholly covered with short setae; genitalia as figured (Fig. 1). Variation.—The females vary in length from 7.5 to 9.5 mm, forewing length from 6.8 to 8.3 mm. In the larger females the black stripes on the front reach the trans- verse band across the vertex and the black streak on the outer side of the hind femora is absent. A paratype male is slightly larger than the allotype, measuring 8.4 mm, fore- wing 8.0 mm. There are no structural dif- ferences worthy of note. Type material.— Holotype 2 and allotype 6, Costa Rica: Heredia, near Puerto Viejo, La Selva, 50 m. Reared from nest of Au- VOLUME 89, NUMBER 3 plopus semialatus Dreisbach, May 1986, W. Eberhard (Museum of Comparative Zool- ogy, Cambridge, Massachusetts). Paratypes: 1 2 and | 6, same data as type; 2 2, same data except dated April 1986 (Museum of Comparative Zoology and U.S. National Museum). Remarks.—This species resembles J. ich- neumonoides Ducke closely both in color and structure; these are the only known American species having deep punctures on the front. Structural differences from ich- neumonoides are as follows; smaller (7.3- 9.5 mm as compared to 9-14 mm in ich- neumonoides), labrum about twice as wide as high (barely wider than high in ichneu- monoides);, scape without a sharp edge be- neath; punctures of front coarser and fewer in number; mesoscutum without a median ridge; male genitalia with parameres much exceeding digiti and parapenial lobes, re- curved tips of digiti less prolonged. Major color differences from ichneumonoides are as follows: antennal flagellum black except apically; front with a pair of black streaks but without a median streak; abdomen yel- 561 low, prominently banded with black; inner surfaces of mid and hind tibiae streaked with black. Undoubtedly both species are mim- ics of species of Ste/opolybia (Vespidae), but judging from the differences in size and ab- dominal coloration 1t seems probable that they mimic different species of that genus. Etymology.—It gives me pleasure to name this species for William G. Eberhard, of the Universidad de Costa Rica, who collected the type series, in recognition of his many contributions to the biology of Neotropical insects and spiders. LITERATURE CITED Evans, H. E. 1969. Studies on neotropical Pompili- dae (Hymenoptera) VII. Jrenangelus Schulz. Stud. Entomol. 12: 417-431. Weislo, W. T., M. J. West-Eberhard, and W. G. Eber- hard. In press. Natural history and behavior of a primitively social wasp, Auplopus semialatus, and a parasite, /renangelus eberhardi (Hymenop- tera: Pompilidae). J. Insect Behav. Williams, F. X. 1919. Philippine wasp studies. Bull. Exp. Stn. Hawaii. Sugar Plant. Assoc. (Entomol.) 14: 1-186. PROC. ENTOMOL. SOC. WASH. 89(3), 1987, pp. 562-570 A REVISION OF THE SHORE FLY GENUS ELEPHANTINOSOMA BECKER (DIPTERA: EPHYDRIDAE) WAYNE N. MATHIS AND JOHN C. DEEMING (WNM) Department of Entomology, NHB 169, Smithsonian Institution, Washington, D.C. 20560; (JCD) Department of Zoology, National Museum of Wales, Cathays Park, Cardiff CFl 3NP, United Kingdom. Abstract.—The shore fly genus Elephantinosoma Becker is revised and now includes the type species, E. chnumi Becker, and a new species, FE. cogani, from Nigeria. The genus is Afrotropical and Mediterranean in distribution and is apparently related to A/llotri- choma, sensu lato, and Hecamede, although a sister-group relationship with either has not been established. The main characters for distinguishing between species are those of the male terminalia, of which figures are provided. The genus Elephantinosoma Becker (1903) was proposed to accomodate E. chnumi and E. perspiciendum, two new species that Becker described in the same paper. The specimens Becker studied came from the Isle of Elephantine, an island in the Nile River, and the island’s name, ev- idently, is the basis for the generic name, not the size or shape of the flies. Indeed, the small size of these shore flies, with body lengths of less than three mm, and their shape, which is typically fly-like, does not allude to an elephant-like body. When Becker described the genus, and in his later treatments of the genus (1922, 1926), he did not designate a type species, and his characterization of the genus was inaccurate for some of the characters he used. Cresson (1946) subsequently designated E. chnumi as the type species for the genus and earlier (1929: 176) had transferred perspi- ciendum from Elephantinosoma to Allotri- choma Becker. Unfortunately, the basis for Cresson’s transfer was a misidentification of specimens he was studying from the type locality in Egypt. Later, Cresson (1946: 249- 250) discovered his error, transferred per- spiciendum back to Elephantinosoma, and described his misidentified specimens as a new species, A//otrichoma aegyptium. The back and forth placement of perspi- ciendum did not abate with Cresson, how- ever. The distinguished English dipterist, J.E. Collin (1949: 204) noted certain anom- alies in the characterization of Elephanti- nosoma and was certain that the two orig- inally included species belonging to “*. . . two quite distinct genera... .”’ Consequently he followed Cresson’s earlier precedent in transferring perspiciendum back to Allotri- choma and retaining E. chnumi as the only included species in Elephantinosoma. Col- lin’s placement of perspiciendum in Allotri- choma was tentative, however, and he fur- ther suggested that this species and a new one he described, A. agens, were possibly related to Pseudohecamede Hendel, a genus known only from the Western Hemisphere. Not having adequate material then avail- able to him for further study of these genera, he left the matter partially unresolved. Some years later, Soika (1956) proposed the sub- genus Eremotrichoma for A. perspiciendum, its type species, A. agens Collin, and A. sim- VOLUME 89, NUMBER 3 plicior Collin, and that subgenus was re- cently revised (Mathis, 1986a). In recent catalogues of the Afrotropical and Palaearctic regions, Elephantinosoma has either included chnumi and perspicien- dum (Cogan, 1980) or just chnumi (Cogan, 1984), with perspiciendum in Allotrichoma. The purpose of this paper is to resolve better some of the remaining problems. The genus is recharacterized, FE. chnumi 1s re- described, and a new species is described from Nigeria. The monophyly of E/ephan- tinosoma within the tribe Atissini is also discussed briefly. The methods used generally in this study were explained previously by Mathis (1985, 1986a, 1986b). The descriptive terminolo- gy, with the exceptions noted in the above- cited papers, follows that published in the recent Manual of Nearctic Diptera, Vol. 1 (McAlpine, 1981). Two venational ratios are used common- ly in the descriptions and are defined here for the convenience of the user (all ratios are averages of three specimens). 1. Costal vein ratio: the straight line dis- tance between the apices of R,,, and R,,;/ distance between the apices of R, and R,,;. 2. M vein ratio: the straight line distance along M basad of crossvein dm-cu/distance apicad of crossvein dm-cu. Descriptions are composite. For the most part, information given in the generic de- scription is not repeated in the species de- scription. Genus Elephantinosoma Becker Elephantinosoma Becker, 1903: 179 [type species: Elephantinosoma chnumi Beck- er, by designation of Cresson, 1946: 249]; 1926: 94-95 [rev.].— Cresson, 1946: 248- 249 [review].—Collin, 1949: 203-206 [disc.].—Cogan, 1980: 657 [Afrotropical cat.]; 1984: 131 [Palaearctic cat.]. Diagnosis.—Small to moderately small shore flies, length 1.80 to 2.65 mm. Head: Wider than high; frons entirely and 563 mostly densely microtomentose, with me- sofrons undifferentiated except for color- ational differences; ocellar setae inserted slightly in front of anterior ocellus; pseudo- postocellar setae moderately well devel- oped, about '2 to *s length of ocellar setae; only reclinate fronto-orbital seta present, no proclinate setae, although sometimes with a minute setula; both inner and outer ver- tical bristles present; ocelli arranged to form isosceles triangle, with distance between posterior pair slightly greater than between anterior ocellus and either posterior ocellus. Antenna dark colored, black, generally within shallow facial groove; aristal length subequal to antennal length and bearing S— 6 dorsal rays, with basal 3-4 rays longer than apical 2-3; 2nd antennal segment with short, proclinate dorsal seta. Eye subround to slightly ovate, bare of setulae. Face be- tween antennal grooves carinate, but not with tuberculate prominence below level of antennal grooves, otherwise face in profile more or less vertical; oral margin broad, wider than narrowest distance between eyes, ventral margin more or less flat, not emar- ginate; facial setae weakly developed, usu- ally only | seta inserted near parafacial; gena moderately high, about 2 eye height and bearing | genal seta; labella broad, fleshy, shorter than mediproboscis; maxillary pal- pus dark, mostly blackish. Thorax: Mesonotum usually light tan to faintly golden, pleura more gray to whitish gray, usually with broad stripe through ven- ter of notopleuron; chaetotaxy generally weakly developed, setulae much reduced and sparse, arranged in setal tracks as follows: acrostichal setulae minute, generally incon- spicuous, in 2—4 rows, setulae of median rows slightly better developed, lateral rows attenuated anteriorly; prescutellar acrosti- chal setae well developed, subequal in length to posteriormost dorsocentral bristles, dis- tance between less than that between either seta and posteriormost dorsocentral seta; dorsocentral track terminated posteriorly with | larger seta; intra-alar setulae irregu- 564 larly seriated; 1 postpronotal seta; presu- tural seta reduced or lacking; | postalar seta; 2 scutellar bristles and with sparse, scattered setulae on scutellar disc; 2 notopleural setae, insertion of posterior seta elevated above level of anterior one; 2 anepisternal bristles along posterior margin and several small, pale setulae on dorsal '; katepisternal seta lacking. Wing: membrane coloration milky white; veins behind costa pale, whitish yellow; vein R,,, extended beyond level of crossvein dm- cu, 3rd costal section about '/ that of 2nd section; alular marginal setulae short, length less than 2 alular height. Legs: not bearing prominent setae or with distinctive conformation; femora mostly gray, microtomentose, tibiae grayish yellow to yellow, tarsi yellow, apical tarsomere darker, browner. Abdomen: Fifth tergum of male visible dorsally, as long as 3rd, not telescoped with- in 4th, and lacking dorsal apodemes that project into 4th but with a distinct oval area toward posteroventral margin (function un- known); 5th sternum poorly sclerotized, U-shaped or as 2 rod-shaped sternites. Male terminalia as follows: cerci fused ventro- laterally with epandrium; surstyli appar- ently lacking; aedeagus long and narrow in lateral view, slightly enlarged apically; go- nite roughly triangular in shape, sheathing base of aedeagus; hypandrium a long nar- row process, slightly longer than aedeagus, that lies between the arms of 5th sternum. Distribution.—Old World. Southern Palaearctic (Mediterranean: Egypt, Israel to Morocco), and Afrotropical (Niger, Nigeria, and Sudan). Cresson (1948: 26) reported a damaged female from India (Punjab). We have examined this specimen and can re- port that it is an undetermined species of Allotrichoma (Eremotrichoma). Natural history.— The few specimens that we have collected of this genus were taken from an exposed, bare shore next to a slight- ly brackish aquatic environment. The spec- imens of E. cogani were collected on a nar- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON row sandy beach having patches of Typha in places and bounded inland by Salvadora scrub. The shore was littered with mammal and fish bones and carrion, including a very ripe dead horse, and had a waveline of drowned tree locusts (Anacridium sp.) and Typha flotsam. Dead chironomids from vast swarms of Tanytarsus spadiceonotatus Freeman blanketed the shore in places. On account of large populations of herons and wading birds, the inshore waters were rich in nutrients from bird lime. The site had a considerable range and quanity of decaying organic matter! Discussion.—The monophyly of Ele- phantinosoma is indicated by the following apomorphies (The outgroups are Allotri- choma, sensu lato, and Hecamede of the tribe Atissin1): 1. One reclinate fronto-orbital seta and no proclinate setae. Usually there 1s at least one proclinate fronto-orbital seta in mem- bers of the tribe Atissini, and its absence in Elephantinosoma is considered to be a de- rived state. 2. Katepisternal seta lacking. Typically the katepisternal seta is present and conspicu- ous. There is no evidence, in a reduced state or otherwise, of this seta in Elephantino- soma, and thus this state is considered to be apomorphous. 3. Facial setae weakly developed, usually only one seta inserted near parafacial. Most atissine taxa have two to three well-devel- oped facial setae. The reduced number of these setae, as in Elephantinosoma, 1s con- sidered an apomorphy. 4. Presutural seta reduced or lacking. The loss of this seta has occurred more than once in the tribe Atissini, each apparently inde- pendently. This character by itself, for this reason, is not strong evidence for the mon- ophyly of Elephantinosoma. As it corrobo- rates the other characters presented here, however, we consider it to be a derived character. 5. Reduced size and number of mesonotal setae. Like character four, this character oc- VOLUME 89, NUMBER 3 565 Figs. 1-4. Elephantinosoma chnumi. |, Head, anterior view. 2, Same, lateral view. 3, Same, dorsal view. 4, Thorax, dorsal view. curs elsewhere in Atissini, but in taxa that are not closely related to Elephantinosoma. Thus, we consider its occurrence in this ge- nus to be an apomorphy and to be conver- gent with the other atissine taxa. A sister group for Elephantinosoma has not been specifically identified, and we are deferring detailed discussion of this matter until a more definitive classification of the tribe has been presented (Mathis, in prep- aration). For this study it is sufficient to know that the genus is related to A//otri- choma, sensu lato, and Hecamede. There is considerable intraspecific vari- ation in color that seems to be related in part to the age of specimens. Younger spec- 566 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 5, 6. imens are lighter colored, especially on the dorsum, and are relatively intact. Older specimens tend to be tattered, and the dor- sum is uniformly darker and with a duller sheen. As there are only two species presently known in this genus, we have not provided a key. The diagnoses and figures for the two should suffice for their identification. Elephantinosoma chnumi Becker Figs. 1-7 Elephantinosoma chnumi Becker, 1903: 180:— Becker, 1922: 72 [rev.]; 1926: 95 [review; figs. of thorax and head].—Cres- Male terminalia of Elephantinosoma chnumi. 5, Lateral view. 6, Posterior view. son, 1929: 179 [rev.]; 1946: 249 [rev., designated as type species of genus].— Co- gan, 1980: 657 [Afrotropical cat.]; 1984: 131 [Palaearctic cat.]. Diagnosis.—Small to moderately small shore flies, length 1.8 to 2.65 mm. Head (Figs. 1-3): Frons mostly light tan to brown, sometimes with fronto-orbits grayish to whitish gray. Antenna blackish; arista with 4—5 dorsal rays. Face mostly white to silvery white; dorsal facial carina concolorous with frons or slightly more golden. Gena concolorous with face. Thorax (Fig. 4): Mesonotum mostly con- colorous with frons to dorsum or notopleu- VOLUME 89, NUMBER 3 AFRICA GOODE BASE MAP SERIES DEPARTMENT OF GEOGRAPHY THE UNIVERSITY OF CHICAGO HENRY M. LEPPARD, EDITOR Fig. 7. ron, sometimes with darker stripes through major setal tracks (acrostichal and dorso- central); area from postpronotum through notopleuron whitish gray; dorsum of anepi- sternum concolorous with mesonotum, oth- erwise pleura uniformly whitish gray. Wing with ratios as follows: costal ratio averaging 0.41; vein M ratio averaging 0.43. Legs with femora mostly whitish gray, concolorous with pleura, tibiae slightly lighter, and tarsi yellowish except for apical 1-3 (some spec- imens with all tarsomeres except for baso- tarsomere dark colored). Abdomen: Dorsum of Ist and 2nd terga lightly tannish, remaining terga light tan- nish gray to silvery gray depending on angle of view. Male terminalia (Figs. 5, 6) as fol- Prepared by Henry M. Leppard © 1964 by The University of Chicago Distribution Map of Elephantinosoma chnumi. lows: ventral apex of epandrium more broadly pointed in posterior and lateral Views. Type material.— The lectotype male of E. chnumi Becker is labeled ““Assuan I 44556 6 [handwritten]/Lectotypus [red; black sub- margin].” The lectotype is double mounted, is in fair condition (right antenna 1s miss- ing), and is in the Humboldt Universitat collection. Although this specimen 1s la- beled as the lectotype, we are not aware of its designation earlier. Other specimens examined.—EGYPT. Aswan (2 °; ANSP, paralectotypes). Sinai: Ofira (sewage), 22 Mar-—21 May 1981, A. Freidberg, W. N. Mathis (2 4, 5 9; TAU, USNM). ISRAEL. Bet Zayada, NE shore 568 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 8, 9. of Lake Kinneret, 5 Aug 1986, W. N. Mathis (5 6, 9 2; USNM); Ma’agan Michael, 29 Oct 1980, A. Valdenberg (1 2°; TAU); Mash’abbé Sade (2 km W), 13 Aug 1986, W. N. Mathis (2 6; USNM); Nahal Iddan, 22 Mar 1980, W.N. Mathis and A. Freidberg (1 2; USNM); Ne’ot Ha’Kikar, 21 Mar 1980, W. N. Math- is and A. Freidberg (1 ¢; USNM); Tel Aviv, li6:SepulO772 Ac Freidberg:(1témlees T AW). MOROCCO. nr. Figuig, Defilia, 5-20 Apr 1966, A. M. Hutson (4 6, 4 9; BMNH). N/- Male terminalia of E/ephantinosoma cogani. 8, Lateral view. 9, Posterior view. GER. Air Massif, Wadi Iberkom, 18°55'’N 8°40’E, 23 Aug 1983, P. C. Matteson (1 34; NMW). SUDAN. Atbara, 3 May 1914, Eb- ner (1 6; ANSP). Distribution (Fig. 7).—Israel, Egypt, Mo- rocco, Niger, and Sudan. Remarks.—This species is very similar to N. cogani and is distinguished from the lat- ter by the shape of the male terminalia, es- pecially the more broadly rounded apices of the epandrium as seen in posterior and lat- VOLUME 89, NUMBER 3 eral views. In addition, the color of abdom- inal terga three through five of N. chnumi is duller, with a slight bluish tinge. The coloration of the dorsum exhibits some age dimorphism, as noted previously for the genus generally. In addition, how- ever, some specimens have distinct, darker brown stripes through the major setal tracks of the mesonotum. As the genitalia of these specimens do not differ from those that lack stripes and because we have not discovered other distinguishing characters, this varia- tion is considered to be intraspecific. Elephantinosoma cogani Mathis and Deeming, New SPeEcIES Figs. /8, 9 Diagnosis. — Moderately small shore flies, length 2.24 mm. Description as in E. chnu- mi except as follows: Thorax: Wing with ratios as follows: cos- tal ratio averaging 0.44; vein M ratio av- eraging 0.38. Abdomen: Terga silvery gray. Male ter- minalia (Figs. 8, 9) as follows: Ventral apex of epandrium narrowly pointed in posterior and lateral view. Type material.— The holotype male is la- beled ““N. NIGERIA: Shore of Lake Chad at Malamfatori, 4-12. iv. 1967. J. C. Deem- ing./Elephantinosoma sp. n. nr. chnumi Becker 2 6 (apex scutellum w. 3-4 fine pale hairs beneath; tergites silvery grey. det. J. C. Deeming 1984/NMWZ. 1981-001.” The holotype is double mounted (glued to a pa- per point), is in good condition, and is in the National Museum of Wales. A male paratype has the same locality data as the holotype but was collected on 13 Apr 1967 (USNM). Etymology.—It is a pleasure to name this species after Brian H. Cogan, who has made numerous contributions to our knowledge of Diptera, the family Ephydridae in par- ticular. Remarks.—This species is distinguished mostly by the shape of the epandrium, es- pecially the ventral margin, which is more 569 sharply pointed. In addition, abdominal terga three through five have a shinier and more silvery gray sheen. ACKNOWLEDGMENTS We are grateful to several individuals and their institutions for the opportunity of studying their collections or having speci- mens loaned to us (we have also listed our home institutions and their acronyms as used in the text). They are as follows: Don- ald Azuma, The Academy of Natural Sci- ences of Philadelphia, Pennsylvania (ANSP); Adrian C. Pont, British Museum (Natural History) (BMNH), London, England; H. Schumann, Humboldt Universitat (HU), Berlin, DDR; National Museum of Wales (NMW), Cardiff, Wales; Amnon Freidberg, Tel-Aviv University (TAU), Tel-Aviv, Is- rael; National Museum of Natural History (USNM), Smithsonian Institution. Elaine R. S. Hodges prepared the illustrations and Holly B. Williams prepared the map. LITERATURE CITED Becker, Th. 1903. Aegyptische Dipteren. Mitt. Zool. Mus. Berl. 2(3): 67-195. 1922. Wissenschaftliche Ergebnisse der mit Unterstutzung der Akademie der Wissenschaften in Wien aus der Erbschaft Treit! von F. Werner unternommenen zoologischen expedition nach dem Anglo-Agyptischen Sudan (Kordofan) 1914. VI. Diptera. Denkschr. Acad. Wiss. Wien. 98: 57- 82. . 1926. Ephydridae und Canaceidae: Families 56a, 56b. 7n Lindner, E., ed., Die Fliegen der pa- laarktischen Region, 6(1): 1-115. Stuttgart. Cogan, B. H. 1980. 71. Family Ephydridae, pp. 655- 669. In Crosskey, R. W., ed., Catalogue of the Diptera of the Afrotropical Region. BM(NH), 1437 pp. London. 1984. Family Ephydridae, pp. 126-176. Jn Soos, A., ed., Catalogue of the Diptera of the Pa- laearctic Region. Hungarian Acad. Sci., 402 pp. Budapest. Cogan, B. H. and W. W. Wirth. 1977. Family Ephyd- ridae, pp. 321-339. In Delfinado, M. D. and D. E. Hardy, eds., A catalog of the Diptera of the Oriental Region. Volume III. Suborder Cyclor- rhapha (excluding Division Aschiza). Univ. Press of Hawaii, 854 pp. Honolulu. Collin, J. E. 1949. Diptera Empididae, Dolichopod- 570 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON idae, Aschiza and Acalypterae. Jn Results of the Armstrong College Expedition to Siwa Oasis (Lib- yan Desert), 1935, under the leadership of Prof. J. Omer-Cooper. Bull. Soc. Fouad I. Entomol. 33: 175-225. Cresson, E. T., Jr. 1929. Studies in the Dipterous Family Ephydridae. Paper II. Trans. Am. Ento- mol. Soc. 55: 165-195. 1946. A systematic annotated arrangement of the genera and species of the Ethiopian Ephydri- dae (Diptera). I. The subfamily Psilopinae. Trans. Am. Entomol. Soc. 72: 241-264. 1948. A systematic annotated arrangement of the genera and species of the Indo-australian Ephydridae (Diptera). II. The subfamily Noti- philinae and supplement to Part I on the subfamily Psilopinae. Trans. Am. Entomol. Soc. 74: 1-28. Mathis, W. N. 1985. Studies of Parydrinae (Diptera: Ephydridae), II: Revision of the Genus Pelinoides Cresson. Smithson. Contrib. Zool. 410, iv + 46 Pp. 1986a. A revision of the subgenus Eremo- trichoma Soika of the shore fly genus A//otrichoma Becker (Diptera: Ephydridae). Israel J. Entomol. 19: 127-139. 1986b. Studies of Psilopinae (Diptera: Ephydridae), I: A revision of the shore fly genus Placopsidella Kertész. Smithson. Contrib. Zool. 430, iv + 30 pp. McAlpine, J. F. 1981. Morphology and Terminolo- gy—Adults [chapter], pp. 9-63. Jn Manual of Nearctic Diptera. Vol. 1. J. F. McAlpine et al., eds., Research Branch Agri. Canada. Monograph 27, 674 pp. Ottawa. Runyan, J. T. and D. L. Deonier. 1979. A compar- ative study of Pseudohecamede and Allotrichoma (Diptera: Ephydridae), pp. 123-137. Jn First Sym- posium on the Systematics and Ecology of Ephy- dridae (Diptera). Deonier, D. L., ed. North Amer- ican Benthological Society 111 + 147 pp. Oxford, Ohio. Soika, A. G. 1956. Contributo allo studio del po- polamento del Sahara: Diptera Ephydridae. Boll. Mus. civ. Storia Nat. Ven. 9: 95-114. PROC. ENTOMOL. SOC. WASH. 89(3), 1987, pp. 571-580 SYSTEMATIC NOTES ON UROPHORA (DIPTERA, TEPHRITIDAE) SPECIES ASSOCIATED WITH CENTAUREA SOLSTITIALIS (ASTERACEAE, CARDUEAE) AND OTHER PALAEARCTIC WEEDS ADVENTIVE IN NORTH AMERICA IAN M. WHITE AND STEPHEN L. CLEMENT (IMW) CAB International Institute of Entomology, 56 Queen’s Gate, London, SW7 5JR, United Kingdom. (SLC) United States Department of Agriculture, Biological Control of Weeds Laboratory (Rome, Italy), American Embassy, Agriculture, APO New York 09794, USA; Present address: USDA, ARS, Regional Plant Introduction Station, 59 Johnson Hall, and Department of Entomology, Washington State University, Pullman, Washington 99164-6402. Abstract.—The choice of Centaurea solstitialis plants of differing origin by Urophora jaculata and U. sirunaseva is described and the distribution of the flies is mapped. A key is provided to separate U. jaculata and U. sirunaseva from other species of current interest to North American weed biocontrol projects, namely U. cardui, U. jaceana, U. quadri- faciata, U. solstitialis and U. stylata. The identity of each species was verified by com- parison with type specimens. A lectotype is designated for U. jaculata Rondani. Three species of Urophora Robineau- Desvoidy (Diptera: Tephritidae) are asso- ciated with Centaurea solstitialis L. (yellow starthistle, YST or St. Barnaby’s thistle) in the Palaearctic region, namely U. jaculata Rondani, U. sirunaseva (Hering) and U. quadrifasciata (Meigen). Yellow starthistle (Centaurea solstitialis) is widespread in the Mediterranean area (Dostal, 1976; Wage- nitz, 1975), southern European USSR (Bes- sarabia, Black Sea coast, parts of the Dnie- per and Don valleys, Crimea, Armenia and Caucasus) (Cherepanov, 1963), Iraq and Iran (Wagenitz, 1980). Yellow starthistle 1s ad- ventive in western and central Europe, North and South America, South Africa, Austra- lia, and New Zealand (Dostal, 1976; Mad- dox, 1981; Maddox et al., 1985; Maddox and Mayfield, 1985). In western United States yellow starthistle is a noxious weed that infests over 3 million hectares of land, especially rangeland. There is considerable interest in using phytophagous arthropods from the Palaearctic region for its biological control (Maddox and Mayfield, 1985; Mad- dox et al., 1985). Moore (1972) lists a fur- ther 28 species of Centaurea (sensu lato) that occur in North America, only two of which are native; the remaining species are all adventive weeds of Palaearctic origin. A seedhead fly from Italy, named as “‘U. sirunaseva,” was the first biological control agent found safe to introduce into western U.S. for yellow starthistle control (Zwé6lfer, 1969; Sobhian and Zwolfer, 1985), but in- troductions of this fly into California in 1969, 1976 and 1977 did not lead to its establishment (Maddox, 1981; Julien, 1982; Sobhian and Zwdlfer, 1985). This failure to establish on Californian populations of yel- low starthistle was attributed to host-plant incompatibility, because “the fly would ovi- Sz PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig: I. jaculata and U. sirunaseva. Spots indicate sites for U. jaculata; stars, U. sirunaseva (old records for Moldavian SSR and Turkey are not included). Due to the close proximity of some U. jaculata sites, some spots represent more than one site. posit on the young buds, and the eggs would hatch, but the larvae would not develop” (D. M. Maddox, pers. comm. in Ehler and Andres, 1983). In 1981, however, a popu- lation of flies, also reported as U. sirunaseva, was discovered at Thermi, near Thessalon- iki, Greece, that will attack yellow starthis- tle of U.S. origin (Sobhian and Zwédlfer, 1985). After one of us (IMW) compared flies from C. solstitialis from seven sites in Italy and eight sites in Greece with the type specimens of U. jaculata and U. sirunaseva, it was dis- covered that the flies from Italy and much of Greece were U. jaculata; only the flies from Thermi were the true U. sirunaseva (Fig. 1). This confirms the suggestion by Steyskal (1979) that the Italian flies released in California were U. jaculata, not U. si- runaseva, and dispels the notion of some weed biocontrol workers (see Maddox et al., Locations of populations of Centaurea solstitialis in Italy and Greece that supplied specimens of U. 1985; Sobhian and Zwéolfer, 1985) that they were dealing with different biotypes or strains of “U. sirunaseva” in Italy and northern Greece. The third species of Urophora associated with yellow starthistle 1s morphologically indistinguishable from U. quadrifasciata (Meigen), a species associated with the Cen- taurea subgenera Jacea (Miller) Hayek, Ac- rolophus (Cass.) Dobrocz. and Phalolepis (Cass.) Dobrocz. Urophora quadrifasciata was released in Canada where it is now es- tablished on C. (Acrolophus) biebersteinii DC. (= C. maculosa: auctt. N. America, nec Lam.) and C. (4.) diffusa Lam. (Harris and Myers, 1984), and this fly has now spread into Montana (Story, 1985b). Yellow star- thistle in North America is not a known host for U. quadrifasciata, suggesting that the Mediterranean population on experimental plantings of Palaearctic and Nearctic yellow VOLUME 89, NUMBER 3 starthistle in Italy may be a distinct host- race or a separate biological species of Uro- phora. There are other Mediterranean pop- ulations that very closely resemble U. quad- rifasciata, these possibly distinct species differ in their aculeus shape and length, and in the colour of their first flagellomere, and many of the host-plant records for U. quad- rifasciata may therefore refer to unde- scribed species. Data on the suitability of differing pop- ulations of yellow starthistle as breeding hosts for U. jaculata, U. sirunaseva, and U. quadrifasciata are presented here. This in- formation was generated by garden plot ex- periments in Italy and Greece set up to mea- sure overall levels of insect herbivory on various thistle species. A key is also pre- sented which separates U. jaculata, U. si- runaseva and U. quadrifasciata from five other Urophora species that have been, or are likely to be, introduced to North Amer- ica for biological weed control. Notes on the systematics, biology and distribution of these eight species are also presented. GARDEN PLOT EXPERIMENTS Methods.—Garden plots with different populations of yellow starthistle and up to three related thistle species were established in the grounds of the USDA Biological Con- trol of Weeds Laboratory, Rome, Italy in 1983, 1984 and 1985, and on the Agricul- tural Research Farm, University of Thes- saloniki, Greece in 1985. The first Rome garden was established on March 25, 1983 when 24 greenhouse-grown rosettes, rep- resenting six populations of yellow starthis- tle (source of seed was Brindisi, Italy; south- ern Spain; Walla Walla and Yakima, Washington; Concord and Tehama County, California), were transplanted into six rows ina plot 12 x 12 m. Representatives of the Six populations (treatments) were arranged ina 6 X 6 Latin square with | m between plants. The plot was surrounded by several hundred plants of the local yellow starthistle population. 573 The 1984 and 1985 Rome garden plots occupied the same plot space (12 x 12 m), which was about 50 m east of the 1983 plot position. In both years greenhouse-grown rosettes were transplanted into the plots be- tween April 3-5. In 1984, representatives of seven yellow starthistle populations (Rome; southern Spain; Walla Walla and Yakima, Washington; Lapwai, Idaho; Sac- ramento and Contra Costa Counties, Cali- fornia) and one cultivar of Carthamus tinc- torius L. were arranged in an 8 x 8 Latin square with 1.5—2.0 m separating the plants. The 1985 plot contained representatives of five populations of yellow starthistle (Rome; southern Spain; Thermi; Lapwai; Sacra- mento) and one cultivar of C. tinctorius in each of four rows, using a randomized com- plete block design for a total of 24 plants. Row and plant spacing was 1.5 m. Urophora colonizing the 1984 and 1985 gardens came mostly from the nearby (50 m) planting of several hundred plants of the local yellow starthistle population, as no yellow star- thistle could be found in the vicinity (0.5- 1 km) of the laboratory. The Thermi, Greece garden (12 x 12 m) contained representatives of three popula- tions of yellow starthistle (Thermi; Lapwai; Sacramento), Cirsium creticum (Lam.) D’Urv., Cynara scolymus L., and Cartha- mus tinctorius in each of six rows, using a randomized complete block. The 18 yellow starthistle rosettes were transplanted into the garden on March 11, 1985. Row and plant spacing was 1.0-1.5 m. There were a few wild plants of yellow starthistle near the plot (within 5 m) and a group of about 40 plants about 10 m away. The ground around and between the Rome garden plots was periodically hoed or machine-tilled to remove weeds. Weeding by hand or with a hoe was done to thin-out aggressive weeds in the Thermi garden. Oc- casional rain was the only source of water for the yellow starthistle plants in all of our garden-plots. Sampling was done by harvesting capitula 574 Table 1. Number of adults of Urophora quadrifas- ciata and U. jaculata that emerged from capitula of yellow starthistle of differing origin grown together in experimental garden plots, Rome, Italy 1983-85. Number of Flies per Year U. quadrifasciata U. jaculata Source of Seed for the YST Plants “1983 1984 1985 1983 1984 1985 Europe Central and southern Italy ue D238 ] SG Southern Spain OF 30 2 On 20.70 Northern Greece - — 2 — 0 United States Washington State 9 1 — 0 O = Idaho State —s 25. IW, .1—. 0:~ 40 California State 0 4 2, Ora 1@ — Plants not grown. in the flower head and seed formation stages (see Maddox, 1981, for description of stages) on each yellow starthistle plant at 3-10 day intervals. Samples were returned to the lab- oratory where Urophora were reared and pinned for identification. All capitula (n = 46,694) were dissected; the presence or ab- sence of Urophora galls in the various pop- ulation groups in the Rome garden was recorded, but only total numbers were recorded from Thermi. Although there was some temporal variation in bud formation and flowering among yellow starthistle pop- ulations in all of the plots, such was not the case between plants of the local strain inside and outside of the plots (Clement, unpub- lished data). Results. —The number of adults of U. jac- ulata and U. quadrifasciata that emerged from the capitula of several different pop- ulation groups of yellow starthistle in the Rome garden plots is shown in Table 1. Flowerhead receptacle galls of U. jaculata were found only in Italian yellow starthistle plants, from which 10 flies emerged. When this small data base is considered along with the statement by Ehler and Andres (1983) that the Italian fly released in California was PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON unable to develop on yellow starthistle in California, our conclusion is that U. jacu- lata (called “U. sirunaseva”’ by Ehler and Andres, 1983) is restricted to its own local populations of yellow starthistle in southern Europe. Table | also shows that yellow star- thistle of several origins is a suitable host for a population of flies that are morpho- logically indistinguishable from U. quadri- fasciata (emerged mid July to mid August, 1983-1985). Although it is possible that U. quadrifasciata 1s a species complex (see ear- lier in this paper), what is important for biological weed control is that North Amer- ican yellow starthistle is a suitable host-plant for some populations of the fly presently known as U. quadrifasciata. Urophora sirunaseva was the only Uro- phora species that attacked the three pop- ulation groups of yellow starthistle in the Thermi, Greece garden plot. The percentage of galls in the capitula of the control (Greek) and Idaho plants that were harvested be- tween June 22 and July 4 averaged (x + SE), 5.63 + 1.70% (n = 6 plants) and 2.92 + 0.95% (n = 6), respectively. These average values are not significantly different (F = 1.72, P > 0.05, data arcsine trans- formed). During the same time period, only two out of five California plants were at- tacked by U. sirunaseva, with 5.10% (n = 98 capitula) and 25.0% (n = 12) of the flow- erheads of these plants containing galls. Overwintering larvae in galls were found in four capitula, which were collected between July 25 and August | on two Idaho plants and one Greek plant. The number of adults that emerged in the laboratory from July I- 17, per population group of yellow star- thistle, were as follows: 17, Greek control; 13, Idaho; 5, California. Sobhian and Zwolfer (1985) stated that the percentage of attack of yellow starthistle heads by U. sirunaseva in northern Greece varied in different yellow starthistle popu- lation groups, but they presented no data to support this claim. From our data we can- VOLUME 89, NUMBER 3 not conclude that U. sirunaseva showed a preference for any one population of yellow starthistle. SEPARATION OF SPECIES Eighty-six nominal species of Urophora are listed in the recent Catalogue of Pa- laearctic Diptera (Foote, 1984), of which 65 species and one subspecies are regarded as good species group taxa. Most of these species are separated by Steyskal (1979), but his paper is largely a compilation of scat- tered published data rather than a revision. In particular, Steyskal notes that the section of his key including U. algira (Macquart), U. jaculata, U. jaceana (Hering), U. siru- naseva and U. solstitialis (Linnaeus) is ten- tative; Steyskal (1979) also presented a sep- arate key to the North American species of Urophora. Steyskal (1979) noted that a full revision of Urophora is needed, using characters of the male and female terminalia dissected from reared specimens. One of us (IMW) has started to gather data for a revision of Palaearctic Urophora species and about 50 species have so far been examined, includ- ing 10 that may be new to science. It has been found that the form of the male ter- minalia is of value for diagnosing the species groups associated with Anthemideae, Car- dueae and Inuleae, but they are not of prac- tical value for separating species associated with Cardueae. Conversely, the shape of the female’s aculeus tip, used in combination with other characters, does provide a reli- able method of identification. As the acu- leus characters have not previously been de- scribed for most species, the aculeus shape and other diagnostic features of the species of current interest to North American weed biocontrol projects are described. For con- venience of use, these data are arranged as a key which includes enough diagnostic in- formation to separate these species from all of the Palaearctic species so far studied; suf- ficient characters are also given to safeguard 5 against the possibility of any of the native North American species being confused with the introduced Palaearctic species. The ter- minology used in the following diagnoses and key follows White (1987a) and is based on that of McAlpine (1981) plus the wing crossband terms of Steyskal (1979). Diagnostic characters of the subfamily Myopitinae.—Cell cup closed by a convex vein CuA,, so that there is no cup extension; head with | pair of orbital setae; dorsocen- tral setae present. Diagnostic characters of Urophora Ro- bineau-Desvoidy, as defined by Steyskal (1979).—Proboscis elongate, with narrow reflexed labella; lower facial margin not pro- truding; fold of proboscis and palpi not ex- tending beyond lower facial margin; vein M ending at or close to wing tip. Diagnostic characters of the Urophora species group associated with Cardueae. — Colour predominantly black; scutellum yel- low; legs and usually antennae mostly or- ange; labellum about 1.5 times as long as Ist flagellomere; palpi (of species included here) orange; wings (of species included here) banded (Fig. 2); wing base yellow or hyaline (except in species associated with Echi- nops); male distiphallus reduced to a narrow membraneous sack, with no sclerotized areas; female spermathecae not sclerotized. KEY TO FEMALES OF SELECTED SPECIES OF UROPHORA Abbreviations: AL = aculeus length; WL = female wing length. 1. Wing with subbasal and discal crossbands fused from C to, or almost to, R,,;. Femora black, except apically. (Preapical and apical cross- bands fused from C to, or almost to, Ry,;.) . 2 — Wing often without a subbasal crossband; if subbasal crossband present, it is separated from the discal crossband by a yellow area. Femora orange, sometimes striped with black ....... 3 2. Discal and preapical crossbands fused from midway between M and CuA,, to hind margin of wing. Ist flagellomere orange, with black apex. Large; WL = 4.9-5.3 mm. Aculeus trun- 576 subbasal, discal, preapical and apical. cate and with a single pair of subapical steps (Fig: 3): (Ale =" 1 -7/-mm)ip. a cardui (Linnaeus) Discal and preapical crossbands not fused. Ist flagellomere orange on inner surface; usually grey on outer surface. Small; WL = 2.3-2.8 mm. Aculeus truncate, without subapical steps (Bigs 74) (ANE = sl e5 2 OMI) arse aati eus, a Cae ME Seer 8 Se Se ee quadrifasciata (Meigen) Aculeus pointed, without subapical steps (Fig. 5). (Subbasal crossband present, from C to A,. Aculeus narrowed well before apex, and again just before apex; AL = 1.3-2.1 mm. WL = 3.4— SUGMOnIGH) ae ae Gen ea Fee eee eee jJaculata Rondani Aculeus truncate, with | or 2 pairs of subapical SUGDS tee eee ee epee eta eae 4 Aculeus with 2 distinct pairs of subapical steps (Figs. 6, 7). (Wing with a subbasal crossband.) Aculeus with | distinct pair of subapical steps (Figs. 8-10), often with a trace of a second pair Of subapicalisteps (Bies..9. lO) css saecee eer 6 Aculeus secondary steps (the pair nearest the apex) placed about three-quarters of the way between the primary steps and the apex (Fig. 6). (WL = 3.1-4.5 mm. AL = 2.6-3.4 mm) SPICE RET ae solstitialis (Linnaeus) Aculeus secondary steps placed about half way between the primary steps and the apex (Fig. 7). (Wing, Fig. 2. WL = 2.6-3.3 mm. AL = [eA DIORA ah sno estos 4 eae sirunaseva (Hering) . Aculeus without secondary steps and margin of aculeus straight between primary steps and apex (Fig. 8). (Wing without, or at most with PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Wing of U. sirunaseva. From wing base to wing apex the four crossbands are named as follows: a trace, of a subbasal crossband. WL = 4.0- 4.8 mm_ Ale, — 2-9=3.5) MM)! 1... rafal Me rence Pete Bat lig Spe aR stylata (Fabricius) — Aculeus with a trace of secondary steps (Figs. DN) ER ties eet Pn oe eee ae 7 7. Small; WL = 2.7-3.2 mm.; AL = 1.4-1.8 mm. Femora yellow. Subbasal crossband reduced, usually only from R, to A, +CuA,. Preapical crossband only one-third to half as broad as its proximal hyaline area. Preapical and apical crossbands separate. (Aculeus, Fig. 9) ...... See OE ee ee bah eh ee affinis (Frauenfeld) — Large; WL = 3.5-4.3 mm.; AL = 2.0-2.6 mm. Fore and hind femora usually with some black markings. Subbasal crossband distinct, from C to A,+CuA,. Preapical crossband usually slightly broader than its proximal hyaline area. Preapical and apical crossbands usually joined, sometimes to beyond R,,;. (Aculeus, Fig. 10) SA ee PO te Oe Jaceana (Hering) BIOLOGY AND DISTRIBUTION Urophora affinis (Frauenfeld, 1857) is a native of central and southern Europe where it attacks Centaurea (Acrolophus) diffusa Lam., C. (A.) maculosa Lam., C. (A.) val- lesiaca (DC.) Jordan and C. (Phalolepis) sterilis Steven. Middle Eastern populations on C. (Calcitrapa) iberica Trev. ex Sprengel are larger and may represent another species (WL = 3.3-4.2 mm; AL = 2.0-2.4 mm). VOLUME 89, NUMBER 3 eT = SS ee a So a ee Ra ce eee Figs. 3-8. Aculei of Urophora spp.: dorsoventral outline with detail of apex. 3, U. cardui; 4, U. quadrifasciata; 5, U. jaculata; 6, U. solstitialis; 7, U. sirunaseva; 8, U. stylata. In the western U.S. and Canada U. affinis has been successfully established on C. (A.) diffusa and C. (A.) biebersteinii DC. (= C. maculosa: auctt. N. America) (Julien, 1982; Piper, 1985). Unfortunately, there are no Palaearctic records of any tephritids asso- ciated with C. biebersteinii, but this is prob- ably a result both of its great rarity and the difficulty of distinguishing it from C. ma- culosa. Following the use of U. affinis in combination with U. quadrifasciata, seed production of Canadian C. biebersteinii and meters Sane cee Figs. 9-10. Aculei of Urophora sp.: dorsoventral outline with detail of apex. 9, U. affinis; 10, U. jaceana. 578 C. diffusa has been reduced, and in certain cases the total plant biomass has now de- clined (Harris, 1984; Harris and Myers, 1984). Urophora affinis forms lignified uni- locular galls of receptacle tissue and the eggs hatch first instar larvae (P. Harris, in litt.). Urophora cardui (Linnaeus, 1758) is a na- tive of cooler areas of the Palaearctic region as far east as Kazakh SSR where it attacks Cirsium arvense (L.) Scop. Urophora cardui has been released in both western and east- ern Canada; it died out in the west, and al- though established in the east, it has failed to reduce the area of its host (Harris, 1984; Peschken, 1984). The fly is established on C. arvense at a few sites in Montana and Washington (Story, 1985b; Piper, 1985). Urophora cardui forms a lignified multiloc- ular stem gall (Lalonde and Shorthouse, 1982) and the eggs hatch second instar lar- vae (Peschken and Harris, 1975). Urophora jaceana (Hering, 1935) is a na- tive of western and central Europe where it attacks C. (Jacea) debeauxii Gren. & God- ron. CG: () jacea L.. Ca(J) nigra Leand GC. (J.) phrygia L. In eastern Canada, where it was first found in 1923 (Shewell, 1961), it has become established on C. (J.) nigra. Urophora jaceana forms a lignified multi- locular gall ofreceptacle tissue (Varley, 1947) and the eggs hatch second instar larvae (Varley, 1937). Urophora jaculata Rondani, 1870 occurs in Italy and Greece (Fig. 1) where it forms lignified unilocular galls of receptacle tissue in C. (Solstitiaria) solstitialis L. Urophora quadrifasciata (Meigen, 1826) is a native of western, central and southern Europe (Middle Eastern and north African records possiby refer to other species). Urophora quadrifasciata attacks the capit- ula of a wide range of Centaurea species and possibly related genera; it forms its gall from the ovary wall (Harris and Myers, 1984) and the gall is not lignified (Varley, 1937). In Canada, U. quadrifasciata has been suc- cessfully established on the same plants as U. affinis (see above) and it has extended PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON its range into Montana and Washington (Story, 1985a; Piper, 1985). Urophora quaa- rifasciata 1s primarily bivoltine, unlike oth- er central European species of known bi- ology, and its eggs hatch as first instar larvae (P. Harris, in litt.). Urophora sirunaseva (Hering, 1938) oc- curs in Moldavian SSR, Turkey and north- eastern Greece on C. (Solstitiaria) solsti- tialis, and in Crete on a related plant which is believed to be C. (S.) idaea Boiss. & Heldr. It forms lignified unilocular galls of recep- tacle tissue which we were unable to distin- guish from those of U. jacu/ata in the same host-plant species. Urophora solstitialis (Linnaeus, 1758) oc- curs in the Palaearctic as far east as Kazakh SSR; its confirmed hosts are all Carduus L. species in which if forms lignified multiloc- ular galls of receptacle tissue. The possibil- ity of using U. solstitialis for the control of Carduus acanthoides L. and C. nutans L. in North America is now being investigated by colleagues at the CAB International In- stitute of Biological Control. Urophora stylata (Fabricius, 1775) occurs throughout Europe (some Asian species may be synonyms) where it attacks Cirsium Mil- ler species, Carduus species and Galactites tomentosa Moench. In British Columbia, U. stvlata has substantially reduced seed production by Cirsium vulgare (Savi) Ten.; it was also released in Quebec, but it died out after the release site was mowed (Harris, 1984; Harris and Wilkinson, 1984). The fly has become established at all release sites in Washington state (Piper, 1985). Uro- Phora stylata forms lignified multilocular galls of receptacle tissue (Harris and Wil- kinson, 1984) and its eggs hatch second in- star larvae (Redfern, 1968). Watson and Harris (1984) list U. maura (Frauenfeld, 1857) and U. kasachstanica (Rikhter, 1964) as potential biocontrol agents of Acroptilon repens (L.) DC. in Can- ada. Unfortunately, the two species asso- ciated with A. repens were not available for study and they could not be included in the VOLUME 89, NUMBER 3 above key; this reference to U. maura is almost certainly based on a misidentifica- tion, as the true U. maura attacks Inula L. species (Inuleae). Moore (1972) lists several other Centaurea species that are attacked by Urophora species in Europe; these are not included in the above key because these Centaurea species are not serious weeds in North America and their associated tephri- tids are unlikely to be required as biological control agents. These other Centaurea species and their associated Urophora species are as follows: U. cf. quadrifasciata on C. (Calcitrapa) iberica Trev. ex Sprengel: undescribed Urophora sp. on C. (Manti- salca) salmantica L.; U. cuspidata (Meigen) on C. (Lopholoma) scabiosa L. Types EXAMINED Depositories of specimens referred to in this paper are as follows: BMNH, for British Museum (Natural History), London, En- gland; LS, Linnaean Society, London, En- gland; MNHP, Museum National d’His- toire Naturelle, Paris, France; MZF, Museo Zoologico dell’Universita degli Studi di Fi- renze, Florence, Italy; NHMV, Naturhis- torisches Museum, Vienna, Austria; ZMUC, Zoologisk Museum, Universitets Copen- hagen, Copenhagen, Denmark. Type local- ities are listed by Foote (1984). Trypeta affinis Frauenfeld, 1857.—Syn- typic male bearing a handwritten label “Frfld 1856” plus 2 female and 2 male possible syntypes; | female dissected (NHMV). Musca cardui Linnaeus, 1758.—Lecto- type female, based on an illustration, des- ignated by White (1987b); specimens pre- sumed destroyed. Euribia jaceana Hering, 1935.—Seven male and 8 female syntypes; | female dis- sected (BMNH). Urophora jaculata Rondani, 1870.—Lec- totype female here designated; dissected (MZF). Two other female specimens against the name “‘jaculata” in the Rondani collec- tion (MZF) are another, possibly unde- scribed, species. 579 Trypeta quadrifasciata Meigen, 1826.— Syntypic female examined, but not dissect- ed (MNHP). Euribia sirunaseva Hering, 1938.—Three male and 4 female syntypes; | female dis- sected (BMNH). Musca solstitialis Linnaeus, 1758.—Pos- sible male syntype examined by White (1987b) (LS). Musca stylata Fabricius, 1775.—Pre- sumed syntype, without an abdomen, ex- amined (ZMUC). ACKNOWLEDGMENTS We are grateful to K. M. Harris, R. A. I. Drew, J. McCaffrey, and A. L. Norrbom for reading drafts of the manuscript, T. Mim- mocchi for her general assistance, A. C. Pont for Russian translation, and to the following for arranging loans of type specimens: R. Contreras-Lichtenberg (NHMV), M. G. Fit- ton (LS), S. Mascherini (MZF), L. Matile (MNHP), V. Michelsen (ZMUC) and A. C. Pont (BMNH). LITERATURE CITED Cherepanov, S. K. 1963. 1624. Centaurea, 20. Sol- stitiaria, pp. 570-573. In Bobrov, E. G. and S. K. Cherepanov, eds., Flora URSS (Flora Unionis Re- rumpublicarum Socialisticarum Sovieticarum). 28. Moscow (in Russian). Dostal, J. 1976. 138. Centaurea. Pp. 254-301. In Tutin, T. G. et al., eds., Flora Europaea 4. Ehler, L. E. and L. A. Andres. 1983. Biological con- trol: Exotic natural enemies to control exotic pests, pp. 395-418. Jn Wilson, C. L. and C. L. Graham, eds., Exotic Plant Pests and North American Ag- riculture. New York. Fabricius, J. C. 1775. Systema entomologiae. Flen- sbvrgi et Lipsiae. 832 pp. Foote, R. H. 1984. Tephritidae (Trypetidae), pp. 66- 149. In Soos, A. and L. Papp, eds., Catalogue of Palaearctic Diptera. 9. Budapest. Frauenfeld, G. R. von. 1857. Beitrage zur Naturge- schichte der Trypeten nebst Beschreibung einiger neue Arten. Sber Akad. Wiss. Wien. 22: 523-557. Harris, P. 1984. 26. Current approaches to biological control of weeds, pp. 95-104. Jn Kelleher, J. S. and M. A. Hulme, eds., Biological Control Pro- grammes Against Insects and Weeds in Canada 1969-1980. Slough. Harris, P. and J. H. Myers. 1984. 31. Centaurea dif- 580 fusa Lam. and C. maculosa Lam. s. lat., diffuse and spotted knapweed (Compositae), pp. 127-137. In Kelleher, J. S. and M. A. Hulme, eds., Biolog- ical Control Programmes Against Insects and Weeds in Canada 1969-1980. Slough. Harris, P. and A. T. S. Wilkinson. 1984. 33. Cirsium vulgare (Savi) Ten., bull thistle (Compositae), pp. 147-154. In Kelleher, J.S.and M. A. Hulme, eds., Biological Control Programmes Against Insects and Weeds in Canada 1969-1980. Slough. Hering, E.M. 1935. Drei neue Bohrfliegen-Arten aus der Mark Brandenburg. (Dipt. Trypetidae). Mark. Tierwelt 1: 169-174. 1938. Neue palaearktische und exotische Bohrfliegen. Dtsch. Entomol. Z. 1938: 397-417. Julien, M. H. 1982. Biological Control of Weeds: A World Catalogue of Agents and their Target Weeds. 108 pp. Slough. Lalonde, R.G. and J.D.Shorthouse. 1982. Exit strat- egy of Urophora cardui (Diptera: Tephritidae) from its gall on Canada thistle. Can. Entomol. 114: 873- 878. Linnaeus, C. 1758. Systema Naturae. Ed. 10. 1: l- 824. Holmiae. Maddox, D. M. 1981. Introduction, Phenology, and Density of Yellow Starthistle in Coastal, Inter- coastal, and Central Valley Situations in Califor- nia. Agric. Res. Serv. U.S. Dep. Agric. ARR-W- 20 July. 33 pp. Maddox, D. M. and A. Mayfield. 1985. Yellow star- thistle infestations are on the increase. Calif. Agric. 39(11/12): 10-12. Maddox, D. M., A. Mayfield, and N. H. Poritz. 1985. Distribution of yellow starthistle (Centaurea sol- stitialis) and Russian knapweed (Centaurea re- pens). Weed Sci. 33: 315-327. McAlpine, J. F. 1981. Morphology and terminolo- gy—adults, pp. 9-63. Jn McAlpine, J. F. et al., eds., Manual of Nearctic Diptera. Vol. 1. Monogr. Biosyst. Res. Inst., Ottawa 27. Meigen, J. W. 1826. Systematische Beschreibungen der bekannten europaischen zweifliigeligen Insek- ten. Hamm. 5: 1-412. Moore, R. J. 1972. Distribution of native and intro- duced knapweeds (Centaurea) in Canada and the United States. Rhodora. 74: 331-346. Peschken, D. P. 1984. 32. Cirsium arvense (L.) Scop., Canada thistle (Compositae), pp. 139-146. /n Kel- leher, J.S.and M. A. Hulme, eds., Biological Con- trol Programmes Against Insects and Weeds in Canada 1969-1980. Slough. Peschken, D. P. and P. Harris. 1975. Host specificity and biology of Urophora cardui (Diptera: Tephriti- dae), a biocontrol agent for Canada thistle (Cir- sium arvense). Can. Entomol. 107: 1101-1110. Piper, G. L. 1985. Biological control of weeds in Washington: Status report, pp. 817-826. /n Proc. VI Int. Symp. Biol. Contr. Weeds, 19-25 August PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1984, Vancouver, Canada. Delfosse, E. S. (ed.). Agric. Can. Redfern, M. 1968. The natural history of spear this- tle-heads. Field. Stud. 2: 669-717. Rikhter, V. A. 1964. Fruit flies (Diptera) in Kazakh- stan. Trudy Zool. Inst. Leningr. 34: 286-290 (in Russian). Rondani, C. 1970. Ortalidinae Italicae collectae, dis- tinctae et in ordinem depositae. Dipt. Ital. Prod- romus, Pars VII-Fasc. 4. Bull. Soc. Entomol. Ital. 2: 5-31, 105-133. Shewell, G. E. 1961. Notes on three European Dip- tera recently discovered in Canada. Can. Entomol. 93: 1044-1047. Sobhian, R. and H. Zwolfer. 1985. Phytophagous insect species associated with flower heads of yel- low starthistle (Centaurea solstitialis L.). Z. An- gew. Entomol. 99: 301-321. Steyskal, G. C. 1979. Taxonomic studies on fruit flies of the genus Urophora (Diptera: Tephritidae). Misc. Publs. Entomol. Soc. Wash. 1979: 1-61. Story, J. M. 1985a. First report of the dispersal into Montana of Urophora quadrifasciata (Diptera: Te- phritidae), a fly released in Canada for Biological Control of spotted and diffuse knapweed. Can. Entomol. 117: 1061-1062. 1985b. Status of biological weed control in Montana, pp. 837-842. Jn Proc. VI Int. Symp. Biol. Control Weeds, 19-25 August 1984, Van- couver, Canada. Delfosse, E. S. (ed.). Agric. Can. Varley, G.C. 1937. The life history of some trypetid flies with descriptions of the early stages (Diptera). Proc. R. Entomol. Soc. Lond. (A) 12: 109-122. . 1947. The natural control of population bal- ance in the knapweed gall-fly (Urophora jaceana). J. Anim. Ecol. 16: 139-187. Wagenitz, G. 1975. 79. Centaurea, pp. 465-585. In Davis, P. H., ed., Flora of Turkey and the east Aegean Islands. 5. Edinburgh. 1980. 39. Centaurea, pp. 313-420. In Re- chinger, K. H. ed., Flora Iranica (Flora des Ira- nischen Hochlandes und der Umrahmenden Ge- birge). 139(b). Graz. Watson, A. K. and P. Harris. 1984. 27. Acroptilon repens (L.) DC., Russian knapweed (Compositae), pp. 105-110. Jn Kelleher, J. S. and M. A. Hulme, eds., Biological Control Programmes Against In- sects and Weeds in Canada 1969-1980. Slough. White, I. M. 1987a. Tephritid flies (Diptera: Te- phritidae). Handbk. Ident. Br. Insects. X. (In press.) 1987b. The Linnaean species of the family Tephritidae (Insecta: Diptera). Zool. J. Linn. Soc. (In press.) Zwolfer,H. 1969. Urophorasiruna-seva (HG.) (Dipt.: Trypetidae), a potential insect for the biological control of Centaurea solstitialis L. in California. Tech. Bull. Commonw. Inst. Biol. Control 11: 105- 154. PROC. ENTOMOL. SOC. WASH. 89(3), 1987, pp. 581-586 A NEW ELACHIPTERA FROM SALT MARSHES, WITH REDESCRIPTION OF E. PENITA AND PARTIALLY REVISED KEY TO NEARCTIC ELACHIPTERA (DIPTERA: CHLOROPIDAE) CurTis W. SABROSKY AND K. R. VALLEY (CWS) Systematic Entomology Laboratory, BBII, Agricultural Research Service, USDA, % National Museum of Natural History NHB 168, Washington, D.C. 20560; (KRV) Bureau of Plant Industry, Pennsylvania Department of Agriculture, Harrisburg, Penn- sylvania 17110. Abstract. —Elachiptera salinaria, new species, is described from adults taken in salt marshes in eastern states from Massachusetts to Florida and distinguished from E. penita, a similar species that does not occur in salt marshes. A lectotype is designated for E. penita, and a partially revised key to Nearctic Elachiptera is presented. The genus Elachiptera Macquart is rep- resented in America north of Mexico by 18 species (Sabrosky, 1965). Sabrosky’s (1948) synopsis of the Nearctic species included descriptions of ten new species and a key; he indicated that the larvae feed on decaying plant material or are secondary invaders in stems of grasses. Sabrosky (1935) also re- ported that adults of two species are known to overwinter. In North America the genus is widespread, and adults are commonly taken in general collecting and in some sur- veys, especially for monocots. During the past decade we have accu- mulated numerous adults of a species from eastern salt marshes from Massachusetts to Florida. These specimens run to E. penita (Adams) in Sabrosky’s (1948) key. Exami- nation of syntypes of EF. penita indicated that the species from salt marshes is new. In this paper we redescribe FE. penita and designate a lectotype, describe E. salinaria new species, the first of its genus known to occur in salt marshes, and revise the appro- priate couplets in Sabrosky’s (1948) key. Elachiptera penita (Adams) Figs. 3, 9 Crassiseta penita Adams, 1908, J. N.Y. Entomol. Soc. 16: 152 (Wisconsin). The species was described from two “males” from Wisconsin. The two syn- types, now in the C. F. Adams Collection in the University of Arkansas, show that the species was misinterpreted in Sabrosky’s (1948) revision of the genus because it was not known to him at that time. It is distin- guished by having the arista broad through- out its length (Fig. 3), ocellar tubercle mi- crotomentose, and scutum chiefly shining, microtomentose only narrowly on sides and before the scutellum. Elachiptera penita of the 1948 revision is the new species de- scribed herein. The male specimen bearing Adams’ handwritten label lacks the hind legs, one middle leg, most of the abdomen, and one wing, but the antennae are complete and show the characteristic broad and flat arista. The second specimen, a female, is complete 582 except for lacking both aristae. Inasmuch as the aristae are the particularly unique fea- ture of this species, the first specimen 1s se- lected as the lectotype, even though it is somewhat incomplete. The second speci- men is labeled as paralectotype. In their present condition, the two specimens must be studied together to realize the complete characterization for the species. The only other specimens known to us are six 2, Washburn Co., Wis., (R. H. Jones); 6, Montcalm Co., Mich., May 15, 1954 (R. R. Dreisbach); 2, Wexford Co., Mich., May 14, 1955 (R. R. Dreisbach); 4, 6 miles east of Kent, Ohio, May 16, 1967 (T. Krys- towski); and 2, 10.5 miles southeast of Kent, Ohio, May 24, 1967 (T. Krystowski). The last specimen is in the Department of Bi- ological Sciences, Kent State University, the others are in the National Museum of Nat- ural History, Washington, D.C. The species may be confused with the common E. nigriceps (Loew) because of the large frontal triangle and shining scutum, but penita has a microtomentose ocellar tu- bercle and incrassate hind femora. The rath- er large hind femora suggest E. formosa (Loew) but the latter has gray mesoscutal stripes, as does EF. vittata Sabrosky, which is also a species with broad arista and gray microtomentose ocellar tubercle. Adams’ description is rather brief and generalized; we supply a more detailed de- scription: Frons nearly square, length barely greater than width; frontal triangle large, approxi- mately as figured for E. angustifrons (Sa- brosky, 1948: fig. 14) but the frons slightly wider, surface of triangle smooth and pol- ished except for marginal groove on each side; ocellar tubercle gray microtomentose; face and cheeks blackish; palpi yellow; an- tennae yellow, aristae black, the latter broad and flat nearly to apex (cf. Sabrosky 1948, Figo). Thorax polished black except for rela- tively inconspicuous thin gray microtomen- tum as a narrow prescutellar band, scutel- lum, postalar declivity, metapleural areas PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON and postscutellum, notopleura, and upper margins of meso- and pteropleura, the mesopleural band barely attaining the hu- meri, the latter chiefly polished; scutum sca- brous on the usual three dorsocentral and acrostical lines and the prescutellar slope; scutum slightly longer than broad, with nu- merous long hairs. Bristles very long: | hu- meral, 1 + 1 notopleural, 1 postalar, 1 pos- terior dorsocentral. Scutellum short and broad, length to breadth as 19:25, rough- ened, dull, disk flat with a number of long hairs, apex blunt: cruciate apical scutellar bristles on well separated small tubercles, one pair of subapical marginals arising on small tubercles, a second pair of marginals close to and only slightly anterior to the subapicals and slightly shorter, but not on tubercles; apical bristles separated at bases by 1.3 times the distance from each to a subapical, the subapicals arising at 0.65 times the length of scutellum (i.e., closer to the apicals than to base of scutellum); pro- sternum black. Abdomen with first segment elongate so that segments | and 2 are subequal, these two fused and occupying nearly one half the length of abdomen, but yellow and weakly sclerotized on broad mesal area, remainder of abdominal dorsum shining brown. Legs predominantly bright yellow, the fore tibia, fore tarsus, and hind tibia brown; hind femur slightly enlarged and elongate, its di- ameter 1.67—2.12 times the diameter of hind tibia. Wing veins pale yellow, venation of nor- mal Elachiptera type, 2nd costal sector: 3rd sector as 13:10, veins 3 and 4 diverging slightly on distal third, crossvein r-m _ be- yond middle of discal cell. Elachiptera salinaria Sabrosky and Valley, NEw SPECIES Figs. do Elachiptera decipiens Loew of Weiss, 1924: 2 UNES.): Elachiptera penita (Adams) of Sabrosky 1948: 380 (Mass.). VOLUME 89, NUMBER 3 583 ieee a 6 9 Figs. 1-9. Elachiptera spp. 1, Head of salinaria. 2, Head of formosa. 3, Third antennal segment and arista of penita. 4-6, Hind femora. 4, Male formosa. 5, Female formosa. 6, Male vittata. 7. Male genital complex of salinaria (AA, aedeagal apodeme; Bph, basiphallus; Dph, distiphallus; Hyp, hypandrium; Pgn, postgonite). 8, 9, Surstyli in left lateral view. 8, Surstylus of sa/inaria. 9, Same of penita. 584 Male and female.— Head: Frons orange- yellow to red, sometimes darkening poste- riorly, approximately as long as broad; fron- tal triangle shining, polished, extending nearly to anterior margin of frons, with gray microtomentum along sides and on ocellar tubercle; occiput (Fig. 1) convexly devel- oped, appearing well rounded when viewed at right angle to frons; occiput, cheeks, and vertex black, microtomentose; face black, yellow to orange between antennal bases and below black band; palpi yellow; antennae orange-yellow, third segment slightly infus- cated at insertion of the black, plumose, slender and attenuated arista. Thorax: Scutum black, microtomentose along perimeter, in dorsocentral areas, and on posterior fourth; humerus (postpronotal lobe) and notopleuron microtomentose; pleural sclerites largely shiny, with small amount of microtomentum on upper bor- der of mesopleuron (anepisternum), on up- per half of pteropleuron (anepimeron), and on laterotergite and hypopleuron (meron). Scutellum black, microtomentose, 2 pairs of marginal bristles on short tubercles, whit- ish hairs on rugulose disc. Legs yellow to dark orange-yellow to nearly black, diam- eter of hind femur |.71—2.0 times diameter of hind tibia. Wings hyaline, 2.4-—2.8 mm long, 0.8-1.0 mm wide. Abdomen: Microtomentose; straw-col- ored basally, apical segments brown, darker along posterior margins. Male genitalia (Fig. 7): cerci rounded at apex, more or less tri- angular, separated by a V-shaped cleft; hy- pandrium closed, with deep, U-shaped notch anteromesally; postgonites broad, aedeagal apodeme with ventral projection that ap- pears fused to hypandrium; aedeagus long, slender, basiphallus lightly sclerotized, ex- panding apically; surstyli (Fig. 8) more elon- gate and slender than in penita, curved and tapering to a point. Type Material.— Holotype 4, allotype, and 12 paratypes (6 4, 6 2), Oceanville, N.J., Oct. 3, 1949 (R. T. Mitchell), common in salt marshes. In the U.S. National Museum of PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Natural History, Washington, D.C. Other paratypes (in Washington unless otherwise noted): Mass.: 2, Falmouth Heights, Aug. 13, 1924 [New York]; 4, 2, Eastham, Gov. Prence Road, May 25, 1969 (K. R. Valley), salt meadow [Ithaca]; 2 6, 1 2°, Cape Cod, Great Sippewissett salt marsh, July 9, 1974 (Susan Vince), on low marsh, dominant Spartina alterniflora. N.Y .: 2, Oak Id., July (A. L. -Melander (Collection); Nain 28c Oceanville, Oct. 4, 1949 (R. T. Mitchell); @, Ocean City, Sept. 1, 1949 (R. T. Mitchell), salt marsh; 8 6, 1 2, Ocean Co., Tuckerton, July 1, 1975 (M. J. Raupp) [New Bruns- wick]; 2 6, 2 2, Ocean Co., 2 mi E of Man- ahawkin, Apr. 1, 1974 (R. F. Denno), on Spartina alterniflora, 6, Morgan, Aug. 7 (Weiss, West) [New York]. Del.: 4, Kent Co., Route 113 at St. Jones River, July 10, 1982 (K. R. Valley), salt marsh [Harrisburg]. N.C.: 6, 2, Beaufort, Sept. 15, 1959 (L. V. Davis); 6, 2 2, Pea Id. National Wildlife Refuge, Oct. 15, 1982 (B. A. Foote) [Kent]; 2 3, 2 2, Ons- low Co., Ashe Id., June 4, 1975 (J. C. Dukes), on Spartina alterniflora; 2 8, 1 °, Carteret Co., Bogue Id., Oct. 17; 1974 (G..CxStey skal); 2, Southport, June 5, 1949 (C. W. Sa- brosky). Ga.: Sapelo Id., on Spartina: 6 °, 7 4, June 1963 (H. Kale); 6, Mar. 21, 1964 (T. Marples); 2 6, May 1963 (E. P. Odum); $, May 17, 1963.(E. P: Odum); andi; 253 Sept. 10, 1963 (E. P. Odum) [Athens]; 4, Glynn Co., Route 50, 0.2 mi W of Jekyll Id., Mar. 29, 1969 (K. R. Valley) [Ithaca]. Fla.: 6, Levy Co., Cedar Key (Lois Wood), salt marsh [Gainesville]; 2 , same locality, Apr. 29, 1984 (A. G. Wheeler, Jr.), sweeping halophytes [Harrisburg]; 3 4, | 2, Wakulla Co., Oyster Bay, July 12, 1981 (C. D. Little), collected as larvae in decaying Spartina al- terniflora [Tallahassee]. Remarks.—Specimens from North Car- olina have infuscated femora and tibiae, es- pecially on the hind legs, and for a time it appeared that there were two species or sub- species, one yellow-legged and northern, the other blackish-legged and southern. How- ever, variation and overlapping of the two VOLUME 89, NUMBER 3 forms, and absence of any observed differ- ences in the male genitalia or other char- acteristics have led us to consider the series before us to represent a single species. Flor- ida specimens have a more extensively mi- crotomentose scutum, but otherwise they too seem conspecific. The specific name is an adjective derived from the Latin sa/inus, salty, referring to the salt marsh habitat of the species. The location of type material has been indicated in the above list by the name of a city in brackets. Following is a key to those localities. The names of the persons to whom we are indebted for the loan of material are in parentheses. Athens: Department of Zo- ology, University of Georgia, Athens, Ga. (E. P. Odum). Gainesville: Florida State Collection of Arthropods, Division of Plant Industry, Florida Department of Agricul- ture & Consumer Services, Gainesville, Fla. (H. V. Weems, Jr.). Harrisburg: Pennsyl- vania Department of Agriculture, Harris- burg, Pa. Ithaca: Department of Entomol- ogy, Cornell University, Ithaca, N.Y. Kent: Department of Biological Sciences, Kent State University, Kent, Ohio (B. A. Foote). New Brunswick: Department of Entomol- ogy & Economic Zoology, Rutgers Univer- sity, New Brunswick, N.J. New York: American Museum of Natural History, New York, N.Y. (C. H. Curran, D. Grimaldi, R. T. Schuh). Tallahassee: Department of Bi- ology, Florida State University, Tallahas- see, Fla. (C. D. Little). Washington: Na- tional Museum of Natural History, Washington, D.C. Comments.—Larval feeding habits of E. salinaria appear similar to those reported for other members of the genus. Recent studies by C. D. Little (1981), Florida State University, indicate this species feeds on decaying shoots of salt water cord grass, Spartina alterniflora Loisel. In addition, Strong et al. (1984) listed salinaria (as Ela- chiptera sp.) as a saprovore associated with S. alterniflora. The key in Sabrosky’s revision (1948) is 585 still serviceable in general, and we revise only part of it. A few notes may be made. 1. Elachiptera aliena Becker has since been referred to Oscinisoma Becker. 2. E. punctulata Becker is now known to be Afrotropical. 3, Couplet 15. second” choice: read’ “or with two or three stripes of gray microto- mentum.” The median or acrostical stripe is always narrower than the dorsocentral stripes and sometimes is absent. The term microtomentum, here and elsewhere in the key and descriptions, is used instead of pol- len or pollinosity, or pruinescence (Sabros- ky, 1983). 4. E. pechumani: After the description of pechumani, G. E. Shewell subsequently called to Sabrosky’s attention another char- acteristic of pechumani 1.e., a spot of gray microtomentum in the lower posterior cor- ner of the mesopleuron (anepisternum). This is not found in nigriceps or angustistylum, nor in the previously keyed angustifrons. Although seeming to be a trifle, 1t 1s appar- ently consistent. 5. Couplet 18, second choice: The black markings are on the mid and hind femora, not tibiae as inadvertently stated. 6. Couplet 20: The “strongly incrassate”’ hind femur of E. formosa is more striking in males than in females. REVISED KEY TO COUPLETS 20-25 OF SABROSKY (1948) 20. Occiput convexly developed, the back of head well rounded (Fig. 1), as viewed from above at right angle to frons; salt marsh species, Massachusetts) toi kloridayere emo acne soe. A Sey ee Elachiptera salinaria, n. sp. — Occiput not convexly developed, back of head straight or barely rounded (Fig. 2) as viewed from above at right angle to frons ......... 21 21. Disk of scutum not vittate, with only a narrow prescutellar crossband of gray microtomen- tum plus gray lateral areas — Disk of scutum distinctly vittate, with stripes of gray microtomentum in dorsocentral po- sitions and a narrower but usually evident median stripe in addition to gray lateral areas that extend from humeral to postalar calli . 23 . Arista broad and flat (Fig. 3); femora deep i) to 586 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON yellow to orange yellow; Wisconsin, Michi- gan, Ohioks — fo.) S ro) = i) O° S ) rs) 7) Q. oO fo} a c (e) {e) = evens > bs ~ ~ oO 14) Qa on pes ” _ o =) {e) oO oO pa 612 0.3 in all cases) except in 1984 when there was a Slight female bias (457:569, binomial test, P < 0.001). However, adults were al- ready emerging when the 1984 sample was collected and, therefore, the protandrous emergence pattern already described may account for the smaller number of males. Adults lived 2-3 weeks in laboratory cages with only water supplied. Nothing 1s known about adult dietary requirements for this species. Females had relatively large ovaries at emergence and mature eggs by 5-6 d old. Matings occurred in the laboratory between newly emerged individuals, but no sperm transfer could be documented until females were 5 d old. The emergence of Procecidochares sp. adults in late September and October is un- usually late for temperate zone, univoltine tephritid gall-formers. By the end of Octo- ber the average nighttime low temperature in Albuquerque is 37°F (2.7°C). Eutreta diana Osten Sacken has a similarly late adult season in Texas (Benbow and Foster, 1982). The 13 species studied by Wangberg (1978, 1980, 1981) in Idaho, including three species of Procecidochares, as well as the six re- maining gall-formers reported herein, emerge earlier in the year. An undescribed species of Procecidochares is facultatively bivoltine in the milder southern California climate (Silverman and Goeden, 1980). Upon emergence, adult Procecidochares sp. leave their larval host plant and fly to a nonhost plant, Atriplex canescens, which serves as the arena in a lek mating system (Dodson, 1986). Sex ratios at the A. canes- cens plants are heavily male biased. After mating, females and eventually males dis- appear from A. canescens. The status of Pro- cecidochares sp. for the next six months 1s unknown. Adults appear again in May in large numbers on C. n. bigelovii. Again sex ratios are highly skewed towards males (three censuses revealed male-female ratios of 20: 4, 71:36, and 69:31; binomial tests, P < 0.001 in all cases). One to three flies per PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON census were also found on A. canescens at this time. Half of the 36 females observed on C. n. bigelovii on 22 May, 1985, were either mating or ovipositing when first seen. At least two hypotheses may account for the period from November to May. Flies might be overwintering as adults in an un- known location, or females might be ovipos- iting in the fall on an undetermined second host. In the latter case the new generation presumably would overwinter in the egg or larval stage, emerge as adults from the sec- ond host in the spring and return to C. n. bigelovii. Either of these scenarios would be unique among tephritids from temperate climates where relatively cold winters are the norm. Most tephritid gall-formers stud- ied in the western United States (e.g. Wang- berg, 1978, 1980, 1981; Silverman and Goeden, 1980; Benbow and Foster, 1982; Dodson, 1987) overwinter as eggs or larvae with continous generations on the same host. Overwintering adults (e.g. 7rupanea bise- tosa, Cavender and Goeden, 1982) are known only from milder climates. The mul- tivoltinism of Procecidochares australis Al\- drich results from a sequential change from galling flowerheads to galling stems, but on the same host (Huettel and Bush, 1972). A potentially parallel example to the one dis- cussed here concerns Myopites cypriacus Hering, which galls the flowerheads of one host in the spring and summer and then galls flowerheads of a different host in the fall in Israel (Friedberg, 1979). Regardless of which life history pattern turns out to be accurate for Procecidochares sp., intriguing consequences are posed for male mating behavior. If the flies observed in fall and spring are separate generations, then male mating strategies must alternate between generations. Adult males emerging in the fall must compete for mates within leks (Dodson, 1986), while spring emerging males search for females at oviposition sites. Alternatively, if spring males are the same generation as the fall (as a result of adult VOLUME 89, NUMBER 3 613 Table 2. Biological comparison of Procecidochares sp. in New Mexico and Procecidochares sp. C in Idaho (Wangberg, 1980). Procecidochares sp. (Dodson, this Paper) Procecidochares sp. C (Wangberg, 1980) Gall morphology Larvae/gall Gall length (mm) All monothalamous Identical Monothalamous/polythalamous Average: 20.8 eS 12.6 Range: 18.5-23.5 7.0-13.5 8.0-18.0 (n = 21) (n = 15) (n = 17) Gall width (mm) Average: D5 4.5 6.9 Range: 4.5-5.5 3.0-6.0 5.0-9.0 (n = 21) (n = 15) (n = 17) Larval length (mm) Average: 3.6 not given Range: 2.5-4.15 2.4-3.1 (7) Pupal length (mm) Average: 3:5 not given Range: 2.7-3.9 2.8-3.5 (n = 20) Gall initiation Adult emergence overwintering), then the same individuals possess dual mating strategies contingent upon time of year. In either case a relatively complex behavioral repertoire is implicat- ed: Procecidochares sp. in New Mexico may be the same species as Wangberg’s (1980) Procecidochares sp. C in Idaho. Table 2 compares biological information on the two populations. Externally the galls of each ap- pear identical (see Fig. 7 in Wangberg, 1980), although there may be some variation in size. Wangberg (1980) described and figured sp. C larvae and pupae with a dorsal patch tanned dark brown. This condition was ev- ident for some but not all of the New Mexico species (49 of 74 larvae, 20 of 75 pupae). Perhaps the most distinctive biological dif- ference was the number of larvae occupying the galls. Wangberg (1980) found both poly- thalamous and monothalamous galls. I dis- Mid-summer Late September—Middle October Mid-summer Late August—Middle September sected 236 galls and never found more than one tephritid larva or evidence of more than one larval chamber. Wangberg (1980) did not observe females Ovipositing on the larval host plant but as- sumed, based on knowledge of two locally occurring congeners, that oviposition did occur and the species overwintered as eggs. He posed two alternatives: that adult flies overwintered or that rapid development took place allowing overwintering in the lar- val stage. As stated above, another possi- bility is a sequential host system. SPECIES REARED FROM FLOWERHEADS Tephritis spp. The following two species were reared from puparia in the flowerheads of their re- spective host plants. Data include the fly species, number of specimens, host plant, 614 and the locality and date of flowerhead col- lection. Information on the numbers of males and females in the Tephritis araneosa rearings are not available at this time. Tephritis araneosa (Coq.); 16 adults, Chrys- othamnus nauseosus ssp. graveolens; mile marker 29 on NM68, Taos Co., 23.VAI Sl: T. signatipennis Foote; 11 males, 4 females; Machaeranthera aquafolia; 16 mi N Je- mez Springs on NM4, Sandoval Co., 30. VIII.1981. These morphologically similar species are both widespread throughout the western United States and Baja California, but this is only the second New Mexico record for T. signatipennis (F. L. Blanc, personal com- munication). Pemberton et al. (1985) list Machaeranthera canescens and Saussurea americana as hosts for T. signatipennis, and Senecio serra is the only other host known (F. L. Blanc, personal communication). By contrast, 7. araneosa has been reared from a wide variety of host species including C. nauseosus (Wasbauer, 1972), but no pre- vious report exists for the subspecies grave- olens. REMAINING SPECIES RECORDS On 5 August, 1981, sixteen species of te- phritids were hand collected off sunflowers, Helianthus annuus, at the Sandia Peak Ski Area, Bernalillo Co., New Mexico. Many of these species were undoubtedly transients on the sunflowers. Among these tephritids were two uncommon species, Mylogym- nocarena apicata (Thomas) (2 females, 1 male) and Procecidochares flavipes Aldrich (1 female). These are the first New Mexico records for both species. M. apicata 1s ex- tremely rare, collected previously only in Colorado and Mexico (Foote, 1960; R. H. Foote and A. L. Norrbom, personal com- munications), whle P. flavipes records are limited to California and Mexico (Foote, personal communication). Wasbauer (1972) lists Brickellia sp. (cited as ““Huettel, un- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON published record’’) as the only host record for P. flavipes. R. D. Goeden (personal com- munication) has reared P. flavipes from two species of Brickellia. There is no host in- formation to date for M. apicata. The final species, Rhagoletis ribicola Doane, likewise has been reared from only one plant genus, Ribes (Wasbauer, 1972). However, this record represents an extreme range extension for R. ribicola. Previously known only from British Columbia (Can- ada), California, Idaho, Oregon, Washing- ton and Wyoming (Bush, 1966; F. L. Blanc, G. L. Bush, and R. H. Foote, personal com- munications), there is at least one state be- tween New Mexico and all other collection sites. The New Mexico population was dis- covered on currant plants (Ribes sp.) in the Sandia Mountains outside of Albuquerque. The host plant no doubt is restricted to me- sic, mountainous habitats in the mainly arid climate of this region. It is not known whether this R. ribicola population repre- sents an isolated, relic population left from a more favorable climatic period in the southwestern U.S., or part of an undocu- mented southerly migration from ancestral populations in the northwestern U.S. In either case, it would be of interest to check for R. ribicola populations during the fruit- ing season of Ribes spp. in Arizona, Colo- rado, and Utah. ACKNOWLEDGMENTS I thank Richard H. Foote and George Steyskal of the Systematic Entomology Lab- oratory, Agricultural Research Service, Washington, D.C., for commenting on the manuscript and for their help in verifying (and sometimes correcting) my tephritid identifications. I am grateful to F. L. Blanc, Richard D. Goeden, and Allen L. Norrbom for constructive criticisms of the manu- script and to my wife, Jill Jereb, who showed unfaltering patience when drives became extra long because of the many “host plant stops.” I also thank my friend Luke George, whose willingness to hike off-trail with me VOLUME 89, NUMBER 3 resulted in the discovery of the Rhagoletis ribicola population, and Durant McArthur for sharing his knowledge of Aciurina galls in Utah. Finally I wish to dedicate this paper to Richard H. Foote in honor of his past and ongoing career in tephritid taxonomy. This paper and others could not have been written without his frequent assistance, which he gave under no obligation and with no expectation of reciprocity. I know that I echo the appreciation of many others whom he has encouraged through the years. LITERATURE CITED Benbow, S. M. and D. E. Foster. 1982. Biology of Eutreta diana Osten Sacken on sand sagebrush Artemisia filifolia Torr. (Diptera: Tephritidae). Pan- Pac. Entomol. 58: 19-24. Blanc, F. L. and R. H. Foote. 1987. Taxonomic ob- servations on United States Tephritidae (Diptera), with seven new species. Proc. Entomol. Soc. Wash. 89: 425-439. Bush, G. L. 1966. The taxonomy, cytology and evo- lution of the genus Rhagoletis in North America. Harvard Univ. Mus. Comp. Zool. Bull. 134: 431- 562. Cavender, G. L.andR.D.Goeden. 1982. Life history of Trupanea bisetosa (Diptera: Tephritidae) on wild sunflower in southern California. Ann. Entomol. Soc. Am. 75: 400-406. Cockerell, T. D. A. 1900. Note on Trypeta notata. J. N.Y. Entomol. Soc. 8: 198. Dodson, G. 1986. Lek mating system and large male aggressive advantage in a gall-forming tephritid fly (Diptera: Tephritidae). Ethology 72: 99-108. 1987. Biological observations on Aciurina trixa and Valentibulla dodsoni (Diptera: Tephriti- dae) in New Mexico. Ann. Entomol. Soc. Am. (in press). Dodson, G. and S. B. George. 1986. Examination of two morphs of gall-forming Aciurina (Diptera: Te- phritidae): Ecological and genetic evidence for species. Biol. J. Linn. Soc. 29: 69-79. Foote, R. H. 1960. Notes on some North American Tephritidae, with descriptions of two new genera and two new species (Diptera). Proc. Biol. Soc. Wash. 73: 107-118. Foote, R. H. and F. L. Blanc. 1959. A new genus of 615 North American fruit flies (Diptera: Tephritidae). Pan-Pac. Entomol. 35: 149-156. 1963. The fruit flies or Tephritidae of Cali- fornia. Bull. Calif. Ins. Surv. 7: 1-117. Freidberg, A. 1979. On the taxonomy and biology of Myopites (Diptera: Tephritidae). Isr. J. Entomol. 13: 13-26. 1984. Gall Tephritidae (Diptera), pp. 129- 167. In Ananthakrishnan, T. N. (ed.). Biology of Gall Insects. Edward Arnold, London. Huettel, M. D. and G. L. Bush. 1972. The genetics of host selection and its bearing on sympatric spe- ciation in Procecidochares (Diptera: Tephritidae). Entomol. Exp. Appl. 15: 465-480. Martin, W. C. and C. R. Hutchins. 1981. A Flora of New Mexico. J. Cramer: Braunschweig, Germany. McArthur, E.D. 1986. Specificity of galls on Chryso- thamnus nauseosus subspecies. U.S. Dept. Agric. For. Serv. Gen. Tech. Rep. INT-200: 205-210. McArthur, E. D., C. E. Tiernan, and B. L. Welch. 1979. Subspecies specificity of gall forms on Chryso- thamnus nauseosus. Great Basin Natur. 39: 81- 87. Pemberton, R. W., C. E. Turner, and S. S. Rosenthal. 1985. New host records for tephritid flies (Dip- tera) from Cirsium and Saussurea thistles (Aster- aceae) in California. Proc. Entomol. Soc. Wash. 87: 790-794. Silverman, J. and R. D. Goeden. 1980. Life history of a fruit fly, Procecidochares sp. on the ragweed, Ambrosia dumosa (Gray) Payne, in southern Cal- ifornia. Pan-Pac. Entomol. 56: 283-288. Steyskal,G.C. 1984. Asynoptic revision of the genus Aciurina Curran, 1932 (Diptera: Tephritidae). Proc. Entomol. Soc. Wash. 86: 582-598. Wangberg, J. K. 1978. Biology of gall-formers of the genus Valentibulla (Diptera:Tephritidae) on rab- bitbush in Idaho. J. Kans. Entomol. Soc. 51: 472- 483. 1980. Comparative biology of gall-formers in the genus Procecidochares (Diptera: Tephriti- dae) on rabbitbrush in Idaho. J. Kans. Entomol. Soc. 53: 401-420. 1981. Gall forming habits of Aciurina spp. (Diptera: Tephritidae) on rabbitbrush (Composi- tae: Chrysothamnus spp.) in Idaho. J. Kan. Ento- mol. Soc. 54: 711-732. Wasbauer, M.S. 1972. An Annotated Host Catalog of the Fruit Flies of America North of Mexico (Diptera: Tephritidae). California Dept. Agric., Sacramento. 172 pp. PROC. ENTOMOL. SOC. WASH. 89(3), 1987, pp. 616-621 HOLCODRYOPS MOULI, AN ANOMALOUS NEW GENUS AND SPECIES OF BEETLE FROM ECUADOR (COLEOPTERA: DRYOPIDAE) PAUL J. SPANGLER Department of Entomology, Smithsonian Institution, Washington, D.C. 20560. Abstract.—A new genus and species, Holcodryops mouli, are described and illustrated with pen and ink drawings. The relationships of the new taxa with the Dryopidae and Limnichidae are discussed. The habitat is described and the collection site is illustrated. The new beetle described below was col- lected 12 years ago in Ecuador during an Ecuador-Peace Corps-Smithsonian Insti- tution Aquatic Insect Survey of that coun- try. Although I hoped that more adults and, perhaps, larvae of this odd genus would be- come available before it was described, that has not happened, and I believe the anom- alous characters of this taxon warrant its description without further delay. The family Dryopidae is diverse, con- sisting primarily of tropical beetles. Adult dryopids may be fully aquatic forms (e.g. Helichus), semiaquatic (e.g. Pelonomus, El- moparnus), or strictly terrestrial (e.g. Pro- toparnus, Sostea, Quadryops), however, all known dryopid larvae are terrestrial. Brown (1981) reported 18 genera and 234 species of Dryopidae for the world; and two addi- tional genera, the monotypic Uenodryops Sato (1981) and Quadryops Perkins and Spangler (1985) with three new species as- signed to it, have been described. Although Dajos (1973) established a new family Chi- loeidae based on his new genus and species Chiloea chilensis from Chile, his Chiloea 1s a synonym of Sosteamorphus and Lawrence (1982) cited Chiloeidae as a synonym of Dryopidae. Therefore, with the new genus described below, the family Dryopidae now includes 21 genera and 240 species. The new genus and species, Holcodryops mouli, described below keys easily to the superfamily Dryopoidea in Crowson’s (1955) classification of the Coleoptera. Be- cause of its distinct metasternal suture, it also fits readily into the Heteroceridae-Lu- trochidae-Limnichidae-Dryopidae-Elm1- dae-Psephenidae lineage as discussed by Lawrence and Newton (1982). In that lin- eage, the anomalous characters of Holco- dryops indicate closest relationships with the families Dryopidae and Limnichidae and suggests three alternatives—this beetle could be assigned to the Dryopidae, to the Lim- nichidae, or to a family of its own. The last alternative has been eliminated because other important diagnostic characters such as those provided by immature stages and internal structures are not available for un- equivocal familial assignment of this taxon. Lacking these additional character states I have centered my discussions on the rela- tionships of this new genus with the Dry- opidae and Limnichidae. The close relationship between the Dry- opidae and Limnichidae has long been rec- ognized by coleopterists. Hinton (1939) in his study of the superfamily Dryopoidea stated, ““The family Limnichidae is closer to the Dryopidae than to any other known family when both the internal and external VOLUME 89, NUMBER 3 anatomy of the adults and larvae are taken into account... .”” Although the larval evi- dence needs to be reevaluated because no true limnichid larvae had been described at that time, the close relationship of the two families is substantial. Referring to the adults, Hinton further stated, ““The only dif- ference between the Dryopidae and Lim- nichidae I have been able to discover which applies to all the forms examined, is a dif- ference between the antennae. In the Lim- nichidae the antennae are filiform or with the apical segments forming a club. In the Dryopidae the segments beyond the second or third are always pectinate and form a club.”” Although that antennal description fits most taxa now in the Dryopidae, it does not fit his unusual genus, Ceradryops Hin- ton (1937) (from Sri Lanka), with only three antennal segments, none of which 1s pecti- nate, nor Uenodryops Sato (from Nepal) which has only six non-pectinate antennal segments. When observed cursorily, the new genus Holcodryops appears to have the typical an- tennae and habitus of a member of the fam- ily Dryopidae; however, closer examination shows that the antennae are atypical and the genus shares some character states with both the Dryopidae and the Limnichidae. The antennae of the Dryopidae vary from 3 to 13 segments with 9 to 11 being the more common number; usually, the first antennal segment is not expanded, the second seg- ment greatly expanded and followed by pectinate segments. However, apomorphic 3-segmented antennae without pectinate segments in Ceradryops Hinton (1937) and 6-segmented, non-pectinate antennae in Uenodryops Sat6 (1981) have been de- scribed for the family. In Holcodryops, the basal antennal segment is greatly expanded; the second is ovoid, smaller than the basal segment, and followed by subpectinate seg- ments 3-7; and the terminal broad compact club is formed by segments 8-11. In his couplet separating the Limnichidae from the Dryopidae, Crowson (1955) used, 617 in addition to the antennal differences, the nearly contiguous metacoxae and last seg- ment of tarsi shorter than the combined length of the other tarsal segments as charac- ters distinctive for the Limnichidae. He also mentioned the peculiar ovipositor of Dryops in connection with its egg-laying habits but did not compare it with that of the limnich- ids. The last tarsal segment of Holcodryops is much longer than the combined length of the other tarsal segments as Crowson (1955) described for the Dryopidae. The ovipositor of Holcodryops has short wide coxites that are very similar to those of Uenodryops but unlike the usual long slender coxites of the typical dryopids such as Dryops and Pelon- omus. In a later discussion of the dryopid-lim- nichid line, Crowson (1978) stated that members of the Dryopidae have five anal veins 1n the metathoracic wing and frequent elytral striae or regular rows of punctures, and lack a frontoclypeal suture —characters not found in the Limnichidae. The meta- thoracic wing of Holcodryops is very similar in venation to that of Helichus, Dryops, and a composite wing for the Dryopidae as il- lustrated by Wallace and Fox (1980). I have examined the wing venation of three lim- nichid genera— Limnichites, Euthryptus, and Eulimnichus—and have found that all three are different than the dryopid wing and have reduced anal veins and the anal cell absent. I have not been able to detect a frontoclypeal suture on the head of Holco- dryops because the surface is densely punc- tate, but I cannot be certain that it is absent. Crowson (1978) also mentions several larval characteristics that distinguish dry- opid larvae from limnichid larvae but not having larvae of this new genus precludes a comparison of larval characters. As noted, examination of the antenna and ovipositor of Holcodryops reveals that they are different from the characteristic anten- nae and ovipositors of most other described dryopid genera. With the noted previous documentation of aberrant departures for 618 the family it seems reasonable that the dif- ferences found in Holcodryops are evidence of another aberration within the family. Although the evidence for placing Hol- codryops in the Dryopidae is incomplete be- cause males and the immature stages are not available, the evidence favors the action I have taken. Holcodryops Spangler, NEw GENUS Figs. 1-6 Body form subrectangular, moderately convex dorsally. Head partially retracted into pronotum; disc slightly depressed; fine carina across head between eyes; anteriorly shallowly recessed for reception of basal an- tennal segment. Maxillary palpus, 4-seg- mented. Labial palpus, 3-segmented. An- tenna, | 1-segmented; basal segment large, subtriangular; second segment almost round; last 4 segments forming compact club (Fig. 3). Clypeus coarsely, densely punctate. Frontoclypeal suture absent (?). Labrum short, broad, straplike, smooth. Pronotum with 2 gibbosities side by side on disc. Pro- sternum about 1.5 times as long as procoxa and trochanter combined. Prosternal pro- cess long triangle between procoxae; apex acute. Scutellum minute. Elytron with 9 rows of coarse punctures and 7 elongate gibbos- ities on intervals as illustrated (Fig. 1). Metathoracic wing (Fig. 4) without radial cross vein; with an anal cell and 5 anal veins. Prosternum, hypomeron, sides of metaster- num, epipleuron, and abdominal sterna | and 2 deeply grooved for reception of legs. Legs with visible portion of procoxae trans- verse and trochantin visible. Tarsal formula 5-5-5; last segment swollen and almost twice as long as segments 1-4 combined. Tarsal claws robust, without teeth. Tibiae slightly expanded apicolaterally and furrowed sub- apically for reception of tarsal segments 1—4. Type species of the genus: Holcodryops mouli Spangler, new species. Etymology.—Holcodryops from holkos, G., meaning furrow; referring to the deep PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON furrows on the prosternum, hypomeron, epipleuron, metasternum, and abdominal sterna | and 2 for reception of the legs; plus Dryops, the nominotypic genus of the fam- ily; gender, masculine. Holcodryops mouli Spangler, NEw SPECIES Figs. 1-6 Holotype female.—Body form and size: Subrectangular, moderately convex dorsal- ly (Figs. 1, 2). Length, 3.52 mm; width, 1.66 mm. Color: Plumbeous dorsally with cupreous metallic reflections on head and pronotum more evident than those on elytra. Ventral surface and appendages rufobrunneous. Head: With coarse, dense, seta-bearing punctures; punctures separated by about half their diameter. Eyes narrow from above. Antenna (Fig. 3) with enlarged subtriangu- lar basal segment; second segment almost round; segments 3-7 narrow, subpectinate; segments 8-11 forming compact club. La- brum very shallowly emarginate anteriorly. Last segment of maxillary and labial palpi enlarged and about twice as broad as pen- ultimate segment. Thorax: Pronotum 0.66 mm long, 1.27 mm wide; widest at base; sides arcuate, di- verging posteriorly, and subserrate; antero- lateral angles obtuse; posterolateral angles in form of right-angles; surface moderately convex medially; lateral margins explanate; disc with 2 adjacent gibbosities and base with 2 prescutellar foveae; coarsely punctate on explanate sides and in front of discal gibbosities; punctures separated by 1 to 4 times their diameter, denser on anterior margin. Elytron with 9 rows of coarse punc- tures, punctures separated by one-half to one times their diameter; intervals micro- reticulate; with 7 low, elongate gibbosities as illustrated (Fig. 1); gibbosities with fine, dense, seta-bearing punctures; side deeply emarginate near midlength for reception of apex of metafemur and base of metatibia VOLUME 89, NUMBER 3 619 i oF ie oe ; e ~ e}e a efe B 7 the ete. ee a ee et ve Ry 1.0mm A a Ko Xe ios al ay 0.2mm Figs. 1-5. Holcodryops mouli, new species. 1, Habitus, Dorsal view. 2, Same, Ventral view. 3, Antenna. 4, Hindwing. 5, Ovipositor. 620 ’ A. aie * vy "ee oe * | i 7 ae bad : . Fig. 6. (Fig. 2). Prosternum 1.5 times as long as combined length of procoxa and trochanter combined. Prosternal process an elongate triangle with sides rimmed on apical third and apex acute. Mesosternum deeply de- pressed medially for reception of prosternal process. Metasternum with disc moderately depressed, deeper posteriorly; with longi- tudinal groove on midline; surface coarsely moderately densely punctate, punctures separated by | to 3 times their diameter. Legs with coarse, dense, seta-bearing punc- tures on ventral surface. Femora when re- tracted into ventral impressions with exten- sive, 1mpunctate, dull areas on upper (opposing) surface; lower surfaces (when re- tracted) punctate and shiny. Tibiae mod- erately expanded apicolaterally and lateral margins of expansion with fringe of dense, short, stout setae. Tarsi of 5 segments; last segment swollen and almost twice as long as segments 1—4 combined. Tarsal claws ro- bust, without teeth. Abdomen: First sternum deeply furrowed for reception of metafemur and metatibia. Second sternum furrowed for reception of metatarsi (Fig. 2). Surface, except furrows, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Holcodryops mouli, new species, biotope; 38 km east of Puerto Viejo, Manabi, Ecuador. coarsely, moderately densely punctate. Punctures on sterna | and 2 larger and sep- arated by one-half to one times their di- ameter; those on sterna 3 and 4 smaller and sparser; those on sternum 5 as on sternum 4 but much denser. Female genitalia: As illustrated (Fig. 5). Male. — Unknown. Type-data.— Holotype 9: ECUADOR: MANABI: Puerto Viejo (38 km E), 11 May 1975, P. J. Spangler; deposited in the U.S. National Museum of Natural History, Smithsonian Institution. Etymology.—The specific epithet, mouli, is a patronym for Edwin T. Moul, my in- spirational high school biology teacher who shared and encouraged my entomological interests. Dr. Moul became a marine phy- cologist, is now retired, and lives in Woods Hole, Massachusetts. Habitat.—The unique holotype was col- lected in the vicinity of a small waterfall (Fig. 6) where I was searching for aquatic beetles in the water, on plants in the splash zone, and along the brook draining from the small pool below the waterfall. This beetle was collected from a plant about 12 m from VOLUME 89, NUMBER 3 the splash zone as I swept an aerial net through plants bordering the brook, and I believe it is a terrestrial genus. ACKNOWLEDGMENTS I thank Andrea Langley Armstrong, Jef- frey H. Cohen, and Ashley B. Gurney for their assistance and pleasant companion- ship during the field work in Ecuador when this specimen was collected. The specimen was collected during the Ecuador-Peace Corps-Smithsonian Insti- tution Aquatic Insect Survey of Ecuador. The field work in Ecuador was partially funded through a grant from the National Geographic Society, a Smithsonian Re- search Award grant and partial support by the Peace Corps program in Ecuador. I also thank Young T. Sohn, biological illustrator, for the pen and ink drawings; Harley P. Brown and Philip D. Perkins for kindly and constructively reviewing the manuscript; and Phyllis M. Spangler for typing the manuscript into the word pro- cessor. LITERATURE CITED Brown, H. P. 1981. A distributional survey of the world genera of aquatic dryopoid beetles (Coleop- tera: Dryopidae, Elmidae, and Psephenidae sens. lat.). Pan-Pac. Entomol. 57: 133-148. 621 Crowson, R. A. 1955. The Natural Classification of the Families of Coleoptera. Nathaniel Lloyd, Lon- don. 187 pp. . 1978. Problems of phylogenetic relationships in Dryopoidea (Coleoptera). Entomol. German. 4: 250-257. Dajos, R. 1973. Description du Coléoptére Chiloea chilensis, n. g., n. sp., type d’une nouvelle famille: Chiloeidae. Ann. Soc. Entomol. France (N.S.) 9: 173-179. Hinton, H.E. 1937. Ceradryops punctatus, new genus and species of Dryopidae from Ceylon (Col.). Proc. Entomol. Soc. Wash. 39: 79-81. 1939. An inquiry into the natural classifi- cation of the Dryopoidea, based partly on a study of their internal anatomy (Col.). Trans. R. Ento- mol. Soc. Lond. 89: 133-184. Lawrence, J. F. 1982. Coleoptera, pp. 482-553. In Synopsis and Classification of Living Organisms. McGraw-Hill Publishing Company, New York. Lawrence, J. F. and A. F. Newton. 1982. Evolution and Classification of Beetles. Ann. Rev. Ecol. Sys- tem. 13: 261-290. Perkins, P. D. and P. J. Spangler. 1985. Quadryops, new genus, and three new species of arboreal Dry- opidae (Insecta: Coleoptera) from Panama and Ec- uador. Proc. Biol. Soc. Wash. 98: 494-510. Sato, M. 1981. Dryopoidea (Coleoptera) of Nepal I. Family Dryopidae. Bull. Nat. Sci. Mus. (A)7: 51- 56 [Tokyo, Japan]. Wallace, F. L. and R. C. Fox. 1980. A comparative morphological study of the hindwing venation of the order Coleoptera, part II. Proc. Entomol. Soc. Wash. 82: 609-654. PROC. ENTOMOL. SOC. WASH. 89(3), 1987, pp. 622-624 Book REVIEW Biological Control in Agricultural IPM Systems. Marjorie A. Hoy and Donald C. Herzog, Eds. Academic Press, Inc., Or- lando, Florida 32887. 1985. 600 pp. (Pro- ceedings of a symposium held in 1984 at the Citrus Research and Education Cen- ter, University of Florida, Lake Alfred, Florida) $49.50. The 46 contributors to this extensive symposium have produced an excellent compendium of information relating to the interaction of biological control and inte- grated pest management. In the concise and thought-provoking preface, the editors apt- ly set the stage for the general theme of the papers that follow. Drs. Hoy and Herzog pose four general questions relating to the context and future of biological control vis- a-vis the practice of IPM. Paraphrased, in part, the following questions present the problems and goals that are the central themes of the symposium: |. Is biological control an integral part of IPM in agricul- ture and, as such, is it ‘“‘a foundation upon which IPM can rest?” 2. Is more research and funding needed to implement and uti- lize biological control in U.S. agriculture? 3. Why isn’t biological control in agriculture expanding —are new technologies needed to fully exploit this science or is it funding alone that limits the application of biological con- trol? 4. What is the actual level of interdis- ciplinary cooperation and coordination within the framework of agricultural IPM? Do we and can we communicate with each other? During the 3-day symposium, most of the presentations addressed, directly or at least indirectly these basic consider- ations. In a discussion of the entomological per- spectives of IPM, Huffaker cites examples in which biological control was a compo- nent part of an IPM program and, at times was the principal controlling factor. Con- versely, the paper presented by Cate reviews numerous attempts to integrate biological control into the IPM strategies for pest con- trol on cotton. However, at present biolog- ical control is not a major functioning com- ponent for IPM in cotton. This variation in the involvement of biological control differs in the research approaches and the disci- plines involved. Questions 2 and 3 address in part the question of budgets and funding. In brief, are more funds required? Peruse the nu- merous recommendations of many of the authors and the answer, as expected, is yes, regardless of the area of study. The broad- based field of biological control, as pre- sented in this book, is expanding but funds are essential to the development of basic research and implementation. Without ad- equate budgets existing technologies will have only limited use and the development of new technologies will suffer. Frisbie and Adkisson briefly summarize the problems and indicate approaches and philosophies in their paper, ““The Future of IPM.” Those two experts believe it is our responsibility as scientists to develop new strategies and, with the aid of agricultural economists, to justify the allocation of more funding to ex- pand IPM and biological control. “The need for effective communication, education, interdisciplinary research and ef- fective interaction between scientist and users’’ was a concern expressed in the dis- cussion section led by Barfield for the plant resistant and cultural practices presenta- tions. But, are the scientists speaking among themselves in interdisciplinary communi- cation? I do not see enough of it in my field of biological control. At least during this symposium the researchers were commu- VOLUME 89, NUMBER 3 nicating. Maybe this type of symposium and the published proceedings is the course to follow to counteract the limited or, at times, non-existent interdisciplinary communica- tion. The major divisions of the book are: A. Executive Conference Summary—1 pa- per, 7 pages, 3 contributors. Subjects: A brief overview of the status and future of IPM in agriculture; research needs; recommendations for a national policy and for strengthening biological control in IPM programs. B. General Introduction—4 papers, 51 pages, 5 contributors and 3 discussion leaders. Subjects: A view from an en- tomological perspective; synopsis of bi- ological control/plant pathology defini- tions; IPM status and definitions; cost benefit analysis and research needs. C. Biological Control of Arthropod Pests— 14 papers, 277 pages, 22 contributors and 5 discussion leaders. Subjects: Overview of plant resistance and cul- tural practices; behavior modifying chemicals; modification of chemical control as a part of ecological selectivity; update on development of selective pes- ticides; problems in establishing natural enemies; genetic improvement of nat- ural enemies; consideration of biotypes; natural enemy augmentation; microbial insecticides; manipulation of diseases and hosts; use of nematodes; statistical analyses and models for natural enemies and crops interaction. D. Biological Control of Weeds—2 papers, 45 pages, 2 contributors and 2 discus- sion leaders. Subjects: Use of arthro- pods, and natural and genetically altered pathogens for weed control. E. Biological Control of Plant Pathogens— 4 papers, 68 pages, 8 contributors and 1 discussion leader. Subjects: Status of and prospects for foliar and soil antag- onists and for molecular genetics of bi- 623 ological agents; control of soilborne pathogens with antagonists. F. Biological Control of Nematodes—1 pa- per, 12 pages, | contributor and | dis- cussion leader. Subject: Status and pros- pects within IPM system. G. Status and Current Limits to Biological Control in Selected Commodity IPM Systems—5 papers, 94 pages, 9 contrib- utors and | discussion leader. Subjects: Status of biological control on (a) citrus in Florida, (b) vineyards in California, (c) alfalfa in eastern U.S., (d) cotton in Texas and Arkansas, and (e) soybeans in the southeastern U.S. The comprehensive yet review approach of the proceedings reminded me of a volume of an Annual Review series. However, the beauty of this book is that all the papers revolved about the central theme of biolog- ical control and IPM which, considering the high level of the papers and the reputation of the contributors, produced a highly in- formative work. I liked the arrangement of the book. The inclusion of the many short but incisive comments of the discussion groups was a positive feature. The book is hardbound on good paper. However, for my tired eyes the print could have been larger. The subjects require slow and attentive reading and I found the thin, smaller print to be tiring. The book 1s worth having especially for those who wish to have an overview of bi- ological control, regardless of one’s defini- tion of this amorphous field of research. Many of the papers will be, and should be, the starting point for future research in view of the reviews conducted and the recom- mendations proposed by the distinguished authors. The editors comment that “Overt utilization of biological control agents has only just begun; much yet remains to be done to make biological control . . . [a] re- liable and accepted component in agricul- tural IPM programs” is an excellent sum- 624 mation of the problems facing this science and its adherents. The use of natural ene- mies for pest control is a science and yet many who give it lip service do not recog- nize the past successes and the great poten- tial of this field of research. Drs. Hoy and Herzog have presented a text that will help PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON define the problems, promises, and path- ways for biological control. John J. Drea, Beneficial Insects Labora- tory, BBIT, Agricultural Research Service, USDA, Beltsville, Maryland 20705. PROC. ENTOMOL. SOC. WASH. 89(3), 1987, pp. 625-631 BYLAWS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON! Article I—Name Article VIII— Meetings Article I]— Objectives Article [LX — Publications Article III— Membership Article X — Dissolution Article IV — Fees Article XI—Limitation and General Prohi- Article V—Officers bitions Article VI—Executive Committee Article XII—Amendments Article VII—Standing Committees Article I.—Name The name of this organization is ““The Entomological Society of Washington.” Article II].—Objectives Section 1.—The objectives of the Society are to promote the study of entomology in all its aspects and to cultivate mutually advantageous relations among those in any way interested in entomology. The Society is organized and operated exclusively for scientific and educational purposes. Article I]. —Membership Section 1.—Members shall be persons who have demonstrated interest in the science of entomology. Section 2.—Applications for membership shall be made to the Membership Committee in writing. They shall be accompanied by brief statements of qualifications. Section 3.—Election of members shall be the responsibility of the Membership Com- mittee in accordance with the procedure outlined in Article VII, Section 2. Section 4.—Any member whose fees are paid in full may become a Life Member by paying $200.00. Fees for a Sustaining Member are $750.00 for a 5-year period. Life Members are exempt from paying further fees. Section 5.—Members of 15 years or more standing, not in arrears for dues or otherwise indebted to the Society when retiring from income-producing employment, may request, and be continued by, the Executive Committee as Emeritus Members without further payment of dues. Members thus relieved of the payment of dues will not be sent copies of the Proceedings. Section 6.—The Society may elect Honorary Members in recognition of long and mer- itorious effort to advance entomological science. Individuals so recognized shall be ap- proved unanimously by the Executive Committee and by two-thirds vote of members present at any regular meeting. Honorary Members shall be elected for life, shall pay no fees, and shall be accorded all privileges of members. The number of Honorary Members ' The last revision of the Bylaws was published in the Proceedings, vol. 81(2): 336-344, April 1979. Amend- ments since then are noted in vols. 84(4): 867, October 1982; 86(4): 974, October 1984; and 88(4): 796, October 1986. Further amendments were adopted at the regular meetings of Dec. 4, 1986, and Mar. 5, 1987, of which the Minutes have not yet been published. — Ad hoc Committee on Bylaws, Donald R. Davis, T. J. Spilman, and Curtis W. Sabrosky, Chairman. 626 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON carried concurrently on the membership roll shall not exceed three, except when an Honorary Member is chosen Honorary President, in which case there may be four. Section 7.—A membership list shall be published at least once every three years in the Proceedings of the Society. Article IV.— Fees Section 1.—The annual fees, which include membership dues and subscription fee, shall be payable by January |. Annual membership dues shall be two dollars and the member’s subscription rate shall be 18 dollars. Members residing outside of the United States may be charged a supplement to cover the increased cost of postage; the amount and nature of this supplement to be determined by the Treasurer and approved by the Executive Committee. Members elected prior to June shall receive all numbers of the Proceedings for that year. Members elected after June shall be exempt from fees for the calendar year in which they are elected and shall receive the Proceedings beginning with January of the next year. Section 2.—The Treasurer shall notify those members one year in arrears for fees. If fees have not been paid one month after notice has been sent, the member’s name shall be removed from the mailing list for the Proceedings. The member shall be notified of such action by the Corresponding Secretary. A member who 1s two years in arrears for fees may be dropped from membership by vote of the Executive Committee two months after a final notice has been sent. Section 3.—A member shall be considered to be one year in arrears if he/she has not paid his/her fees by January of the year in which they are payable. Section 4.—Members dropped for non-payment of fees, or those who have resigned, may be reinstated by payment of fees for the current year and by payment at membership rate for all copies of the Proceedings for which they are delinquent. Article V.— Officers Section 1.—The elected Officers of the Society shall be a President, a President-Elect, a Recording Secretary, a Corresponding Secretary, a Treasurer, an Editor, an Associate Editor, a Custodian, a Program Chairman, and a Membership Chairman. These shall be chosen from resident members. Section 2.—The President or, in his/her absence, the senior officer present (as listed in the previous Section) shall preside at all meetings of the Society and of the Executive Committee. The President or his/her substitute shall have authority to and shall appoint any standing or special committees whose services are required in the interests of the Society. He/she shall conduct such correspondence as should appropriately bear his/her signature as presiding officer. He/she shall represent the Society on the Board of Managers of the Washington Academy of Sciences; or he/she may appoint a substitute for him/her. The President shall deliver an address on some subject pertinent to the objectives of the Society at the first or second meeting subsequent to the completion of his/her term of office. Section 3.—The President-Elect shall automatically succeed to the Presidency at the close of the annual meeting, or whenever the President is unable to complete his/her term of office. Section 4.—The Recording Secretary shall make a record of the Proceedings of the Society of the meetings of the Executive Committee, and shall submit a record of the Society’s Proceedings to the Editor for publication. VOLUME 89, NUMBER 3 627 Section 5.—The Corresponding Secretary shall conduct all official correspondence of the Society except as otherwise provided, and shall keep a list of all members and their addresses. Section 6.—The Treasurer shall have charge of and be responsible for all funds and investments of the Society, shall make routine disbursements, shall maintain the mailing list of members and subscribers together with their addresses, and shall be responsible for the mailing of the Proceedings. Unusual disbursements and investments shall be made only at the direction of the Executive Committee. He/she shall collect all sums due to the Society from any source, notify all members and subscribers who are in arrears, and shall present to the Executive Committee an annual report on the financial status of the Society, and conduct such correspondence as is necessary to carry out these duties. The fiscal year of the Society shall be November | to the following October 31. The Treasurer shall close his/her books at the end of the fiscal year so the accounts of the Society may be audited prior to its annual meeting. Section 7.—The Editor shall be responsible for editing all publications of the Society and shall conduct such business as is necessary to carry out this responsibility. Section 8.—The Associate Editor, who shall automatically succeed to the Editorship, shall normally be elected one year prior to the termination of service by the Editor, which will occur with the completion of a volume of the Proceedings. The Associate Editor shall be responsible for the issues of the next volume of the Proceedings, which will be entirely his/her responsibility as the incoming Editor. Section 9.—The Custodian shall have charge of the reserve stock of the Society’s pub- lications, shall make such sales as lie within the interests of the Society, and shall be responsible for preserving such records, papers, or items of the Society as shall be deemed necessary by the Executive Committee. Section 10.—The Program Chairman shall be responsible for arranging, with the as- sistance of the members of the Program Committee, the program of each meeting of the Society and for notifying the resident members of the Society of all meetings. Those members living in metropolitan Washington and nearby areas shall be considered resident members. Section 11.—The Membership Chairman shall be responsible for activities of the Mem- bership Committee as provided for in these Bylaws. He/she shall notify the Corresponding Secretary of the names and addresses of new members, and cooperate with that officer and the Treasurer in maintaining an accurate membership list. Section 12.—At the annual meeting, having before it the list of candidates submitted by the Nominating Committee, the membership present may make other nominations from the floor. A separate election by written ballot shall be held for each officer for which there are two or more candidates, the ballots being distributed, collected and counted by tellers appointed by the President. When only one candidate for an office is before the Society, election shall be viva voce on motion and second from the floor and in that case two or more offices may be treated in one motion. Section 13.—The officers shall serve for one year and until their successors are elected. They shall assume their duties at the end of the annual (December) meeting except the Treasurer who shall assume his/her duties as soon as arrangements can readily be made with his/her predecessor and the banks for transfer of Society funds. The Executive Committee may ask for such reports of officers as are deemed necessary. Except for the President and President-Elect, who cannot be reelected to these offices in consecutive years, there shall be no limitation as to the number of terms to which an officer may be elected. 628 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Section 14.— Vacancies in any office except President and President-Elect shall be filled through appointment by the Executive Committee. Members selected to fill such vacancies shall hold office only until their successors are elected. If the office of President-Elect becomes vacant, the position will be filled by a special election in accordance with regular nomination procedures. Article VI.—Executive Committee Section 1.—The activities of the Society shall be guided by an Executive Committee. The Committee membership shall consist of all officers and the last available past pres- ident. Section 2.—The Executive Committee shall assume the responsibility for and shall conduct the activities of the Society, direct finances, and provide for meetings and pub- lications. As provided elsewhere in these Bylaws, the Committee shall report fully to the Society once each year, on its conduct of the Society’s business, either through the different officers or by a specially approved representative. The report shall include an approved audit of the Treasurer’s accounts. The Committee shall also consider and present to the Society proposals for change or improvements, and shall transact all other business re- quiring attention and not otherwise assigned. Section 3.—The Executive Committee shall hold such meetings as are required to transact the business of the Society during the year. One of these shall be sufficiently prior to the annual meeting to permit consideration and approval of a summary report for presentation by the President at the annual meeting on the state of the Society and the work of the officers. Other meetings of the Executive Committee may be called at any time by the President or his/her substitute and shall be called promptly by the presiding officer on request of any three members of the Committee other than the presiding officer. The presence of five members of the Executive Committee at any meeting shall establish a quorum. Article VII.—Standing Committees Section 1.—The standing committees of the Society shall consist of a Membership Committee, a Program Committee, a Publications Committee, a Finance Committee, and a Nominating Committee. New non-elective members of these committees shall be ap- pointed by the incoming President each year. The Committees shall report to the Society at one of its meetings or to the Executive Committee as may be required. Section 2.—The Membership Committee, consisting of the elected Chairman and four appointed members, shall search for prospective new members of the Society among professional workers, students and amateurs in entomology, see that their applications are properly executed and by majority vote approve the candidates’ qualifications for membership. The name of each new electee shall be reported by the Membership Com- mittee at a regular meeting; and, in the absence of adverse notice from the members, the electee’s name shall be read as a new member at the next regular meeting, and if feasible he/she shall be introduced by a sponsoring member. If the committee receives an adverse notice concerning an electee, the candidate’s name shall be referred to the Executive Committee and final approval shall be by majority vote of that committee with formal announcement of membership to be made in the usual manner at the next regular meeting following action by the Executive Committee. Section 3.—The Program Committee, with the elected Program Chairman serving as head, shall arrange for the programs and meeting places of all regular meetings of the VOLUME 89, NUMBER 3 629 Society. This Committee shall consist of the Chairman and three members, the latter to be appointed each year. Section 4.—The Publications Committee shall consist of the Editor as Chairman, the Associate Editor, and at least three appointed members. The appointed members shall assist the Editor. The committee shall consider and put into execution plans for promoting the sales of Society publications. It shall make recommendations to the Executive Com- mittee regarding publication of Memoirs and any other special publication. Terms for appointed members of the Publication Committee shall be at the discretion of the Editor and President. Section 5.—The Finance Committee, consisting of the Treasurer as Chairman, the Editor, the Custodian, and the Program Chairman, shall assist the Treasurer in matters of finances of the Society and make recommendations to the Executive Committee relative to these matters. It shall be a particular duty of this Committee to prepare for the Executive Committee at the beginning of each year a statement of the income and expenditures of the preceding year and to prepare a budget based on the estimated receipt and disburse- ments of the coming year, with such recommendations as seem desirable. No financial obligation against the Society may be contracted by any officer or member except as specified in the annual budget or as provided for by special action of the Executive Committee upon recommendation of the Treasurer and Finance Committee. Section 6.—The Nominating Committee of three members shall prepare a list of can- didates comprising one nominee for each office, except the office of President, for pre- sentation to the membership at the regular meeting one month before the annual meeting. The committee shall secure the acquiescence of each candidate before presentation. Section 7.—The Auditing Committee of three persons shall be appointed by the Pres- ident not later than October of each year. It shall inspect the accounts of the Treasurer and report to the Society at its next annual meeting. Article VIII.— Meetings Section 1.—The regular meeting of the Society shall be held, unless otherwise ordered by the vote of the Society or of the Executive Committee, on the first Thursday of each month except June, July, August and September. The annual meeting for the election of officers shall be the regular meeting for the month of December. Special and field meetings may be called by the Executive Committee. The Program Committee or committees appointed for special meetings may, with the approval of the Executive Committee, incur reimbursable expenses. Twenty members shall constitute a quorum. Section 2.—The suggested order of business at the regular meetings, except the annual meeting, shall be as follows: Reading and approval of minutes. Reports of officers and committees. Introduction of new members. Miscellaneous business. Presentation of notes and exhibition of specimens. Presentation of announced topics. Introduction of visitors. Adjournment. Section 3.—The suggested order of business at the annual meeting in December shall be as follows: ——————————— 630 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1. Reading and approval of minutes. 2. Introduction of new members. 3. Presentation by the President of a summary report on the state of the Society and the work of the retiring officers. 4. Election of new officers. 5. Miscellaneous business. 6. Presentation of notes and exhibition of specimens. 7. Presentation of announced topics. 8. Introduction of visitors. 9. Installation of newly elected President. 0. Adjournment. Section 4.—Either the first or the second regular meeting following the annual meeting shall be set aside for the delivery of the annual address of the retiring President. Article [X.— Publications Section 1.— Publications of the Society shall consist of a periodical to be known as the Proceedings of the Entomological Society of Washington which shall contain the Pro- ceedings of the Society and such papers as are accepted for publication in it, a series of Memoirs, and such miscellaneous handbooks or other special publications as may be deemed desirable. Each member in good standing, except Emeritus Members relieved of payment of fees, is entitled to one copy of each issue of the Proceedings (see Art. IV, Sec. py Section 2.—Financial support of the Proceedings shall be provided from subscription revenues, from the sale of complete or partial sets of the Proceedings, from the fees of life and sustaining memberships, from editorial charges, and from such other funds as the Executive Committee shall determine. Section 3.—The Society shall maintain a separate fund to be known as the Special Publication Fund. At the discretion of the Executive Committee, any unrestricted portion of the Special Publication Fund may be used for publishing Memoirs, handbooks, or other special publications. In any one year, a sum not exceeding the previous five years’ income from interest on the Fund monies may be taken from this Fund and applied toward the publication of the Proceedings, such sum to be returned to the Special Publication Fund at the discretion of the Executive Committee. The Special Publication Fund will be derived from bequests and gifts, from the sale of Memoirs, handbooks, or other special publi- cations, and from such other funds as the Executive Committee shall determine. Section 4.—The publications of the Society shall be of the highest quality and content. To ensure this the Editor shall require review of all manuscripts before their acceptance for publication. Article X.— Dissolution If the Society should be dissolved or terminated, all its assets remaining after payment of all liabilities of the Society shall be transferred by the Executive Committee, upon recommendation of the Executive Committee and approved by the membership of the Society, to one or more nonprofit organizations that have scientific and educational pur- poses, preferably with the same object as the Society, and that qualify as exempt orga- nizations under Section 501 (c) (3) of the Internal Revenue Code, or under corresponding successor provisions. VOLUME 89, NUMBER 3 631 Article XI.— Limitation and General Prohibitions Section |.—The objectives of the Society are listed in Article II of the Bylaws. Lobbying or activities specifically designed to influence legislation are not among the objectives of the Society and no official group within the Society shall engage in such activity. Section 2.— Notwithstanding any provision of the Bylaws which might be susceptible to a contrary construction: a. no part of the net earnings of the Society shall or may under any circumstances inure to the benefit of any private shareholder or individual: b. no substantial part of the activities of the Society shall consist of carrying on prop- aganda, (or otherwise attempting to influence legislation); c. the Society shall not participate in, or intervene in (including the publishing or distributing of statements) political campaigns on behalf of any candidate for public office; d. the Society shall not be organized or operated for profit; e. the Society shall not: 1) lend any part of its income or corpus, without the receipt of adequate security and a reasonable rate of interest, to; 2) pay any compensation, in excess of a reasonable allowance for salaries or other compensation for personal services actually rendered, to; 3) make any part of its services available on preferential bases, to; 4) make any purchase of securities or any other property, for more than adequate consideration in money or money’s worth from; 5) sell any securities or other property for less than adequate consideration in money or money’s worth to; or 6) engage in any other transactions which result 1n substantial diversions of its income or corpus to any officer, member of the Executive Committee, or substantial contributor to the Society. The prohibitions contained in this subsection (e) do not imply that the Society may make such loans, payments, sales or purchases to anyone else, unless such authority be given or implied by other provisions of the Bylaws. Article XII.—Amendments These Bylaws may be amended at any regular meeting by a two-thirds vote of the members voting, if the total number voting represents a quorum, provided that such amendment has been passed by a two-thirds vote of the Executive Committee and pre- sented to the Society in written form at the meeting prior to the meeting at which the vote is taken. PROC. ENTOMOL. SOC. WASH. 89(3), 1987, p. 632 ANNOUNCEMENT OF A NEW MEMOIR No. 13 of the Memoirs of the Entomological Society of Washington, An Identification Manual for the North American Genera of the Family Braconidae (Hymenoptera), was published in April. The authors are P. M. Marsh, S. R. Shaw, and R. A. Wharton. The memoir is 98 pages long and is spiral bound so that it will lie perfectly flat when opened. The key treats 233 genera. Structural characters referred to in the key are completely illustrated with scanning electron micrographs and line drawings on 51 plates. References to taxonomic and other studies are given in the key. Copies can be purchased for $18.00 each from V. L. Blackburn, Custodian, Entomological Society of Washington, % USNM NHB-168, Washington, D.C. 20560. ANNOUNCEMENT TO CONTRIBUTORS The October issue of the Proceedings is complete. Please send all new manuscripts to: Dr. Hiram G. Larew, Associate Editor, Entomological Society of Washington, Bldg. 470, BARC-East, ARS-USDA, Beltsville, MD 20705. : PUBLICATIONS FOR SALE BY THE ENTOMOLOGICAL SOCIETY OF WASHINGTON / MISCELLANEOUS PUBLICATIONS _ Cynipid Galls of the Eastern United States, by Lewis H. Weld 0 etn $ 5.00 @ymipid Galls of the Southwest, by Lewis,\H. Weldii 2) 3.00 HEtHEpADETS (ON) CYNIPIG Pals Ue A knee VIE POU EIA ee Ei ak Bae ee) We P| 6.00 - Identification of Alaskan Black BlyVarvae; by, Kathryn Mi Sommerman iui iii iha er Le URE 1.00 Unusual Scalp Dermatitis in Humans Caused by the Mite Dermatophagoides, by Jay R. eS ECAR SR US A A AN A 1.00 _ A Short History of the Entomological Society of Washington, by Ashley B. Gumey.. 1.00 _ Pictorial Key to Species of the Genus Anastrepha (Diptera: Tephritidae), by George C. RS URE PT RSS BS Pa AY TR PRN NOE hh A 1.50 _ Taxonomic Studies on Fruit Flies of the Genus Urophora (Diptera: Tephritidae), by George C. BREE VSK Al move ey LPs BRULEE i SAUER DCR RR PO MOB WEEE EEG SEC WN GOS EOE bs BE 2.00 ; ie MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON No.1. The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939... $15.00 . No.2. A Classification of Larvae and Adults of the Genus Phyllophaga, by Adam G. Boving. (out of | 2731) yoy op OA MEY: PAB t Sd SOLON Tc LM SN Sa 2 PMA TS A Od YA RY SE lata LG BD print) No. 3. The Nearctic Leafhoppers, a Generic Classification and Check List, by Paul Wilson Oman. e 228 OWLS SSC ATI 28 2 SO Did aT OP a 15.00 . No. 4. A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 1952.00. 15.00 No A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp. USA Ls OE ES Ci SN A AOR A RE VD Op ad CE 15.00 : No. 6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi 5) Makahasi“ 2 S0¢pp FOO heel ee COE by LRA DANE Ce OUR OL Be LID eee BUR CUASn) RN fl PDD 15.00 No. 7. Ant Larvae: Review and Synthesis, by George C. Wheeler and Jeanette Wheeler. 108 pp. 1h ES OS A 8 11.00 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. 1979.0 AND 12.00 No.9. The Flower Flies of the West Indies (Diptera: Syrphidae), by F. Christian Thompson. 200 / Daoist ee okey UNCURE RUE RAPA: PRP rR E EV RN i ae RUT RU Ug Sm MOE OD BA ET 10.00 2 No. 10. Recent Advances in Dipteran Systematics: Commemorative Volume in Honor of Curtis W. | \, Sabrosky. Edited by Wayne N. Mathis and F. Christian Thompson. 227 pp. 1982.0. 11.00 No. 11. A Systematic Study of the Japanese Chloropidae (Diptera), by Kenkichi Kanmiya. 370 pp. H FORSTER INCRE NC Ea OL A I YD A 18.00 . No. 12. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff. f NAIM LS? UN ASSO A Nd Se ME ana Oe 5.00 | 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 _... 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 1e Custodian, Entomological Society of Washington, c/o Department of Entomology, NHB 168, Smithsonian Institution, Washington, D.C. 20560. «' i i CONTENTS (Continued from front cover) MATHIS, W. N. and J. C. DEEMING—A revision of the shore fly genus Bop hliusispave rie Becker (Diptera? Ephydridae) wis) 2) A Ea ee a Ta TNE a i ie aL ee wa MATTA, J. F. and D. E. PETERSON— The larvae of two North American diving beetles of the} genus) Acilius' (Coleoptera: Dytiscidae)) ) Lie sue PO Te Pa a PURRINGTON, F. F. and D. G. NIELSEN — Discovery of the T. W. Harris collection at Haraid a , University and designation of a lectotype for Podosesia syringae Harris (Lepidoptera: Sesiidae) ie yee UR St TA ee Ee a a a YSIS REEVES, R. M.—A new arboreal Carabodes four eastern North America iran Oribatida: Warabodrdae) ty pee Rae AIR AN Tas VR ANT OR UREA oe) wy ae Ape aed nae SABROSKY, C. W. and K. R. VALLEY—A new Elachiptera from salt marshes, with rede- scription of E. penita and partially revised key to Nearctic Elachiptera (Diptera: i: i Chioropidae) iis Lak Say pA AE PRN a A RT ly OSL ee ns ee SCHAEFER, C. W. and I. AHMAD— A cladistic analysis of the genera of the Lestonocorini (Hemiptera: Pentatomidae:, Pentatominae)! 2 3). fk 2 ee a a ee SELANDER, R. B. and A. A. LAURENSE—On the immature stages of Psalydolytta fusca f' ; (Coleoptera: /Meloidae))).0 NAb Ayer ae Guta PU RE DAR Te a 2) a Ya a ay. SLATER, J. A. and D. A. POLHEMUS—Two new species of Botocudo from vertical rock faces ini Indonesia) (Hemiptera: Liyeacidae) fark SOM REN aT A ene ee ee pee ee SPANGLER, P. J.—Holcodryops mouli, an anomalous new genus and species of beetle from is Ecuador'(Coleoptera; Dryopidae) iia Mae dee oe ie Le sO 9 rae ere ie ; STARK, B. P. and R. F. SURDICK—A new Kathroperla species from western North America (Piceoptera: |Chloroperlidae) ee A We Be Te a . TABER, S. W., J.C. COKENDOLPHER, and O. F. FRANCKE-—Scanning electron microscopic study of North American Pogonomyrmex (Hymenoptera: Formicidae) ................. TODD, J. L. and B. A. FOOTE— ‘Spatial and temporal distribution of shore flies i ina freshwater marsh.( Diptera: (Ephiydridae)) yj.) f.3 hyd AU ORL EMR See Ae oe eS UN 6 Se WALLER, D. A., J. P. LAFAGE, R. L. GILBERTSON, and M. BLACK WELL—Wood- decay fungi aeoeinien with subterranean termites (Rhinotermitidae) in Louisiana ............ WHITE, I. M. and S. L. CLEMENT —Systematic notes on Urophora (Diptera, Tephritidae) = species associated with Centaurea solstitialis (Asteraceae, Cardueae) and other Palaearctic / weeds adventiveiin| North) America (SU e Pare ta Ale Et) ae ON Oe a aaa BOOK REVIEW DREA, J. J.—Biological Control in Agricultural IPM Systems .........0.000 000000 e cece eens 6 2 i BYLAWS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ...... EVR 9) ANNOUNCEMENT OF: A‘ NEW MEMOER Aut eu h ee a eae Bio i ANNOUNCEMEN E TO CONTRIBUIGORS "pine rte TUN tad 08 Be oe te a OCTOBER 1987 NO. 4 (ISSN 0013-8797) | PROCEEDINGS INTOMOLOGICAL SOCIETY 4 WASHINGTON PUBLISHED QUARTERLY CONTENTS DLER, P. H.—A new North American species in the Simulium vernum group (Diptera: Si- muliidae) and analysis of its polytene chromosomes ................ 00.000 e cece ences 673 iM KER, G. T.—Apical sensilla on the adult and larval labial and maxillary palpi of Odonto- i Dhigenius disjunctus \(lliger) (Coleoptera: Passalidae) |) 3.0. eS VA ee 682 bs y SAWN A AONUMA WSR CL IA Um AUN TA OY NG MOE IAT VP 825 HN). Abr. P. E.—Host specificity and laboratory rearing studies of Megacyllene mellyi (Coleop- y | i ‘tera: Cerambycidae), a potential biological control agent of Baccharis neglecta Britt. (As- , a oi teidests) AUB CALLUS OPEN ie at ae A A LL AA aged SE 665 i, DENNO, R. F., M. E. SCHAUFF, S. Ww. WILSON, and K. L. OLMSTEAD— Practical diagnosis _ and natural history of two sibling salt marsh-inhabiting planthoppers in the genus Prokelisia aa aaa Beiphaeidaay eye yw te MG EN eh eR Ue Wk eR MRR Ree 687 " Coccinellidae) i in PEN MUG States jp the AU Ee BRNO PEARL PERU eC Se ROTEL DE Um Ba 821 Hh the GeSch PON! Ob MINEWASPECIES eh pi bee Selina tat DUMP gee aah acne Y Os SOT Be fa i Me pace te 811 _ Cerambucidae) on nearby goldenrod, Solidago spp. (Asteraceae), at a forest-field edge ... 706 } ; Redescription of the male, female, and nymph and aie ofvthe larva SY hb 646 . 3 Biba haad, Nebo eed bltiels) BELAY EEE NRL | BLY MSE BS ALE BU ESB HL STS Oe ON ES BO Bt 701 ¥ K URCZEWSKI, F. E.—Nesting behavior of Tachysphex laevifrons and T. crassiformis, with a AW note bn ws Krombeinth (Hymenoptera: Sphecidae))) Wi) .\ hoe A SUP a ee Week eRe Be 715 x LEGNER, E E. F. and R. D. GOEDEN ~ Larval parasitism of Rhagoletis completa (Diptera: OT MRO a a a AS a Le ee Se RU ee 739 te INGAIR, R. W.—Mating behavior at floral resources in two species of Pseudomasaris (Hy- Ne i} menoptera: Vespidae: Masarinae) SR EAT AA END Me AOA UE AA CD La Rd ea A 2 759 i (Continued on back cover) ame THE \ ENTOMOLOGICAL SOCIETY OF WASHINGTON Bs ORGANIZED MARCH 12, 1884 4 OFFICERS FOR 1987 4 THOMAS E. WALLENMAIER, President MICHAEL J. RAupP, Program Chairmar F. EUGENE Woop, President-Elect GEOFFREY B. WHITE, Membership Chairman PAUL M. MarsH, Recording Secretary Victor L. BLACKBURN, Custodiar RICHARD G. RosBINs, Corresponding Secretary MANYA B. STOETZEL, Delegate, Wash. Acad. Sci NORMAN E. WOoDLEY, 77reasurer RAYMOND J. GAGNE, Editor Publications Committee DAVID R. SMITH THEODORE J. SPILMAN GEORGE C. STEYSKAL Associate Editor HirRAM G. LAREW Honorary President C. F. W. MUESEBECK Honorary Members ASHLEY B. GURNEY THEODORE L. BISSELL All correspondence concerning Society business should be mailed to the appropriate officer at the followin address: Entomological Society of Washington, c/o Department of Entomology, NHB 168, Smithsonian Insti tution, Washington, D.C. 20560. R MEETINGS. — Regular meetings of the Society are held in the Natural History Building, Smithsonian Institue on the first Thursday of each month from October to June, inclusive, at 8 P.M. 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(a Please see p. 730 of the July 1984 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 Wash- ington, c/o Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash- ington, D.C. 20560. My Editor: Raymond J. Gagné, Systematic Entomology Laboratory, c/o U.S. National Museum NHB 168, A ington, D.C. 20560. Managing Editor and Known Bondholders or other Security Holders: none. 4 This issue was mailed 23 October 1987 Second Class Postage Paid at Washington, D.C. and additional mailing office. } PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA f Gly fem oh \ PROC. ENTOMOL. SOC. WASH. VEU Y J C 89(4), 1987, pp. 633-645 BANDED ADMIRA N MARYLAND: ANALYSIS OF THE LIMENITIS (BASILARCHIA) ARTHEMIS-ASTYANAX COMPLEX (LEPIDOPTERA: NYMPHALIDAE) AT GREEN RIDGE STATE FOREST , AUSTIN P. PLATT Department of Biological Sciences, University of Maryland Baltimore County, Ca- tonsville, Maryland 21228. Abstract. — The various forms of butterflies comprising the Limenitis arthemis-astyanax complex are reviewed and their genetic relationships established. Suggestions are made for distinguishing forms albofasciata Newcomb and proserpina Edwards. The species status of arthemis and astyanax is reviewed from an historical perspective. The occurrence of banded butterflies (forms arthemis and albofasciata) at Green Ridge State Forest in western Maryland is documented. Hardy-Weinberg analyses demonstrate that the observed and expected numbers of banded, partially banded, and unbanded insects closely approximate each other in three consecutive annual samples (1982-84) and in the total sample as well (n = 586). Analyses of “beak tear” frequencies among quantitative samples of banded, partially banded, and unbanded butterflies in the UMBC and Milwaukee Public Museum Collections, suggest that bird predation is an important factor driving the evolution of the disruptively banded and unbanded mimetic wing patterns north and south of the blend zone, respectively. Within the blend zone, similar data suggest that selection is “relaxed.” Wing fragments obtained from 47 butterflies eaten by a single gray catbird at Green Ridge State Forest in 1982 and 1984, suggest that localized predation can be quite heavy, with a single bird consuming an average of 11 butterflies per day. The banded purple and red-spotted pur- ple butterflies of the northeastern United States form a fascinating complex of blend- ing populations which are of extreme inter- est to students of evolution and natural se- lection. Despite the facts that the two parental forms look very different from one another, and that they occupy different but adjacent geographic areas, they are none- theless extremely similar to each other in many features, and can be judged to rep- resent merely forms within a single inter- breeding species complex (Platt, 1983). This paper will review the important literature relating to these butterflies, and will ex- amine the genetics, population biology, and selective adaptations of the intergrade pop- ulation located at Green Ridge State Forest (Allegany Co.) in western Maryland. The region is part of the “Ridge and Valley” biotic area of Maryland (Fales, 1974). This population is important because it repre- sents a southern protrusion of the north- eastern L. arthemis-astyanax blend zone, which runs southward along the ‘‘back- bone” of the Appalachians, extending into western Virginia. Apparently, Green Ridge is the only locality in Maryland where the banded admirals occur in any numbers. FORMS OF THE ADMIRAL BUTTERFLIES The L. arthemis-astyanax complex con- sists of four major forms (Table 1), each of 634 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON which will be discussed and described in turn: 1) Form arthemis (Drury, 1773).—This is the common and widespread northern banded purple butterfly. It 1s believed to be ancestral to the other forms and species within the genus (Chermock, 1950; Platt et al., 1970). The white bands are wide and conspicuous on all wings. Its geographic range extends from Alaska across Canada to Nova Scotia and southward into Penn- sylvania and New England. The medial wing bands provide a disruptive color pattern of alternating dark and light fields (Platt and Brower, 1968; Platt, 1983). 2) Form astyanax (Fabricius, 1775).— This is the common black and iridescent blue-green butterfly, lacking bands, but hav- ing large and conspicuous red-orange spots along the ventral hindwing (VHW) margins. It is a known Batesian mimic of the unpal- atable blue swallowtail (Battus philenor L., Papilionidae), with which its geographic range is widely co-incident (Brower and Brower, 1962; Platt et al., 1971). The sub- species L. a. astyanax is distributed from central New England southward to the Gulf Coast, into central Mexico, and westward across the Great Plains to Arizona. 3) Form proserpina (Edwards, 1865).— This butterfly is distributed across two de- grees of latitude (41°-43°N) within the ar- themis-astyanax blend zone (Edwards, 1884; Saunders, 1932; Hovanitz 1949; Platt and Brower, 1968). The type locality is the Catskill Mountains of eastern New York State (Edwards, 1865, 1873, 1877, 1891; Smith, 1891). In the central region of the blend zone, this partially banded hetero- zygous form usually is the most prevalent one. The form is subject to considerable phenotypic variation, running the gamut from conspicuous partial white banding on all wing surfaces to individuals having vir- tually no expression of the band at all. These forms have been classified as white band (WB) categories 3, 4, and 5, respectively by Platt and Brower (1968) and Platt (1975). Hence, proserpina continuously grades into the astyanax phenotype. It is distributed from Maine across southern Ontario to ex- treme northeastern South Dakota, and southward into New York and Pennsylva- nia, as well as throughout Maryland (where it is rare), and into western Virginia. It is most prevelant within the central region of the arthemis-astyanax blend zone, but can occur wherever the geographic ranges of the two parental forms overlap. 4) Form albofasciata (Newcomb, 1907).— This form was first described to represent a white banded “‘ursula’’ (sic astyanax). The type localities of the original male and fe- male specimens are in eastern Massachu- setts. Unfortunately, the two types (pres- ently in the USNM collection, Smithsonian Institution) now are known to represent ge- netically distinct phenotypes. The male (a reared specimen) possesses a distinctive, very straight and narrow, medial white band, especially evident on the hindwings. On the other hand, the female type has a somewhat wider, bowed white band, which, although narrower than normal, cannot be genetically distinguished from form arthemis per se. Thus, I suggest restricting the usage of form albofasciata (Newcomb) to apply only to those individuals having the distinctive nar- row banding of the male type specimen. Such a form occurs in both sexes and breeds true. The differences between form arthemis and form albofasciata are shown in Figs. 1-4. Figs. 2 (arthemis) and 3 (albofasciata) are very close to Newcomb’s original type spec- imens of albofasciata. The form albofascia- ta iS rare in most populations. It occurs mainly throughout the southern one-half of the L. arthemis-astyanax blend zone (Platt, 1983, fig. 6, p. 18). Itis common among the banded butterflies at Green Ridge in west- ern Maryland, and it has been taken in Nel- son County, and at Mountain Lake (Giles Co.), Virginia (Clark and Clark, 1951). SPECIES STATUS OF THE L. ARTHEMIS-ASTYANAX COMPLEX Because they possess distinctive color patterns and geographic ranges (see above) VOLUME 89, NUMBER 4 Dorsal Ventral Figs. 1-4. Lab-reared F, specimens of banded admirals, showing the wide and narrow banded forms. The specimens are in the UMBC collection. They represent the central Massachusetts stock, but are nearly identical to those found in western Maryland. 1) & 2) 6 & 2 of form L. a. arthemis (Drury), 3) & 4) 6 & 2 of form L a. albofasciata (Newcomb). The two central specimens [2) 2 arthemis (with white band slightly narrowed and bowed) & 3) 4 albofasciata], closely approximate Newcomb’s (1907) types of form a/bofasciata. The latter are illustrated life-sized in color on the plate accompanying the original description in Psyche. Both types are now in the USNM collection in Washington, D.C. arthemis and astyanax were first considered to be distinct species. This antiquated view is in agreement with the original “type species’ concept, based exclusively on mor- phology. However, the more recent (and ad- vanced) species concepts include 1) the “non-dimensional” concept, which empha- sizes the “discontinuities of nature” (e.g. that all natural species are distinctive, or differ- ent, from one another), and the “‘multi-di- mensional” (or biological) species concept, which questions whether or not two forms are either potentially, or actually, capable of interbreeding in the natural environment (Mayr, 1963, 1970). By either of these two latter concepts, the L. arthemis-astyanax butterflies must be judged to represent a sin- gle species for reasons I shall demonstrate. Grey (1879), Edwards (1884, 1891) and Scudder (1889) recognized the seeming plasticity of these butterflies. Likewise, Field (1904, 1910) and Poulton (1909) stressed the importance of their species relation- ships, and what these, in turn, can tell us about the evolution of mimicry. Nakahara (1924) provided evidence that suggested that 636 arthemis and astyanax were the same species, based on his genitalic studies of all of the North American Limenitis. Saunders (1932) and Gunder (1934) discussed the L. arthemis-astyanax complex at length, stressing both the abundance and variable nature of the intergrading forms of proser- pina, but Klots (1953) accepted the single species viewpoint. Other authors, Hovanitz (1949) and Remington (1958, 1968) have re-emphasized that they are two species, whereas, Platt and Brower (1968), Platt et al. (1970), and Platt (1975; 1983) have fur- ther stressed the single species concept. Fisher (1958: 145-146) provided an ini- tial interpretation of this butterfly complex, as follows: ““An example of a species in process of fission, in which sexual preference is evi- dently playing an important part, occurs in butterflies of the genus Limenitis (Basilar- chia) in the Eastern United States. ... The interpretation of the data is facilitated by the circumstance that the conspicuous white band in L. arthemis is due to a single Men- delian factor, in which that form differs from astyanax .... The white band is incom- pletely recessive and hybrid females caught wild and yielding 50 per cent arthemis-like young, have thus shown themselves to be fertile with the one parental form... .” ““A most important feature, the signifi- cance of which has been pointed out by the late Professor Poulton, is that the rarity of the heterozygotes, in the zone in which they occur, implies that the butterflies in this zone display a strong preference in mating, each for its own kind... . It is probable, in fact, that heterozygosis is here never maintained for more than a few generations, and that almost the entire supply of heterozygotes comes from stray matings of the subspecies, which are thus on the verge of complete genealogical separation.” The above interpretation is most certain- ly wrong, as data provided by Platt and Brower (1968), Bergman and Masters (1971), and by Platt (1983) point out. Across PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON the zone of intergradation between arthemis and astyanax, a north-south transect in New England (four localities) revealed that the heterozygotes (form proserpina) were abun- dant (not rare, as presumed by Fisher), and that the butterflies were, in fact, breeding at random, in accordance with the Hardy- Weinberg principle (Platt and Brower, 1968). Annual studies of the northern Ver- mont and central Massachusetts popula- tions (1965-1975) have confirmed these findings. Likewise, other populations within the blend zone (e.g. those in northern and central Pennsylvania, central Wisconsin, Minneapolis, and northeastern South Da- kota) also evidence “‘free”’ interbreeding just as was found for the New England localities (Platt, 1983). Thus, the original scenario presented by Fisher (1958) is incorrect: arthemis and as- tyanax apparently are not “on the verge of complete genealogical separation.”’ In fact, Platt and Brower (1968) have suggested that their relationship is one of primary inter- gradation, in which the two parental but- terflies never have been geographically sep- arated from each other, rather than one of secondary intergradation, following isola- tion, as Fisher’s hypothesis assumes. A more simple and straightforward mod- el is that the differences between arthemis and astyanax are the result of natural se- lection, which factors disruptive coloration (wing banding) in the north, and mimicry (without wing banding) of Battus philenor in the south. The northeastern blend zone, then, can be viewed as a steep cline of se- lective reversal between these two forces, with the frequencies of the morphs (and al- leles) of the white band (WB) adjusting themselves to the relative strengths of the two forces at any one locality, thus effec- tively neutralizing the “net” selection across the blend zone from north to south. This intergrade zone then may be considered to be a region of “relaxed” selection, thus, ac- counting for each local population being in Hardy-Weinberg equilibrium. It is, in es- VOLUME 89, NUMBER 4 sence, a valley (or trough) between two adaptive peaks. A detailed examination of this phenomenon, as it relates to the inter- grade population at Green Ridge State For- est in western Maryland follows. METHODS AND MATERIALS The Hardy-Weinberg (H-W) Principle. — This principle is based on a number of as- sumptions almost all of which do not hold precisely true when applied to natural pop- ulations of organisms. Thus, the H-W prin- ciple often is used as an hypothetical “‘yard- stick” against which to assess mating systems in natural populations, to determine just how much they deviate from theoretical H-W predictions as a result of selective forces. Among these assumptions are the follow- ing: 1) a single locus having two alleles con- trols the trait (or traits) in question; 2) ran- dom mating (panmixia) occurs between the sexes in the population; 3) no migration or immigration is occurring in the population; 4) a constant environment is assumed; and, finally, 5) no “‘net”’ selection is taking place. All of these assumptions hold only approx- imately at best, when applied to the L. ar- themis-astyanax butterflies in nature. Procedures for the H-W analyses.—The first step necessary for carrying out an H-W analysis 1s to obtain a quantitative “‘ran- dom” sample (or samples) of butterflies from one of the blending populations. This means that all insects encountered must be col- lected, regardless of phenotype, wing con- dition, or sex. Equal effort must be expend- ed to collect each and every individual (no matter how common or rare), in the order in which they are encountered. Usually about 70% of the Limenitis seen can be tak- en with hand nets. Such samples most often are biased in favor of males, which, unlike females, tend to be territorial, and often sit in roadways for a day or so following eclo- sion, presumably to obtain nutrients and salts. Samples of between 30 to 100+ in- dividuals should be collected in as brief a time period as possible. Each sample should 637 be obtained from as small a localized area as possible, so that it will be representative of a single deme, or localized interbreeding group. Next the sample must be pinned and fully labeled. Each specimen is catalogued by specimen number and its complete phe- notype (of four independently inherited wing color pattern characters) is recorded (see re- sults section). The following steps are done to perform the H-W analysis: 1) Calculation of the allelic frequencies for the major gene controlling wing banding. This genetic locus will be termed the ‘‘B”’ (for ““banding’’) lo- cus. We assume that it possesses two alter- native alleles, having incomplete domi- nance: the B, allele codes for wing banding and the B, allele codes for the unbanded condition. The H-W principle algebraically states that p + q = 1, and, hence, that (p + q)’ = 1, as well. In this case, p equals the frequency of the ““dominant” allele (e.g. B,) and q equals the frequency of the “‘reces- sive”’ allele (e.g. B,); 2) Expansion of the H-W binomial: If (p + q) = 1 (by definition), then p? + 2pq + q°’ = 1, also. Here p’ represents the pheno- typic frequency of the “dominant”? homo- zygotes (e.g. B,B,, or unbanded astyanax) in the population, 2pq equals the pheno- typic frequency of the heterozygotes (B,B,, or partially banded proserpina), and q? equals the phenotypic frequency of the “‘re- cessive” homozygotes (B,B, or banded ar- themis, including form albofasciata); 3) Next these three phenotypic frequen- cies are used to generate the expected num- bers of butterflies of each phenotype to be found in a sample of known size, by mul- tiplying them by the total number of insects collected in the original sample. [In so doing, one statistical degree of freedom (df) is used up, leaving only a single degree of freedom for the chi-square analysis which follows (Srb and Owen, 1952)]. 4) Finally, the observed and expected (based on the above H-W assumptions) numbers of butterflies belonging to each 638 Phenotypic and genetic relationships between the four major forms of the L. arthemis-astyanax complex. Table 1. astyanax (Fabricius) proserpina (Edwards) albofasciata (Newcomb) arthemis (Drury) Morphs 3, 4,5 1 Relative phenotypic categories (Platt & Brower, 1968) Wing banding phenotypes Absent Partial (visible dorsally, Full, narrow Full, wide to trace beneath forewings only) B,B,,—,— PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON BB = B,B,,m,m,,m,m, B,B,,M,—,M,— Genotypes (incompletely dominant (heterozygote) homozygote) (double, or triple recessive homozygote) (incompletely recessive homozygote, modifier loci dominant) Note: The phenotypic relationships are as follows: albofasciata (B,B,,M,—,M,—) (B,B,,m,m,,m,m,) > arthemis > (B,B,) astyanax > proserpina (B,B,) phenotypic class are compared using chi- square analysis. If the two groups of num- bers closely approximate one-another, a low chi-square value will result, leading to the inference that they do not differ significantly from one another. If, on the other hand, the observed and expected numbers are dis- parate, a large chi-square value will result. Iflarge enough (chi-square, = 3.84), then the null hypothesis (H,) that the observed and expected numbers do not differ significantly must be rejected at the 5% level of confi- dence. Then, an alternative hypothesis (H,), e.g. that the numbers do differ significantly, will be chosen. This latter result 1s the one to be expected if the heterozygous pheno- type (proserpina) 1s truly rare in natural pop- ulations of the L. arthemis-astyanax but- terflies. See Speiss (1977) for further in-depth considerations of these procedures. RESULTS Genetic relationships of the forms.—The genetic relationships of the four common forms of the L. arthemis-astyanax complex are shown in Table 1. White wing banding (WB) is controlled by a single major auto- somal gene having two alleles, banded (B,) and unbanded (B,). The alleles exhibit in- complete dominance, in and of themselves. However, the astyanax phenotype quite likely is evolving dominance (Clarke and Sheppard, 1960; O’Donald and Barrett, 1973; Ford, 1975) in regions where the un- banded morph prevails, through the accu- mulated action of unlinked modifier alleles at separate autosomal loci. The penetrance of the partial white banding in the hetero- zygous form proserpina (B,B,) 1s less on av- erage in the southern areas of the blend zone than it is further north, in the central regions of the zone. Earlier breeding studies with insect strains from Shutesbury (Franklin Co.), Massachusetts (see Figs. 1-4) dem- onstrated that form albofasciata bred true and was recessive to form arthemis, as well (Platt, 1983). The differences between the two forms resulted from the action of two VOLUME 89, NUMBER 4 separate unlinked autosomal modifier loci (M, and M.,, respectively). However, recent breeding experiments using insects from the Green Ridge, Maryland population, in which form albofasciata was recovered from heterozygous (“‘trace”’ and partially banded) parents (1984-85 unpublished data), sug- gest that only a single modifier locus is seg- regating in that strain. Perhaps the other modifying locus has become locally fixed, so that it no longer segregates. This has hap- pened either just in my laboratory strains alone, or in the entire western Maryland population itself. Further study will be re- quired to resolve these two possibilities. THE INTERGRADING POPULATION AT GREEN RIDGE STATE FOREST, MARYLAND Through verbal and written correspon- dence with several members of the Mary- land Entomological Society (M.E.S.) I have located 21 records of banded admirals col- lected in Green Ridge State Forest (Allegany Co.), Flintstone Township, Maryland and the immediate surrounding areas. Most of these insects were collected along Fifteen Mile Creek Road near bridges and other areas where the adjacent stream and drain- age ditches adjoin the gravel roadway. Sim- mons and Andersen (1961) reported the first banded admiral from Maryland. A speci- men in the Carnegie Museum of Natural History in Pittsburgh, taken by the late Har- ry Clench, was collected in 1970 (J. E. Raw- lins, pers. comm.). between 1981-1984 at least 19 other specimens have been taken, a number of them by several M.E:S. collec- tors, including E. Cohen, J. Fales, W. Grooms, P. Kean, and J. Zeligs. All of the specimens were collected between 10-30 June, except for one in early August. The specimens include 20 males and one wide banded female, as well as 11 arthemis (wide banded) and 10 a/bofasciata (a near 1:1 ra- tio). The chronological collection dates of these specimens are as follows: 1) VI.30.60, 2) VI.19.70, 3-5) VI.14, 15, and VHI.6.81, 6- 639 14) VIUSs 16:(2)) 175,.2013)223,cand'30.82, 15-18) VI.10, 15, 19, and 26.83, and 19- 21) VI.20 (3).84. These 21 specimens have been phenotypically classified for three oth- er wing color characters (in addition to wing banding), using the six relative category val- ues defined by Platt and Brower (1968), and Platt (1975). These are given below: 1) Dorsal forewing marginal red-orange spotting (FWR): 2) dorsal hindwing marginal red-orange spotting (HWR); 3) dorsal hindwing blue-green iridescence (IB). For FWR, HWR, and wing banding (WB), the six relative phenotypic categories are: Category | = full expression (large, or wide); Category 2 = full expression (small, or nar- row); Category 3 = partial expression (greater, or dorsal); Category 4 = partial expression (lesser, or ventral only); Category 5 = trace expression (remnant only); and Category 6 = expression lacking For IB the relative expression among the six phenotypic categories 1s exactly the reverse of those given for the three traits above. The xX + SE values of these four traits among the 21 wild-collected banded insects from Maryland are: FWR: 5.1 + 0.4, HWR: 5.0 + 0.3 1B 3:3-2210)2 and W Beil osee Onlk All three of these other traits are con- trolled by autosomal polygenic loci which are not correlated either to white banding, or to each other (Platt, 1975). However, the dorsal hindwing bands of one unusual ex- ample of albofasciata taken by W. Grooms and J. Zeligs on 23 June, 1982, are markedly suffused with iridescent blue scales, closely resembling partially banded form ‘“‘cerulea” (Ehrmann, 1900) which exhibits this same condition ventrally. In addition to these collected specimens, I am aware of four additional “‘sight”’ rec- 640 Table 2. Observed and expected numbers (the lat- ter in parentheses) of butterflies in consecutive annual samples obtained from Green Ridge State Forest, Maryland. A) observed and expected numbers in an- nual samples. B) allelic frequencies and results of H-W analyses. C) G statistics: R x C tests of independence [methods of Sokal & Rohlf (1981: 745)]. A) Observed and expected numbers in the annual samples: Banded (arthemis; Partially Annual albo- Banded Unbanded Sample fasciata) (proserpina) (astyanax) Totals 1982 72 33 2229 264 (3 days) (1.30) (34.37) (228.33) 1983 l 19 62 82 (1 day) G3) (18.29) (62.4) 1984 3 32 205 240 (6 days) (1.44) (35.04) (203.52) Totals 6 84 496 586 (10 days) (4.10) (87.90) (494.00) B) Allelic frequencies and results of the H-W analyses: Allelic Frequencies Chi Annual Unbanded _ Banded scala Significance Sample (p) (q) (df = 1) (P = 0.33) 1982 .930 .070 0.43 0.52 (NS) 1983 .872 .128 0.10 0.75 (NS) 1984 921 .079 1.97 0.18 (NS) Overall 918 .082 1.06 0.33 (NS) C) G tests: G. Significance Test No. & Variables Value df (P = ) 1) 3 x 3 contingency test 2934 (3 phenotypes vs. 3 0.57 (NS) annual samples) 2) 2 x 3 contingency test Osea (observed vs. expected 0.71 (NS) H-W values in overall sample) NS = Not significant. ords of banded admirals in Maryland. Three of these were made by A. Platt and P. Kean, while collecting at Green Ridge [VI.2.82, and VI.23.83(2)]. The fourth was made near Cunningham Falls (Frederick Co.) in June, 1982 by W. Cooper. This latter record sug- gests that banded admirals may occur else- where in the state as well. These rare banded admirals always occur in company with large numbers of unband- ed astyanax, and the somewhat less com- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON mon partially banded individuals referrable to proserpina (the heterozygous form). The majority of the latter in Maryland have a mere trace of the partial white band. These are referrable to WB phenotypic category 5 of the Platt and Brower (1968) system. That these “trace” banded insects are true het- erozygotes (genotype B,B,) 1s attested to by the fact that I have been able to recover banded forms arthemis and albofasciata from them during selective laboratory breeding studies carried out at UMBC dur- ing 1984-85. (The results of these recent studies will be reported in a future paper.) Other proserpina specimens possess clear ventral expression of the partial band (WB category 4 individuals), but only a very few insects exhibit partial banding dorsally (WB category 3). However, a fine example of this latter phenotype was collected by P. J. Kean on VII.3.83 at Green Ridge. These various forms of proserpina are found (uncommon- ly) flying with astyanax throughout the state. Examples in my collection (and those of other M.E.S. members) come from Alle- gany, Frederick, Montgomery, Calvert, Carroll, Baltimore, and Dorchester coun- ties, but proserpina 1s probably even more widespread in Maryland than these records indicate. H-W analyses of the Green Ridge popu- lation.—Results of H-W analyses per- formed on the Green Ridge samples ob- tained between 1982 and 1984 are given in Table 2. The L. arthemis-astyanax butter- flies were exceedingly common during 1982 and 1983. It was during these years that 13 of the 21 banded specimens were collected. Table 2A lists the observed and H-W ex- pected numbers in each annual sample, and among the total of 586 butterflies captured. The allelic frequencies and the chi-square test results (2B), clearly indicate that the observed and expected numbers do not dif- fer significantly from one another. > x?, = 0.18 in all cases, with the overall > x7,, value being 1.06, with P = 0.33). Since the banded butterflies are so rare in VOLUME 89, NUMBER 4 the western Maryland population, their ex- pected numbers in the H-W analyses are below five. This means that there is a high likelihood of the observed and expected numbers deviating from one another simply by chance alone. However, the data them- selves show that this has not occurred. Fur- thermore, two G tests of independence have been run on these data (Table 2C). Test 1 shows that the three annual samples are alike, and so can be pooled. Test 2 indicates that the observed and expected numbers are very close approximations of one another. Neither of these two tests yielded significant G values, with P = 0.57 in each test. The G tests are independent of the numbers in each subcategory, yet their results support the findings of the above chi-square anal- yses. Thus, I conclude that the L. arthemis- astyanax butterflies are breeding at random (and hence exhibit panmixia) at Green Ridge, just as they were at all of the New England localities studied earlier by Platt and Brower (1968), and at other more northern localities within the blend zone (Platt, 1983). Among the 84 proserpina collected at Green Ridge between 1982 and 1984 were four WB category 3 (5%), 28 WB category 4 (33%), and 52 category 5 (62%). The H-W analyses indicate that there is no statistical deficiency of these partially banded hetero- zygotes. Thus, they form a stable part of the astyanax populations along the southern margin of the L. arthemis-astyanax blend zone, and the banded alleles are commonly carried by individuals closely approaching the astyanax phenotype. Assuming panmixia and utilizing the ge- notypic frequencies of the three forms [ar- themis (including albofasciata = 0.007), proserpina = 0.150, and astyanax = 0.843], it is possible to calculate the frequency with which matings are expected to occur (by chance) between the various morphs. Of all possible matings, 71.0% should occur be- tween unbanded (astyanax) individuals. Another 25.3% should take place between 641 partially banded and unbanded insects. Only 2.3% will involve two heterozygotes (pro- serpina), and another 1.2% would be be- tween banded and unbanded individuals (yielding all partially banded progeny). Be- cause of the low genotypic frequencies, only one in every 500 matings will involve a banded x partially banded cross (0.20%), and the banded individuals are so rare that only one in 20,000 matings should involve two banded individuals (0.05 x 10~ 3%). The rare banded individuals occurring at Green Ridge mainly result from crosses between partially banded heterozygotes (form pro- serpina). Such crosses are expected to yield approximately 25% banded individuals. Selection and bird predation.— When quantitative samples of L. arthemis-astya- nax butterflies are obtained, certain indi- viduals possess large, jagged chunks missing from their wings. The wing condition of many of these insects is otherwise fresh. Such damage is assumed to represent “beak tears” caused by insectivorous birds chasing and attempting to catch the live butterflies (Bowers and Wiernasz, 1979). Similar dam- age has been found on wings of 47 arthemis- astyanax butterflies from western Mary- land, which were killed and eaten presum- ably by a male gray catbird (Mimidae: Du- metella carolinensis) at Green Ridge. Beak tears result from a butterfly (or moth) strug- gling and escaping from a bird that has caught it, and is holding one or more wings in its bill (Sargent, 1973, 1976; Robbins, 1980). On the other hand, “beak marks” are crisp impressions of the outlines of bird beaks, which result from a bird biting and then gaping, thus, releasing the insect (Ed- monds, 1974). Beak marks seldom are found on wings of admiral butterflies (except on wings of those that have been eaten, referred to above). They are more likely to be found on wings of unpalatable species, such as model butterflies (Danaus, Heliconius, etc.), which birds attack and catch, but often fail to kill or eat (Brower, 1984). The frequency of beak tears among L. 642 Table 3. ‘Beak tears” among wild-collected Li- menitis arthemis-astyanax butterflies in the UMBC and Milwaukee Public Museum collections— 1982 data. No. and Percent of Individuals with “Beak Tears” Sample Locality and Form Size So. Quebec and No. Vermont arthemis 263 = 81 : (30.8%) Central Mass. intergrades arthemis 155 =10 (6.5%) proserpina 376 =. 228 (7.5%) astyanax 351 34 (9.7%) Subtotals 882 72 (8.2%) Central Wisconsin proserpina (M.P.M.) 76 Southern astyanax (CT RIENJ; VA; W.MD, IL) 16 (21.1%) 373) %42(11E3%) arthemis-astyanax butterflies in the UMBC collection, and in a sample of proserpina borrowed from the Milwaukee Public Mu- seum, is given in Table 3. The Vermont, Massachusetts, and southern (astyanax) samples have been collected “at random” by me during June and early July. They are comparable in terms of butterfly condition and age. I assume that the beak tear fre- quencies shown in the table reflect differ- ential predator intensity between localities. These analyses reveal that about 30% of the disruptively banded northern arthemis but- terflies exhibit beak tears, whereas only about 11% of the mimetic unbanded south- ern astyanax have similar damage. In cen- tral Massachusetts (within the central region of the zone of intergradation) only 8.2% of 882 butterflies examined exhibit such dam- age, and the frequency of this damage is not significantly different between the three in- Sectforms (x2, — 1-812 P= 033) Incen- tral Wisconsin, 21.1% of a sample of form proserpina (n = 76) showed beak tears. Among 234 beak tears on the wings of L. arthemis-astyanax butterflies analyzed (all samples pooled), 68% were on the hind- wings (Type I damage), 27% were on the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON forewings (Type II damage), and only 5% of the beak tears involved both fore- and hindwings (Type III damage) together. This system of categorization 1s modified from one used by (Sargent 1976) for noctuid moths (Catocala spp.). Some individuals in Table 3 contained several separate beak tears. Attacks occurred from either the left or right sides with equal frequency. My interpretations of the beak tear data are based upon the assumption that the fre- quency of beak tears on the insect wings reflects true attack rates of birds on the but- terflies themselves. If this is true, then the differences in beak tear damage shown by the samples in Table 3 represent actual dif- ferences in the attack rates by birds on the butterflies in the different geographic re- gions. What such damage is measuring, in reality, are successful butterfly escapes from bird attacks. The missing information is just how many butterflies of each form are caught and eaten by birds. Some light can be shed on this problem by examination of wing fragments collected by P. J. Kean and myself at Green Ridge on VI.20 and VI.23, 1982. Separate wings of 13 males (four proserpina, eight astya- nax) were collected from a single locality on the first day. A male gray catbird was flitting about the adjacent shrubbery, near where a large ““mud-puddle club” of male butterflies had aggregated at the edge of the roadway. Crisp beak marks, sharp and V-shaped, as well as numerous beak tears, were found on the insect wing fragments. The outline of the beak marks, including a sharp pin-like hole at the tip of the “V,” matched the outline of a catbird’s beak per- fectly. Three days later, wing fragments of 33 additional insects were recovered at the same area, and the male catbird still was very much in evidence. The bird apparently was feeding heavily on a locally abundant and easily obtainable food supply, and was consuming an average of 11 butterflies per day. The butterflies evidently were being caught while sitting on the gravel roadway, VOLUME 89, NUMBER 4 where they could easily be closely ap- proached, even by humans. Two years later, VI.20.1984, wing fragments of two addi- tional butterflies, one an astyanax and the other a banded form albofasciata, were col- lected at precisely the same locality. This evidence indicates that birds at times will prey on the adult admiral butterflies, es- pecially when they are common, and it sug- gests that the damage termed “beak tears” is, in fact, caused by birds attacking the live butterflies. Such attacks can occur either when a butterfly is perching or sitting on a roadway (as above), or when the insect is flying. The author has witnessed three un- successful bird pursuits of flying arthemis butterflies over the years. Two occurred in northern Vermont, and involved a gray cat- bird and a barn swallow (Hirundinidae: H- rundo rustica). The other occurred in central Massachusetts, and involved a blue jay (Corvidae: Cyanocitta cristata). DISCUSSION AND CONCLUSIONS The evidence taken in its entirety sup- ports the contention that the L. arthemis- astyanax zone of intergradation 1s main- tained by the counter-balancing selective forces of disruptive coloration and mimic- ry, as postulated by Platt and Brower (1968). The driving forces behind the evolution of both the banded and unbanded morphs is very likely differential predation by birds. Throughout the blend zone selection is “‘re- laxed,”’ since these two opposing forces are in balance with one another. The banded and unbanded allelic frequencies adjust themselves to the relative strengths of the two forces at any one locality, resulting in a situation of no “net” selection. The but- terflies clearly are breeding at random with one another throughout the blend zone. Both males and females apparently totally dis- regard wing banding phenotype in choosing their mates. Because no “net” selection oc- curs across the zone of intergradation, local populations remain in Hardy-Weinberg equilibrium. The entire complex, therefore, 643 must be regarded as a single polymorphic, biological species. North of the blend zone the banded arthemis form prevails, where- as, southward, where the model species is abundant, unbanded mimetic astyanax are favored. As the unbanded morph evolved, the banded alleles have become “‘masked,”’ or covered up, by the action of unlinked modifier loci. That the alleles have not been “lost” from the astyanax populations is im- portant. Within the blend zone, the partially banded form proserpina more or less “‘strad- dles the fence.’ It provides a genetic link between the two unlike parental morphs. The presence of variable phenotypes within the intergrade populations, possibly serves to confuse bird predators, by keeping them from becoming specialists on any one morph, and perhaps, causing them to seek alternative prey species a higher proportion of the time. ACKNOWLEDGMENTS P. J. Kean of Catonsville, Maryland has provided considerable assistance with the field sampling at Green Ridge. He has se- cured five of the wild-collected banded in- sects himself. Without his many additional contributions, this study could not have been done. I thank also the various Maryland Entomological Society members and others who have shared with me records of banded admirals collected or sighted in Maryland. I thank Tim Ford, graphics illustrator at UMBC for preparing the final copy of the figures. LITERATURE CITED Bergman, W. A. and J. H. Masters. 1971. A study of intraspecific hybridization of Limenitis arthemis in Minnesota. Mid-Continent Lepid. Ser. 2, No. 31: 1-11. Bowers, M. D. and D. C. Wiernasz. 1979. Avian predation on the palatable butterfly, Cercyonis pe- gala. Ecol. Entomol. 4: 205-209. Brower, L. P. 1984. Chemical defense in butterflies. In Vane-Wright, R. I., and P. R. Ackery, eds., Biology of Butterflies, Sympos. No. 11. R. Ento- mol. Soc. Lond. Ch. 12, pp. 109-134. Brower, L. P. and J. V. Z. Brower. 1962. The relative 644 abundance of model and mimic butterflies in nat- ural populations of the Battus philenor mimicry complex. Ecology 43: 154-158. Chermock, R. L. 1950. A generic revision of the Li- menitini of the world. Am. Midl. Nat. 43: 513- 569. Clark, A. H. and L. F. Clark. 1951. The Butterflies of Virginia. Smithsonian Misc. Colls. 116(7). 239 pp. Clarke, C. A. and P. M. Sheppard. 1960. The evo- lution of dominance under disruptive selection. Heredity 14: 163-173. Drury, D. 1773. Nym. Phal. arthemis. Illustrs. Nat. Hist.-Papilio arthemis. White, London. Vol. 2, pl. X, Figs. 3 & 4; index & p. 17. Edmonds, M. 1974. Significance of beak marks on butterfly wings. Oikos 25: 117-118. Edwards, W. H. 1865. Descrption of a new species of Limenitis (L. proserpina). Proc. Entomol. Soc. Phila. 5: 148. 1873. Remarks on Limenitis proserpina and arthemis. Can. Entomol. 5: 232-233. . 1877. Notes on Limenitis proserpina and ar- themis. Can. Entomol. 9: 114-115. . 1884. Butterflies of North America. Privately printed, Vol. 1: 111-147. 1891. On the position of Limenitis proser- pina. Edw. Can. Entomol. 23: 49-56. Ehrmann, G. A. 1900. Limenitis ursula var. cerulea Ehrm. Can. Entomol. 32: 348. Fabricius, J. C. 1775. Papilio astyanax. Syst. Ento- mol., p. 447. Fales, J. H. 1974. Check-list of the skippers and but- terflies of Maryland. Chesapeake Sci. 15(4): 222- 229: Field, W. L. W. 1904. Problems in the genus Basi- larchia. Psyche 11: 1-6. Field, W. L. W. 1910. The offspring of a captured Basilarchia proserpina. Psyche 17: 87-89. Fisher, R. A. 1958. The Genetical Theory of Natural Selection, 2nd revised edition. Dover Publs. Inc., N.Y. 291 pp. Ford, E. B. 1975. Ecological Genetics, 4th edition. Chapman and Hall, Ltd., London. 442 pp. Grey, R. M. 1879. Remarks critical and suggestive, on the genus Limenitis east of the Mississippi. Can. Entomol. 11: 16-17. Gunder, J.D. 1934. Achecklist revision of the Genus Basilarchia Scud. (Lepid: Rhopalocera). Can. Entomol. 66: 39-48. Hovanitz, W. 1949. Increased variability in popu- lations following natural hybridization, pp. 339- 355. In Jepsen, G. L., E. Mayr, and G. G. Simpson, eds., Genetics, Paleontology, and Evolution. Princeton Univ. Princeton. NJ. Klots, A. B. 1951. A Field Guide to the Butterflies. Houghton Mifflin Co., Boston, pp. 115-116. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Mayr, E. 1963. Animal Species and Evolution. Bel- knap Press of Harvard University Press, Cam- bridge, MA. vii and 797 pp. 1970. Populations, Species, and Evolution: An Abridgement of Animal Species and Evolu- tion. Belknap Press of Harvard Univ. Press, Cam- bridge, MA. 453 pp. Nakahara, W. 1924. A revision of the genus Basi- larchia (Rhopalocera: Nymphalidae). Bull. Brook- lyn Entomol. Soc. 19: 166-180. Newcomb, H. H. 1907. Description of a new variety of Limenitis ursula. Psyche 14: 89-91. O’Donald, P. and J. A. Barrett. 1973. Evolution of dominance in polymorphic Batesian mimicry. Theor. Pop. Biol. 4: 173-192. Platt, A. P. 1975. Monomorphic mimicry in nearctic Limenitis butterflies: experimental hybridization of the L. arthemis-astyanax complex with L. ar- chippus. Evolution 29: 120-141. . 1983. Evolution of North American admiral butterflies (Limenitis: Nymphalidae) Bull. Ento- mol. Soc. Am. 29:10-22. Platt, A. P. and L. P. Brower. 1968. Mimetic versus disruptive coloration in intergrading populations of Limenitis arthemis and astyanax butterflies. Evolution 22: 699-718. Platt, A. P., R. P. Coppinger, and L. P. Brower. 1971. Demonstration of the selective advantage of mi- metic Limenitis butterflies presented to caged avi- an predators. Evolution 25: 692-701. Platt, A. P., S. D. Frearson, and P. N. Graves. 1970. Statistical comparisons of valval structure within and between populations of North American Li- menitis (Nymphalidae). Can. Entomol. 102: 513- 593) Poulton, E. B. 1909. Mimicry in butterflies of North America. Ann. Entomol. Soc. Am. 2: 203-242. Remington, C. L. 1958. Genetics of populations of Lepidoptera. Proc. Tenth Internat. Congr. Ento- mol. 2: 787-805. 1968. Suture-zones of hybrid interaction be- tween recently joined biotas. Evol. Biol. 2: 321- 428. Robbins, R. K. 1980. The Lycaenid “false head” hypothesis: Historical review and quantitative analysis. J. Lepid. Soc. 34: 194-208. Sargent, T. D. 1973. Studies on the Catocala (Noc- tuidae) of southern New England. IV. A prelimi- nary analysis of beak-damaged specimens, with discussion of anomaly as a potential anti-predator function of hindwing diversity. J. Lepid. Soc. 27: 175-192. 1976. Legion of Night—The Underwing Moths. Univ. Mass. Press, Amherst, MA. 222 pp. Saunders, A. A. 1932. Butterflies of the Allegany State Park. N.Y. State Mus. Handbook 13. University. State of N.Y. 270 pp. VOLUME 89, NUMBER 4 Scudder, S. H. 1889. The Butterflies of the Eastern United States and Canada, with special reference to New England. Publ. by the author, Cambridge, MA. Vol. 1: 250-305. Simmons, R. S. and W. A. Andersen. 1961. Notes on five new butterfly records for the state of Mary- land. J. Lepid. Soc. 15: 99-101. Smith, J.B. 1891. Limenitis arthemis, etc. Can. Ento- mol. 23: 104-105. 645 Spiess, E. B. 1977. Genes in Populations. John Wiley and Sons, N.Y. 780 pp. Sokal, R. R. and F. J. Rohlf. 1981. Biometry—the Principles and Practice of Statistics in Biological Research. W. H. Freeman and Co., N.Y. 859 pp. Srb, A. M. and R. D. Owen. 1952. General Genetics. W. H. Freeman Co., San Francisco. 561 pp. PROC. ENTOMOL. SOC. WASH. 89(4), 1987, p. 645 NOTE New records of Brachydeutera (Diptera: Ephydridae) in Malaysia During a recent trip to Malaysia I was able to collect several small series of Brachy- deutera from throughout the country. As the genus was recently revised for the Oriental Region (Mathis, W. N. and K. D. Ghor- padé. 1985. Smiths. Contr. Zool. 406: 1- 25) it is appropriate to add some additional distributional information. Three species were encountered with B. pleuralis Malloch representing a new record for Malaysia. Brachydeutera hardyi Wirth was found both in peninsular and East Malaysia, but in low numbers. In Sabah State (2 44, 6 99; Tawau, pond shoreline, 6 Nov 1986) it was collected from the surface of a shallow drainage pool which was subject to periods of extreme flooding. The pool had a grass base. In Perak State (MARDI-Hilir Perak, 16 mi W Telok Anson, 26 Nov 1986) only two females were obtained from the surface ofa small, rain-fed drainage ditch. This ditch had a mud and sand base with little emer- gent vegetation. Brachydeutera hardyi shared this habitat with B. /ongipes Hendel. Brachydeutera longipes Hendel was col- lected in Perak State (1 4, 1 2; same as above except 25 Nov) and in large numbers in Melaka State (46 66, 83 99; Kg. Gadek, nr Alor Gajah, 19 Nov 1986). The Melaka habitat consisted of numerous small, rain- fed depressions which were 3-10 cm in depth. The pools had a grass base and also were susceptible to periods of drying and flooding. Brachydeutera pleuralis Malloch repre- sents a new record for Malaysia but was collected only in Kelantan State (13 36, 22 22; Bachok, shallow pool, 16 Nov 1986). This species was taken from the surface of numerous sandy, shallow pools which were several hundred meters from the coastline. The pools were 2-30 cm deep and contained no macro-vegetation. There was little mac- ro-vegetation surrounding the pools that were bordered on both sides by large dunes. The water showed no signs of salinity. I thank Azhar Ismail and Yahaya Hussain for serving as my hosts and guides during my trip to Malaysia. This paper has been assigned number 7708, Agricultural Re- search Center, College of Agriculture and Home Economics, Washington State Uni- versity, Pullman, Washington 99164. Work conducted under projects 9043 and 9902. Richard S. Zack, Department of Ento- mology, Washington State University, Pull- man, Washington 99164-6432. PROC. ENTOMOL. SOC. WASH. 89(4), 1987, pp. 646-659 AMBLYOMMA BABIRUSSAE SCHULZE (ACARI: IXODIDAE): REDESCRIPTION OF THE MALE, FEMALE, AND NYMPH AND DESCRIPTION OF THE LARVA JAMES E. KEIRANS AND RICHARD G. ROBBINS Department of Health and Human Services, Public Health Service, National Institutes of Health, National Institute of Allergy and Infectious Diseases, Department of Ento- mology, Museum Support Center, Smithsonian Institution, Washington, D.C. 20560. Abstract.—The male, female and nymph of Amblyomma babirussae Schulze are rede- scribed and the larva is described for the first time. Adults and immatures are illustrated with black and white and color drawings and with scanning electron photomicrographs of specimens collected primarily on artiodactyl mammals and from vegetation on the island of Sulawesi, Indonesia. Amblyomma babirussae was originally described from two collections. The first contained 3 males, 7 females and 2 nymphs from Sus babirussa, “Vaterland Boeroe oder Celebes.” This might be the island of Boeroe (03.24S, 126.40E) or some locality un- known to us but named Boeroe on present- day Sulawesi. The second collection con- tained 3 nymphs from Minahassa, northern Celebes, collected by O. Warburg. The only other taxonomic study of 4. babirussae was by Anastos (1950) who saw 15 males, 9 fe- males, 18 nymphs and | larva in three col- lections, all from Sulawesi. When the Hoogstraal tick collection was transferred from Cairo, Egypt, to the Mu- seum Support Center, Smithsonian Insti- tution, in November of 1986, we found 95 collections of A. babirussae, all from Sula- wesl. Most of these specimens were col- lected by members of U.S. Naval Medical Research Unit No. 2, Jakarta Detachment, especially W. P. Carney, J. F. Duncan, S. W. Joseph and P. F. D. Van Peenen. We thank these collectors for donating their ma- terial to Dr. Hoogstraal’s tick collection. A. babirussae remains an unfamiliar species to most tick workers. For this rea- son, we have decided to redescribe in detail the male, female and nymph and to describe the hitherto unknown larva. In the follow- ing account, all measurements are in mil- limeters. A range is given followed by a mean in parentheses. Ticks were prepared for scanning electron microscopy by the meth- od of Corwin et al. (1979). Amblyomma babirussae Schulze 1933 Amblyomma babirussae Schulze 1932: 525 [nomen nudum]. Amblyomma babirussae Schulze 1933: 317, figs. 1-5, original description. Amblyomma cyprium Kriygsman & Ponto 19312 140 143219322 1222 igs sa. 26 [nec Neumann, 1899]. Amblyomma cyprium Toumanoff 1944: 110, pl. 78, figs. c, d (6 only, not 2) [nec Neumann, 1899]. Male (Figs. 1—5).—Length from scapular apices to posterior body margin 2.88-3.52 (3.13); width 2.38—3.00 (2.72), widest at level VOLUME 89, NUMBER 4 647 Fig le of spiracular plates, outline broadly oval. Scutum (Fig. 1) ornate, typically with a large iridescent reddish-orange spot encircled with a band of yellow and green located at the posterior angle of the pseudoscutum. The remaining ornamentation is found periph- erally, ranging from an almost continuous orange/yellow stripe encircling the scutum, including spots on all festoons, to almost no ornamentation except the central spot. Or- namentation in Fig. | is typical except most males lack the slight coloration on the dor- sum of the basis. Punctations large and deep Amblyomma babirussae 6 SULAWESI (RML 99981). Dorsal view. marginally, a few slight, shallow puncta- tions centrally. Scutum with roughened ap- pearance due to raised pseudoscutal area and central extension toward festoon six. Marginal groove absent. Cervical grooves narrow, extending laterad from emargina- tion to short cervical pits. Eyes small, flat, level with scutal surface. Capitulum (Figs. 2, 3): Length from pal- pal apices to cornua apices 0.755—-0.952 (0.851), width 0.465-0.635 (0.535); basis capituli dorsally subrectangular, ornamen- tation found on one of ten specimens; pos- 648 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON f . 3 ie at UG bela ; ai Figs. 2-5. Amblyomma babirussae 6 SULAWESI (RML 118497). 2, Capitulum, dorsal view (95 x). 3, Ca- pitulum, ventral view (106 x). 4, Coxae I-IV (74x). 5, Spiracular plate (212 x). terior margin straight between short, broad- ly triangular cornua. Palpi length 0.332- 0.621 (0.521), width 0.189-0.247 (0.214); average lengths of segments II and III, 0.292 and 0.202 respectively. Palpal segment II dorsally with a narrow posterointernal ridge; suture between segments II and III distinct. Hypostome (Fig. 3) length 0.382-0.534 (0.468) including teeth, crenulations, and stalk to base, bluntly rounded apically; den- tal formula 4/4, teeth of files 1-4 number 6-7 (6.40), 6-8 (6.40), 5-7 (6.20) and 5-8 (6.50) respectively; apex with a large corona of fine denticles. Legs (Fig. 4) moderate in length, tro- chanter, femur and tibia often with a green- ish-blue ring at segment’s distal end. Coxa I with a moderately long, broadly rounded VOLUME 89, NUMBER 4 internal spur and a long, lanceolate external spur; coxa II with a very small internal pro- tuberance, arising anterior to but not reach- ing the posterior margin of coxa II, and an elongate narrowly triangular external spur; coxae III and IV each with an external spur similar to that on coxa II, internal spurs absent. Trochanters without spurs. Tarsus I abruptly tapered distally, 0.680-0.858 (0.763) long, 0.191-0.253 (0.222) wide. Tarsus IV gradually tapered distally, 0.5 13- 0.691 (0.609) long, 0.144-0.194 (0.171) wide. Genital aperture (Fig. 4) at level of coxae II. Spiracular plate (Fig. 5) with a narrowly elongate dorsal prolongation. Descriptive statistics for principal char- acters of males are summarized in Table 1. Female (Figs. 6—10).—Lerigth (unfed, ex- clusive of capitulum) 3.16—-3.93 (3.64), width 2.62-3.30 (3.01). Scutum (Fig. 6) length 1.55-1.91 (1.76), width 1.91-2.25 (2.10), outline as illustrated; scapulae large, bluntly rounded. Ornamentation variable but al- most always a reddish-orange spot encircled by a narrow yellow band and an outer green- ish band located in the posterior scutal an- gle. This spot is of the same color and in the same location as that of the male (Fig. 1). Diffuse ornamentation of the same red- dish-yellow color in cervical area from scap- 649 Fig. 6. Amblyomma babirussae 2? SULAWESI (RML 99967). Scutum. ula to eye, background color brown. Eyes flat, at lateral angles. Cervical grooves nar- rowly elongate, comma-shaped, usually containing punctations. Punctations large and numerous Over scutum except centrally Table 1. Amblyomma babirussae: Descriptive statistics for measurements from 10 unengorged males. Measurement Mean with Standard Error Coefficient of Standard Deviation Variation Body: length 3213) 10107 0.22 7.03 width 2.72 + 0.06 0.20 35 Capitulum: length OMS TEE OL017 0.054 6.35 width 05355-01015 0.048 8.97 Palps: length 0521. 01025 0.078 14.97 width 0.214 + 0.006 0.020 Ss) Hypostome: total length 0.468 + 0.015 0.047 10.04 Tarsus I: length 0.763 + 0.021 0.065 8.52 width 0.222 01007 0.021 9.46 Tarsus IV: length 0.609 + 0.018 0.058 9.52 width 0.171 + 0.005 0.015 8.77 650 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 7-10. Amblyomma babirussae 2? SULAWESI (RML 99967). 7, Capitulum, dorsal view (64 x). 8, Ca- pitulum, ventral view (74 x). 9, Genital aperture (318 x). 10, Coxae I-IV (64 x). and posterolaterally. Dorsum of body with numerous large white setae; marginal groove absent. Venter with fewer, smaller white setae than on dorsum. Genital aperture (Figs. 9, 10) U-shaped, without thickened areas or flanges, situated medially at posterior mar- gin of coxae II or between coxae II and III. Spiracular plates essentially as in male. Capitulum (Figs. 7, 8) length from palpal apices to posterior margin of basis capituli 1.09-1.37 (1.27), width 0.75 1-0.892 (0.837); porose areas deep, elongate in anterior—pos- terior plane, diameter of one area 0.090- 0.124 (0.111); interporose area 0.245-0.379 (0.278); 4 of 10 specimens with slight or- namentation in the interporose area; exter- nal margin of basis broadly convex; cornua VOLUME 89, NUMBER 4 absent or indicated by slight protuberances. Ventrally, basis as figured (Fig. 8) with sev- eral minute white setae at each posterolater- al angle. Palpi length 0.724—-0.980 (0.823), width 0.216-0.287 (0.258); average lengths of segments II and III, 0.494 and 0.271 re- spectively; palpal segment II dorsally with an anteriorly broadening mesal ridge. Hy- postome (Fig. 8) total length 0.732-0.832 (0.781); dental formula 4/4; teeth of files 1- 4 number 7-9 (8.00), 7-9 (7.67), 6-9 (7.67), 6-9 (7.78) respectively; apex indented, with a large corona of fine denticles. Legs (Fig. 10) moderate in length, tro- chanter, femur and tibia often with green- ish-blue markings at segment’s distal end. Spurring of coxae I-IV as in 6 except exter- nal spurs of II-IV are broadly rounded rath- er than pointed. Some 2 specimens lack the very small internal protuberance on coxa II. Trochanters without spurs. Tarsus | abrupt- ly tapered distally, 0.83 1—1.060 (0.932) long, 0.212-0.298 (0.260) wide. Tarsus IV grad- ually tapered distally, 0.585-0.780 (0.673) long, 0.157-0.210 (0.189) wide. Descriptive statistics for principal char- acters of females are summarized in Ta- ble 2: 651 Fig. 11. Amblyomma babirussae Nymph SULA- WESI (RML 99968). Scutum. Nymph (Figs. 11-15).—Measurements from 15 unengorged specimens. Body. Length from scapular apices to posterior body margin 1.315-1.562 (1.453); width 1.215-1.426 (1.318), widest at level of coxa Table 2. Amblyomma babirussae: Descriptive statistics for measurements from 10 unengorged females. eee Coefficient of Measurement Mean with Standard Error Standard Deviation Variation Body: length 3.64 + 0.08 0.24 6.59 width 3.01 + 0.06 0.19 6.31 Scutum: length 1.76 + 0.04 OAS 12) width 2 AOE O103 0.11 5.24 Capitulum: length 127 10103 0.10 7.87 width 0.837 + 0.014 0.044 5.26 Palps: length 0.823 + 0.028 0.090 10.94 width 0.258 + 0.006 0.020 Tags Porose area: diameter OTT = 0/003 0.011 9.91 Interporose area 0.278 + 0.013 0.040 14.39 Hypostome: length 0.781 + 0.010 0.030 3.84 Tarsus I: length 0.932 + 0.023 0.074 7.94 width 0.260 + 0.009 0.028 10.77 Tarsus IV: length 0.673 + 0.016 0.051 7.58 width 0.189 + 0.005 0.016 8.47 ee 652 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 12-15. IV; dorsum of idiosoma with 35-50 (41) long, stout setae, concentrated laterally and posteriorly. Scutum (Fig. 11). Length 0.658- 0.786 (0.725), width 0.928-1.083 (1.017); cervical grooves short, comma-shaped, with several large, deep punctations; iridescent golden red/green patches more or less well developed between borders of eyes and cer- vical grooves and at posterior angle of scu- Amblyomma babirussae Nymph SULAWESI (RML 99968). 12, Capitulum, dorsal view (159 x). 13, Capitulum, ventral view (138 x). 14, Coxae I-IV (127). 15, Spiracular plate (424 x). tum; punctations larger and more numerous laterally than mesally, those in posterior ir- idescent patch ranging in number from | to mG): Capitulum (Figs. 12, 13) length from posterior margin of cheliceral sheaths to posterior capitular margin 0.172-0.230 (0.206); width at level of scapulae 0.377- 0.436 (0.399). Basis capituli dorsally (Fig. VOLUME 89, NUMBER 4 653 Table 3. Amblyomma babirussae: Descriptive statistics for measurements from 15 unengorged nymphs. Coefficient of Measurement Mean with Standard Error Standard Deviation Variation Body: length 1.453 + 0.018 0.068 4.68 width 1.318 + 0.015 0.059 4.48 Scutum: length 0.725 + 0.007 0.029 4.00 width 1.017 + 0.009 0.034 3.34 Basis capituli: length 0.206 + 0.004 0.017 8.25 width 0.399 + 0.004 0.016 4.01 Palps: length 0.449 + 0.006 0.025 Dail width 0.089 + 0.002 0.006 6.74 Hypostome: length of toothed portion 0.197 + 0.002 0.007 BE) Tarsus I: length 0.470 + 0.008 0.030 6.38 width ONS 701002 0.008 6.11 12) subquadrate, without ornamentation; cornua present as faint rounded extensions of posterolateral margins; posterior margin between cornua straight. Ventrally (Fig. 13), basis with posterior margin convex but in- dented before joining lateral margins. Palpi ca. 5x longer than wide; 0.406-0.490 (0.449) long, 0.079-0.100 (0.089) wide; seg- ment II ca. 2 as long as segment III; seg- ments decreasing in size in the order II, III, I, IV; setae as illustrated. Hypostome round- ed and notched apically; length of toothed portion (minus crenulations) 0.180—-0.207 (0.197); dentition 3/3 throughout; 7-8 teeth Table 4. Amblyomma babirussae: Descriptive statistics for measurements from 25 unengorged larvae. Measurement Body: length width Scutum: length width Capitulum: length width Palps: length width Hypostome: length of toothed portion Tarsus I: length width Distance between posthypostomal setae Setal lengths Sr SC; PA, PA, MV, MD, MD, CD} CD, Standard Coefficient of Mean with Standard Error Deviation Variation 0:72 ==10!003 0.013 1.80 0.720 + 0.003 0.016 D222 0.291 + 0.001 0.007 2.41 0.563 + 0.003 0.013 2.31 0.342 + 0.002 0.009 2.63 0.215 + 0.001 0.006 2.79 0.205 + 0.001 0.006 2.93 0.052 + 0.000 0.002 3.85 0.098 + 0.001 0.004 4.08 0.252 + 0.001 0.006 2.38 0.089 + 0.000 0.002 Duns 0.049 + 0.001 0.004 8.16 0.026 + 0.000 0.002 7.69 0.015 + 0.000 0.001 6.67 0.025 + 0.000 0.002 8.00 0.027 + 0.000 0.001 3.70 0.032 + 0.001 0.003 9.38 0.029 + 0.001 0.003 10.34 0.028 + 0.001 0.003 HORT 0.021 + 0.000 0.002 9252 0.025 + 0.001 0.003 12.00 : 3 mj it 4 Be, . Bs ees .. as ~ . ee ¥ 2 et aes a 3 ie ye a PARQ ee Mla ‘Baie PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 16-19. Amblyomma babirussae Larva SULAWESI (RML 99902). 16, Dorsal view (127 x). 17, Capit- ulum, dorsal view (318 x). 18, Capitulum, ventral view (318 x). 19, Coxae I-III (318 x). in file 1, 6-7 teeth in file 2, 5—7 teeth in file 3; all files diminishing to crenulations ba- sally; numerous minute denticles of corona in 2 or 3 transverse rows arranged 5/5 or 6/6. Legs (Fig. 14): Coxa I with a broadly rounded internal spur and elongate external spur; coxae II, III] and IV each with a round- ed external spur; external spurs gradually decreasing in size posteriorly; setae: 4 on coxa I, 3 on I-IV. Trochanters without spurs. Tarsus I 0.423-0.520 (0.470) long; 0.119-0.144 (0.131) wide. Spiracular plate (Fig. 15) sunken at macula. Descriptive statistics for principal char- acters of nymphs are summarized in Ta- ble 3: Larva (Figs. 16-19).—Measurements from 25 unengorged specimens. Chaetotax- ic terminology is that of Clifford and An- VOLUME 89, NUMBER 4 astos (1960). Body (Fig. 16) nearly circular, widest just posterior to coxa III. Length from center of emargination to posterior body margin 0.689-0.749 (0.721); width 0.686- 0.756 (0.720). Dorsal body setae number 10 pairs: 2 pairs central dorsals, CD, 0.016- 0.027 (0.021), CD, 0.022-0.031 (0.025); 8 pairs marginal dorsals, MD, 0.024-0.035 (0.029), MD, 0.022-0.035 (0.028). Ventral setae usually 15 pairs: 3 pairs sternals, ST, 0.023-0.030 (0.026); 2—3 pairs (2.06) prean- als, PA, 0.022-0.028 (0.025), PA, 0.024— 0.030 (0.027); 4-5 pairs (4.02) premargin- als; 5 pairs marginal ventrals, MV, 0.024— 0.038 (0.032); 1 pair anals. Festoons longer than wide, separated by prominent grooves. Sensilla auriformia 7—9 pairs (8.00) dorsal- ly, 12-14 pairs (12.70) ventrally. Four pairs dorsal sensilla sagittiformia: one pair near posterior margin of scutum at level of MD,; one pair just anterior to MD,; one pair sit- uated midway between CD, and festoons 2; one pair within festoons 5. Eight pairs ven- tral sensilla sagittiformia: one pair between coxa I and body margin; one small pair be- tween internal spur of coxa I and ST,; two pairs posterior to each of coxae II and III, the internal sensillum smaller in both cases; one pair between PM, and festoons 3; one pair within festoons 4. Scutum (Fig. 16): Length 0.280-0.304 (0.291), width 0.534— 0.578 (0.563); 3 pairs of setae, SC; 0.011- 0.016 (0.015); cervical grooves deep and parallel, extending to level of SC,; one pair of conspicuous auriform-like sensilla be- tween cervical grooves, posterior to SC,; median sensillum between SC;,; eyes con- vex, length more than twice width, deeply pigmented and with an orange border in preserved specimens. Capitulum (Figs. 17, 18): Length from apex of hypostome to posterior ventral mar- gin of basis 0.320-0.359 (0.342); width 0.204-0.228 (0.215). Basis capituli dorsally (Fig. 17) subtriangular, lateral margins broadly rounded, posterior margin shallow- ly concave or indented; cornua absent. Ven- 655 trally (Fig. 18), basis with rounded lateral and posterior margins. One pair of minute posthypostomal setae, distance between them 0.044-0.062 (0.049). Palpi 0.188- 0.216 (0.205) long, 0.050-0.056 (0.052) wide; sutures between segments distinct; segments II and III each more than twice as long as I and IV; segment I with a ventral ridge; segment II with a dorsal ridge along internal margin; setae: 0 on segment I, 4 dorsally and 2 ventrally on both II and III, 12 on IV. Hypostome rounded and notched apically; length of toothed portion (minus crenulations) 0.090-0.106 (0.098); denti- tion 2/2 throughout; 5-6 teeth in file 1, 4— 5 teeth in file 2; minute denticles of corona in 2 or 3 transverse rows, each arranged 3/3. Legs (Fig. 19): Coxa I with a broadly rounded internal spur and a longer external spur bisecting posterior coxal margin; coxa II with a single wide spur near center of posterior margin; coxa III with a low, some- what triangular spur skewed mesad and barely overlapping posterior margin; setae: 3 on coxa I, 2 on II, 2 on III. Trochanters without spurs. Tarsus I 0.236—0.260 (0.252) long, 0.084—0.094 (0.089) wide. Descrip- tive statistics for salient characters of larvae are summarized in Table 4. Schulze (1933) described A. babirussae from two collections: 3 males, 7 females, 2 nymphs collected on “Sus babirussa” (= Babyrousa babyrussa), ‘““Vaterland Boeroe oder Celebes,” and 3 nymphs collected by O. Warburg at Minahassa (= Minahasa pen- insula), northern Celebes. He deposited the first collection in the Zoologisches Museum, Greifswald, and selected one male as the ““type.”’ The collection of 3 nymphs was de- posited in the Zoologisches Museum, Ham- burg. However, when Frau Helene Schulze donated her husband’s tick collection to the Rocky Mountain Laboratory, it contained one slide-mounted female of 4. babirussae (RML 118517) without any collecting data and 2 nymphs of A. babirussae (RML 46433) in alcohol, collected by O. Warburg, Min- 656 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 5. Amblyomma babirussae examined from Sulawesi, Indonesia. ee SEITE ETE EEE ETESI IIIS Number of Ticks é Q N JE Locality Month Remarks eT Mammalia (Human) 1 Trail to Kulawi Aug. Riding horse 1 Tomado Aug. On leg; trail on mountain behind Tomado 3 1 Tokalosi Sep. In forest Perissodactyla: Equidae Domestic horse l Lake Lindu, Tomado Aug. 6 horse 3 Ales Lake Lindu, Langko Oct. 6 horse Artiodactyla: Suidae Sus scrofa PAG 1 Lake Lindu (nr) Aug. 4 9 2 Soroako Aug. From two pigs 2 Gumbasa Valley, Gumbasa Jan. 12 Lake Lindu, Tomado Oct. G Lake Lindu, Luo Oct. Wild pig 1 3 Lake Lindu, Lembo Nov. Wild pig 4 8 10 Lake Lindu, Palili-Luo Dec. Wild pig 4 9 Lake Lindu, Bamba Mar. Wild pig (2 colls.) 4 yu Lake Lindu, Bamba Mar. ie 20, 1 Lake Lindu, Anca May 4 pig 14 21 1 Lake Lindu, Bamba Jun. 4 pig (2 colls.) Artiodactyla: Cervidae Cervus timorensis 6 3 Lake Lindu (NE side), Luo Jul. ex 1 4, 1 2 deer l 4 Lake Lindu, Palili Jul. deer (2 colls.) Selo 1 Soroako Aug. (2 colls.) 9 Lake Lindu, Palili Aug. 33 ile Lake Lindu, Palili Dec. G Py Bil Lake Lindu, Bamba Dec: 1 Luwa Kabupaten, Tedeboe Jul. 4 2 Lake Lindu, Luo May deer (2 colls.) 1 1 Lake Lindu, Wongkodono Mar. 1 4 Lake Lindu, Anca Mar. 3 5 4 Lake Lindu, Anca Nov. 1 Dae ee Lake Lindu, Anca May 4 Lake Lindu, Kati May Artiodactyla: Bovidae Bubalus (Anoa) depressicornis Gy 28 6 Lake Lindu, Palili Dec. 1 6 8 Lake Lindu, Paku Oct. (2 colls.) 29 4 Lake Lindu, Tomado May (2 colls.) Artiodactyla: Bovidae Domestic buffalo 4 7 7 Lake Lindu, Tomado Jul. 8 ! D Trail from Lake Lindu to Kulawi Aug. ye NG Soroako Aug. 1 2. 1 Lake Lindu, Paku Aug. 4 buffalo (2 colls.) 1 Lampesue Sep. eee eee en een SS oe VOLUME 89, NUMBER 4 Table 5. Continued. Number of Ticks 6 aay. 2 1 1 Dae Carnivora: Viverridae Viverra tangalunga 1 Rodentia: Muridae Rattus exulans tO Mn Rattus hoffmanni 2 Miscellaneous Vegetation 2 2 1 5 4 6. 12 5 3 >, 215 3 | 5 5 4 6 1 5 4 l Gls ay 20 1 9 1 2 2 15 1 l 2 5 Ay lit 12 2 3 1 Locality Lake Lindu, Paku Lake Lindu, Tomado Lake Lindu, Anca Lake Lindu, Paku Parigi Lake Lindu, Paku Lake Lindu, Malo Lake Lindu, Paku Lake Lindu, Puro Lake Lindu (W side), Wongkodono Lake Lindu Lake Lindu, Dono Lake Lindu, Bamba Gumbasa Valley, Gumbasa Lake Lindu, Lomba Lake Lindu, Tomado Donggala, trail from Kulawi to Lake Lindu Lake Lindu, nr. Tomado Lake Lindu, Antja Lake Lindu, Tomado Trail from Lake Lindu to Kulawi Lake Lindu, Paku Lake Lindu, Anca Lake Lindu, Tomado Lake Lindu, Paku Lake Lindu, Langko Lake Lindu, Anca Lake Lindu, Langko Nuara Gumbasa Nuara Gumbasa Malapi Tokalosi Lake Lindu, Tomado Lake Lindu, Lembo Gunung Mogogonipa Month Remarks Oct. (4 colls.) Oct. (2 colls.) Oct. Mar. Jun. cow Aug. Sep. 2 cows May cow Jan. Jul. Aug. rat (3 colls.) Oct... @ rat May @rat Jan. rat (2 colls.) May May Jul. Jul. (2 colls.) Jul. Forest pasture Near wallow of Babyrousa babyrussa Jul. Aug. Aug. (2 colls.) Sep. Sep. Sep. (2 colls.) Sep. Oct. (2 colls.) Aug. Sep. Aug. Sep. Oct. (7 colls.) Oct. Aug. SS 658 ahassa, northern Celebes, 24 Nov. 1890. G. Miller of the Zoologisches Museum, Greifswald, has informed us that his staff found no type specimens of A. babirussae in their tick collection, nor was there ever an entry recorded for A. babirussae in their type-card index file. Furthermore, Gisela Rack of the Zoologisches Museum, Ham- burg, assures us that no nymphs of A. ba- birussae are in their tick collection, so un- doubtedly 2 of the 3 nymphs collected by O. Warburg are now in our collection at the Museum Support Center. Our slide-mounted female 4. babirussae (RML 118517) is in all probability one of the 7 females from Babyrousa babyrussa and we designate RML 118517 as lectotype fe- male Amblyomma babirussae, so mark it and deposit it in the Museum Support Cen- ter, Smithsonian Institution. The 2 nymphs in alcohol (RML 46433) are designated paralectotypes. SPECIES RELATIONSHIPS Anastos (1950), in his review of the In- donesian Ixodidae, listed three species of Amblyomma from Sulawesi: A. babirussae, A. testudinarium and A. cyprium cyprium. The presence of a triangular pointed spur on coxae II and III separates adults of both A. babirussae and A. c. cyprium from A. testudinarium, which has a broad rounded spur on coxae II and III. Adults of A. ba- birussae can be separated from A. c. cyprium by the presence in A. babirussae of large deep scutal punctations and an external spur on coxa | that is much longer than the in- ternal, whereas in A. c. cyprium the scutal punctations are small and shallow and coxa I has an external spur only slightly longer than the internal. Since the Anastos review, two additional species of Amblyomma have been reported from Sulawesi, A. cordiferum and A. hel- volum (Lazell et al., 1987). Female A. cor- diferum have 2 spurs on coxae II and II and an inornate scutum. Males of A. cor- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON diferum are undescribed. Female 4. babi- russae have a single spur on coxae II and III and an ornate scutum. Adult Amblyom- ma helvolum are small Aponomma-like ticks with indistinct eyes and a 3/3 hypostomal dentition. Adult A. babirussae are rather large robust ticks with obvious eyes and 4/4 hypostomal dentition. DISTRIBUTION AND Hosts Specimens of A. babirussae are relatively rare in museum collections and thus far are known only from Sulawesi and perhaps Boeroe Island. This tick has been collected from most areas of Sulawesi and at altitudes from sea level to over 1800 meters. Data presented in Table 5 indicate that A. babirussae can be collected throughout the year and apparently prefers to feed on larger artiodactyls. ACKNOWLEDGMENTS We extend our special thanks to George Venable for his color rendering of the male of A. babirussae and to Taina Litwak for the pen and ink illustrations of the female and nymph. We thank also G. Miiller and G. Rack of the Zoologisches Museums of Greifswald and Hamburg, respectively, for informing us of the absence of type speci- mens of A. babirussae in their tick collec- tions. LITERATURE CITED Anastos, G. 1950. The scutate ticks, or Ixodidae, of Indonesia. Entomol. Am. 30: 1-144. Clifford, C. M. and G. Anastos. 1960. The use of chaetotaxy in the identification of larval ticks (Ac- arina: Ixodidae). J. Parasitol. 46: 567-578. Corwin, D.; C. M. Clifford, and J. E. Keirans. 1979. An improved method for cleaning and preparing ticks for examination with the scanning electron microscope. J. Med. Entomol. 16: 352-353. Krijgsman, B. J. and S. A. S. Ponto. 1931. Die Ver- breitung der Zecken in Niederlaéndisch-Ostindien. Z. Parasitenkd. 4: 140-146. . 1932. De teken van den Oost-Indischen Ar- VOLUME 89, NUMBER 4 chipel. Veeartsenijk. Meded. (79): 62 pp., maps 1-5. Lazell, J. D., J. E. Keirans, and G. A. Samuelson. (In press.) A remarkable ectoparasitic aggregation on the little-known Sulawesi black racer, Coluber (‘“Ptyas’’) dipsas. Copeia. Schulze, P. 1932. Uber das Zustandekommen des Zeichnungsmusters und der Schmelzfarbung in der Zeckengattung Amblyomma Koch nebst Bemer- 659 kungen uber die Gliederung des Ixodidenk6rpers. Z. Morph. Oekol. Tiere 25: 508-533. . 1933. Neue Ixodiden aus den Gattungen Am- blyomma und Aponomma. Zool. Anz. 104: 317- 323: Toumanoff, C. 1944. Les Tiques (Ixodoidea) de I’In- dochine. Institutes Pasteur de Il’ Indochine, Saigon. 220 pp. PROC. ENTOMOL. SOC. WASH. 89(4), 1987, p. 659 NOTE Baetis caelestis Allen and Murvosh, an available name for Baetis sp. A of Morihara and McCafferty (Ephemeroptera: Baetidae) Morihara and McCafferty (1979. Trans. Am. Entomol. Soc. 105: 139-221) in their revision of the Baetis larvae of North Amer- ica included three unnamed but otherwise comprehensively described species as Bae- tis sp. A, Baetis sp B [= Baetis magnus McCafferty and Waltz (1986. Proc. Ento- mol. Soc. Wash. 88: 604)], and Baefis sp. C. These were initially unnamed because, although distinctive, the possibility re- mained that they were associated with for- merly named adults since they had not been reared. Baetis sp. A was based on larvae from Castron and Otero counties in New Mexico and Los Angeles County, California. We have just recently acquired the original ma- terial of Baetis from Baja California de- scribed by Allen and Murvosh (1983. Ann. Entomol. Soc. Am. 76: 425-433). From our comparative examinations we have deter- mined that Baetis caelestis Allen and Mur- vosh is an available name for Baetis sp. A of Morihara and McCafferty. The Allen and Murvosh type material was made up of a mixture of larvae, some of which we have identified as Baetis tricaudatus Dodds. The holotype, however, is clearly identifiable as Baetis sp. A. Our finding is based in part on the presence of robust setae on the serrate margins of the gills. We call attention to the equivalency of Baetis sp. A to Baetis caelestis because Bae- tis caelestis, based on the original descrip- tion and discussion that followed, cannot be distinguished from many other species of Baetis or even placed in the rhodani species group, to which it belongs. If ref- erence is made to Baetis sp. A, however, Baetis caelestis larvae can be readily distin- guished by the description, figures, and key of Morihara and McCafferty (1979). Un- fortunately, due to an inadvertent inversion of letters in that key, users should note that Baetis caelestis will key to sp. B and Baetis magnus will key to sp. A. We thank W. J. Pulawski and N. D. Pen- ny of the California Academy of Science for the loan of specimens. This paper is Purdue Experiment Station Journal No. 11,155. W. P. McCafferty and R. D. Waltz, De- partment of Entomology, Purdue University, West Lafayette, Indiana 47907. PROC. ENTOMOL. SOC. WASH. 89(4), 1987, pp. 660-664 ALACHUA FLORIDENSIS, A NEW GENUS AND SPECIES OF ENTEDONINAE (HYMENOPTERA: EULOPHIDAE) PARASITIC ON THE FLORIDA CARPENTER ANT, CAMPONOTUS ABDOMINALIS (FORMICIDAE) MICHAEL E. SCHAUFF AND ZDENEK BOUCEK (MES) Systematic Entomology Laboratory, BBI, Agricultural Research Service, USDA, % U.S. National Museum of Natural History, NHB 168, Washington, D.C. 20560, USA; (ZB) CAB International Institute of Entomology, % British Museum (Natural History) Cromwell Road, London SW7 SBD, UK. Abstract. —Alachua floridensis, new genus and new species (Hymenoptera: Eulophidae), is described and illustrated. Its relationship to other entedonine genera is discussed. This species is a gregarious endoparasitoid in the pupae of the Florida carpenter ant Camponotus abdominalis (Hymenoptera: Formicidae), in southeastern United States. Over the last 15 years several collections of a eulophid parasite on ant pupae have been made in various localities throughout Florida. The identity of the host species was not confirmed, although the ant was be- lieved to be a species of Camponotus. Re- cently a series of specimens was reared from the pupae of Camponotus abdominalis Fa- bricius at Gainesville, Florida. This species is the dominant Camponotus in Florida and also occurs in neighboring states (Nickerson and Harris, 1985). Some pupae that were collected before the parasites emerged were preserved and the parasites are clearly vis- ible inside. One of these host pupae was dissected and contained 21 pupae of the parasite. Although parasitization of ants is quite common in some groups of Chalcidoidea (e.g. eucharitids), it is rare for the Eulophi- dae and especially for the subfamily Ente- doninae. A species of ant has been recorded as a host for only one other genus in that group (Myrmokata (Bouéek, 1972) from prepupae of an unidentified species of Cre- matogaster in Africa). Given the economic injury caused by Camponotus species, the presence of this parasite may be of some use in biological control. In addition, the un- usual host relationship and the need for a name for this taxon for phylogenetic studies has encouraged us to describe and figure this interesting species. Alachua Schauff and Bouéek, New GENUS Type species. — Alachua floridensis Schauff and Boucéek, New Species. Description. — Head in dorsal view about 2x as broad as long, frontal grooves X-shaped with upper branches diverging to- ward top of eye (Fig. 1). Antennal formula 1:1:1:3:2 (Fig. 10), scape not reaching me- dian ocellus. Clypeus not clearly delimited, the ventral margin slightly upturned. Eyes heavily setose, eye height 74 head height; dorsal and lateral margins of occiput cari- nate (Fig. 2); frontovertex with abundant pilosity (Fig. 1). Pronotum in dorsal view a narrow band, with transverse carina ante- VOLUME 89, NUMBER 4 riorly, and carinate laterally. Scutum (Fig. 3) with 2 pairs of submedian setae, notauli visible in anterior half as weak lines, pos- teriorly visible as a very narrow, broad depression, postero-medial margin sinuate. Transscutal groove widened into a narrow fovea medially. Axillae slightly advanced forward of anterior margin of scutellum, with a single large seta and a small fovea on the postero-lateral margin. Scutellum con- vex, slightly broader than long, with single pair of strong setae at lateral margin, ante- rior margin slightly angulate medially, with- out medial or lateral grooves. Dorsellum a narrow band, slightly overlapping propo- deum. Propodeum 3x as wide as medial length (Figs. 4, 5), gently sloping, with a linear fovea along the anterior edge that ex- tends slightly past the edge of the metano- tum laterally, with a small finger-like pro- jection anteromedially, without a median carina, with several strong setae scattered medially, spiracle small and without ob- vious grooves laterally or anteriorly. Pre- pectus triangular, with an anterior fovea, posterior edge interrupted by anterior edge of the mesosternum (Fig. 5). Mesepister- num with mesofemoral depression. Mes- epimeron divided by a straight, complete suture. Metapleural protuberance well de- veloped and with carina along anterior side (Fig. 9). Gaster petiolate, with first tergite greatly enlarged. Forewing (Fig. 11) with marginal vein equal to submarginal; sub- marginal with 2 strong dorsal setae; stigmal about equal in length to postmarginal; basal and cubital vein setae present (speculum closed); marginal fringe present. Remarks.—The relationship of this genus to other described entedonines will be dis- cussed in detail in a paper in preparation by the senior author. However, some fea- tures possessed by this new genus do enable us to indicate tentative relationships. Sev- eral unique features clearly set Alachua off from other entedonines, in particular, the presence of several strong setae on the me- dian and submedian propodeum and the 661 heavily setose vertex. The presence of a ca- rina extending down the side of the head behind the eyes is also very rare for this group of genera. A nearly complete lack of sculpturing on the dorsal thorax is unusual for entedonines but not unique to Alachua. Without additional species it is difficult to assess how widespread some of these char- acters may be. The indentation of the pos- terior edge of the prepectus by the meso- sternum is shared with Horismenus and the presence of an anterior tooth-like projection medially on the propodeum is also shared with that genus. Species of Horismenus also generally have a sinuate margin postero- medially on the scutum, which is another indication of a link to Alachua. The X-shaped frontal forks of the head in which the scrobes do not unite before reaching the upper forks, the lack of a median scutellar groove, and the smooth non-furrowed me- dian propodeum separate this genus from Horismenus. Etymology.—The generic name is based on the locality of the type series, Alachua County, Florida. It is considered feminine. Alachua floridensis Schauff and Bouéek, NEw SPECIES Female.—Length 0.9-1.4 mm. Color as follows: head, thorax, gaster, and coxae dark brown to black. Antennae, femora, apex of last tarsomere light brown to yellow. Tibiae, first three tarsomeres and basal half of fourth yellow. Antennae (Fig. 10) with scape 3 x as long as wide, inner surface convex. First funicular article slightly longer than wide (3.5:4.0), second as long as wide, third slightly wider than long. Club (including ter- minal spine) 2x as long as wide. Face smooth, oral fossa 2x malar space. Eye height 3x malar space. Mandibles with 2 large ventral teeth and 5-6 smaller teeth (Fig. 8). Lower margin of clypeus projecting slightly forward, frons above fork at a dis- tinct angle from vertex. Ocelli with POL 2.5x OOL. Pronotum smooth posterior of carina, with 4—5 pairs of large setae, lightly 662 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON at » eee -_ st 8 oF oe r r VOLUME 89, NUMBER 4 /| | '/ I) / 4| !/ wi 1, Zit by Ve W1Yi| / Figs. 10, 11. alutaceous on anterior slope. Propleura and prosternum alutaceous laterally becoming lightly reticulate medially. Scutum smooth, twice as broad as long. Propodeum smooth dorsally, callus with about 15 setae. Meta- sternum below and between hindcoxae re- ticulate (Fig. 6), with two carinae laterad of the petiolar foramen which converge below. Petiole slightly longer than wide, reticulate, with a laterally projecting point posteriorly. First gastral tergum in dorsal view about '2 length of gaster, laterally about 7 of length (nearly entire length of gaster in air dried and collapsed specimens), dorsum smooth, bare medially, laterally with numerous se- tae, sides smooth except for small reticulate patch in anterior '4, with only a few setae anteriorly. Remaining terga subequal in length, each with a row of setae and a narrow — 663 Female A. floridensis. 10, Antenna. 11, Forewing. reticulate band of sculpture along posterior margin. Ovipositor sheaths short, only about ’; length of gaster, not extending past tip. Sterna with a single pair of setae postero- medially, last reticulate laterad of oviposi- tor sheaths. Forewing 2.2 x as long as wide; ratio submarginal : marginal: stigmal : postmarginal 70:70:5:5, marginal fringe: wing width 12:95. Hindwing rounded ap1- cally, marginal fringe: wing width 15:40. Forefemur slightly swollen, 3.5 x as long as wide, with a double row of long setae along ventral edge. Tibia equal in length to femur. Hindfemur 3 as long as wide, hindtibia with numerous long setae, each about as long as width of tibia. Male.—Similar to the female except for the following: Length 1.1—1.3 mm; ventral edge of scape with an elongated ridge (Fig. Figs. 1-9. Scanning electron micrographs of A. floridensis. 1, Head, anterior view. 2, Head, lateral view. 3, Thorax, dorsal view. 4, Propodeum. 5, Thorax, lateral view. 6, Propodeum and metasternum. 7, Male scape. 8, Mandibles. 9, Metapleural protuberance. 664 7); petiole without postero-lateral projec- tions. Types. — Holotype 2 on point with the fol- lowing data: “‘Florida, Alachua Co., Gaines- ville, 19 June, 1986. L. Davis and L. Morel. Ex. Camponotus abdominalis F. Deposited in the U.S. National Museum of Natural History (USNM). Paratypes: 58 2 and 15 ¢ with same data as holotype, deposited in USNM, British Museum of Natural His- tory, and Canadian National Collection. Other specimens examined.—Several se- ries from the following localities and hosts: FLORIDA: Key, ex. Formicidae pupa; Ala- chua Co., Gainesville, ex. Camponotus flor- idanus Buckley [now considered a synonym of C. abdominalis F., D. R. Smith, pers. comm.]; Alachua Co., 5.5 mi. W of Gaines- ville, T9S,R19E, section 4. From colony of Camponotus floridanus. Dade Co., Home- stead; Noname Key, ex. ant pupae; Miami, ex. Camponotus pupa. Etymology.—The species epithet refers to PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON the state of Florida, where all the specimens of this species were collected. ACKNOWLEDGMENTS We thank L. Davis (Agricultural Re- search Service, USDA, Gainesville) for the reared specimens and D. R. Smith (System- atic Entomology Laboratory, Agricultural Research Service, USDA) for confirming the identification of the host. We are also grate- ful to J. Huber, S. Heydon, S. Shaw, T. Hen- ry, and E. E. Grissell for reviewing the manuscript and for their helpful comments. LITERATURE CITED Bouéek, Z. 1972. Descriptions of new eulophid par- asites (Hym., Chalcidoidea) from Africa and the Canary Islands. Bull. Entomol. Res. 62: 199-205. Nickerson, J. C. and D. L. Harris. 1985. The Florida Carpenter Ant, Camponotus abdominalis flori- danus (Buckley) (Hymenoptera:Formicidae). Fla. Dept. Agric. Consumer Serv. Div. Plant Indus. Entomol. Circ. 269. PROC. ENTOMOL. SOC. WASH. 89(4), 1987, pp. 665-672 HOST SPECIFICITY AND LABORATORY REARING STUDIES OF MEGACYLLENE MELLYI (COLEOPTERA: CERAMBYCIDAE), A POTENTIAL BIOLOGICAL CONTROL AGENT OF BACCHARIS NEGLECTA BRITT. (ASTERACEAE)! PAUL E. BoLpT Grassland, Soil and Water Research Laboratory, Agricultural Research Service, USDA, ©), Box, 6112, Temple: Texas 76503-6112. Abstract.—The stem borer Megacyllene mellyi (Chevrolat), originally from Brazil and imported from Australia, was studied in quarantine as a possible biological control agent of Baccharis halimifolia L. and B. neglecta Britt. (Asteraceae) in the United States. Larvae were reared for 30-45 days on stems of Baccharis before being transferred to artificial diet containing milled Baccharis stems. Developmental times for colony-reared insects at 26°C were: eggs, 10.7 days; larvae, 99.2 days; pupae, 15.2 days; and adults, 10-29 days. There were 3 generations/year. Host specificity tests were conducted on 35 species of plants. Oviposition occurred on Baccharis, a few weedy Astereae shrubs and large, rough- barked stems of some trees of other plant families. Larvae, however, successfully developed only on Baccharis. Female borers were attracted to a volatile water-soluble chemical in the phloem layer of stems of B. neglecta and preferred to oviposit on large stems with rough bark. The genus Baccharis (Asteraceae: Aster- eae) consists of 511 species of unisexual shrubs, most of which occur in Brazil, Ar- gentina, or Chile (Malagarriga, 1976; Bar- roso, 1976). Approximately 21 species oc- cur in southern United States (Anon., 1982), including three undesirable weeds of range- lands, pastures, and low areas: B. halimi- folia L., B. neglecta Britt., and B. salicifolia (R&P) Pers. These native species attain heights of 2 to 4 m, are prolific seed pro- ducers, tolerate a wide range of soil types and salinity conditions, and regrow from basal stem buds (Scifres, 1980; Westman et al., 1975). Control methods such as shred- ding and burning are only temporary, ' Mention of a proprietary product does not consti- tute an endorsement or a recommendation for its use by the U.S. Department of Agriculture. whereas herbicide applications are effective but expensive (Mutz et al., 1979). Baccharis halimifolia occurs from Texas to Florida and as far north as Massachusetts (Tarver et al., 1979) and is unpalatable and not grazed by livestock (White, 1936). The leaves contain a cardiotoxic glucoside that iS poisonous to cattle (Manley et al., 1982) and poultry (Duncan et al., 1957). The pol- len causes hay fever in humans (Wode- house, 1971). Another species, Baccharis neglecta, has infested over 330,000 ha of range and pasture land in Texas (unpub- lished brush survey, 1982, USDA Soil Con- servation Service, Temple, Texas) and pro- gressively has become a management problem on productive grassland soils and abandoned fields (Mutz et al., 1979; Scifres, 1980). Baccharis salicifolia is an undesir- able phreatophyte and is also unpalatable 666 as forage as well as forming dense thickets along streams (Parker, 1972). The stem borer, Megacyllene mellyi (Chev.) (Coleoptera: Cerambycidae), is na- tive to South America. In Brazil, its pre- ferred host plant is B. microdonta DC., but it occasionally attacks B. dracunculifolia DC., B. spicata (Lam.) Baillon, B. tridentata Vahl and Baccharidastrum sp. (Bennett, 1963; McFadyen, 1979). Adults are black in color with red markings on the elytra and white setae scattered on the legs, head and thorax. The larvae bore through the bark and feed beneath the surface. Stems which contain several larvae are weakened (McFadyen, 1979). The host specificity of M. mellyi was de- termined by McFadyen (1983) for use in Australia as a potential biological control agent against B. halimifolia. He tested 55 species of economically important plants and 20 species of plants that were hosts or closely related to plants attacked by other Megacyllene spp. Females deposited most eggs on B. halimifolia, but a few were also deposited on the leaf surfaces of corn, peach, pear, grape, and pineapple; however, larvae did not survive on any of the test plants. The borer was approved for introduction and released in Queensland, Australia in 1978 (McFadyen, 1983). To be considered for release in the United States, we tested M. mel/lyi for ovipositional preference and larval survival on various plants of economic importance in North America, native asters which are closely re- lated to Baccharis, and host plants of the native Megacyllene. The results of this test- ing and the data collected from laboratory- rearing studies are presented in this report. MATERIALS AND METHODS A laboratory colony of M. mellyi was maintained from June 1982 to September 1984 under strict quarantine conditions at Temple, Texas. The colony was started and replenished with shipments of larvae and pupae from the Alan Fletcher Research Sta- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tion, Queensland Department of Lands, Sherwood, Australia. Rearing methods for the colony were similar to those used by A. Tomley and P. McFadyen (pers. comm.) of the Alan Fletcher Research Station. Cut stems (3-6 cm diam. x 40-50 cm length) of either B. neglecta or B. halimifolia were exposed to ovipositing females for 3-5 days depending on the number and age of the adults in the cage. Ends of the stems were dipped in hot parafin wax to conserve moisture. Females oviposited under 2 cm wide strips of cotton cloth tightly spiraled around the stems. These cloth strips in- creased the number of oviposition sites, en- couraged females to oviposit over a larger area of the stem and provided a source of eggs attached to cloth for some of the ex- periments. The Baccharis stems were then placed in a polyethylene bag, sprayed with water from 3 to 6 times weekly to maintain high humidity, and held at 26°C for 30 to 50 days. Stems of B. halimifolia were pre- ferred for rearing larvae because they did not split and dry as rapidly as did stems of B. neglecta. After 30 to 50 days the stems were dissected and each larva was placed in a hole in a | cm? piece of artificial diet in a glass vial. The diet was replaced ca. once every 2 weeks. For mature larvae the diet was provided in granular form. The diet used was similar to the diet of Harley and Willson (1968) except the cellulose was re- placed with an equal weight of milled Bac- charis stem. Pupae and teneral adults were held in empty vials for 3-5 days after emergence. Teneral adults used in the experiments were held for an additional 5 days before testing and were given only water and honey so- lution on cotton pads. All tests were conducted between October and June 1982 to 1984, in the quarantine greenhouse (Boldt, 1982), with a natural photoperiod and at temperatures of ca. 30°C in the day and 20°C at night. An entire ex- periment could not be done at the same time but there was one control treatment each VOLUME 89, NUMBER 4 Table 1. Plant species used in the host specificity testing of M. mellyi. 667 Family Species Common Name Gramineae Saccharum officinarum L. sugarcane Gramineae Sorghum vulgare Pers. sorghum Juglandaceae Carya illinoinensis (Wang) K. Koch pecan Juglandaceae Juglans nigra L. black walnut Ulmaceae Celtis laevigata Willd. Texas sugarberry Ulmaceae Ulmus crassifolia Nutt. cedar elm Moraceae Ficus carica L. fig Rosaceae Prunus persica (L.) Batsch. peach Leguminosae Albizia julibrissin Durazz. mimosa-tree Leguminosae Gleditsia triacanthos L. honey locust Leguminosae Phaseolus vulgaris L. bean Leguminosae Prosopis glandulosa Torr. honey mesquite Leguminosae Sophora affinis T. & G. Eve’s necklace Balsaminaceae Impatiens balsamina (L.) Lam. impatiens Vitaceae Vitis vinifera L. Niagara grape Malvaceae Gossypium hirsutum L. cotton Convolvulaceae Ipomoea batatas (L.) Lam. sweet-potato Cucurbitaceae Cucurbita pepo L. pumpkin Asteraceae* Astereae Baccharis halimifolia L. Astereae Baccharis neglecta Britt. Astereae B. pilularis DC. coyote bush Astereae B. brachyphylla Gray Astereae Gutierrezia microcephala DC. threadleaf snakeweed Astereae Isocoma drummondii (T. & G.) Drummond’s goldenweed Astereae Ericamerica austrotexana M.C. Johnst. false broomweed Astereae Aster novae-angliae L. New England aster Astereae Aster sp. Michaelmas daisy Astereae Erigeron aurantiacus Regel. midsummer aster Astereae Chrysothamnus nauseosus (Pall.) Britt. rubber rabbitbrush Astereae Solidago altissima L. tall goldenrod Astereae Conyza canadensis (L.) Cronq. horsetail conyza Heliantheae Chrysanthemum morifolium Ramat. florists mum Heliantheae Helianthus annuus L. sunflower Inuleae Antennaria fallax Greene large leaf pussy’s toes Anthemideae Artemisia filifolia Torr. sand sagebrush Eupatorieae Eupatorium compositifolium Walt. yankee weed 2 iribes. time any plants were tested. The control plant, B. halimifolia, was replaced by B. ne- glecta in the multiple choice oviposition and attractants tests because it was locally com- mon and fresh stems were readily available. Plants tested are listed in Table 1. During the oviposition tests, cage interiors were sprayed twice daily with water. Means and standard deviations were calculated for the data. No-choice oviposition test.—In this test, ten unsexed adults were confined in a screen cage (20 cm diam. x 50-90 cm ht.) for 10 days on each individually potted plant. At the end of the test, the plant was dissected to record the number of eggs. Multiple-choice oviposition tests. —In the first test, 40 unsexed adults were confined for 10 days in a 1 m® screen cage with 4 woody stems of each of 3 species of shrubs or trees listed in Table 2 and 4 stems of B. neglecta. All stems (ca. 3.5 cm diam. x 40 668 cm length) were cut in the field 4 to 24 hours before the start of the test. A hole (7.6 cm deep x 0.32 cm diam.) drilled in one end of each stem enabled the stem to stand up- right over a nail driven up through the wood floor. Cut ends of each stem were coated with a thin layer of paraffin wax to reduce moisture loss. Stems were arranged in the cage in a Latin-square design. Ten species of plants plus B. neglecta were tested in this experiment. At the end of each test the stems were dissected and the number of eggs pres- ent and their location on the stem were re- corded. In the second test, 15 unsexed adults were also confined in each cage for 10 days with only one stem of each of 3 plants plus | stem of B. neglecta. One replicate consisted of 3 cages of 3 plant stems each and | B. neglecta stem in each cage. The experiment was replicated 4 times using all combina- tions of test plants. Larval survival test.—A strip of cotton cloth from the rearing colony, containing 10 (0-24-h old) attached eggs, was taped to each stem used in the experiment. The stems were cut in the field, 4-24 h before the test and soaked in water for ca. 3 h; the cut ends were dipped in hot paraffin wax to maintain moisture content. Eggs and stems were then held in moist peat moss in a polyethylene plasticscage:(2ax ex’ Mean + standard deviation. cubation period at 26°C was 10.7 + 0.65 days (n = 99). Percent hatch for 456 eggs was 92.1%. Young larvae fed in the wood just below the bark for 30 to 45 days before being transferred to diet. Mean duration of the larval stage was 99.2 + 21.6 (n = 44) and mean duration for the pupal stage was 15.2 + 6.2 (n = 43) days. No-choice oviposition test.— Twenty-five species of plants, representing nine families, were included in the single plant no-choice Oviposition test (Table 2). Of the total of 279 eggs laid, 81.0% were placed on B. ne- glecta and 12.5% on B. pilularis DC.; the remainder were distributed between six species of shrubby Astereae and one com- mercial plant, sugarcane, Saccarum offici- narum L. Eggs were usually laid singly in crevices and narrow depressions or under loose pieces of bark except on sugarcane and yankee weed, Eupatorium compositifolium Walt., where they were laid on the smooth upper surface of the leaves. Several eggs were also found on the screen of a cage containing a threadleaf snakeweed plant, Gutierrezia microcephala DC. Multiple-choice oviposition test.—Elev- en species of shrubs and trees, representing 8 families, were included in the multiple- choice ovipositional test (Table 3). Ovipo- sition in both tests was greatest on stems of B. neglecta although eggs were also laid on 5 other plants in the first and 8 other plants in the second test. Of ihe total of 1400 eggs laid in the two tests, 87.4% were laid on B. neglecta and 4.2% were laid on honey mes- 670 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 3. Number of eggs laid by !. mellyi on woody stems in multiple-choice tests. Test | Test 2 No. of Bees No. of Fees Plant Stems No. Laid No./Stem?* Stems No. Laid No./Stem* Baccharis neglecta 16 765 47.8 + 33.8 12 459 38.3 + 22.6 Honey mesquite 8 34 a3) ae 0) 4 DS) 6:35 IES Eve’s necklace 8 10 137 == 10 4 9 Mesiae Oh Texas sugarberry 4 5) INS} sie De) p) 3 15 =a Walnut 8 4 OFS-EM el 4 1 025) ==05 Pear 4 4 Oias 2X0) 4 2, (Osea {0 Pecan 4 0 _ 4 7 14.3 + 10.4 Honey locust 4 0 = 4 9 ppsyar W116 Fig 4 0) 4 5 jesse. S) Mimosa-tree 4 0 = 4 0 _ Cedar elm — — 2 8 4.0 + 2.8 aMean + standard deviation. quite. No eggs were laid on pecan, Carya illinoinensis (Wang) K. Koch, in the first test but 20.9% of the total were on pecan in the second test. The only plant with no eggs in either experiment was mimosa-tree, A/bizia Julibrissin Durazz. Larval survival test.—Twelve species of trees and shrubs representing 9 families were included in the larval survival test (Table 4). Eclosion ranged from 70 to 90% on each plant species. Feeding damage by newly hatched larvae was observed in the bark of all plants and was recognizable by the pres- ence of a small pile of sawdust around a small entrance hole in the bark. Of the 38 larvae which survived the average 43-day duration of the tests (ca. one-half of the nor- mal larval developmental period of 99 days), 37 had tunneled into the Baccharis spp. and 1 larva had tunneled into a fig, Ficus persica L. The tunnel of the latter larva was small and located between the bark and wood; it was also smaller than any larval tunnel in Baccharis. The fig stem appeared to have a high moisture content which may have aid- ed survival of the larva by preventing des- iccation. The damage in the remainder of the plants was limited to one short tunnel per larva in the bark which did not extend to the wood. Attractancy tests.—In the test for chem- ical attractancy, adults were exposed to cloth strips that had previously been soaked in water extract from the phloem layer of a stem of B. neglecta; control strips were soaked in deionized water. Females laid a mean of 59.6 + 43.5 eggs per replicate on stems with the Baccharis extract and 1.0 + 1.5 eggs on the control. In the test for physical attractancy, adults were given a choice of 3 types of stems of B. neglecta for oviposition. Females laid 1.0 + 1.4, 4.0 + 2.8, and 40.0 + 25.4 eggs/ replicate on small, medium, and large stems, respectively. The largest stems had bark with the most ridges and deepest grooves. DISCUSSION The oviposition tests indicate that Bac- charis is the preferred host plant (Tables 2 and 3). Oviposition occurred only on Bac- charis and some rough-stemmed woody pe- rennials and did not occur on commercial asters or on non-woody plants outside of the tribe Astereae. A few eggs were also loosely laid on the leaf surface of yankee weed, sugarcane, and the screen of the cage. However, this was probably the result of the artificial condi- tions of the test. Eggs laid on leaves in the field would have little chance of survival due to normal predation. VOLUME 89, NUMBER 4 671 Table 4. Summary of survival of larvae of M. mellyi for 5 to 7 weeks in stems of various woody plants, 1983-84. Eggs Larvae Plant Total No. Hatched (%) No. Fed in Bark (%) No. Survived (%) Baccharis halimifolia 66 65 (98) 44 (67.7) 28 (43.1) B. neglecta 70 63 (90) 32 (50.8) 9 (14.3) Fig 40 28 (70) 7 (25.0) 1 (3.6) Pecan 40 36 (90) 20 (55.5) 0 Mimosa-tree 40 33 (82) 20 (60.6) 0) Walnut 40 40 (100) 14 0 Cedar elm 30 30 (100) 21 (70) 0 Sugarberry 30 23H) 1 0 Honey mesquite 30 30 (100) 6 0 Honey locust 30 29 (97) 5 0 Eve’s necklace 30 30 (100) 11 0) Pear 10 9 (90) 4 0 Females are attracted to their host plant or are stimulated to oviposit by both chem- ical and physical means. The presence of water-soluble chemicals which attracted M. mellyi or stimulated oviposition was dem- onstrated to exist in the phloem layers of B. neglecta and probably exists in B. halimi- folia as well, although this plant was not tested. Females searching the surface of the stem preferred to oviposit on large stems with rough bark and deep grooves. These grooves were simulated in the rearing cage by wrapping a strip of cloth around a stem to increase the number of oviposition sites. Results of the larval survival test (Table 4) as well as those of McFadyen (1983) in- dicate that VM. mellyi is restricted to plants in the genus Baccharis and after additional testing on the commercial shrub B. pilularis should be considered as a potential biolog- ical control agent for the United States. Al- though neonate larvae fed to varying de- grees in the bark of most of the test plants, they neither penetrated the phloem layer nor fed into the woody portion of the stem. Bark is apparently a neutral material which does not contain stimulatory chemicals and is tolerated by the larvae only in an effort to reach the wood. Larvae survived for 5— 7 weeks only on B. halimifolia, B. neglecta, and fig. The single larva on fig, however, was small and in poor condition at the time the stem was dissected. It probably could not have survived much longer. Larvae of M. mellyi can cause severe damage to Baccharis, but the borer’s effec- tiveness as a biological control agent is dif- ficult to evaluate. In Brazil, stems that con- tain several larvae are weakened and plants less than one meter in height may be killed (McFadyen, 1979). In Australia, releases made on B. halimifolia in 1978 are estab- lished but populations are slow to increase and damage is inconsistent (McFadyen, 1983). ACKNOWLEDGMENTS I thank Bill Haseler, Paul McFadyen and Alan Tomley, of the Alan Fletcher Research Station, Queensland Department of Lands, Sherwood, Australia, for their generous sup- port and cooperation and for shipments of M. mellyi. 1 thank Don Foster, Olin E. Teague Veterans’ Center, Temple, Texas, for special assistance with the attractant studies. I also thank Lisa Bruggman, Ray Ullrich, and Tom Robbins for their assis- tance in rearing the borers and conducting some of the experiments. I especially thank C. J. DeLoach, USDA Argricultural Re- 672 search Service, Temple, Texas, for his many helpful suggestions and continued encour- agement. LITERATURE CITED. Anonymous. 1982. National List of Scientific Plant Names. 1. List of Plant Names. U.S. Dept. Ag. SCS-TP 159. 416 pp. Barroso, G. 1976. Compositae—Subtribo Bacchari- dinae Hoffman. Estudo das species ocorrentes no Brasil. Rodriguesia 28: 1-273. Bennett, F. 1963. Final report on surveys on the in- sects attacking Baccharis spp. in the S.E. United States of America and Brazil. 1960-63. Comm. Inst. Biological Control Rep. 27 pp. Boldt, P. E. 1982. Quarantine facility for exotic phy- tophagous insects. FAO. Plant Prot. Bull. 30: 73- Wile Duncan, W., P. Piercy, S. Feut, and R. Starling. 1957. Toxicological studies of southeastern plants. II. Compositae. Econ. Bot. 11: 75-85. Harley, K. L. and B. W. Willson. 1968. Propagation of a cerambycid borer on a meridic diet. Can. J. Zool. 46: 1265-1266. Malagarriga, R. 1976. Nomenclator Baccharidina- rum omnium. Mem. Soc. Cienc. Nat. La Salle 107: 129-224. McFadyen, P. J. 1979. South American insects in- troduced into Australia for the biological control PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON of Baccharis halimifolia L. Ph.D. Dissertation. Univ. Queensland. 398 pp. 1983. Host specificity and biology of Me- gacyllene mellyi (Col.: Cerambycidae) introduced into Australia for the biological control of Bac- charis halimifolia (Compositae). Entomophaga 28: 65-72. Manley, G., G. Edds, and S. Sundlof. 1982. Cattle deaths from poisonous plants. Fl. Vet. J. 11: 20. Mutz, J., C. Scifres, W. Mohr, and D. Drawe. 1979. Control of willow baccharis and spiny aster with pelleted herbicides. Tex. A&M Exp. Stn. Bull. B-1194. Parker, K. F. 1972. An Illustrated Guide to Arizona Weeds. Univ. Arizona Press, Tucson. 338 pp. Scifres, C. J. 1980. Brush Management. Texas A&M Press, College Station. 360 pp. Tarver, D., J. Rodgers, M. Mahler, and R. Lazor. 1979. Aquatic and Wetland Plants of Florida, p. 72. Fl. Dept. Nat. Res. Westman, W., F. Panetta, and T. Stanly. 1975. Eco- logical studies on reproduction and establishment of the woody weed, groundsel bush (Baccharis hal- imifolia) L.: Asteraceae). Aust. J. Agric. Res. 26: 855-870. White, C. T. 1936. Groundsel-bush in southeastern Queensland (Baccharis halimifolia). Qd. Agric. J. ASST: Wodehouse, P. 1971. Hayfever Plants. Hafner Pub. Co., N.Y. 280 pp. PROC. ENTOMOL. SOC. WASH. 89(4), 1987, pp. 673-681 A NEW NORTH AMERICAN SPECIES IN THE SIMULIUM VERNUM GROUP (DIPTERA: SIMULIIDAE) AND ANALYSIS OF ITS POLYTENE CHROMOSOMES PETER H. ADLER Department of Entomology, Clemson University, Clemson, South Carolina 29634. Abstract.—Simulium loerchae Adler, new species, a North American member of the Simulium vernum group, is described from South Carolina, Pennsylvania, and Tennessee. Larvae, pupae, males, females, the polytene chromosomes, and the habitat are described. The species is chromosomally monomorphic and can be derived from the standard se- quence of S. vernum by three fixed inversions. Affinities with North American and Eu- ropean vernum group members are given. The Simulium vernum Macquart group in North America consists of eleven de- scribed species, nearly all of which are pri- marily northern in distribution. The group is characterized by the triangular flange of the male gonostylus. Taxonomic and chro- mosomal treatments of the group members were given by Stone and DeFoliart (1959), Davies et al. (1962), Brockhouse (1985), Hunter and Connolly (1986), and Adler and Currie (1986). Immatures of most species inhabit small, cool, woodland streams, and the adults are principally ornithophilic. I describe all life stages and the polytene chromosomes of yet another new species in the S. vernum group. Larval descriptions are based on material preserved in Carnoy’s fix- ative (1 part glacial acetic acid : 3 parts ab- solute ethanol). Pupal descriptions are based on specimens in Carnoy’s fixative and exu- viae in glycerin. Measurements of adults are taken from freshly pinned material; colors are those of dried specimens. Life-stage de- scriptions follow the format and terminol- ogy used by Adler and Currie (1986). Chro- mosomes were prepared by the Feulgen technique (Rothfels and Dunbar 1953), and both temporary and permanent mounts (in Euparal®) were examined under oil im- mersion (1250 x). The chromosomal com- plement is described relative to the vernum standard (Knebworth cytotype) sequence of Brockhouse (1985). Fixed inversions are italicized and numbered in order of their discovery (within an arm), beginning after the last numbered inversion of Hunter and Connolly (1986). All material in Carnoy’s fixative was transferred to 80% ethanol for permanent storage. The holotype and some paratypes are deposited in the United States National Museum of Natural History, Washington, D.C. Additional paratypes are deposited in the Canadian National Collection, Biosys- tematics Research Centre, Ottawa; the Brit- ish Museum (Natural History), London; the Clemson Entomological Museum, South Carolina; and the Frost Entomological Museum, Pennsylvania State University. Photographic negatives and contact sheets accompany paratypic chromosome prepa- rations. Simulium loerchae Adler, NEw SPECIES Figs. 1-13 Larva (final instar).— Length 5.4-6.7 mm (x = 6.0 mm). Head capsule (Fig. 6) rather 674 uniformly pale yellowish brown; headspots brown, distinct, occasionally surrounded by infuscation posteriorly (as in Fig. 1); eye spots rather large; line over eye spots thin, brown. Antenna with distal article brown, median article translucent or brown dor- sally, proximal article brown dorsally, translucent ventrally or entirely translucent; apex of median article not reaching end of labral-fan stalk; proportions of articles (dis- tal to proximal) 1.1:1.0:1.0. Labral fan with 37-51 (X = 45) primary rays in South Car- olina specimens [50-54 (x = 52) in Ten- nessee specimens; 48-55 (X = 52) in Penn- sylvania specimens]. Hypostomal teeth (Fig. 8) with median tooth and lateral teeth rel- atively large and subequal in length and prominence; sublateral teeth variously smaller, with outermost or innermost sub- lateral teeth longest and median sublateral teeth shortest; lateral margin of hypostoma with 0-2 paralateral teeth and 3-5 lateral serrations per side; hypostoma with 3-4 prominent and 1-2 small lateral setae per side. Postgenal cleft (Fig. 7) about 1.5 times as long as wide, extending about 7 distance to hypostomal groove, widest at midpoint, rounded apically; subesophageal ganglion lightly pigmented. Maxillary palpus 4.0—4.4 times as long as basal width. Inner subapical ridge of mandible with 2—4 small teeth basal to | large, subtriangular tooth. Lateral plate of thoracic proleg moderately sclerotized, elongate, extending almost entire length of apical article. Body (Figs. 1, 2) reddish; pig- mentation heaviest dorsally on segment 8 and dorsolaterally on segments 6-7; seg- ment 5, and sometimes 1, with a single con- spicuous, reddish pigment spot per side; in- tersegmental bands clear, distinct; ventral tubercles conspicuous, about '4 depth of ab- domen at attachment points; posterior 4 ab- dominal segments dorsally with widely scat- tered, short, yellowish brown, simple setae. Anterodorsal arms of anal sclerite broadly connected to and subequal in length to or slightly shorter than posteroventral arms. Rectal setulae present between and at apices PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON of anterodorsal arms of anal sclerite. Pos- terior proleg bearing 7-12 hooks in 65-68 rows. Anal papillae of 3 compound lobes. Pupa.—Length 2.6-3.2 mm. (x = 2.9 mm). Head projecting downward, with many minute granules as on thorax; anten- nal sheath of female extending slightly be- yond posterior margin of head; antennal sheath of male extending about '2 distance to posterior margin of head. Gill (Fig. 3) about 0.4 mm longer than pupa, consisting of 4 filaments; base short, giving rise to 2 petioles that diverge vertically at an angle of 46-73° (x = 58°); ventral petiole up to twice as long as, subequal in thickness to, and (in dorsal view) subparallel to dorsal petiole; ventral petiole giving rise to 2 fil- aments bifurcating in horizontal plane; dor- sal petiole giving rise to 2 filaments bifur- cating 1n vertical plane (plane of bifurcation variable for filaments of both petioles in Pennsylvania specimens); filaments long, thin, tapering, with numerous ridges and shallow furrows; surface sculpturing of base weakly differentiated. Thorax with abun- dant, closely set, uniformly spaced, minute, rounded granules (Figs. 4, 5); trichomes simple, long, very slender, pale, 7 or 8 on each side of thorax. Tergite I with | pair of setae; tergite II with 5-7 anteriorly directed setae on each side of midline, and 1-2 mi- nute setae laterally; tergites II] and IV each with 4 anteriorly directed hooks on poste- rior margin on either side of midline, | small seta between and anterior to 2 outermost hooks, and 1—2 small setae laterally; tergites V to VIII each with row of fine, posteriorly directed spines along anterior margin on either side of midline (spines of tergite V often numbering O-1 per side); tergite IX with pair of very short, stout, slightly curv- ing, dorsally directed terminal spines. Pleu- ral membrane of segments II to VII usually with at least 1 minute seta per side. Sternite III with pair of minute setae anteriorly and 2 pairs of minute setae posteriorly; sternite IV posteriorly with 2 pairs of heavy, simple or bifid, anteriorly directed setae and at least VOLUME 89, NUMBER 4 ° @ - 9527: fo git? , of. 2 On Pon 20 oo? ° Figs. 1-S. Simulium loerchae. Figs. 1, 2. Larval habitus. |, Dorsal view. 2, Ventral view. Figs. 3-5. Pupa (scanning electron micrographs). 3, Gill (lateral view), scale bar represents 50 um. 4, Thorax (dorsal view), scale bar represents 38 um. 5, Granules on portion of thorax (dorsal view), scale bar represents 10 um. 2 pairs of fine setae; sternite V posteriorly with 2 pairs of closely set, anteriorly di- rected, bifid or trifid, hook-like setae and at least 1 pair of fine setae; sternites VI and VII posteriorly with 2 pairs of distantly set, bifid to quadrifid, hook-like setae, and at least | pair of fine setae; sternites VIII and IX with at most | pair of fine setae; sternites III to VIII with numerous tiny rows of ex- tremely fine microspines. Cocoon well formed, with anterodorsal projection broad and accounting for about 16.9-26.5% (x = 676 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 22.2%) total cocoon length (in lateral view); anterior margin and anterodorsal projection reinforced. Female.—Generally brown with gray pruinosity and pale golden pile (some spec- imens with pile more silvery). Length: body, 2.7-3.0 mm (X = 2.8 mm); wing, 2.7-3.1 mm (xX = 2.9 mm). Frons at vertex about 2 times broader than at narrowest point, slightly less than 4 width of head, with decumbent, pale golden pile laterally. Clypeus slightly longer than wide, with sparse, pale golden pile. Occiput with long, pale golden pile reaching poste- rior margin of eye; postocular setae black. Antenna with fine silver pubescence; first flagellomere longest; pedicel and scape brown to light brown; flagellum dark brown. Mandible with 38—40 serrations. Lacinia with 23 retrorse teeth. Palpus dark brown, with stout, golden setae; palpomere V 1.3- 1.6 times as long as III. Sensory vesicle lo- cated subcentrally, occupying 2 to % of pal- pomere III; neck long, thin, arising near anterodorsal margin, opening to exterior through rounded, slightly expanded mouth. Median proximal space of cibarium U-shaped. Postpronotum dark brown; pile long, erect, pale golden. Scutum dark brown, hu- meral angle brown; pile recumbent, pale golden. Scutellum brown, with long, pale golden pile mixed with black setae. Post- notum brownish black. Anepisternum and katepisternum dark brown; membrane and mesepimeron lighter brown; mesepimeral tuft of long, golden setae. Wing veins pale yellowish brown. Setae on stem vein and costal base dark brown with golden reflec- tions; setae on other veins primarily brown; subcosta setose ventrally; fringe of calypter and alar lobe pale golden. Halter tan, with line of golden pile. Coxae, tarsi, and apices of femora and tibiae primarily brown, re- mainder of legs light brown; pile on coxae pale golden, golden on femora and tibiae, primarily brown on tarsi; hind basitarsus 6.5-6.7 times as long as broad; calcipala and pedisulcus well developed; claws each with large, basal, thumb-like lobe. Abdominal sclerites brown; pile sparse, pale golden; additional sparse, long, black setae on tergites VIII-IX; membranous areas gray to tan, with sparse, mixed black and pale golden pile. Basal fringe of very long, pale golden pile. Anal lobe subtriangular to subquadrate in lateral view, with acute pos- terodorsal extension, and small, anteroven- tral nipple. Cercus subrectangular, about twice as broad as long, posterior margin straight, corners well rounded. Hypogynial lobe subtriangular, somewhat acute poste- riorly, space between lobes rather wide. Genital fork (Fig. 11) with stem moderately long and slender; lateral arms rather broad basally, forming subquadrangular space in region of bifurcation, posteromedial areas well developed and with bluntly squared an- gles. Spermatheca of moderate size, longer than broad, with surficial pattern of sub- equal polygons. Male. —Generally velvety black with gray pruinosity and golden-brown pile. Length: body 2.5-3.0 mm (X = 2.7 mm); wing, 2.3- 2.6 mm (X = 2.5 mm). Frons and clypeus with erect, brown pile. Occiput with long, erect, brown pile. An- tenna dark brown, with fine, light brown pile; scape and pedicel paler. Palpus dark brown, with brown pile; palpomere V about twice as long as palpomere III. Sensory ves- icle subspherical, about 4 length of its seg- ment; neck long, slender, opening to exte- rior through rounded mouth. Postpronotum brown, with golden pile. Scutum black, with recumbent, golden pile. Scutellum brown, with brown or mixed gold and brown pile. Postnotum brownish black. Anepisternum dark brown, lighter poste- riorly; katepisternum blackish brown; membrane and mesepimeron brown; mes- epimeral tuft of long, mixed golden-brown and brown pile. Wing veins pale yellowish brown. Setae on stem vein and costal base VOLUME 89, NUMBER 4 677 Figs. 6-11. Simulium loerchae. Figs. 6-8. Larva. 6, Head capsule (dorsal view). 7, Head capsule (ventral view). 8, Hypostoma (ventral view). Figs. 9, 10. Male. 9, Terminalia (ventral view with left gonocoxite and gonostylus removed). 10, Dorsal plate (dorsal view). Fig. 11. Female genital fork. 678 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON brown; setae on other veins brown to golden brown; fringe of calypter and alar lobe gold- en to golden brown. Halter tan to gray, with darker edging and golden-brown pile. Legs dark brown; midsection of femora and tib- iae and base of basitarsi brown to light brown; pile golden to brown. Hind basitar- sus 4.1—4.6 times as long as broad. Abdominal tergites dark brown, with golden-brown pile; membranous areas gray- ish, with long, golden-brown setae laterally on segments III and IV; sternites brown, with long, brown pile. Basal fringe of very long, brown to golden-brown pile. Termi- nalia as in Fig. 9. Gonocoxite about 1.5 times longer than broad. Gonostylus about as long as gonocoxite, about 2.5-3.0 times as long as breadth at midpoint, expanded apically into flattened, subtriangular, medially di- rected flange bearing | apical spine. Ventral plate in ventral view subrectangular, about 2 times as broad as long, tapering poste- riorly, with posterolateral corners well rounded and posterior margin slightly con- cave; anterior margin slightly convex; arms strongly bowed, with tips curving inward; lip in terminal view pronounced, broadly rounded; stem of median sclerite long, slen- der, forked about 4—'» distance from distal end; dorsal plate (Fig. 10) large, well scler- otized, subrectangular, with short projec- tions at each corner and rounded protuber- ance distally; paramere in lateral view moderately narrow basally, broadening me- dially and bearing | long, slender, strongly sclerotized spine-like process. Chromosomes (from larval salivary glands, 44 preparations examined).—n = 3; chromocenter absent; B chromosomes lack- ing; chromosome I with expanded centro- mere region; IS standard for vernum se- quence; IL (Fig. 12) with inversions /L-2 of Hunter and Connolly (1986) (limits 40B4— 42C1 inclusive) and JL-8 (limits 26C2- 37A2 inclusive), and with secondary nu- cleolar organizer expressed in section 41C— 42B; IIS standard for vernum sequence; IIL (Fig. 13) with inversion //L-8 (limits 60B— 63B inclusive); IIIS and IIIL standard for vernum sequence; sex chromosomes undif- ferentiated; floating inversions lacking. Types.— Holotype: ¢ with exuviae, trib- utary of Indian Creek, 1.0 km from entrance of Clemson University Experimental For- est, Pickens County, South Carolina (34°44'40’N x 82°50'51”W), 22-IT-86, col- lected by C. R. L. Adler. Allotype: 2 with exuviae, same data as holotype, 22-II-86, collected by C. R. L. Adler? Paratypes: SOUTH CAROLINA: PICKENS COUN- TY: unnamed spring-fed stream, Willow Wood subdivision, 5.5 km NE of center of Clemson University campus (34°42'25’N x 82°47'22”W), 17-XI-85, 5 larvae, C. R. L. Adler; 11-I-86, 17 larvae, 2 chromosome preparations (4 larvae), 1 6 with exuviae, | 2 with exuviae, C. R. L. and P. H. Adler; 19-I-86, 3 larvae, 2 pupae, | pupal exuviae, 1 2 with exuviae, C. R. L. and P. H. Adler; 25-I-86, 31 larvae, 3 pupae, | 2 with exu- viae, C. R. L. and P. H. Adler; 2-II-86, 15 larvae, 6 chromosome preparations (5 ¢ lar- vae, | 2? larva), 3 66 with exuviae, | 2 with exuviae, C. R. L. and P:; He Adler fi6-ie 86, 14 larvae, 8 chromosome preparations (2 6 larvae, 6 2 larvae) with photographic negatives and contact sheet, 3 6¢ with exu- viae, 3 9° with exuviae, C. R. L. and P. H. Adler; 2-III-86, 2 46 with exuviae, | ? with exuviae, C. R. L. and P. H. Adler; 16-III- 86, 1 larva, 2 66 with exuviae, C. R. L. and P. H. Adler; 19-X-86, 4 larvae, C. R. L. and P. H. Adler; Indian Creek, 0.5 km from entrance of Clemson University Experi- mental Forest, 22-II-86, 7 larvae, C. R. L. and P. H. Adler; 19-XII-86, 7 larvae, 1 chromosome preparation (2 larva), C. R. L. Adler; 6-I-87, 16 larvae, 3 chromosome preparations (2 larvae), 2 pupae, C. R. L. Adler; tributary of Indian Creek, 1.0 km from entrance of Clemson University Ex- perimental Forest, 22-II-86, 3 larvae, 1 chromosome preparation (4 larva), | pupa, C. R. L. and P. H. Adler; first stream past VOLUME 89, NUMBER 4 1g vl fe * if he 12 %, ‘es | ce 1 98 ba celave chap: \ ue Rigss 2= 13: He A “ee AWS ‘i Kwon: 679 , 40 Simulium loerchae, chromosomes of larvae from Pennsylvania (18 June 1986). 12, IL sequence with fixed inversion limits indicated by brackets, A = apolar diffuse group, C = centromere, NO = nucleolar organizer, 2NO = secondary nucleolar organizer, Z = ‘Z’ marker. 13, I[L sequence with fixed inversion limits indicated by bracket, P = DNA puff, PB = parabalbiani. Willow Spring, 2.2 km from entrance of Clemson University Experimental Forest, 22-II-86, 1 larva, C. R. L. Adler; second entrance to Dalton Road, 2.6 km from en- trance of Clemson University Experimental Forest, 22-II-86, 2 larvae, C. R. L. Adler; Hemlock Picnic Area, Table Rock State Park, 11-V-86, 84 larvae, 11 chromosome preparations (5 6 larvae, 6 2 larvae) with photographic negatives and contact sheet, 2 66 with exuviae, | 2 with exuviae, P. H. Adler; 21-III-87, 12 larvae, 5 pupae, 2 pupal exuviae, C. R. L. and P. H. Adler; White Oaks Picnic Area, Table Rock State Park, 18-V-86, 1 larva, P. H. Adler; OCONEE COUNTY: tributary of Chauga River, county road TU-37, 4-II-86, 3 larvae, | pu- pal exuviae, C. R. L. Adler; ANDERSON COUNTY: W tributary of Watermelon Creek, 0.4 km N of S-4-148, 11-XII-80, 5 larvae, J. S. Weaver and E. R. Taylor; 18- V-79, | larva, J.S. Weaver and E. R. Taylor; W tributary of Watermelon Creek, S of S-4- 63, 26-III-81, 1 larva, J. S. Weaver and E. R. Taylor; 15-XII-80, 2 larvae, J.S. Weaver and E. R. Taylor; Watermelon Creek, 0.6 km N of S-4-63, 3-IV-80, 1 larva, J. S. Weaver and E. R. Taylor; Browns Creek (E Rock Creek), 0.4 km N of S-4-147, 14-III- 79, 1 larva, J. S. Weaver and E. R. Taylor; E tributary of Rock Creek, N of S-4-670, 27-XI-80, 1 larva, J. S. Weaver and E. R. Taylor; jct. Rt. 412, W of Starr, 1.0 km E of Rt. 187, 8-II-87, 1 larva, 1 chromosome preparation (¢@ larva), P. H. Adler; Tangle- wood Spring, Pendleton, 9-II-87, 3 larvae, M. W. Heyn and C. Watson; PENNSYL- VANIA: CENTRE COUNTY: Sand Spring, 680 Shingletown Gap on Tussey Mountain, ca. 5 km SSE of State College, 31-V-86, 5 lar- vae, 2 chromosome preparations (1 ¢ larva, 1 2 larva) with photographic negatives and contact sheet, | pupa, S. Tessler; 18-VI-86, 11 larvae, 5 chromosome preparations (3 4 larvae, 2 2 larvae) with photographic neg- atives and contact sheets, S. Tessler; 16-IX- 86, 20 larvae, 2 chromosome preparations (2 larvae) with photographic negatives and contact sheet, 4 pupae (1 with pharate 3), S. Tessier: » TENNESSEE SULEIVAN COUNTY: East Inlet Creek, Bays Mt. Park, near Kingsport, 28-II-87, 3 larvae, 2 chro- mosome preparations (4 larvae), C. Watson; 15-III-87, 8 larvae, C. Watson. Additional specimens examined.— PENNSYLVANIA: CARBON COUNTY: Hickory Run State Park, 12-V-87, P. H. Adler, 3 larvae (third—fourth instars). Etymology.—This species is named in honor of my wife, Cynthia R. Loerch Adler, whose many collections of simuliids over the years resulted in discovery of the type locality. She subsequently collected and reared many of the specimens in the type series. Biology.—Simulium loerchae has been collected from the mountains of central Pennsylvania and eastern Tennessee and the mountains, foothills, and upper piedmont of South Carolina. In South Carolina, im- matures were collected from October to mid- May in permanent and temporary, sandy- bottomed, woodland streams less than a meter in width and ranging in temperature from 6.5° to 17°C (¥ = 12°C, n= 24). Stream conductivity averaged 22.7 + 10.4 uS (stan- dard deviation) and pH averaged 6.86 + 0.30 (n = 7). Pupae were found as early as January; adults emerged from these pupae within 24 h at room temperature. Larvae and pupae were collected most frequently from fallen leaves. Pupae were often ob- scured by sediment covering the leaves. Im- matures were collected with those of S. tuberosum Lundstr6m cytospecies FGH at all 1985-1986 sites, with Prosimulium mix- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tum Syme & Davies at half of the sites, with S. tuberosum cytospecies A at two of the sites, and once with S. impar Davies, Pe- terson, & Wood. One larva was infected with the fungus Coelomycidium simulii Debais- ieux. In Pennsylvania, S. /oerchae is apparently bivoltine. Immatures were collected in Centre County from a permanent, sandy- bottomed, woodland stream ranging in tem- perature from 9° to 15.5°C and not more than 0.5 m in width. Larval substrates in- cluded leaves, pebbles, and detritus. Pu- pation occurred on pebbles and on both sides of fallen leaves. Immatures were collected with those of P. fontanum Syme & Davies, S. gouldingi Stone, and S. venustum Say cy- tospecies CG. In Tennessee, larvae were collected from fallen leaves in a permanent stream ap- proximately 1.0 m wide, up to 30 cm deep, and with a temperature of 9.5°C. Associated simuliids included Stegopterna mutata (Malloch) triploid cytospecies and Prosi- mulium rhizophorum Stone & Jamnback. Females emerged with immature eggs and little stored nutrient. Ornithophily is in- ferred from the well-developed mouthparts and bifid claws. Remarks.—The pupa and female of S. loerchae are very similar to those of S. ca- ledonense Adler and Currie. The pupa of S. loerchae may be differentiated by the abun- dance of closely and uniformly set granules on the head and thorax. The female may be distinguished with difficulty from that of S. caledonense by the wider space between the hypogynial valves and the shape of the gen- ital fork. The larva may be identified prin- cipally by the color and distribution of ab- dominal pigment and the shape of the postgenal cleft. The large, subrectangular dorsal plate of the male is unique. Chromosomally, S. /oerchae is mono- morphic, possessing a primitive X,Y, Sex- chromosome system and lacking autosomal rearrangements. It can be derived from the vernum standard by three inversions, and VOLUME 89, NUMBER 4 can be recognized most easily by the DNA- puff inversion (limits 60B-63B). Simulium loerchae shares the fixed inversion /L-2 with North American S. gouldingi, S. croxtoni Nicholson & Mickel, and S. sp. of Hunter and Connolly (1986), as well as with Eu- ropean S. carpathicum Knoz and S. cry- ophilum Rubzov. Inversion JL-8, which re- orients the ‘Z’ marker and places the apolar diffuse group 26C2-27A (inclusive) in the distal half of IL, and inversion //L-S are both autapomorphic. ACKNOWLEDGMENTS I thank Fiona Hunter (University of To- ronto) for critically reviewing the cytologi- cal section, J. C. Morse (Clemson Univer- sity) for reviewing the manuscript, J. R. Brushwein (Clemson University) for pho- tographing the larva, S. Tessler (Pennsyl- vania State University) for providing Penn- sylvania specimens and accompanying biological information, and C. Watson (Clemson University) for providing Ten- nessee material and information. Technical 681 Contribution No. 2734 of the South Caro- lina Agricultural Experiment Station, Clem- son University. LITERATURE CITED Adler, P. H. and D. C. Currie. 1986. Taxonomic resolution of three new species near S7mulium ver- num Macquart (Diptera: Simuliidae). Can. Ento- mol. 118: 1207-1220. Brockhouse, C. L. 1985. Sibling species and sex chro- mosomes in Eusimulium vernum (Diptera: Sim- uliidae). Can. J. Zool. 63: 2145-2161. Davies, D. M., B. V. Peterson,and D.M. Wood. 1962. The black flies (Diptera: Simuliidae) of Ontario. Part 1. Adult identification and distribution with descriptions of six new species. Proc. Entomol. Soc. Ont. 92: 70-154. Hunter, F. F. and V. Connolly. 1986. A cytotaxo- nomic investigation of seven species 1n the Eusi- mulium vernum group (Diptera: Simuliidae). Can. J. Zool. 64: 296-311. Rothfels, K. H. and R. W. Dunbar. 1953. The sali- vary gland chromosomes of the black fly Simulium vittatum Zett. Can. J. Zool. 31: 226-241. Stone, A. and G. R. DeFoliart. 1959. Two new black flies from the western United States (Diptera, Sim- uliidae). Ann. Entomol. Soc. Am. 52: 394-400. PROC. ENTOMOL. SOC. WASH. 89(4), 1987, pp. 682-686 APICAL SENSILLA ON THE ADULT AND LARVAL LABIAL AND MAXILLARY PALPI OFODONTOTAENIUS DISJUNCTUS (ILLIGER) (COLEOPTERA: PASSALIDAE) GERALD T. BAKER Drawer EM, Mississippi State University, Mississippi State, Mississippi 39762. Abstract.—Three distinct types of sensory receptors are situated in the sensory field on the apex of the adult and larval labial and maxillary palpi of Odontotaenius disjunctus (Illiger) (Coleoptera: Passalidae). The coronal pegs are located on the periphery of the apical sensory field, and each one consists of a central disc and notched collar. There are short and long uniporous sensilla, which have a large apical pore, and short and long multi-porous sensilla. There are no differences between adults and larvae in the types and distribution of these sensilla, except in the total number of sensilla and the number of chemoreceptors on the apex of the palpi. Members of the family Passalidae exhibit a primitive type of social behavior and are large and easy to rear. These beetles have generated a considerable amount of re- search interest (Pearse et al., 1936; Gray, 1946; Krause and Ryan, 1953; Reyes-Cas- tillo and Jarman, 1980, 1983; Schuster and Schuster, 1985; Schuster, 1983). Both the larvae and adults are able to stridulate (Reyes-Castillo and Jarman, 1980, 1983), and this aspect and others of their behavior have received considerable attention (Gray, 1946; Mullen and Hunter, 1973; Schuster and Schuster, 1985). Although it is well doc- umented that cuticular sensilla play an im- portant role in the behavioral repertoire of insects (Kaissling, 1977; Seabrook, 1978; Zacharuk, 1980; Chapman, 1982; Bell and Cardé, 1984), only one study deals with the thin walled and porous sensilla on the an- tennal flagellum of a passalid beetle (Slifer and Sekhon, 1964). At the present time there are no papers that compare the sensilla on the larval and adult palpi of O. disjunctus or other beetle species. It is known that these sensilla on other beetles are involved in gus- tatory and olfactory responses (Bassler, 1958; Klinger, 1966; Kellogg, 1970; White et al., 1974). This paper compares the mor- phology, number, and distribution of sen- silla on the apex of the labial and maxillary palpi of larval and adult Odontotaenius dis- Junctus (Illiger). MATERIALS AND METHODS Adults and larvae were obtained from rot- ting logs. Labial and maxillary palpi were cut from the larval specimens and placed in fixative. The adults had to be dissected to enable sexing of the specimens before the palpi were fixed. The fixative contained 5% glutaraldehyde in 0.2 M Na-cacodylate buffer, pH 7.2, and the palpi were fixed for 24 h at 4°C. The palpi were washed in the same buffer and then post-fixed in 3% OsO, for 24 h at 4°C. The specimens were de- hydrated, critical-point dried and attached to stubs which were then coated with gold/ palladium. The material was examined with a Hitachi HHS-2R scanning electron mi- croscope at 20 kV. In order to identify the presence of porous VOLUME 89, NUMBER 4 683 Figs. 1-7. Palpal sensilla of Odontotaenius disjunctus. 1, Adult maxillary palpus. 2, Larval maxillary palpus. 3, Adult labial palpus. 4, Larval labial palpus. 5, Apex of short uniporous sensilla (arrows) and short multiporous sensilla. 6, Coronal sensillum. 7, Apex of a long uniporus sensillum (arrow = pore). C = collar; CD = central disc; CS = coronal sensillum; PP = multi-porous sensillum; UP = uniporous sensillum. cuticular areas, live adults and larvae were stained with the reduced silver nitrate tech- nique of Schafer and Sanchez (1976) and the crystal violet method of Slifer (1960). The labial and maxillary palpi of 10 adults and larvae were cleared and mounted in Hoyer’s medium. Measurements were ob- tained by using a light microscope with an ocular micrometer and are given as an av- erage plus the range. The data on the num- 684 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Comparison of the total number of sensilla on the apex of the maxillary and labial palpi. Mean + SD Adult maxillary palpus 32.30 + 0.82 a Adult labial palpus 28.20 + 0.79 b Larval maxillary palpus DOES OOPS 2iC Larval labial palpus 18.70 + 0.67 d Means not followed by the same letter are signifi- cantly different (P < 0.01) as determined by ANOVA followed by Student-Newman-Keuls Test (n = 10). ber of apical sensilla were subjected to anal- ysis of variance followed by Student- Newman-Keuls Test (P < 0.01). RESULTS AND DISCUSSION The apical sensilla on the labial and max- illary palpi of O. disjunctus can be divided into three distinct types, coronal sensilla, uniporous sensilla and multi-porous sensil- la. The first type, coronal sensilla, are sit- uated on the periphery of the sensory field on the apex of the labial and maxillary palpi (Figs. 1—4, 6), and encircle the other sensilla. This sensillum is 5.5 wm wide at the base and 7.1 um (6.6—7.4) long. There are two distinct morphological features of this sen- sillum; the first is the central disc (Fig. 6) and the second 1s the collar which surrounds the disc. The disc is 2.2 um (1.9-2.4) wide and 3.1 wm (2.8-3.2) long, and the basal portion is rounded whereas the apex Is nip- ple-like (Fig. 6). The collar of the coronal sensilla is higher in the inner surface than the outer surface where it has a well-defined notch (Figs. 1, 3, 6). The collar portion dif- fers greatly from the collar found on coronal sensilla of other beetles which have a collar of uniform length and no frontal notch (Doane and Klinger, 1978; Whitehead, 1981; Hallberg, 1982). From ultrastructural information (Whitehead, 1981; Hallberg, 1982) and their non-porosity as indicated by the lack of staining with reduced silver nitrate and crystal violet, this type of sen- sillum is considered to be a mechanorecep- tor. The coronal sensillum is not found on Table 2. Comparison of the total number of chem- oreceptors on the apex of the maxillary and labial palpi. Mean + SD Adult maxillary palpus 21.90 + 0.74a Adult labial palpus 70S 057 b Larval maxillary palpus 14.20 + 0.63 ¢ Larval labial palpus 11.00 + 0.67 d Means not followed by the same letter are signifi- cantly different (P < 0.01) as determined by ANOVA followed by Student-Newman-Keuls Test (n = 10). all beetle larvae and adults, for example Lyctus brunneus (Stephens) (Iwata and Nishimoto, 1981) and Sitophilus oryzae (L.) (Speirs et al., 1986), but when it is situated on the palpi, this receptor is found on the periphery of the apex (Doane and Klinger, 1978; Hallberg, 1982). Within the circle of coronal sensilla there are two other types of sensilla, uniporous and multi-porous pegs (Figs. 1, 2, 4). The uniporous pegs can be further subdivided into short 9.5 wm (9.3-10.2) and long 19.7 um (19.4—20.7) pegs that are 5.3 wm (5.0- 5.5) wide at the base. These sensilla have a tapered apex with a large, single pore (Figs. 1, 3, 5, 7). Uniporous sensilla are found on the palpal apex of scolytids. Their ultra- structure indicates that this type of sensil- lum is a mechano- and contact chemo-re- ceptor (Whitehead, 1981; Hallberg, 1982). The multi-porous pegs may be divided into short 8.9 um (8.7-9.4) and long 19.3 um (18.8-19.6) and are 4.9 um (4.7-5.3) wide at the base. This type of sensillum has a blunt, rounded apex as compared to the un- iporous pegs (Figs. 1, 5). The ultrastructural information on this sensillum from other beetles indicates that it may be an olfactory receptor (Whitehead, 1981; Hallberg, 1982). Both types of the above-mentioned sensilla stain with crystal violet and reduced silver nitrate. This demonstrates their porosity and is an indication of a possible chemorecep- tor. The sensilla on the apex of the palpi of other beetles are known to respond to a wide variety of olfactory and gustatory stimuli VOLUME 89, NUMBER 4 (Mitchell and Schoonhoven, 1974; White et al., 1974; Doane and Klinger, 1978). There are no differences between adult females and males in the morphology, num- ber and distribution of the apical sensilla. There is no difference in the morphology and distribution of the sensilla between adults and larvae, but there is a distinct dif- ference in their number (Tables 1, 2). The adult maxillary palpus has the greatest total number of apical sensilla and the larval la- bial palpus the least (Table 1). This is also observed for the total number of chemo- receptors (Table 2). The order for the total number of sensilla and chemo-sensilla would be the following: adult maxillary palpus > adult labial palpus > larval maxillary pal- pus > larval labial palpus. This distinct di- chotomy may be due to the more complex behaviors of the adults, such as mating, host finding, aggression and brood care (Mullen and Hunter, 1973; Schuster and Schuster, 1985). ACKNOWLEDGMENTS I thank S. B. Ramaswamy, G. Tyson, D. Adamski, and an anonymous reviewer for reviewing the manuscript and B. Perrigin for typing it. This paper is assigned No. 6630 of the Mississippi Agriculture and Forestry Experiment Station. LITERATURE CITED Bassler, U. 1958. Ubersuche zur Orientierung der Stechmuken die Schwarmbildung und die Bedeu- tung des Johnstonschen Organs. Z. Vgl. Physiol. 41: 300-330. Bell, W. J. and R. Cardé. 1984. Chemical Ecology of Insects. Chapman and Hall Ltd., London. 524 pp. Chapman, R. F. 1982. Chemoreception: The signif- icance of receptor numbers. Adv. Insect Physiol. 16: 247-356. Doane, J. and J. Klinger. 1978. Location of CO,- receptive sensilla on the larvae of the wireworms Agriotes lineatus-obscurus and Limonius califor- nicus. Ann. Entomol. Soc. Am. 71: 357-363. Gray, I. E. 1946. Observations on the life history of the horned passalus. Am. Mel. Nat. 35: 728-746. Hallberg, E. 1982. Sensory organs in [ps typographus (Insecta: Coleoptera). Fine structure of the sensilla 685 of the maxillary and labial palps. Acta Zool. 63: 191-198. Iwata, R. and K. Nishimoto. 1981. Observations on the external morphology and the surface structure of Lyctus brunneus (Stephens) (Coleoptera: Lyc- tidae) by scanning electron microscopy. Kontyu (Tokyo) 49: 542-557. Kaissling, E. 1977. Control of insect behavior via chemoreceptor organs, pp. 45-65. Jn Shorey, H. H. and J. J. McKelvey, eds., Chemical Control of Insect Behavior: Theory and Application. John Wiley & Sons, Inc., New York. Kellogg, F. 1970. Water vapour and carbon dioxide receptors in Aedes aegypti. J. Insect Physiol. 16: 99-108. Klinger, J. 1966. Uber den Sitz der CO,— Rezeptoren bei der Larve von Otiorrhynchus sulcatus. Ento- mol. Exp. Appl. 9: 271-277. Krause, J. and M. Ryan. 1953. The stages of devel- opment in the embryology of the horned Passalus beetle, Popilius disjunctus Iliger. Ann. Entomol. Soc. Am. 47: 1-20. Mitchell, B. and L. Schoonhoven. 1974. Taste re- ceptors in Colorado beetle larvae. J. Insect Phys- iol. 20: 1787-1793. Mullen, V. and P. Hunter. 1973. Social behavior in confined populations of the horned passalus beetle (Coleoptera: Passalidae). J. Ga. Entomol. Soc. 8: 115-123. Pearse, A., M. Patterson, J. Rankin, and G. Wharton. 1936. The ecology of Passalus cornutus F., a bee- tle which lives in rotting logs. Ecol. Monogr. 6: 456-490. Reyes-Castillo, P. and M. Jarman. 1980. Some notes on larval stridulation in neotropical Passalidae (Coleoptera: Lamellicornia). Coleopt. Bull. 34: 263-270. 1983. Disturbance sounds of adult passalid beetles (Coleoptera: Passalidae): Structural and functional aspects. Ann. Entomol. Soc. Am. 76: 6-22. Schafer, R. and T. Sanchez. 1976. The nature and development of sex attractant specificity in cock- roaches of the genus Periplaneta. I, Sexual di- morphism in the distribution of antennal sense organs in five species. J. Morphol. 149: 139-158. Schuster, J. 1983. Acoustical signals of passalid bee- tles: Complex repertoires. Fla. Entomol. 66: 486- 496. Schuster, J. and L. Schuster. 1985. Social behavior in passalid beetles (Coleoptera: Passalidae): Co- operative brood care. Fla. Entomol. 68: 266-272. Seabrook, W. 1978. Neurobiological contributions to understanding insect pheromone systems. Ann. Rev. Entomol. 23: 471-485. Slifer, E. 1960. A rapid and sensitive method for 686 identifying permeable areas in the body wall of insects. Entomol. News 71: 179-182. Slifer, E. and S. Sekhon. 1964. Fine structure of the thin walled sensory pegs on the antenna ofa beetle, Popilius disjunctus (Coleoptera: Passalidae). Ann. Entomol. Soc. Am. 57: 541-548. Speirs, R., G. White, and J. Wilson. 1986. SEM ob- servations of rice weevil larvae, Sitophilus oryzae (L.) (Coleoptera: Curculionidae). J. Kans. Ento- mol. Soc. 59: 390-394. White, R., U. Paim, and W. Seabrook. 1974. Max- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON illary and labial sites of carbon dioxide sensitive receptors of larval Orthosoma brunneum. J. Comp. Physiol. 88: 235-246. Whitehead, A. 1981. Ultrastructure of sensilla on the female mountain pine beetle, Dendroctonus pon- derosae Hopkins (Coleoptera: Scolytidae). Int. J. Insect Morphol. & Embryol. 10: 19-28. Zacharuk, R. 1980. Ultrastructure and function of insect chemosensilla. Ann. Rev. Entomol. 25: 27- 47. PROC. ENTOMOL. SOC. WASH. 89(4), 1987, pp. 687-700 PRACTICAL DIAGNOSIS AND NATURAL HISTORY OF TWO SIBLING SALT MARSH-INHABITING PLANTHOPPERS IN THE GENUS PROKELISIA (HOMOPTERA: DELPHACIDAE) ROBERT F. DENNO, MICHAEL E. SCHAUFF, STEPHEN W. WILSON, AND KAREN L. OLMSTEAD (RFD, KLO) Department of Entomology, University of Maryland, College Park, Mary- land 20742; (MES) Systematic Entomology Lab, BBII, Agricultural Research Service, USDA, % U.S. National Museum NHB 168, Washington, D.C. 20560: (SWW) Depart- ment of Biology, Central Missouri State University, Warrensburg, Missouri 64093. Abstract.—Two salt marsh-inhabiting planthoppers, Prokelisia marginata (Van Duzee) and Prokelisia dolus (Wilson), have been the subjects of extensive ecological research. In the past these species have been confused taxonomically because they are difficult to differentiate, particularly the females and nymphs. In this report we show that males can be separated on the basis of style shape; the styles of P. marginata project dorsally and are caliper shaped while those of P. dolus are flared posteriorly and are avicephaliform in shape. Analysis of morphometric data from laboratory-reared isolines as well as field- collected specimens of each species reveals that females and nymphs are separable using frons morphology. The frons of P. marginata is proportionally longer and narrower than that of P. dolus. Also, macropters have a slightly longer and narrower frons than bra- chypters regardless of species. Last, we review the natural history, host plant relationships, and population biology of these two congeneric planthoppers. Planthoppers in the genus Prokelisia have been the focus of extensive ecological study. For example, Prokelisia marginata (Van Duzee) has been used to study plant-her- bivore interactions (Denno, 1977, 1983; Tallamy and Denno, 1979; Vince et al., 1981; Denno et al., 1986), migration and population dynamics (Denno, 1979, 1983, 1985; Denno and Grissell, 1979; Denno et al., 1980; Denno et al., in prep.; Roderick, 1987), parasite-host interactions (Stiling and Strong, 1982a, b), predator-prey relation- ships (Débel and Denno, in prep. a, b), life history theory (Denno and Dingle, 1981; Denno and McCloud, 1985, in prep.), bio- geography (Rey, 1981), and wing polymor- phism (Denno, 1976, 1978; McCoy and Rey, 1981; Strong and Stiling, 1983; Denno et al., 1985; Roderick, 1987). There are several reasons why these non- pest sapfeeders have received so much sci- entific attention. First, planthoppers in the genus Prokelisia are extremely abundant on mid-Atlantic coast salt marshes. They com- prise 95-99% of all herbivores sampled on their host grass, Spartina alterniflora Loisel (Poaceae) (Denno, 1976, 1977; Tallamy and Denno, 1979). Second, Spartina dominates the vegetation of North American intertidal marshes where it commonly occurs as ex- tensive, pure stands (Adams, 1963; Blum, 1968; Redfield, 1972; Duncan, 1974). These relatively simple, naturally occurring monocultures are attractive for studying 688 plant-herbivore interactions in the absence of mixed vegetational effects. Third, these planthoppers exhibit wing dimorphism; there are fully winged macropterous adults and flightless brachypterous adults, allow- ing for the easy visual identification of mi- gratory and sedentary forms for population studies (Denno, 1976; Denno and Grissell, 1979; Wilson, 1982; Denno et al., 1985). Fourth, intertidal marshes export minerals and organic nutrients and, in doing so, serve as nursery grounds for commercially im- portant coastal fish and shellfish (McHugh, 1966: Gosselink et al., 1973). Thus, it is important to determine if herbivores such as planthoppers significantly reduce Spar- tina production and influence the energetics of the system. Last, the study of planthop- pers in the genus Prokelisia takes on added significance because some species are eco- logically very similar to pest planthoppers like Nilaparvata lugens (Stal.), a major threat to rice production in Asia (Brady, 1979; Kisimoto, 1981; Heinrichs et al., 1982; Kenmore et al., 1984; Wilson and Claridge, 1985). Due to the widespread interest in these planthoppers as subjects for ecological re- search, it is especially important to identify them accurately. Thus, the primary objec- tive of this report is to elaborate on biosys- tematics, provide field identification char- acters, and update the natural history and host plant relationships for the two conge- neric salt marsh-inhabiting planthoppers, P. marginata and P. dolus. The diagnosis of these two species is particularly timely be- cause P. dolus is a recently described species (Wilson, 1982), which has been confused with P. marginata in the past, and females and nymphs in particular are difficult to identify. METHODS To obtain pure laboratory cultures of P. marginata and P. dolus for genitalic and morphometric analysis, we collected adults of both species from short-form S. alter- niflora meadows on June 26, 1986 at Tuck- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON erton, Ocean County, New Jersey. Isolines for each species were established by trans- ferring two males and one female onto a seedling of S. al/terniflora enclosed in a plas- tic tube cage (see Denno et al., 1985, 1986 for details on Spartina culture and cage de- sign). Thirty isolines were set up for each species in the laboratory. Oviposition and development occurred and after five weeks F, adults began to emerge. The genitalia of all emerging male offspring were examined to verify the purity of the lines. Male and female offspring of 26 successful P. margin- ata isolines and 21 P. dolus isolines were preserved in 70% ethanol for electron mi- croscopy and morphometric analysis. Scanning electron micrographs were tak- en of the heads (frontal view) and male gen- italia (posterior and lateral views) of P. mar- ginata and P. dolus. We examined both macropters and brachypters of each species from New Jersey and Florida (Gulf) pop- ulations. Specimens were dissected, mount- ed on metal stubs, and coated with carbon followed by gold-palladium. They were photographed at 10 Kv in a Cambridge 100 Stereoscan microscope. Wilson (1982) reported that the frons of P. marginata was generally longer and nar- rower than that of P. dolus. To quantify this observation, frons measurements for mac- ropters and brachypters of both species were made under a stereomicroscope fitted with a filar ocular micrometer. Specimens were oriented so that the plane of the frons was parallel to that of the microscope stage. Frons length was determined by measuring the distance between the frontoclypeal su- ture and the vertex. Frons width was deter- mined by measuring its widest point. Frons metrics were expressed as a ratio of frons length to width. From the isolines, approx- imately twenty adults of each sex and wing form and ten nymphs were measured for both species. To determine differences in frons proportion between species, wing forms and sexes, data were examined using analysis of variance (ANOVA, General Lin- ear Model; SAS Institute, 1982). To deter- VOLUME 89, NUMBER 4 689 Figs. 1-8. Posterior view of the genital capsule of male Prokelisia marginata (1-4) and P. dolus (5-8). Macropter, New Jersey (1 and 5); Macropter, Florida (2 and 6); Brachypter, New Jersey ( 3 and 7); Brachypter, Florida (4 and 8). Note the caliper shape of the styles of P. marginata and the avicephaliform shape of the styles of P. dolus. Bars on scanning electron micrographs are 0.1 mm. 690 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 9, 10. Lateral view of the styles of male Prokelisia marginata (9) and P. dolus (10). The styles of P. marginata project straight up, while those of P. dolus are posteriously flared. Bars on scanning electron micro- graphs are 0.1 mm. mine if a skilled technician could accurately identify field-collected material to species, approximately 25 adults of each sex and wing form and 25 fifth instar nymphs for both species were culled from field samples (Tuckerton, N.J.) and sorted to species on the basis of frons shape. Frons measure- ments were made on these specimens and the resulting frequency distribution was compared to the isoline distribution by AN- OVA. The origin (field or isoline) of the populations was entered as a source of vari- ation in the ANOVA model. A significant effect of origin would indicate an inaccurate identification of field-collected specimens. RESULTS AND DISCUSSION Genitalic distinctness of males.—Al- though the gross anatomy of P. marginata and P. dolus is very similar, males are easily separated by the shape of their styles (= claspers) (Wilson, 1982). The styles of P. marginata are caliper shaped and taper gradually from the base to the apex (Figs. 1-4). By contrast, the styles of P. dolus are avicephaliform and expand distally with an- gulate outer margins (Figs. 5-8). In lateral view, the styles of P. marginata are straight and project dorsally (Fig. 9), whereas those of P. dolus are posteriorly flared (Fig. 10). These genitalic characters remain distinct throughout the entire geographic distribu- tion of both species and are not influenced by wing form (see New Jersey-Florida and macropter-brachypter comparisons in Figs. 1-8). In the thousands of male specimens examined, we never observed a genitalic in- termediate (Wilson, 1982; this report). Style characters are readily visible under a ste- reomicroscope, allowing for the rapid iden- VOLUME 89, NUMBER 4 Figs. 11-14. Frontal view of the frons of adult male Prokelisia marginata (11) and P. dolus (12). Bars on electron micrographs are 0.5 mm. Frontal view of the frons of the fifth instar nymph of P. marginata (13) and P. dolus (14). Bars on electron micrographs are 0.2 mm. tification of males in field-collected samples of planthoppers. Additional genitalic differ- ences include the shape and dentition of the aedeagus (Wilson, 1982), slight differences in the shape of the pygofer (Figs. 1-10), and rugose sculpturing over the entire surface of the styles of P. dolus (Figs. 1-8). The apical third of styles of P. marginata lack rugose sculpturing (Figs. 1-8). These differences, although subtle, are no less so than those observed in the extensively studied Muel- lerianella complex (Drosopoulos, 1977; Boo, 1981). Morphometric analysis of the frons.— Whereas genitalic characters can be used readily to separate males of P. marginata and P. dolus, they are not useful for distin- guishing between females. Wilson (1982) re- ports that P. marginata generally can be separated from P. dolus by its longer and narrower frons (compare Figs. 11 with 12 and 13 with 14). Here we pursue more rig- orously the ratio of frons length/frons width as a diagnostic character for identifying fe- males, males and fifth instar nymphs of the two Prokelisia species. The frequency distributions of the frons length/frons width ratio overlap very little between P. marginata and P. dolus adults from laboratory isolines (Fig. 15 black box- es only). Mean ratios for females of P. mar- ginata were 1.810 for brachypters and 1.879 for macropters. Mean frons ratios for P. do- lus were 1.552 and 1.628 for brachypters and macropters, respectively, documenting the longer and narrower frons of P. mar- ginata (Table 1). Analysis of variance on individuals from isolines found a highly sig- nificant effect (P < 0.0001) of species and wing form on frons ratio (Table 2). Not only is the frons of P. marginata longer and nar- rower than the frons of P. dolus, but mac- 692 Frequency 15 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON GJ Pmarginata Mo P marginata Bo CO Os P marginata M®@ _— P marginata B® — WwW O Pdolus Mo& — WW Pdolus Bo peneawereer | => 5 3 P dolus MQ 1 Pdolus BQ 5S) 1.6 7. 1.8 19 2.0 Frons Length/ Frons Width Fig. 15. Frequency distribution of the ratio of frons length/frons width for the wing forms (M = Macropter, B = Brachypter) and sexes of Prokelisia marginata and P. dolus from laboratory isolines (black boxes) and field samples collected at Tuckerton, New Jersey (open boxes). ANOVA finds a significant (P < 0.0001) effect of species and wing form on frons ratio corroborating the longer and narrower frons of P. marginata compared to P. dolus, and the slightly longer and narrower frons of macropters compared to brachypters. VOLUME 89, NUMBER 4 693 Table 1. Mean frons length/frons width ratios with standard errors and sample sizes for the sexes and wing forms of Prokelisia marginata and P. dolus from laboratory isolines. The higher ratios observed in P. marginata document its long, narrow face compared to P. dolus. Species Sex Wingform Mean SE n P. marginata male macropter 1.903 .0176 PD| P. marginata male brachypter a2: .0473 4 P. marginata female macropter 1.879 O19] 20 P. marginata female brachypter 1.810 .0237 18 P. dolus male macropter 1.616 .0193 20 P. dolus male brachypter 1.597 .0169 22 P. dolus female macropter 1.628 .0143 20 P. dolus female brachypter 1.552 .0168 20 ropters of both species have a longer and narrower frons than brachypters. Of the ex- plainable variation in adult frons ratio, however, 89% of it is attributable to species effects, while wing form accounts for only 8% (Table 3). Fifth instar nymphs of P. marginata from isolines also have a significantly (P < 0.0001) longer and narrower frons (1.441 + 0.021, xX + SE; n = 10) than nymphs of P. dolus @204° 5°0.017,% "SE; n =-10):(Fig. 16 black boxes only, Table 4). Thus, by using individuals from pure isolines in the labo- ratory we have determined that most fe- males and fifth instar nymphs can be ac- curately identified to species by the relative length and width of the frons. A second objective of our study was to determine if a skilled technician could ac- curately and quickly sort field-collected adults (particularly females) and nymphs to species. The rapid and accurate identifica- tion of field-collected material is essential for population studies of planthoppers. To determine if this objective was possible, we compared the frequency distribution of the frons length/frons width ratio of individual females originating from isolines in the lab- oratory (black boxes in Fig. 15) with the frons ratio distribution of field-collected fe- males (open boxes in Fig. 15) that were sort- ed to species by eye under the microscope. There were no significant effects of origin (isoline vs. eye-sorted field sample) on frons ratio variation, and only species and wing form contributed significantly (P < 0.0001) as before (Table 5). These results strongly suggest that a trained technician can accu- rately sort females of Prokelisia to species. This procedure was also conducted on fifth instar nymphs with the same result. They too can be accurately sorted to species by Table 2. Analysis of variance of the frons length to width ratios of adult Prokelisia marginata and P. dolus reared from laboratory isolines. Source of Variation df MS F P Species 1.5784 233.02 .000 1 ** Sex 1 0006 09 .7698 Species x Sex l 0037 54 .4621 Wingform | 1454 21.47 .000 1** Species x Wingform l 0183 2.70 .1025 Sex x Wingform 1 .0000 .0O 9495 Species x Sex x Wingform 1! .0231 3.41 .0668 Error 137, .0068 ** Highly significant. 694 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 3. Analysis of variance results (% total sums of squares) of the frons length to width ratios of adult Prokelisia marginata and P. dolus reared from labo- ratory isolines. Source of Variation df % Total SS Species 1 SS 512s* Sex | .022 Species x Sex l Ala Wingform 1 5391" Species x Wingform I .007 Sex x Wingform ] .000 Species x Sex 80%) of macropters, while brachyptery predominates ( > 70%) in the P. dolus populaticn. upper surface of blades. In mid-Atlantic populations, active nymphs overwinter in litter or rolled dead leaves of standing vege- tation (Denno, 1977). Tallamy and Denno (1979) reported that if litter is removed in autumn, significantly smaller populations occur the following season, suggesting that litter is a critical resource for winter surviv- al. The most striking ecological difference between P. marginata and P. dolus is the wing form composition of populations along the Atlantic coast where both species are so abundant. Populations of P. marginata are 698 composed of >80% macropters, while the proportion is reversed in P. dolus with macroptery rarely exceeding 20% (Wilson, 1982: Denno, 1985; Denno et al., 1985). The discrepancy in wing form composition of populations reflects a major difference in mobility between the two species. P. mar- ginata is a migratory species on the Atlantic coast, undergoing annual interhabitat movements between overwintering habitats on the high marsh (short-form Spartina) and better summer sites for development on the low marsh (tall-form Spartina) (Denno and Grissell, 1979; Denno, 1983, 1985; Denno et al., 1985). By moving to low marsh sites during the summer, immigrant planthop- pers encounter a much more nutritious host plant than they would had they remained on the high marsh. For P. marginata, feed- ing on a more nutritious host results in larg- er females with increased fecundity (Denno and McCloud, 1985; Denno et al., 1986). Denno et al. (1986) argue that the contrast in Spartina nutrition between high and low marsh habitats as well as differential winter survival in these habitats have been major selective forces behind the evolution of mi- gration in P. marginata. By contrast, P. do- lus is a very sedentary species, remaining mostly on the high marsh in short-form Spartina throughout the entire year (Denno et al., 1985; Denno et al., in prep.). Cur- rently, far less is known about the popula- tion dynamics of P. dolus. Along the Gulf Coast, populations of both species are composed on average of 90% brachypters, a situation very different from that along most of the Atlantic coast (Denno and Grissell, 1979; McCoy and Rey, 1981; Wilson, 1982; Strong and Stiling, 1983; Denno, 1985). Explanations for the high levels of brachyptery in Gulf Coast popu- lations of P. marginata include the persis- tence and homogeneity of Spartina stands (Denno and Grissell, 1979; Denno, 1983, 1985), the isolation of Spartina patches (McCoy and Rey, 1981), and the selective PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON loss of dispersing macropters from islets of Spartina (Strong and Stiling, 1983). For Prokelisia marginata and P. dolus there appears to be a strong environmental component involved in wing form deter- mination, particularly for females (Denno et al., 1985; Denno, unpub. data; Roderick, 1987). Various environmental conditions experienced during nymphal development act as cues, triggering a developmental switch that selects one of several alternative mor- phogenic pathways (Lees, 1961, 1966; Mochida, 1973; Denno et al., 1985). Of the cues investigated, crowding has by far the most significant effect on wing form, al- though host plant nutrition and tempera- ture have an influence as well (Denno et al., 1985; Denno, unpub. data). At high den- sitites on poor quality hosts most adults molt to macropters, enabling them to escape de- teriorating host plants. However, the threshold density that triggers the produc- tion of macropters is much lower for P. marginata than P. dolus. This provides a proximate explanation for the much higher levels of macroptery in P. marginata com- pared to P. dolus populations that occur in the mid-Atlantic states (Denno, 1985; un- pub. data). ACKNOWLEDGMENTS We sincerely appreciate the suggestions of David Dussourd, Eric Grissell, Charles Mitter, George Roderick and Norman Woodley, who reviewed an early draft of this report. This research was supported by National Science Foundation Grants BSR- 8206603 and BSR-8614561. This is Scientific Article Number A-4600, Contri- bution Number 7596 of the Maryland Ag- ricultural Experiment Station, Department of Entomology. LITERATURE CITED Adams, D. A. 1963. Factors influencing vascular plant zonation in North Carolina salt marshes. Ecol. Mon. 44: 445-456. VOLUME 89, NUMBER 4 Blum, J. L. 1968. Salt marsh spartinas and asso- ciated algae. Ecol. Mon. 38: 199-221. Booij, K. 1981. Biosystematics of the Muellerianella complex (Homoptera, Delphacidae), taxonomy, morphology and distribution. Netherlands J. Zool. 31: 572-595. Brady, N. C. 1979. Brown Planthopper: Threat to Rice Production in Asia. International Rice Re- search Institute, Manila, Philippines. Bush, G. L. 1975. Sympatric speciation in phytoph- agous parasitic insects, pp. 187-206. Jn Price, P. W., ed., Evolutionary Strategies of Parasitic In- sects and Mites. Plenum, New York. Bush, G. L.andS.R. Diehl. 1982. Host shifts, genetic models of sympatric speciation, and the origin of parasitic insect species. Pp. 297-306. Jn Visser, J. H.and A. K. Minks, eds., Insect and Host Plants— 5th International Symposium on Insect-Plant Re- lationships, Wageningen, Netherlands. Denno, R. F. 1976. Ecological significance of wing- polymorphism in Fulgoroidea which inhabit salt marshes. Ecol. Entomol. 1: 257-266. . 1977. Comparison of the assemblages of sap- feeding insects (Homoptera— Hemiptera) inhabit- ing two structurally different salt marsh grasses in the genus Spartina. Environ. Entomol. 6: 359- Bis 1978. The optimum population strategy for planthoppers (Homoptera: Delphacidae) in stable marsh habitats. Can. Entomol. 110: 135-142. 1979. The relation between habitat stability and the migration tactics of planthoppers. Misc. Publs. Entomol. Soc. Am. 11: 41-49. 1983. Tracking variable host plants in space and time, pp. 291-341. Jn Denno, R. F. and M. S. McClure, eds., Variable Plants and Herbivores in Natural and Managed Systems. Academic Press, New York. . 1985. Fitness, population dynamics and mi- gration in planthoppers: The role of host plants, pp. 623-640. Jn Rankin, M. A., ed., Migration: Mechanisms and Adaptive Significance. Contri- butions in Marine Science Vol. 27, Marine Science Institute, the University of Texas at Austin, Port Aransas, Texas. Denno, R. F. and H. Dingle. 1981. Considerations for the development of a more general life history theory, pp. 1-6. Jn Denno, R. F. and H. Dingle, eds., Insect Life History Patterns: Habitat and Geographic Variation. Springer-Verlag, New York. Denno, R. F. and E. E. Grissell. 1979. The adap- tiveness of wing-dimorphism in the salt marsh- inhabiting planthopper, Prokelisia marginata (Homoptera: Delphacidae). Ecology 60: 221-236. Denno, R. F., L. W. Douglass, and D. Jacobs. 1985. Crowding and host plant nutrition: Environmental 699 determinants of wing-form in Prokelisia margin- ata. Ecology 66: 1588-1596. 1986. Effects of crowding and host plant nu- trition on a wing-dimorphic planthopper, Proke- lisia marginata. Ecology 67: 116-123. Denno, R. F. and E. S. McCloud. 1985. Predicting fecundity from body size in the planthopper, Pro- kelisia marginata (Homoptera: Delphacidae). En- viron. Entomol. 14: 846-849. . In prep. Reproductive cost of flight capability in the wing-dimorphic planthopper Prokelisia marginata. Denno, R. F., M. J. Raupp, D. W. Tallamy, and C. F. Reichelderfer. 1980. Migration in heterogeneous environments: Differences in habitat selection be- tween the wing-forms of the dimorphic planthop- per, Prokelisia marginata (Homoptera: Delphac- idae). Ecology 61: 859-867. Denno, R. F., K. L. Olmstead, H. Dobel, and L. Hanks. In prep. Comparative population dynamics of two salt marsh-inhabiting planthoppers in the genus Prokelisia. Dobel, H. and R. F. Denno. In prep. a. The functional response of Pardosa spiders to increases in plant- hopper density: A comparison of planthopper species and wing-forms. . In prep. b. Numerical responses of Pardosa spiders to planthopper prey. Drosopoulos, S. 1977. Biosystematic studies on the Muellerianella complex (Delphacidae, Homop- tera, Auchenorrhyncha). Meded. Land. Wagen- ingen 77: 1-133. Duncan, W. H. 1974. Vascular halophytes of the At- lantic and Gulf Coasts of North America north of Mexico, pp. 23-50. Jn Reimnold R. J. and W. H. Queen, eds., Ecology of Halophytes. Academic Press, New York. Gosselink J. G., E. D. Odum, and R. M. Pope. 1973. The value of the tidal marsh. Urban and Regional Development Center, University of Florida. Work Paper No. 3, 32 pp. Gould, F. 1983. Genetics of plant-herbivore systems: Interactions between applied and basic study, pp. 599-653. In Denno, R. F. and M. S. McClure, eds., Variable Plants and Herbivores in Natural and Managed Systems. Academic Press, New York. Heinrichs, E. A., W. H. Reissig, S. L. Valencia, and S. Chelliah. 1982. Rates and effects of resurgence- inducing insecticides on Nilaparvata lugens (He- miptera: Delphacidae) and its predators. Environ. Entomol. 11: 1269-1273. Kenmore, P. E., F. O. Carino, C. A. Perez, V. A. Dyck, and A. P. Gutierrez. 1984. Population regulation of the rice brown planthopper (Ni/aparvata lugens Stal) within rice fields in the Philippines. J. PI. Prot. Trop. 1: 19-37. 700 Kisimoto, R. 1981. Development, behavior, popu- lation dynamics and control of the brown plant- hopper, Nilaparvata lugens Stal. Rev. Plant Prot. Res. 14: 26-58. Lees, A. D. 1961. Clonal polymorphism in aphids. In Kennedy, K. S. ed., Insect Polymorphism. Symp. R. Entomol. Soc. Lond. 1: 68-79. _ 1966. The control of polymorphism in aphids. Adv. Ins. Physiol. 3: 207-277. McCoy, E. D. and J. R. Rey. 1981. Alary polymor- phism among the salt marsh Delphacidae (Ho- moptera: Fulgoroidea) of northwest Florida. Ecol. Entomol. 6: 285-291. McHugh, J. L. 1966. Management of estuarine fish- eries. Am. Fish Soc. Spec. Publ. No. 3, pp. 133- 134. Mitter, C.and D. J. Futuyma. 1983. Anevolutionary- genetic view of host-plant utilization by insects, pp. 427-459. In Denno, R. F. and M. S. McClure, eds., Variable Plants and Herbivores in Natural and Managed Systems. Academic Press, New York. Mochida, O. 1973. The characters of the two wing- forms of Javesella pellucida (F.) (Homoptera: Del- phacidae), with special reference to reproduction. Trans. R. Entomol. Soc. Lond. 125: 177-225. Redfield, A.C. 1972. Development ofa New England salt marsh. Ecol. Mon. 42: 201-237. Rey, J. R. 1981. Ecological biogeography of arthro- pods on Spartina islands in Northwest Florida. Ecol. Mon. 51: 237-265. Roderick, G. 1987. Ecology and evolution of migra- tion and dispersal in a salt marsh insect. Ph.D. Dissertation, Univ. of California, Berkeley. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON SAS Institute Inc. SAS User’s Guide. 1982. SAS Inst. Inc., Caty,NG: Stiling, P. D. and D. R. Strong. 1982a. Parasitoids of the planthopper, Prokelisia marginata (Ho- moptera: Delphacidae). Fla. Entomol. 65: 191- 192. 1982b. Egg density and the intensity of par- asitism in Prokelisia marginata (Homoptera: Del- phacidae). Ecology 63: 1630-1635. Strong, D. R. and P. D. Stiling. 1983. Wing dimor- phism changed by experimental density manipu- lation in a planthopper Prokelisia marginata (Ho- moptera: Delphacidae). Ecology 64: 206-209. Tabashnik, B. E. 1983. Host range evolution: The shift from native legumes to alfalfa by the butterfly Colias philodice eriphyle. Evolution 37: 150-162. Tallamy, D. W. and R. F. Denno. 1979. Responses of sap-feeding insects (Homoptera: Hemiptera) to simplification of host plant structure. Environ. Entomol. 8: 1021-1028. Vince, S. W., I. Valiela, and J. M. Teal. 1981. An experimental study of the structure of herbivorous insect communities in a salt marsh. Ecology 62: 1662-1678. Wilson, M. R. and M. F. Claridge. 1985. The leaf- hopper and planthopper faunas of rice fields, pp. 381-404. In Nault, L. R. and J. G. Rodriguez, eds., The Leafhoppers and Planthoppers. John Wiley and Sons, New York. Wilson, S. W. 1982. The planthopper genus Proke- lisia in the United States (Homoptera: Fulgoroi- dea: Delphacidae). J. Kans. Entomol. Soc. 55: 532- 546. PROC. ENTOMOL. SOC. WASH. 89(4), 1987, pp. 701-705 NOTES ON NORTH AND CENTRAL AMERICAN LOPHOSCUTUS SPP. (HEMIPTERA: PHYMATIDAE, MACROCEPHALINAE) NICHOLAS A. KORMILEV 5924 Gulfport Blvd. S., Gulfport, Florida 33707. Abstract.—Two new species are described: Lophoscutus froeschneri from Mexico and L. haitiensis from Haiti. A key is given to the 17 North and Central American species of Lophoscutus. Macrocephalus gracilis Handlirsch and Macrocephalus falleni Stal are trans- ferred to Lophoscutus. Macrocephalus stali Handlirsch is synonymized with Lophoscutus lepidus (Stal), 1862. The genus Lophoscutus Kormilev, 1951, is distributed in the tropical and sub-trop- ical areas of the Americas. In the north, it penetrates into southern United States; in the south, it extends only to Bolivia and central Brazil. The Caribbean area has many endemic species. This genus, originally de- scribed as a subgenus of Macrocephalus (Kormilev, 1951), was given generic status by Maa and Lin (1956). Kormilev (1984) provided a key to the genera belonging in the sub-family Macrocephalinae, including Lophoscutus. In the past 25 years the number of Central American species has nearly doubled and the only existing key to these species given by Handlirsch (1897) is 90 years old. In this paper, I offer a new key to the North and Central American species. Macrocephalus falleni Stal and Macrocephalus gracilis Handlirsch are transferred to the genus Lo- phoscutus. Lophoscutus gracilis described from ‘‘Amérique du Nord,” is actually a Brazilian species and, therefore, omitted from the key. Lophoscutus stali (Hand- lirsch) in long series, cannot be separated from Lophoscutus lepidus (Stal) and, there- fore, is synonymmized with the latter. All measurements in this paper were tak- en with a micromillimeter eyepiece, 25 units = | mm. The length of abdomen was measured from the anterior border of con- nexivum II to the tip of the abdomen. KEY TO NORTH AND CENTRAL AMERICAN SPECIES OF LOPHOSCUTUS 1. Granulation of head and pronotum setigerous se Ie, LEER eee EE ON ee eee ee 2. — Granulation of head and pronotum not setig- CTOUS§ eres ere ae ne ee a 6 2. Granulation of pronotum spiculoid ....... 3 — Granulation of pronotum rounded ........ 5 3. Large species, (male) over 7 mm; head, prono- tum, scutellum and fore femora with strong, spiculoid, setigerous granulation; venter with round, setigerous granulation; lateral angles of pronotum rounded; Venezuela, also from MI@X1CO 2 ieee nee Peer ee pers eee: asper (Stal). — Small species, (male) less than 7 mm; spicu- loid granulation less strong ............... 4 4. Large species, (male) over 6 mm; lateral an- gles of pronotum rounded and slightly in- Gised-IMIexICOm as ner spiculosus (Champion) — Small species, (male) less than 5 mm; lateral angles of pronotum acute; Mexico Plot distance from forest edge. 3* Denotes that the value is significantly different than the value immediately above it; Chi-square test for goodness-of-fit. 708 Table 3. Locust borer adults, counted in circular plots 2 meters in diameter, at varying distances from wood’s edge. Borers per Plot Distance Date of Count from Woods . Edge (m) Sept,10) ‘Sept 20 Sept 30 Oct 10 Total 0 0 1 6 0 7 20 0 1 l 2 4 40 0 2 0 0 2 ers 0 Sea ae Total 0 4 8 2 14 October 10, 1984. Transects were walked and the presence of locust borers was re- corded, as was distance from the transect beginning point, and borer activity and lo- cation on the plants. The number of borers per goldenrod and occurrence of mating ac- tivity were also recorded. In plots, all goldenrod plants were count- ed and maturity of flowers was recorded at each visit. Borers, if present, and their ac- tivity and location on the plants were re- corded. Weather during the period September 1 0- October 10 became cool, with onset of frost and periods during which weather condi- tions apparently inhibited borer activity and flowering of the goldenrod plants. On each of the observation dates, however, weather conditions, i.e. temperature, precipitation, and winds, were favorable and we observed insect movement among the goldenrod. Statistical comparisons by data collection date were conducted using a Spearman Rank Correlation test for correlation between borer frequency and distance from the wood’s edge. RESULTS Borer adults per transect on each date are shown in Table 1. Borer concentrations were significantly greater (P = 0.003) in the tran- sect nearest the wood’s edge, decreasing far- ther into the field. This trend was consistent for each date and transect, except for Sep- tember 20, when the 20 m transect yielded PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 4. Frequency of groups of adult locust borers on goldenrod stems. Plants with Distance Plants with Plants with Three or from Wood’s One Borer Two Borers’ More Borers Edge (m) per Stem per Stem per Stem Total 0 95 30 5 130 20 21 8 2 31 40 19 4 0 23 60 9 1 0 10 Total 144 43 7 194 fewer insects than the 40 m transect. Borer numbers decreased over time, but varied possibly due to weather. The most notice- able difference in occurrence through the field was observed on the last study date, October 10, when borers were not encoun- tered in the outer three transects (20 m, 40 m and 60 m), but were present in substantial numbers in the | m transect. Numbers and percentages of blooming goldenrods in the plots are shown in Table 2. Collectively, the four plots provided a 483-plant sample. Percentages of fully blooming plants decreased progressively with each 10-day interval from September 10-October 10. The only exception was in plot 2 (20 m from wood’s edge) where the percentage of fully blooming plants was greater on September 30 than on September 20. The data in Table 2 indicate that peak bloom was on or prior to September 10. Percentages of fully blooming plants varied from a maximum of 47% (plot 3, September 10) to 0% (plots 3 and 4, October 10). Too few borers were encountered in the 2 m cir- cular plots for statistical analysis (Table 3). However, overall numbers were greatest near the wood’s edge, as was the case in the transects. The transect counts yielded a total of 194 borer sightings. Each sighting included all borers on a given Solidago plant. Borers per plant per sighting are shown by transect (Ta- ble 4) and sampling date (Table 5). Most sightings (144 of 195) were of one borer per plant (significance, P = 0.001). Two per plant VOLUME 89, NUMBER 4 709 Table 5. Numbers of adult locust borers per goldenrod stem through the season. Plants with Plants with Plants with Three or One Borer Percent Two Borers Percent More Borers Percent Date per Stem of Total per Stem of Total per Stem of Total Total Sept 10 49 Wl 18 26 2 3 69 Sept 20 39 89 5 11 0 0 44 Sept 30 43 Ta 10 18 3 5 56 Oct 10 13 50 10 38 3 11 26 Total 144 43 8 195 followed in frequency (43 of 195 sightings), but more than 2 per plant occurred only rarely (8 of 195 sightings). Of the latter, six were of 3 borers per plant, and two were of 5 borers per plant. Multiple occurrences (more than one borer per plant) were sta- tistically more numerous in the transect nearest the forest edge (P = 0.05) than in the remaining three transects. Multiple oc- currences were recorded on each study date, through October 10, and proportionally in- creased with the total count as the season progressed. The highest percentage of three or more borers per stem occurred on Oc- tober 10. Both sightings of five borers per stem occurred on October 10. Despite these concentrations, no matings were recorded on October 10. DISCUSSION The borer count of October | demon- strated that a substantial number of insects were still active at that time, after the onset of frost (mid-September) and the decline of much of the goldenrod bloom. The distinct- ly uneven borer distribution on October 10, with the 46 sightings in transect | nearest the forest fringe, and none in the other tran- sects, was not explained. Possibly it reflect- ed shorter flights, nearer the trees, and/or fewer surviving adults. The frequency and/ or quality of blooming plants near the forest fringe could also have had an effect, al- though blooming plants still occurred in many parts of the field on October 10. No attempt was made to identify different species of flowering plants usable by the lo- cust borer, but on one occasion during the study a locust borer adult was observed on a garden-growing sunflower Helianthus an- nuus L. In a non-quantified observation, we no- ticed an apparent tendency by the borers to occupy goldenrod plants in full bloom, and seldom before and/or after. The sighting of a borer adult on goldenrods prior to bloom- ing, or on faded or seeding flowers, was rare. Likewise, they appeared to be attracted to the more spectacular flowers in terms of brightness and size. Some clusters of iso- lated goldenrods outside the study area, dis- junct from other goldenrods but near the black locusts, drew especially large numbers of borers (more than 10 per plant). The tendency for the congregation of nu- merous borers per goldenrod plant ap- peared to be slight. One likely reason for this is that the pollen-feeding habit exposes the insects to predation. The flower-feeding habit probably led to the borer’s mimicry, in elytral and abdominal color pattern, of stinging hymenopterans, implying the influ- ence of predation. In this study the only instances of predation observed were by the garden spider Argiope aurantia Lucas, which was common. We observed 1|5 cases of lo- cust borers killed and wrapped in webbing. No cases of avian or insect predation were observed, although the blooming flowers were visited by a large array of insects. No bird species appeared to be active in or around the field during data collection. 710 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Our findings here offer some implications for control of the locust borer. Concentra- tions of the insects near forest-field edges could be the targets of limited chemical spray efforts during an exposed and vulnerable phase of the life cycle. The knowledge of likely concentration sites would also en- hance controls utilizing manual and bait- trapping methods. LITERATURE CITED Berry, F. H. 1945. Effect of site and the locust borer on plantations of black locust in the Duke Forest. J. For. 43: 751-754. Boyce, S. G. and R. W. Merz. 1959. Tree species recommended for strip mine plantations in west- ern Kentucky. USDA Forest Service. Central States Forest Exp. Stn. Tech. Paper 160. 13 pp. Cuno, J. B. 1930. Utilization of black locust. USDA Circ. No. 131. 18 pp. Galford, J. R. 1977. Evidence for a pheromone in the locust borer. USDA Forest Service. Research Note NE-240. 3 pp. . 1979. Preliminary test of booby-trapping for control of two cerambycids. USDA Forest Service. Research Note NE-284. 3 pp. 1980. Use of a pheromone to cause copula- tion between two species of cerambycids. USDA Forest Service. Research Note NE-289. 2 pp. . 1984. The locust borer. USDA Forest Insect and Disease Leaflet 71. 6 pp. Garman, H. 1916. The locust borer (Cyllene robiniae) and other enemies of the black locust. Ky. Agric. Exp. Stn. Bull. 29: 99-135. Hall, R. L. 1942. Control of the locust borer. USDA Circ. 626. 19 pp. Harman, D. M., P. Rudolf, and K. R. Dixon. 1985a. Influence of stand composition on locust borer (Coleoptera: Cerambycidae) attack rates. J. Ento- mol. Sci. 20: 207-211. Harman, D. M., M. VanTyne, and W. A. Thompson. 1985b. Comparison of locust borer Megacyllene robinia Forster (Coleoptera: Cerambycidae) at- tacks on coal strip-mined lands and lands not mined. Ann. Entomol. Soc. Am. 78: 50-53. Hopkins, A. D. 1907. The locust borer and methods for its control. U.S.D.A. Bur. Entomol. Circ. No. 83. 8 pp. St. George, R. A. and J. A. Beal. 1932. New sprays effective in the control of the locust borer. J. Econ. Entomol. 25: 713-721. Wollerman, E. H. 1955. Control of black locust in- sects by systemics. J. Econ. Entomol. 48: 760-761. 1968. A Search for Borer-Resistant Black Locust. Pp. 53-54. Jn Proc. 6th Central States For- est Tree Impr. Conf. PROC. ENTOMOL. SOC. WASH. 89(4), 1987, pp. 711-714 HERNIOSINA VOLUMINOSA: A NEW SPHAEROCERID OF ISOLATED PHYLOGENETIC POSITION DESCRIBED FROM NORTHEASTERN NORTH AMERICA (DIPTERA: SPHAEROCERIDAE) S. A. MARSHALL Department of Environmental Biology, University of Guelph, Guelph, Ontario NIG 2W1, Canada. Abstract.—Herniosina voluminosa, new species, is described from extensive series col- lected using flight intercept traps in northern New Hampshire. This highly autapomorphic species is included in the previously Old World genus Herniosina on the basis of a single putative synapomorphy, a modified male first sternite. The genus Herniosina was described by Rohaéek in 1983 to include two similar Eu- ropean species characterized by a convex bulge on the male synsternite | + 2. The new species Herniosina voluminosa has a similar sternal bulge (Fig. 3) and also resem- bles described Herniosina in the following characters: discal cell long and narrow , al- ula narrow, R,,; weakly sinuate and not bypassed by the costa, male mid tibia with a row of ventral spines, female mid tibia with apical ventral bristles only, female ab- domen narrow and telescoping, and dor- socentral bristles in two pairs with the an- terior pair short. Characters of the male terminalia, usually of great importance in determining phylogenetic affinity of the Sphaeroceridae, do not suggest a relation- ship between H. voluminosa and European Herniosina or any other genus. Despite this difference, voluminosa is included in Herniosina rather than in a new, monotypic genus on the basis of the possible synapo- morphy of the male sternal bulge, a char- acter not known elsewhere in the Sphaero- ceridae. Other similarities between voluminosa and European Herniosina are probable plesiomorphies. Herniosina voluminosa Marshall, NEw SPECIES Description.— Body length 2.2—2.4 mm, dark brown to black, pruinose, minutely punctate; tarsi brown. Interfrontal plate very broad, subequal in height and width; bor- dered by 3 pairs of interfrontal bristles, mid- dle pair cruciate and twice as long as upper pair, lower pair half as long as upper pair. Postocellar bristles present, small. Eye height 3 times genal height at point of maximum eye height; gena entirely pruinose; vibrissa as long as eye height; genal bristle short, equal to subvibrissa. Dorsocentral bristles in 2 pairs, prescutellar pair 3 times as long as anterior pair; acrostichal bristles long, in 6-7 rows, prescutellar pair slightly enlarged. Mid tibia of male with distal ventral row of spinules and a small apical ventral bristle (Fig. 5); mid tibia of female ventrally with apical bristle and apical anteroventral bris- tles only. Mid tibia of both sexes dorsally with one proximal anterodorsal, one distal anterodorsal, one distal dorsal, and one dis- tal posterodorsal bristle (Fig. 5). Wing membrane brownish, dark brown along an- terior margin; second costal sector |.3-1.4 712 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VOLUME 89, NUMBER 4 times as long as third in male, 1.2—1.3 times as long in female; alula narrow, pointed; discal cell long, narrow; R,,; slightly sin- uate, both R,,; and costa ending near wing tip (Fig. 4). Scutellum 0.7—0.8 times as wide as long; surface densely tomentose; margin with 4 bristles and dense tomentum. Preab- dominal tergites uniformly dark, weakly punctate, tomentose medially and bare lat- erally. First sternite of male bulged out to form posteromedial ventral lobe (Figs. 1, 3), first sternite of female not modified (Fig. 7). Male terminalia.—Sternite 5 with a des- clerotized membranous area surrounded by a dark band anteriorly and patches of long bristles laterally (Fig. 1). Sixth sternite large (Fig. 1). Epandrium small, with a long lat- eral bristle (Fig. 2); cerci trapezoidal, con- tiguous at corners, lighter in color than epandrium and with 2 long bristles. Surstyli small, simple, posteriorly setulose, ventrally setose; with a dark inner ventral lobe; left surstylus smaller than right (Fig. 1). Aedea- gal complex unusually large, usually hang- ing well below axis of body. Distiphallus larger than epandrium; with broad, widely separated lateral lobes, right lobe distally broader than left (Fig. 2). Basiphallus elon- gate but without epiphallus or similar pro- jection; ejaculatory apodeme small but well sclerotized. Paramere very broad, distally with a lateral row of setulae. Female terminalia.—Terminalia retrac- tile into preabdomen (Fig. 7); tergite 8 des- clerotized medially, apparently divided (Fig. 8). Epiproct very pale except lateral margins which are fused with the short, setose cerci (Fig. 8). Sternite 8 brown; hypoproct with anterior part bare and transparent, crescent- — TAS shaped posterior part brown and setulose (Fig. 9). Spectacles-shaped sclerites present as very narrow rings, not visible in cleared (KOH) specimens. Spermathecae dark, elongate and wrinkled, sclerotized part of duct very short (Fig. 6). Types.— Holotype ¢ (Canadian National Collection) and 9 paratypes (1 3, 8 2): USA. New Hampshire, Coos Co., 3 mi. NE East Inlet Dam, Norton Pool, 12—24.vi.1986, Flight Intercept Trap, D. S. Chandler. Other Paratypes: USA. New Hampshire, 3 mi. NE East Inlet Dam, Norton Pool, 9—26.v.1986, Flight Intercept Trap, D. S. Chandler (3 6, 3 2); same, 27.v—11.vi (2 4, 1 9), 25.vi- 9.vil (5 6, 3 2), 10—24.vii (4 8, 1 9), 17.ix- 17.x (16); 1 ami, NE. East Inlet Dam: 25.vi-9.v11.1986, 10—24.vi1.1986, 25.vii- 7.vi1.1986, Flight Intercept Trap, D. S. Chandler (2 4, 1 2); Carr Co., 1 mi. N Won- alancet, E. Pk. Spring Brook, 16.iv—8.v.1985, 9-15.v.1985, 24-30.vu1.1985, 29.viii- 5.1x.1985, 6-17.1x.1985, Flight Intercept Traps, D. S. Chandler (5 4); The Bowl, 2.5 mi. NW Wonalancet, 15.1v-8.v.1985, 22.vi— 1.vii.1985, 11-17.vui.1985, 14—21.vii.1985, 2-17.x.1985, Flight Intercept Trap, D. S. Chandler (5 3). Paratypes are in the Uni- versity of New Hampshire Collection (Dur- ham), the University of Guelph Collection and The Canadian National Collection (Ot- tawa). Comments. — Herniosina voluminosa can be easily distinguished from all other Nearc- tic sphaerocerids by the distinctive male first sternite and the unusual interfrontal bris- tles. Other diagnostic characters include the fusion of the cerci with the epiproct (also found in Opalimosina and some Kimosina Figs. 1-9. Herniosina voluminosa. Figs. 1-5. Males. 1, Abdominal sternites and terminalia, aedeagus and associated parts removed. 2, Terminalia, aedeagus and associated parts stippled, paramere pivoted anteriorly into an unnatural position to expose distiphallus (left lateral). 3, Abdomen, aedeagus and associated parts removed (left lateral). 4, Left wing. 5, Left middle femur, tibia and tarsomeres | and 2 (anterior). Figs. 6-9. Females. 6, Spermathecae. 7, Abdomen (left lateral). 8, Terminalia (dorsal). 9, Terminalia (ventral). Abbreviations: aed apo, aedeagal apodeme. epan, epandrium. sur, surstylus. para, paramere. basi, basiphallus. disti, distiphallus. S6, sternite 6. S7, sternite 7. SS, sternite 8. hypo, hypoproct. epi, epiproct. 78, tergite 8. 714 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON and Opacifrons) and the pigmented wing with a long discal cell. Males of H. volu- minosa differ from all other sphaerocerids in their highly characteristic terminalia, in which the aedeagus and associated struc- tures are extremely large and distinctive. Other, less obvious differences between males of H. voluminosa and other Hernios- ina include the presence of an ejaculatory apodeme, short broad cerci, a lateral (not dorsolateral) epandrial bristle, and an un- modified fifth tergite. The females differ markedly in the shape of the spermathecae, other Herniosina having smooth, pear- shaped spermathecae. The entire type series was collected in flight intercept traps maintained by D. S. Chandler for a two year period in mature forest in nothern New Hampshire. These traps also captured several other rarely col- lected species of Sphaeroceridae such as Bitheca caballa Marshall, Spelobia bispina Marshall, Spelobia frustrilabris Marshall, Terrilimosina pexa Marshall, Aptilotus spatulatus Marshall, Pullimosina antennata Duda, Pullimosina hirsutiphallus Marshall, Minilimosina gemella Rohaéek and unde- scribed species in the genera Aptilotus, Ru- dolfia and Spelobia. With the notable ex- ception of Bitheca caballa, which is known only from two specimens from Tennessee and Massachusetts, most of the other species (ca. 50 spp.) of Sphaeroceridae taken in the same traps as H. voluminosa are either northern Holarctic species or have their closest relatives in the northwest. LITERATURE CITED Rohaéek, J. 1983. A monograph and re-classification of the previous genus Limosina Macquart (Dip- tera, Sphaeroceridae) of Europe. Part 11. Beitr. Entomol. Berlin 33: 3-195. PROC. ENTOMOL. SOC. WASH. 89(4), 1987, pp. 715-730 NESTING BEHAVIOR OF TACHYSPHEX LAEVIFRONS AND T. CRASSIFORMIS, WITH A NOTE ON T. KROMBEINI (HYMENOPTERA: SPHECIDAE) FRANK E. KURCZEWSKI Environmental and Forest Biology, State University of New York College of Environ- mental Science and Forestry, Syracuse, New York 13210. Abstract.— Aspects of the nesting behavior of Tachysphex laevifrons, T. crassiformis, and 7. krombeini were studied as part of a larger study on the comparative behavior of species in the Tachysphex pompiliformis group. Tachysphex laevifrons nested in sand and had only a single, vernal emergence of adults per year. Females did not level the tumulus, left the entrance open while hunting and provisioning, and placed one or two paralyzed Melanoplus sp. or M. puer group in a short, shallow, single-celled nest. The acridids were small to relatively large in size and transported to the nest in flight or on the ground, respectively. Larger wasps laid larger eggs. Females of 7. crassiformis exhibited similar behavior, except that they placed from one relatively large Chortophaga sp. (North Car- olina), Trimerotropis sp. (Oklahoma), or Psinidia fenestralis (Florida) to four small Tri- merotropis sp. (Oklahoma) in a one-celled nest in sand. This species of wasp demonstrated seasonal variation in nest dimensions and acridid size and an inverse relationship between prey size and the number of prey per cell. Both 7. crassiformis and T. krombeini were apparently multivoltine in Florida. 7. /aevifrons and T. crassiformis affixed their egg on an acridid in a position typical of most other species in the genus. The nesting behavior and ecology of 7. /aevifrons are compared with those of the closely related 7. tarsatus, while the nesting behavior and ecology of 7. crassiformis and T. krombeini are compared with those of a related species, 7. antennatus. Species in the large Tachysphex pompil- iformis group can be arranged sequentially, using adult external morphology (W. J. Pu- lawski, pers. comm.). They can also be placed arbitrarily in subgroups using com- parative nesting behavior (Kurczewski, 1987), but some behavioral patterns tran- scend subgroup distinctions. For example, T. texanus (Cresson) and 7. psammobius (Kohl), two totally unrelated species, fly with prey and store several or many, small or tiny acridids, respectively, in a one-celled nest (Kurczewski, 1987). 7. ashmeadii Fox, T. montanus (Cresson), T. pauxillus Fox, T. pompiliformis (Panzer), and 7. tarsatus (Say), species belonging to at least two subgroups, usually carry their prey on the ground, often ventral side up, leave the en- trance open during provisioning, and store one or a few, relatively large nymphal, rarely adult, acridids in a single-celled nest (Peck- ham and Peckham, 1900, 1905: Williams, 1914; Newton, 1956; Evans, 1970; Alcock, 1973; Alcock and Gamboa, 1975; Elliott and Kurczewski, 1985; Pulawski, pers. comm.). On the other hand, certain behav- ior and prey type can be used to delimit some subgroups. 7. aethiops (Cresson), T. punctifrons (Fox), and 7. pompiliformis in Europe carry their prey on the ground, dor- 716 sal side up, and build multicellular nests (Adlerz, 1903; Evans, 1970, 1973; Pulaw- ski, 1971; Alcock, 1973; Kurczewski, 1987; M. F. O’Brien, pers. comm.). 7. pechumani Krombein is distinctive in its temporary closure of the entrance and leveling of the tumulus (Kurczewski and Elliott, 1978). 7. semirufus (Cresson) provisions its nests with nymphal Tettigoniidae (Kurczewski and Evans, 1986; Pulawski, pers. comm.). Information on the nesting behaviors of Nearctic species of the pompiliformis group is fragmentary, consisting mostly of isolated prey records and observations of single fe- males and their nests (Kurczewski, 1987). Only 7. ashmeadii, T. pechumani, T. pom- piliformis, and T. tarsatus have been stud- ied in any detail (Kurczewski and Elliott, 1978; Krombein, 1979; Elliott and Kur- czewski, 1985). The following observations on various aspects of the nesting behavior of T. laevifrons (Smith), T. crassiformis Vie- reck, and T. krombeini Kurczewski are pre- sented to elucidate their phylogenetic rela- tionships within this large and diverse group. The wasp and prey specimens from the study have been given ethology note numbers and deposited in the State University of New York College of Environmental Science and Forestry and Cornell University insect col- lections, respectively. Essentially nothing 1s known about the nesting behavior of 7. /aevifrons (Krom- bein, 1979). T. crassiformis has been stud- ied briefly by Williams (1914) as 7. pleno- culiformis Williams and by Krombein (1963) as T. boharti Krombein. One female of 7. krombeini has been observed nesting by Kurczewski (1971). Williams (1914) noted a female of T. crassiformis in Kansas flying with a small acridid. She successively took five small grasshoppers directly into an open nest at intervals of 4-10 min and then closed the burrow with sand, apparently using the fore- legs and end of the abdomen. The nest was one-celled, 1% inches long, and 1*4 inches deep which seems unusually vertical for a PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON species of Tachysphex. The cell contained six paralyzed, nymphal tryxaline acridids. Krombein (1963) presented information on two nests of 7. crassiformis, one dug by him in North Carolina and the other dug by H. E. Evans in Florida. Krombein ob- served prey transport of a nymphal acridid slightly larger than the wasp both on the ground and in short flights. The nest he ex- cavated was single-celled, 5 cm deep, and held three nymphal Psinidia fenestralis (Serville), each 6-7 mm long. The paralyzed grasshoppers exhibited movements of the legs and palpi. The wasp’s egg was placed transversely on the sternum of one prey, near a forecoxa. Evans’ observation was of a relatively large acridid carried slowly on the ground by the bases of its antennae and held venter up and head forward. This prey was placed inside the wasp’s entrance, pulled into the nest from within, and, 1.5 min after entry, the wasp began filling the burrow with sand. The burrow was 3 cm long and led to a cell, 1 cm deep, which held a single, par- alyzed nymph of Scirtetica marmorata picta Scudder. The wasp’s egg was attached to the “throat” of the prey. The female of 7. kKrombeini observed by Kurczewski (1971) left the entrance open and brought five small prey to her nest in 65 min. The individuals were carried in rather high, rapid flights and held beneath the wasp’s body with the legs and mandi- bles. The female released each prey in the entrance, entered, turned around inside, and pulled in the acridid by its antennae with the mandibles. After the full complement of prey had been placed inside the nest, the wasp filled the burrow with sand, using the forelegs and abdomen. The burrow entered the sand obliquely for 4.8 cm, including cell length, and ended in a small cell, 2.8 cm beneath the surface, including cell height. The cell contained six nymphal acridids (Melanoplus sp.) and one nymphal tettigo- niid (Odontoxiphidium apterum Morse), most of them head inward and ventral side up. The wasp’s egg was attached to the left VOLUME 89, NUMBER 4 forecoxal corium of the largest acridid in the cell and extended transversely between the bases of the fore- and midlegs. Tachysphex laevifrons (Smith) T. laevifrons has only a single, vernal emergence of adults per year (Kurczewsk1, 1971). Thirteen females (TX-15) were ob- served nesting on the sand flats along the Peace River at Arcadia, De Soto County, Florida (see Krombein and Kurczewski, 1963, Fig. 4) during March 25—April 4, 1963, March 26-27, 1965, April 9-10, 1971, and March 29-30, 1972. Females nested in flat, loose, moist, white sand containing decum- bent legumes and slopes of bare, white, coarse-grained, firm sand. The nests of three wasps were | 1-24 m apart. Females dug nests, hunted, and provi- sioned on clear, sunny days at ambient (shade) temperatures of 29°-33°C. During periods of cloud cover, they stopped hunt- ing and rested or cleaned. Females were ob- served hunting as early as 0925 and as late asil730 hu(ESi); A female searching for a place to dig walked rapidly forward on the sand, moving in a zigzag manner. She held the wings flat on the dorsum when walking and periodi- cally made a rapid flight of several centi- meters or, rarely, a few meters to a new area to resume searching. Such a wasp tapped her antennal tips alternately on the sand, sampled the sand with the mandibles, but then usually moved elsewhere. Two females of T. laevifrons made false starts 10 and 30 cm apart, respectively. Another wasp made eight trial digs, averaging 4 mm in length, before completing a burrow. One female at- tempted three burrows in shallow depres- sions, entered a hole, 5 mm deep, dug lat- erally, and then also abandoned this excavation. After breaking the sand crust with the mandibles, the female used the forelegs in unison to rake the loosened sand backward. The body, especially the abdomen, lifted up and down with the digging forelegs. The mid- 717 and hindlegs were used only for walking in the burrow and entrance and on the tu- mulus. At intervals the wasp backed out of her excavation and removed sand that had accumulated in the burrow and entrance. The intervals, as measured by the time the female was inside her nest, increased as she dug deeper. One wasp spent only 7-26 s (mean, 13.6) inside between each of her first five sand removals but 10-54 s (mean, 29.0) between individual sand removals 21-25. The female spent 3-8 s (mean, 4.6) working in the entrance and on the tumulus during sand removals 1-5 but only 1-4 s (mean, 2.0) during removals 21-25. She backed onto the tumulus to distances of 0.4-1.8 cm (mean, 1.1; N = 25) from the bottom of the entrance in mostly straight or slightly curved directions but made no attempt to level the mound of accumulated sand. The tumuli in front of 10 entrances measured 2.5—4.4 cm (mean, 3.3) long and 2.1—3.0 cm (mean, 2.5) wide. Females also dug short, shallow burrows in which they rested during rain and spent the night. A wasp exhibited the same com- ponents for digging this burrow as she did when she dug a burrow and cell to contain prey. Two females spent 5.5 and 6 min, respectively, to dig a resting burrow. A wasp rested at the end of the burrow, facing the entrance, after closing the opening with sand from inside. One wasp placed a prey in her nest without ovipositing and then dug a resting burrow. She returned to the nest the following morning and finished provision- ing it. A female never closed her entrance while hunting or provisioning. After digging a burrow, she appeared headfirst in the en- trance, ran around the opening, and then went off hunting in low rapid flights inter- spersed with running. Females sometimes hunted small prey on the stems and leaves of plants as far as 25 m from their entrances. Usually, however, a wasp hunted larger prey on the ground or prostrate vegetation within 5-10 m of the entrance. One such female 718 returned to her nest without prey six times and, each time, entered her burrow, turned around inside in 10-14 s, and exited head- first. Other hunting wasps did not enter their nests upon returning empty-handed. Some hunting females pounced on the dorsum of acridids noticeably larger than themselves only to fly off immediately and resume hunting elsewhere. One wasp, weighing (wet) 17 mg, encountered an ac- ridid about twice her size (13.5 mm long) but much heavier (89 mg). After a brief struggle, she managed to turn the grasshop- per onto its back but did not attempt to sting it. The acridid righted itself, leapt 1 cm onto a shrub, reared forward, and raised both hindlegs in a threatening manner. The wasp paused, faced the grasshopper, moved her head from side to side, and then went off hunting. Another wasp, weighing 14 mg, stung an acridid weighing only 4.5 mg, abandoned it on the sand, and then stung and abandoned a second acridid weighing 28 mg. Females always captured and stung prey on the ground. One wasp stung an acridid, weighing only 7 mg, twice in the venter of the thorax and another, weighing 93 mg, five times near the leg bases. A sting inser- tion averaged 11.3 s (2-20; N = 7). Paralysis of the prey still allowed rhythmic move- ments of the mouthparts, especially palpi, antennae, and abdominal segments. After stinging, sometimes malaxating the prey with the mandibles, and cleaning her- self, a female straddled the acridid head for- ward and often ventral side up but occa- sionally on the side or dorsum up. The wasp grasped the prey’s antennae with the man- dibles and, holding the sides of the grass- hopper’s body with the hindlegs, proceeded forward on the ground or, in the case of smaller prey, grasped the prey’s body with the legs and flew. Manner of transport was related to the relative weights of the wasp and prey: (1) ground, ratio of weight of prey to wasp, R = 1.9-5.8:1, mean, 3.3, N = 7; (2) 2-15 cm-long flights, 1.2:1, N = 1; and, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (3) 1-20 m-long flights, R = 0.4—0.6:1, mean, 0.5, N = 2. One female with prey 2.1 times heavier than herself moved nearly 1 m in 4 s. A wasp with prey 5.8 times heavier than herself grasped the grasshopper far out on its antennae and, with her hindlegs, near the bases of its forelegs. She had great difficulty in transporting this prey and had to pause frequently, Another female overran her en- trance and located it only after making sev- eral circles. Upon reaching the open nest, a female placed the prey ventral side up or on its side with the head just inside the entrance (N = 8) or ran directly inside holding the acridid underneath (N = 2). The size of the prey usually determined the manner in which it was taken into the nest. However, one ac- ridid the same size as the wasp (weight ratio, 1.2:1) was taken in directly and a smaller prey (0.6:1) was released with its head, tho- rax, and first three abdominal segments in- side the entrance. The wasp entered the bur- row, turned around inside, and, using her mandibles, pulled in the prey by its anten- nae. Wasps brought successive prey to their nests in 3-150 min (mean, 68.6; N = 5). After placing the full complement of prey in a cell and laying an egg on one of the acridids, the wasp filled the burrow and en- trance with sand. Females (N = 5) appeared headfirst in their entrances 5-6 min after taking in the last prey for the cell and raked sand backward into their burrows, using the forelegs in unison. During this behavior the abdomen moved synchronously up and down and the wings were held flat on the dorsum. As the wasp backed in with sand, her abdomen vibrated rapidly back and forth to pack the sand in the burrow. The mid- and hindlegs were braced against the lower sides of the burrow. When the burrow was filled flush to the surface the wasp moved her abdomen from side to side, smoothing over the fill. One female averaged 21.3 s (R = 10-40) between her first five trips to the surface to obtain sand but only 8.7 s (R = 5-15) for the next five trips. She took VOLUME 89, NUMBER 4 Fipsale 12 min from entering with her last prey for the cell to filling the burrow flush with the sand surface. The entrances to 10 nests of 7. /aevifrons were 0.4—0.6 cm (mean, 0.5) in diameter. The burrows penetrated the sand at angles of 30°-38° with the horizon and proceeded obliquely downward to the cell in an almost straight line (Fig. 1). Ten burrows averaged 5.3 cm (4.3-6.6) long, including cell length, and terminated in individual cells at a mean depth of 3.4 cm (2.4—4.2), including cell height. The cells averaged 0.58 cm (0.5-0.7) high, 0.63 cm (0.5-0.7) wide, and 1.20 cm (0.8-1.5) long. Wasps preyed on the nymphal acridids Melanoplus sp. (12 records) and M. puer group (4). The number of prey placed in fully-provisioned cells was one or two (mean, 1.6; N = 10). The individual wasps weighed (wet) 11-24 mg (mean, 16.6; N = 8), where- 719 Incomplete nest of Tachysphex laevifrons with burrow and cell containing paralyzed, nymphal Me- lanoplus sp. as their prey averaged 40.8 mg (7-93; N = 16) in wet body weight. The full comple- ment of prey in a cell averaged 67.0 mg (35- 137; N = 10). The acridids were placed in the cells head inward and ventral side up and, in the case of two prey in a cell, laid side by side. In six fully-provisioned cells each with two prey, the egg was attached to the largest and heaviest acridid in the cell. These grasshop- pers averaged 49.2 mg (28-76) in wet body weight. In two cells containing individually marked prey, the egg was affixed to the sec- ond acridid taken into the nest. An egg of 7. /aevifrons was sausage- shaped, white, curved slightly, and tapered at the proximal end. It was affixed to a grass- hopper by the less tapered, distal end at a forecoxal corium and extended transversely between the first two pairs of legs. Seven eggs were afhxed to the prey’s left forecoxal 720 coria and three to the right forecoxal coria. The size of an egg was related to the size of the wasp. Wasps weighing (wet) 12, 14, 23, and 24 mg laid eggs measuring 0.17 x .040, 0.18 x .045, 0.23 x .050, and 0.24 x .055 cm, respectively. Tachysphex crassiformis Viereck There were two or more flight periods of T. crassiformis per year in Florida with dates of collection ranging from March through September. Twenty-nine females were ob- served nesting in loose sand behind the beaches along the Atlantic Ocean at Fort Pierce Beach, St. Lucie County, Florida on July 16, 1962, April 3, 1963, and March 30, 1965 (TX-8) and at Kill Devil Hills, Dare County, North Carolina on July 23-24, 1962 (TX-9); in the sand flats adjacent to the Peace River at Arcadia, De Soto County, Florida on June 30-July 10, 1962, March 23, 1963, April 6, 1966, April 9-10, 1971, and March 28—-April 9, 1973 (TX-12); in a sandy fire- trail at the Archbold Biological Station, Highlands County, Florida on April 12, 1973 (TX-83); and, on the sandy flood plain along the Cimarron River, near Guthrie, Logan County, Oklahoma on July 27 and August 12, 1965 (TX-68, 70). Eight of 22 nests were situated beneath prostrate vege- tation, overhanging twigs, or near the bases of grass clumps. Nine entrances were locat- ed 1.8--55 m apart. Females in Florida began nesting in March-April at 0925 h (EST) and nesting activity was at its height at about 1400 on sunny or partly sunny days when the am- bient temperature in the shade reached 28° 33°C. No wasp hunted, dug, or provisioned below 27°C. The latest observation (daily) was made at 1534 h. A female searching for a place to dig ex- hibited behavior similar to that of 7. /ae- vifrons (see above), except the velocity of the movements was more rapid. The wasps usually made one or more trial digs before PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON completing a burrow. One such false start was 5 mm long. Most excavations were be- gun from the sand surface but one wasp (TX-8) started in a depression 5 mm deep. The manner in which the burrow was dug was similar to that of 7. /aevifrons, except the digging movements of 7. crassiformis were noticeably faster. One female (TX-12) paused every 5-10 s and made a hovering flight of 1-2 s, 10 cm above the entrance, until a depth of 3 mm had been reached. These flights were probably in response to high sand surface temperatures (58°-59°C). Another wasp (T X-12) removed sand from the burrow 26 times, remaining inside 21- 50 s (mean, 35.5) between successive sand removals. This female, when removing sand, backed further from the entrance at the be- ginning of the excavation (2-3 cm) than at the end (1-2 cm). A third wasp (TX-68) removed 27 sand loads from the burrow and entrance, stayed outside only 4-27 s (mean, 11.4) per sand removal, rebacked (Kur- czewski, 1968) 1-12 times (mean, 4.9) on the tumulus during some of the sand re- movals, and raked the sand to distances of 0.8—2.2 cm (mean, 1.6) from the entrance. Her times and distances remained approx- imately the same throughout the excava- tion. Most wasps walked backward with the sand loads in a straight or nearly straight line and made little attempt to level the tu- mulus. The tumuli of 15 nests averaged 3.3 cm (2.0-4.5) long, 2.8 cm (1.7-—4.0) wide, and 0.5 cm (0.3-0.6) high. The largest tu- mulus in Florida, 4.5 cm long, 3.0 cm wide, and 0.6 cm high, was associated with the longest burrow (6.0 cm) and second deepest cell (3.1 cm). One female (TX-12) took 12.5 min while another (TX-68) spent 53 min to construct a burrow and cell. After digging a burrow and cell, a female appeared headfirst in the entrance with the wings held flat on the dorsum. Some wasps (TX-8, 9, 12) then tapped their antennal tips alternately on the tumuli, made four or five runs around an entrance, probably a form VOLUME 89, NUMBER 4 of orientation, and went offhunting in short, low, rapid flights interspersed with running. Other wasps (T X-68) exited headfirst, made 7-13 s long orientation flights, facing the entrances, turned 180° in flight, and flew off. Some females, after exiting and running around the entrances, re-entered their nest one or a few times and backed out of the entrance removing sand with the forelegs. A nest entrance remained open during hunt- ing and provisioning. Females hunted on the ground or low vegetation, often within 10 m of their en- trances. Six wasps (TX-12, 70) captured prey only 0.6—2.1 m from the openings. Wasps (N = 27) unsuccessful in capturing prey re- turned to their entrances 1-5 times, entered their burrows, turned around inside, exited headfirst in 8--50 s (mean, 13), and ran or flew away rapidly. Four females, after entry, removed 1-4 loads of sand from the bur- row. Consecutive returns to the nests with- out prey averaged 18.6 min (2-55; N = 27). A hunting female’s antennal movements, running, pivoting, and zigzagging increased in rapidity when in close proximity (5-10 cm) to a potential prey. If the grasshopper remained motionless, the hunting wasp often passed it by. One female (TX-8) increased the rapidity of her searching movements when she ran across an area of sand where a nymphal acridid had been resting 30 s earlier. Some wasps, which encountered ac- ridid nymphs weighing 25-30 mg, touched them with their antennae or flew onto their dorsum, and then flew away. One female (TX-8), weighing 8.5 mg, approached an ac- ridid nymph, 10.5 mm long and weighing 36 mg, mounted it, turned it onto its back, stung it in the right side of the thorax near a hindleg for 14 s, dismounted, and cleaned herself. The grasshopper immediately right- ed itself and jumped away. The wasp did not follow. She then passed by an acridid nymph, 13 mm long and weighing 84 mg, which raised both hindlegs simultaneously and kicked them backwards. The wasp flew 721 away immediately. Another female (TX-8) was kicked by the hindlegs of a 15-mm long acridid. She flew a short distance, landed, rested for a few seconds, and cleaned. Females always stung their prey on the ground, the acridid being on its venter, dor- sum or side (Fig. 2). The female bent her abdomen underneath the grasshopper until the apex touched the prey’s thoracic venter, presumably the corium surrounding a hind- coxa, and inserted the sting therein. This critical insertion probably prevented the grasshopper from leaping away (Steiner, 1976). The wasp maintained herself dorsum up while attempting to position the longi- tudinal axis of her body at nearly a 90° angle to that of her prey during subsequent stings in the sequence. From one to five stings were administered to a single prey during a period of 8-120 s. Whether this represented a partial or com- plete stinging sequence is unknown (Steiner, 1976). Small prey subsided almost imme- diately after one or two stings, but more than two stings were necessary to subdue larger grasshoppers. A sting insertion av- eraged 13.8 s (5-30; N = 16). In 12 of 17 stinging sequences, females alternated one or more times between stinging the prey in the left and right sides. Prey were stung into a state of incomplete paralysis which per- mitted periodic movements of the mouth- parts, especially maxillary and labial palpi, and antennae. The abdominal segments ex- hibited rhythmic breathing movements. Following the initial sting or, between subsequent stings, the wasp cleaned herself while straddling the prey or, more often, dismounted, walked a few centimeters, and cleaned. This behavior lasted 5—40 s (mean, 11.7; N = 13) and consisted of running the antennae through the strigilis and rubbing the hindtarsi over the wings and abdominal apex. One female, after stinging her prey, dismounted, cleaned herself for 15 s, re- mounted the grasshopper, and malaxated its left forecoxal corium with her mandibles 722 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VOLUME 89, NUMBER 4 (Fig. 3). She then turned 180°, cleaned the prey’s left hindleg for 75 s, turned back, and malaxated the prey’s left forecoxal corium for 26 s. During transport to the nest, she intermittently cleaned and repeatedly malaxated the acridid. Despite this exten- sive malaxation, the prey was not used for Oviposition. After stinging and malaxating the acridid and cleaning herself, a female straddled the prey usually ventral side up or, rarely, dorsal side up or on the side and head forward. The wasp grasped both antennae of the prey if venter or dorsum up, or one antenna, if on the side, with the mandibles. She then either proceeded forward on the ground, clutching the grasshopper with the hindlegs while using the fore- and midlegs for walk- ing (Fig. 4) or grasped the acridid with the legs and flew. The manner of transport was dependent upon the relative sizes (weights) of the wasps and their prey: (1) ground, ratio of weight of prey to wasp, R = 2.2-6.8:1, mean, 4.4, N = 12; (2) 1-4 cm-high and 3- 20 cm-long flights, R = 1.3-2.3:1, mean, 19S Ni =36>,and, G)) 10+ cm high, 1+ m long flights, R = 0.3-1.1:1, mean, 0.7, N = 11. Females varied their position on the prey and the location of the grasp of its body depending upon the acridid’s size and weight. Females atop larger (heavier) prey straddled the acridid more anteriorly in or- der to increase leverage and provide space for the moving fore- and midlegs (Fig. 4). The heaviest prey (R = 4.2-6.8:1, mean, 4.9, N = 4) were grasped with the mandibles by their antennae about one-third of the way from the ends or farther out and clutched with the wasp’s hindtarsi near the throat or the bases of the forelegs. Medium-sized prey (R = 1.9-3.3:1, mean, 2.5, N = 8) were grasped by their antennae about midway 123 out and with the wasp’s hindtarsi near the bases of the midlegs. If a female had difh- culty in ground transport such as ascending an incline or slipping on loose sand, she invariably adjusted her position on the prey further anteriorly. One wasp (TX-70) with prey 6.8 times her weight went 0.5 m in 1.5 min, 0.4 m in 1.5 min, and 0.2 m in 47 s, with several second-long pauses in between. The smallest (lightest) prey (R = 0.3-1.1:1, mean, 0.7, N = 11) were grasped with the mandibles near the bases of their antennae and, during flight, around the body with the legs. Upon reaching the nest a female placed an acridid venter up or on its side, rarely dorsum up, with the head and, sometimes, thorax and abdomen just inside the en- trance (Figs. 5, 6), or, ran or flew directly inside, holding the grasshopper underneath (Fig. 7). The size and weight of the prey often influenced the manner in which it was taken into a nest: (1) head outside entrance, ratio, 6.8:1, N = 1; (2) head inside entrance, R = 0.6—5.7:1, mean, 2.2, N = 9; (3) head and thorax inside entrance, 2.1:1, N = 1; (4) head, thorax, and abdomen inside en- trance, R = 0.3-1.1:1, mean, 0.7, N = 5; and, (5) direct entry with prey, R = 1.6-1.9: 1, mean, 1.8, N = 3. After releasing the prey, entering, and turning around inside the nest, a female appeared headfirst in the entrance in 4-6 s (mean, 5; N = 5), and, with the mandibles, pulled the acridid inside by its antennae. The wasp exited in 8-13 s (mean, 11; N =7)ifshe went in search of additional prey; otherwise, she stayed inside the nest and oviposited. Females brought successive prey to their nests in 3-68 min (mean, 20.3; N = 11). After laying an egg on a prey in the cell, a female filled her burrow with sand. Wasps — Figs. 2-4. 2, Female of Tachysphex crassiformis inserting sting near base of right hindcoxa of nymphal Psinidia fenestralis. 3, Female of Tachysphex crassiformis malaxating left forecoxal corium of recently stung, nymphal Psinidia fenestralis. 4, Female of Tachysphex crassiformis transporting nymphal Psinidia fenestralis, holding grasshopper’s antennae with mandibles and body with hindlegs. 724 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VOLUME 89, NUMBER 4 came out of their entrances headfirst to ob- tain sand 2—7.5 min (mean, 3.8, N = 6) after entering with their last prey. The compo- nents of filling in the nest were essentially as described under T. /aevifrons, except the movements of 7. crassiformis were more rapid. Three females (TX-8) came out of their entrances as far as 2.5-3.0 cm every 5-35 s to obtain loose sand for the fill, whereas four others (TX-68) went only 0.4— 1.3 cm every 1-2 s. Two wasps (TX-8, 70) came out to get sand 14 and 9 times, re- spectively, to complete the fill. After filling the burrow nearly flush with the surrounding sand, some females turned 180° every several seconds and tapped their antennae on the fill. They then turned back and, standing on the filled entrance with the mid- and hindlegs spread and, curving the abdomen underneath, vigorously ham- mered the fill. Five females (TX-8, 12, 70) took from 5 to 14 min (mean, 7.1; N = 5) to fill their burrows and entrances flush with the surrounding sand. Females of 7. crassiformis dug short, shallow burrows which entered the sand obliquely at angles of less than 45° to the surface and terminated in single cells. Bur- row length and cell depth dimensions from Florida, North Carolina, and Oklahoma are presented in Table I. The seasonal differ- ences in mean burrow length and cell depth from Florida and North Carolina nests are highly significant (¢ (length) = 3.660; df = 18a <= 0:01: 2 (depth) = 5.883; df — 13; a < 0.001). Similarly, mean burrow length and cell depth from July and August Okla- homa nests are significantly different (¢ (length) = 11.446; df = 4; a < 0.001; ¢ (depth) = 18.807; df = 4; a < 0.001). En- trance diameters averaged 0.43 cm (R = 0.3-0.8; N= 15), the burrow diameters — Figs. 5-7. U2 being only slightly smaller (R = 0.3-0.5 cm; mean, 0.35). Fourteen March-April cells from Florida were 0.5—0.7 cm (mean, 0.61) high, 0.6-0.7 cm (mean, 0.66) wide, and 1.0-1.2 cm (mean, 1.09) long. In Oklahoma four cells from July were 0.5—0.6 cm (mean, 0.53) high, 0.5-0.6 cm (mean, 0.58) wide, and 0.9-1.0 cm (mean, 0.98) long, whereas two cells from August were 0.7 cm high, 0.8 cm wide, and 1.5—1.6 cm long. The Okla- homa cell differences correlated positively with the use of four small prey per cell in July and one or two much larger individuals per cell in August. Females preyed on the nymphal acridids Chortophaga sp. in North Carolina (1 re- cord), Psinidia fenestralis (Serville) in Flor- ida (20), and, 7Trimerotropis sp. in Okla- homa (17), all members of the subfamily Oedipodinae. The number of prey stored in fully-provisioned March-April Florida cells ranged from | to 4 (mean, 2.5; N = 12) but was only | in Florida and North Carolina cells during July (N = 3). These mean dif- ferences are highly significant (¢ = 4.313; df = 13; a < 0.001). In Oklahoma in July females stocked four cells each with four prey but used only one or two (mean, 1.5; N = 2) acridids per cell in August (¢ = 5.774; df = 4; a < 0.01). Regardless of locality the number of prey per cell was inversely re- lated to the sizes (weights) of the individ- uals: one prey per cell, 21-84 mg (mean, 45.4; N = 5); two prey, 11-81 mg (mean, 24.6; N = 14); three prey, 11-35 mg (mean, 18.9; N = 12); and, four prey, 2-22 mg (mean, 8.3; N = 20). Prey individuals from Florida (March-April) were significantly heavier than those from Oklahoma (July) (t = 5.1227; df = 47; a < 0.0005) but sig- nificantly lighter than those from Oklahoma (August) (¢ = 3.2087; df = 33; a < 0.0025). 5, Female of Tachysphex crassiformis releasing relatively large nymph of Psinidia fenestralis, ventral side up, in entrance. 6, Abdomen and legs of relatively large, nymphal Psinidia fenestralis projecting from entrance after its release. 7, Female of Tachysphex crassiformis entering nest with relatively small nymph of Psinidia fenestralis. 726 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Seasonal variation in nest dimensions of Tachysphex crassiformis. Dee eee eee eee Locality N Range Mean + SD 9IIIII EEE = Burrow Length (cm) Florida, March-April 12 3.8-6.0 VO) ae (Rpt t Florida—North Carolina, July 3 3.0-3.7 33210360" Oklahoma, July 4 6.6-7.6 fl 7320.47 58% Oklahoma, August 2 28-32 3:00:30" Cell Depth (cm) Florida, March-April 12 1:9 =355 P28 04545 Florida—North Carolina, July 3 1.0-1.4 pale 2e=sO0: 20% Oklahoma, July 4 4.3-4.7 { 4:4 10.19*** Oklahoma, August D el 14 ies ee OSE Ae ee *** Data within brackets are statistically significant at 99% confidence level. Prey from Oklahoma (August) were signif- icantly heavier than prey from Oklahoma (July) (t = 5.625; df = 18; a < 0.0005). The mean sum weights of the prey per cell were: one prey per cell, 45.4 mg (R = 21-84; N = 5); two prey, 49.2 mg (R = 30-92; N = 7); three prey, 56.7 mg (R = 45-72; N = 4); and, four prey, 33.2 mg (R = 16-65; N = 5). Tests (¢) applied to all pairwise combi- nations of the means are not significantly different at a = .05. Female wasps from Florida weighed 5-14 mg (mean, 9.1; N = 11), whereas wasps from Oklahoma weighed 6-10 mg (mean, 7.6; N = 5). These mean differences are not significant (¢ = 0.985; df= 14: a > 0.20). In 18 fully-provisioned cells containing from 1 to 4 prey per cell, all acridids were positioned ventral side up and head inward. In four other cells (TX-8, 68), each with four prey, one acridid was placed dorsum up and head inward and three were placed venter up and head inward. In all 22 cells the ac- ridid bearing the wasp’s egg was positioned ventral side up and head inward. In six of the cells, each with two prey placed side by side, the egg was attached to either the left or right acridid equally. In one cell with two acridids laid in tandem, the egg was placed on the prey that was farthest in. In each of four cells with one acridid piled atop another, the egg was placed on the upper- most prey. The egg was affixed to the heavi- est acridid in 12 of 16 cells with two or more prey, an intermediate-sized prey once, and the lightest prey three times. In one cell with two acridids (TX-70) the egg was affixed to a prey weighing only 9 mg while the other grasshopper weighed 68 mg. The mean weight of an egg-bearing prey of T. crassiformis was 26.7 mg (7-84; N = 18) while other prey individuals in the cells averaged only 15.7 mg (2-68; N = 35), a significant difference (¢ = 2.269; df = 51; a < 0.05). In nine examples of two or more prey per cell eggs were laid on the initial acridid taken into the nest (3), second prey taken in (2), or last grasshopper taken in (4). Two egg-bearing acridids had received ex- tensive malaxation by the wasps, whereas three other extensively malaxated acridids were not oviposited upon. An egg of T. crassiformis was similar to that of T. /aevifrons except for being smaller in size. Eight such eggs (TX-8, 9, 12, 68) averaged 0.16 (0.15—0.18) cm long and 0.042 (0.040-0.045) cm wide at the middle. The egg was placed similarly to that of 7. /ae- vifrons but fit more tightly into a depression on the underside of the thorax of the oed- ipodine prey. Although certain females af- fixed their eggs consistently to the left (TX- VOLUME 89, NUMBER 4 68) or right (TX-12, 70) sides of their prey, the sum of the affixation sites was 11 to the left and 10 to the right forecoxal corium. Tachysphex krombeini Kurczewski One female was observed searching for a place to dig a burrow on the sand flats beside the Peace River at Arcadia, Florida on April 10, 1973. She walked rapidly in a zigzag pattern and finally selected a depression in bare sand. She began digging with the man- dibles and then removed the loosened sand backward, using the forelegs in unison. She held the wings flat on the dorsum. The movements of her forelegs were rapid and her forward and backward movements within the entrance were noticeably jerky. The female did not attempt to level the small mound of sand that accumulated in front of the entrance. After 4.5 min of digging, the wasp turned 180°, filled the entrance with sand, using the forelegs in unison, and flew away, abandoning the excavation. DISCUSSION Tachysphex laevifrons is similar morpho- logically and behaviorally to 7. tarsatus. The latter species occurs commonly throughout much of the United States and southern Canada but only sparingly in the Southeast, being absent entirely from southern Georgia and Florida (Pulawski, pers. comm.). 7. /ae- vifrons is essentially a geographic replace- ment of 7. farsatus, inhabiting not only southeastern U.S. but also extending rarely into Texas, Oklahoma and Kansas (Krom- bein, 1979; Pulawski, pers. comm.). Where- as T. tarsatus has two flight periods per year in northeastern U.S. and more than two at southern latitudes, 7. /aevifrons has only a single, vernal flight per year (Kurczewski, 1971). In this regard the latter species re- sembles 7. pechumani (Kurczewski and El- hiott, 1978). Females of both 7. /aevifrons and T. tar- satus occur in sparsely vegetated sandy soil, often constructing nests near the bases of 127 plants. The components involved in search- ing for a nesting site and burrow construc- tion are essentially identical. Both species make little attempt to level the tumulus, run around the entrance for orientation prior to hunting, leave the entrance open during hunting and provisioning, and hunt in prox- imity to their nests. 7. /aevifrons and T. tarsatus capture medium-sized or relatively large, occasionally small, usually nymphal acridids, transport them on the ground or in low flights, depending on their size and weight, take them into the entrance var- iously, based on their size, and store one or a few paralyzed individuals in a single-celled nest. Final closure of the nest contains es- sentially identical components in the two species. The nests of 7. /aevifrons and T. tarsatus are short and shallow with an oblique burrow. Prey of both species are placed in the cell mostly head inward and ventral side up and the wasp’s egg is affixed to a forecoxal corium of the acridid, ex- tending transversely between the bases of the fore- and midlegs. The acridids captured by T. tarsatus comprise members of three subfamilies (Williams, 1914; Krombein, 1979; Elliott and Kurczewski, 1985), where- as those caught by 7. /aevifrons consist only of species of Me/anoplus. The apparent nar- row prey preference of the latter species may be related to its having been studied at only a single locality in Florida. T. crassiformis, T. krombeini, and T. an- tennatus are allied morphologically and be- haviorally. 7. crassiformis has an extensive distribution throughout North, Central, and South America (Krombein, 1979). T. an- tennatus also occurs throughout much of the U.S. and Mexico but is replaced in southeastern U.S. by 7. krombeini (Pu- lawski, pers. comm.). 7. crassiformis and T. Arombeini nest in areas of sand inhabited also by T. laevifrons. T. antennatus inhabits abandoned gravel pits and gravelly paths along the edges of woodlands (pers. ob- serv.). None of the three species nest in ag- 728 gregations and, consequently, the entrances of conspecific nests are often widely scat- tered. Based upon dates of collection 7. crassiformis and T. krombeini have two or more generations per year in Florida (Kur- czewski, 1971), while 7. antennatus has only a single generation per year with a small, partial second generation during optimal summers (Pulawski, pers. comm.; pers. ob- Serv.). In searching for a nesting site and digging anest 7. crassiformis, T. krombeini, and T. antennatus exhibit rapid movements. Those of a T. krombeini or T. antennatus female digging within her entrance are sometimes rather jerky and include much rebacking, as in Plenoculus davisi (Fox) (Kurczewski, 1968) and Diploplectron peglowi Krombein (Kurczewski, 1972). Williams (1914) noted the same behavior for digging females of T. crassiformis (as T. plenoculiformis) in Kan- sas. However, I found the digging move- ments of 7. crassiformis to resemble those of T. tarsatus (Williams, 1914) and T. ash- meadii (Elliott and Kurczewski, 1985). During digging neither 7. crassiformis, T. krombeini, nor T. antennatus level the sand that accumulates in front of their entrances. Females of all three species omit a tem- porary closure of the entrance during hunt- ing and provisioning. 7. crassiformis and T. antennatus capture medium-sized or small acridid nymphs which they carry to their nests in low flights (Williams, 1914; Elliott and Kurczewski, 1985), but the former species also preys upon relatively large ac- ridid nymphs which are transported on the ground (Krombein, 1963). There is a rela- tionship between prey size, method of trans- port, and manner of entry into the nest in T. crassiformis and T. antennatus. Rela- tively large prey are invariably carried close to the ground, released with their head in- side the entrance and pulled into the nest from within, whereas small prey are often flown directly into the entrance. There is seasonal variation in prey storage related to prey size in 7. crassiformis (see also Wil- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON liams, 1914; Krombein, 1963). 7. krom- beini provisions with small prey, carries them in extensive flights, and stores several individuals in a cell, after releasing each one with its head in the entrance during entry (Kurczewski, 1971). Final closure of the nest in all three species involves raking sand backward into the tunnel with the forelegs which are bent me- dially and packing the sand into place with the apex of the abdomen (Williams, 1914; Kurczewski, 1971, pers. observ.). The movements exhibited during this behavior are very rapid. T. crassiformis and T. krombeini dig rath- er straight, short burrows which enter the sand obliquely and terminate in single cells (Williams, 1914; Krombein, 1963; Kur- czewski, 1971). With few exceptions (Krombein, 1963), seasonal variation in burrow length and cell depth is evident in T. crassiformis. In contrast 7. antennatus digs a short, curving, single-celled nest, often beneath a flat stone, and demonstrates only slight variation in nest dimensions (pers. observ.). The majority of prey of 7. crassiformis, T. krombeini, and T. antennatus is placed in cells head inward and venter up, and the wasp egg is laid on an individual so posi- tioned. The egg is affixed as in T. /aevifrons, except that of 7. crassiformis fits tightly against the thoracic sternum of oedipodine prey. Whereas 7. krombeini stores both nymphal acridids and tettigoniids in the cell (Kurczewski, 1971), 7. crassiformis and T. antennatus provision only with nymphal acridids. Both 7. krombeini and T. anten- natus capture a preponderance of Melano- plus spp. (Cyrtacanthacridinae) (Kurczew- ski, 1971; Elliott and Kurczewski, 1985; pers. observ.), whereas 7. crassiformis preys upon Oedipodinae (Krombein, 1963; pers. observ.) and Tryxalinae (Williams, 1914). In conclusion, the morphology and nest- ing behavior of 7. /aevifrons and T. tarsatus are alike. No single behavioral character- istic can be used to separate the two species. VOLUME 89, NUMBER 4 The single, annual spring appearance of 7. laevifrons adults and the restriction of this species essentially to southeastern U.S. pro- vide allochronic and geographic evidence for its distinctness. 7. farsatus is multivol- tine at southern latitudes and occurs throughout the U.S. except for southern Georgia and Florida. Although 7. crassi- formis, T. krombeini, and T. antennatus are morphologically similar, 7. Arombeini can be separated readily from the two other species by its use of tiny acridid and tetti- gonid prey and its extensive provisioning flights. 7. antennatus nests in gravelly soil beneath stones and stores several Melano- plus spp. (Cyrtacanthacridinae) in a cell, whereas 7. crassiformis nests in sandy soil beneath vegetation or in the open and places one or a few Oedipodinae, rarely several Tryxalinae, in acell. 7. crassiformis exnibits much variation in nesting behavior throughout its range. ACKNOWLEDGMENTS I thank K. V. Krombein, Smithsonian In- stitution, and M. G. Spofford, SUNY-CESF, for reviewing the manuscript and assistance with statistical tests, respectively, and M. F. O’Brien, The University of Michigan, for permitting me to examine his unpublished manuscript on the nesting behavior of Tachysphex aethiops. | am indebted to K. V. Krombein and W. J. Pulawski, Califor- nia Academy of Sciences, for confirming the identities of the species of Tachysphex, and, A. B. Gurney (retired), Systematic Ento- mology Laboratory, BBII, Agricultural Re- search Service, USDA, for identifying the prey Acrididae. W. J. Pulawski kindly per- mitted me to examine his unpublished mansucript on the Tachysphex of North and Central America. I am grateful to the late Richard Archbold for providing the excel- lent facilities of the Archbold Biological Sta- tion, Lake Placid, Florida, which was used as a base for many of the studies. I especially thank N. F. R. Snyder, Portal, Arizona for the photographs used in the text. Sigma Xi 729 RESA Grants-in-Aid of Research provided partial funding for this study during 1962, 1963, and 1966. LITERATURE CITED Adlerz,G. 1903. Lefnadsférhallanden och Instinkter inom Familjerna Pompilidae och Sphegidae. K. Svenska Vet.-Akad. Handl. 37: 1-181. Alcock, J. 1973. Notes on a nesting aggregation of digger wasps in Seattle, Washington (Hymeno- ptera). Wasmann J. Biol. 31: 323-336. Alcock, J. and G. J. Gamboa. 1975. The nesting be- havior of some sphecid wasps of Arizona, includ- ing Bembix, Microbembex, and Philanthus. J. Ariz. Acad. Sci. 10: 160-165. Elhott, N. B. and F. E. Kurezewski. 1985. Nesting and predatory behavior of some Tachysphex from the western United States (Hymenoptera: Spheci- dae). Great Basin Natur. 45: 293-298. Evans, H. E. 1970. Ecological-behavioral studies of the wasps of Jackson Hole, Wyoming. Bull. Mus. Comp. Zool. 140: 451-511. . 1973. Further studies on the wasps of Jackson Hole, Wyoming (Hymenoptera, Aculeata). Great Basin Natur. 33: 147-155. Krombein, K. V. 1963. A new Tachysphex from southeastern United States (Hymenoptera, Sphecidae). Entomol. News 74: 177-180. . 1979. Genus Tachysphex, pp. 1627-1632. In Krombein, K. V., P. D. Hurd, Jr., D. R. Smith, and B. D. Burks, eds. Catalog of Hymenoptera in America North of Mexico. Vol. 2, Apocrita (Acu- leata). Smithson. Institution Press, Washington, D:€; Krombein, K. V. and F. E. Kurczewski. 1963. Bio- logical notes on three Floridian wasps (Hymeno- ptera, Sphecidae). Proc. Biol. Soc. Wash. 76: 139- L522 Kurczewski, F. E. 1968. Nesting behavior of Pleno- culus davisi (Hymenoptera: Sphecidae, Larrinae). J. Kans. Entomol. Soc. 41: 179-207. . 1971. Anew Tachysphex from Florida, with keys to the males and females of the Florida species. Proc. Entomol. Soc. Wash. 73: 111-116. 1972. Observations on the nesting behavior of Diploplectron peglowi Krombein (Hymeno- ptera: Sphecidae). Proc. Entomol. Soc. Wash. 74: 385-397. 1987. A review of nesting behavior in the Tachysphex pompiliformis group, with observa- tions on five species (Hymenoptera: Sphecidae). J. Kans. Entomol. Soc. 60: 118-126. Kurczewski, F. E. and N. B. Elliott. 1978. Nesting behavior and ecology of Tachysphex pechumani Krombein (Hymenoptera: Sphecidae). J. Kans. Entomol. Soc. 51: 765-780. 730 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Kurczewski, F. E. and H. E. Evans. 1986. Correct the solitary wasps. Bull. Wisc. Nat. Hist. Soc. 1: names for species of Tachysphex observed by Ev- 85-93. ans (1970) at Jackson Hole, Wyoming, with new 1905. Wasps Social and Solitary. Houghton information on T. alpestris and T. semirufus (Hy- Mifflin Co., Boston, xv + 311 pp. menoptera: Sphecidae). Proc. Entomol. Soc. Wash. Pulawski, W. J. 1971. Les Tachysphex Kohl (Hym., 88: 720-721. Sphecidae) de la Région Paléarctique Occidentale Newton, R.C. 1956. Digger wasps, Tachysphex spp., et Centrale. Panstwowe Wydawnictwo Naukowe, as predators of a range grasshopper in Idaho. J. Wroclaw, 464 pp. Econ. Entomol. 49: 615-619. Williams, F. X. 1914. Monograph of the Larridae of Peckham, G. W. and E. G. Peckham. 1900. Addi- Kansas. Univ. Kans. Sci. Bull. 8: 117-213. tional observations on the instincts and habits of PROC. ENTOMOL. SOC. WASH. 89(4), 1987, pp. 731-738 PHYLOGENETIC PLACEMENT OF TWO GENERA OF HADENINAE FROM SOUTHWEST RUSSIA (LEPIDOPTERA: NOCTUIDAE) Tim L. MCCABE Biological Survey, New York State Museum, State Education Department, Albany, New York 12230. Abstract.—The types of two genera (Lepidoptera: Noctuidae) from southwest Russia have been examined and determined to be synonymous with existing European and American genera: Epipsammia Staudinger (1879), new synonym, is the junior subjective synonym of Hecatera Guenée (1852); Namangana Staudinger (1888) new synonym, is the junior subjective synonym of Trichoclea Grote, 1883. Morphological evidence for this synonymy is discussed. A lectotype is selected for Namangana cretacea Staudinger (1888). The following new combinations are made: Trichoclea cretacea (Staudinger), He- catera deserticola (Staudinger), Hecatera fixseni (Christoph) and Hadena boursini (Wilt- shire). Epipsammia and Namangana previously stood in the Acronictinae and are now formally transferred to the Hadeninae. North American and European lepidop- terists are generally not familiar with the noctuid genera Epipsammia Staudinger (1879) and Namangana Staudinger (1888). Both genera were described from arid des- erts of southwest Russia. My recent studies on Scotogramma Hy. Edwards (1887) have caused me to review these genera to settle questions of synonymy. Hampson (1909) was the first to apply Namangana to many of the Nearctic species. He did this without reference to the geni- talia, although he apparently did have ac- cess to a correctly determined specimen of the type species. Hampson overlooked the hairy eyes and placed several Nearctic Ac- ronictinae species in this genus. Barnes and Benjamin (1924, 1926) erected genera to ac- commodate many of the species then placed in Namangana. Several other species placed in Namangana by Hampson (e.g. Protor- thodes texana consors Smith) are now rec- ognized as Hadeninae. Namangana, in reference to Nearctic species, appeared in our literature between 1909 and 1926, but was never cited in the familiar Nearctic checklists or catalogues. Hampson’s application of the name to our species did not occur until after Smith’s (1893) Catalogue and Dyar’s (1903) List and by the next major checklist, that of Mc- Dunnough (1938), the included species had all been placed in other genera. Barnes and Benjamin (1926) and Suk- hareva (1973) both considered Namangana to be Hadeninae and not Acronictinae as it had been placed. Barnes and Benjamin (1926) considered Namangana to be simi- lar to the Nearctic 7richoclea Grote (1883a) and Scotogramma. Sukhareva (1973) inti- mated Namangana might be considered a subgenus of the Palearctic Hadula Staudin- ger (1889), a related genus. Namangana is currently placed in the Acronictinae (Nye, 1975) even though it had been linked to the Hadeninae on at least two occasions (Barnes and Benjamin, 1926; Sukhareva, 1973). Staudinger as well as Hampson (1909) con- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VOLUME 89, NUMBER 4 sidered the genus to lack hairy eyes and placed it in the Acronictinae. Hair on the eyes is a controversial, not absolute, yet ex- pedient character for differentiating Had- eninae from most other trifid noctuids. I examined the type of this genus and dis- covered it had long, although sparse, hairs on the compound eyes. Barnes and Benja- min (1926) had borrowed the cotypes of Namangana cretacea Staudinger (1888) and one of the included species, N. accurata Christoph (from Armenia) and assigned them to the Hadeninae (accurata was orig- inally described as a Mamestra species in the Hadeninae). They did not examine the genitalia, designate a lectotype, illustrate the adult, or synonymize the genus, but clearly indicated the connection with Scotogram- ma and Trichoclea. They considered it dis- tinct based on vestiture differences. I borrowed the cotypes of N. cretacea from the Museum fiir Naturkunde der Humboldt Universitat. Staudinger did not indicate a type specimen and I hereby designate the male, which I have illustrated (Figs. 2 and 5), as lectotype. It bears the following labels: Namangana Stgr. Cretacea Stgr.; Zool. Mus. Berlin; Origin.; McCabe slide 1157. Namangana cretacea is extremely close to another type species, Trichoclea decepta Grote (1883a), as is apparent by the simi- larity in habitus (Figs. 4 and 5) and genitalia (Figs. 1 and 2). Trichoclea decepta is quite variable genitalically, and I selected as an example (Fig. 1) one which has greatly re- duced harpal elements and consequently approaches cretacea in appearance. Tricho- clea decepta adults are also variable and one of the adults depicted (Fig. 4) was chosen because it compares well with cretacea; Fig. 10 is another example at the other end of a — Figs. 1-3. 733 cline. Trichoclea florida (Smith) (Fig. 11) is given as another example of a Trichoclea species. Notice also the similarity in habitus to the Nearctic Scotogramma submarina (Grote) (Fig. 12). A vesica with a simple branch that ter- minates in a bulbed cornutus is character- istic of Trichocosmia, Trichoclea, Scoto- gramma, Hadula, Cardepia Hampson (1905) and related genera, but subject to convergence because of its simplicity. How- ever, cretacea falls within the range of vari- ation seen amongst the species of Trichoclea sensu stricto, hence Trichoclea and Na- mangana, new synonym, are congeners and Trichoclea has priority. Trichoclea is com- prised of arid-land species that share a sim- ple vesica with one branch near the base; a single, bulbed cornutus; generally have asymmetrical clavi (i.e. right clavus devel- oped into a rounded protuberance); and a normal juxta (as apposed to a carinate juxta—see Fig. 3). This is a combination of features that are subject to convergence, but taken together establishes monophyly. The asymmetrical clavi, in particular, is a de- rived feature of considerable rarity and characteristic of Trichoclea and near rela- tives. Although all the species in these re- lated genera share the single bulbed cor- nutus on the vesica, the shape “and configuration varies, and in this, cretacea agrees very well with decepta (Figs. 1 and 2 vs. Fig. 3). The trend is toward greater asymmetry with eventual development ofa battledore structure on the right valve and a heavily carinate juxta. They are very sim- ilar in habitus as is apparent from the figures (Figs. 4 and 5). Trichoclea cretacea, new combination, is distinct in lacking the heavy spines on the foretarsi typical of most 77/- 1, Trichoclea decepta valves & aedoeagus with vesica everted, McCabe slide no. 700, Miles City, Custer Co., Montana, USA. 2, Namangana cretacea, holotype, valves and aedoeagus with vesica everted, McCabe slide no. 1156, Namangan, USSR. 3, Scotogramma submarina valves and aedoeagus with vesica everted, McCabe slide no. 854, Stockton, Utah, USA. 734 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 4-9. 4, Trichoclea decepta, male, Lone Tree, Uinta County, Utah, USA. 5, Namangana cretacea, holotype male, Namangan, USSR. 6, Epipsammia deserticola, holotype male, Narun sandy region, USSR. 7, Hadena boursini (after Wiltshire, 1957) Shaqlawa, Iraq. 8, 9, Hadena bicruris valves and aedoeagus with vesica everted, McCabe slide 759, Europe. —— ee VOLUME 89, NUMBER 4 12 Figs. 10-15. 15 10, Trichoclea decepta, male, Miles City, Custer County, Montana, USA. 11, Trichoclea florida, male, Great Exuma Island, Bahamas. 12, Scotogramma submarina, male, Montana, USA. 13, Hadena bicruris, female, Vaasa, Finland. 14, Hecatera bicolorata, male, Helsinki, Finland. 15, Hecatera dysodea, male, Barcelona, Spain. choclea species. It differs from Scotogram- ma in that it lacks the carinate juxta (see Fig. 3, Scotogramma submarina) and the well developed flap overlapping the base of the cucullus. Boursin considered Namangana as well as Pseudathetis Boursin to be synonymous with Epipsammia and stated that the new species similar to E. deserticola being de- scribed by Wiltshire was “an interesting modification of the same type of male gen- italia” (Wiltshire, 1957). I disagree with 736 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VOLUME 89, NUMBER 4 Boursin’s appraisal of Namangana, how- ever, and think Namangana goes with Tri- choclea; whereas Epipsammia Staudinger (1879) and Hecatera Guenée (1852) are congeneric and Hecatera has priority. He- catera dysodea (Denis and Schiffermiiller), the type species, is illustrated (Fig. 15 and 17). Pierce (1909) also depicted dysodea, but the valves (which are asymmetrical) were made to appear symmetrical as was often done during that period. Epipsammia, new synonym, has rarely been applied to species other than the type species, FE. deserticola Staudinger (1879). It is a plain, buff-colored moth (Fig. 6) with the ordinary lines ob- scure. The male genitalia of Hecatera de- serticola, new combination, are illustrated (Fig. 16). I borrowed the type from the Mu- seum ftir Naturkunde der Humboldt Uni- versitat and it proved to have hairy eyes and typical Hecatera genitalia: a squat, right- angled cucullus; an unmodified juxta; an apical process off the sacculus; symmetrical development of the clavi; a reinforced costa; an uncus that is adorned with blunt-tipped fixed setae; and a vesica with a bulbed cor- nutus (lacking the distal patch of band-like cornuti that is typical of many, though not all, Hadena). Boursin (1960) later described two noctuids from Afghanistan as ?Fpi- psammia constantialis and ?Epipsammia agrapha under the Amphipyrinae. Boursin placed them after Epipsammia fixseni and Epipsammia deserticola, respectively. They were not illustrated. I have not examined these two species, but presumably they will also prove to be Hadeninae. Boursin syn- onymizes Pseudathetis Boursin with Epi- psammia (Wiltshire, 1957), and I regard Epipsammia as a synonym of Hecatera; hence the sole included species, Pseuda- thetis fixseni Christoph, becomes Hecatera — Figs. 16-18. 137 fixseni (Christoph), new combination. The species still needs to be examined to verify the relationship to Hecatera, and it may ac- tually prove to belong in another genus, per- haps even Hadena. Wiltshires’ species, originally described as Epipsammia boursini, is, in my opinion, a Hadena, which means it should be known as Hadena boursini (Wiltshire) new combi- nation. The male genitalia are illustrated in Wiltshire (1957), and I have shown his il- lustration (Fig. 7). The genotype of Hadena, H. bicruris (Fig. 13) has a spinulose juxta (Fig. 8) with lateral arms that are found in most of the species of Hadena, a derived feature not seen amongst related genera. An extension of the claval region apparently takes over the function of this juxtal mod- ification in Hecatera. Wiltshire’s (1957) species boursini (Fig. 7), has the spinulose juxta and I feel it falls within the limits of Hadena. Although I have illustrated only the type species of Hadena (Figs. 8, 9 and 13), boursini is genitalically more similar to other European Hadena species, most no- tably, Hadena luteago (Denis and Schiffer- miller), which is illustrated in Pierce (1909) (as barrettii (Doubleday)). ACKNOWLEDGMENTS I am indebted to H. J. Hannemann of the Museum fiir Naturkunde der Humboldt Universitat zu Berlin Zoologisches Mu- seum for his cooperation in the loan of the two types and other specimens. Martin Honey of the British Museum called my attention to the Wiltshire reference. Chris Supkis assisted in the photography. Kauri Mikkoli, Hermann Hacker and Gottfried Behounek provided an exchange for several of the specimens used in the illustrations. Voucher specimens are deposited in the New 16, Epipsammia deserticola, holotype male, valves and aedoeagus, Boursin slide no. 364, Narun sandy region, USSR. 17, Hecatera bicolorata, male, valves and aedoeagus with vesica everted, McCabe slide 927, Helsinki, Finland. 18, Hecatera dysodea, male, valves and aedoeagus with vesica everted, Barcelona, Spain. 738 York State Museum. This is contribution number 521 of the New York State Science Service. LITERATURE CITED Barnes, W. and F. H. Benjamin. 1924. Contributions to the Natural History of the Lepidoptera of North America. 5(3): 99-199. 1926. Generic synonymy (Lepid., Phalaeni- dae). Pan-Pac. Entomol. 3: 64-74. Boursin, C. 1960. Nouvelles ‘“Trifinae’’ D’Afghani- stan de L’Expedition Kapperich (3™ note). Bull. Mens. Soc. Linn. Lyon 29: 136-152. Dyar, H. G. 1902 [1903]. A list of North American Lepidoptera and key to the literature of this order of insects. U.S. Nat. Mus. Bull. 52: 1-723. Edwards, H. 1887. Descriptions of new species of North American Heterocera, with notes. Can. Entomol. 19: 146. Grote, A. R. 1883a. On Stiria, with new genera and species of Noctuidae. Papilio 3: 29-35. 1883b. New species and notes on structure of moths and genera. Can. Entomol. 15: 3-13. Guenée, M. A. 1852. 6: 27. In Boisduval, J. A. and M. A. Guenée, Histoire Naturelle des Insectes. Species Géneral des Lépidopteéres, Paris. Hampson, G. F. 1905. Catalogue of the Lepidoptera Phalaenae in the Collection of the British Mu- seum. Taylor and Francis, London. Vol. 5. 634 Pp. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON . 1909. Catalogue of the Lepidoptera Phalaen- ae in the Collection of the British Museum. Taylor and Francis, London. Vol. 8. 583 pp. McDunnough, J. 1938. Checklist of the Lepidoptera of Canada and the United States of America. Pt. 1, Macrolepidoptera. South. Calif. Acad. Sci. Mem. 275) pp: Nye, I. 1975. The Generic Names of Moths of the World. Vol. 1 Noctuidae (part) Noctuidae, Aga- ristidae, and Nolidae. Trustees of the British Mu- seum Public. No. 770. 586 pp. Pierce, F. N. 1909. The Genitalia of the Group Noc- tuidae of the Lepidoptera of the British Islands. E. W. Classey, Feltham, Middlesex. xii + 88 pp. Smith, J. B. 1893. A catalogue, bibliographical and synonymical, of the species of moths of the lepi- dopterous superfamily Noctuidae, found in boreal America. U.S. Nat. Mus. Bull. 44: 1-424. Staudinger, O. 1879. Ueber Lepidopteren des siid6st- lichen europdischen Russlands. Stett. Entomol. Zeit. 40: 315-328. . 1888. Centralasiatische Lepidopteren. Stett. Entomol. Zeit. 49: 1-65. 1889. Centralasiatische Lepidopteren. Stett. Entomol. Zeit. 50: 16-60. Sukhareva, L. L. 1973. The systematics of the subfamily Hadeninae Guenée, 1973 (Lepidoptera, Noctuidae). Entomol. Rev. 2: 277-286. Wiltshire, E. P. 1957. The Lepidoptera of Iraq. Nich- olas Kaye Limited, London. 159 pp. PROC. ENTOMOL. SOC. WASH. 89(4), 1987, pp. 739-743 LARVAL PARASITISM OF RHAGOLETIS COMPLETA (DIPTERA: TEPHRITIDAE) ON JUGLANS MICROCARPA (JUGLANDACEAE) IN WESTERN TEXAS AND SOUTHEASTERN NEW MEXICO E. F. LEGNER AND R. D. GOEDEN Division of Biological Control, University of California, Riverside, California 92521. Abstract. Two hymenopterous parasitoids, Biosteres sublaevis Wharton (Braconidae) and Trybliographa sp. (Cynipidae) caused varying degrees of natural parasitism on Rhag- oletis completa Cresson in western Texas and southeastern New Mexico during four survey years, 1978, 1980, 1981, and 1983. A ‘“‘killing-power”’ analysis in 1980 indicated a sig- nificant (P < 0.05) impact of combined natural mortality on host reduction. Biosteres sublaevis demonstrated the greatest measureable activity as a cause of natural mortality. The walnut husk fly, Rhagoletis completa Cresson, has been a pest of Persian walnut, Juglans regia L., in California since 1926 (Boyce, 1934). It initially invaded a walnut orchard at Chino in southern California in 1925 (Michelbacher and Ortega, 1958; S. E. Flanders, pers. commun.). The founders of the California population may have origi- nated in western Texas (Berlocher, 1976). Natural enemy activity on this species in wild and cultivated Juglans species in Cal- ifornia has been negligible in California (Boyce, 1934; E. F. Legner, unpublished data). We were told of a high incidence of larval parasitism of this species by an opine brac- onid in its native range in the Davis Moun- tains of western Texas in 1974 (S. H. Ber- locher, pers. commun.). This report details field collections of R. completa larvae and subsequent measurement of parasitoid ac- tivity throughout this area during 1978- 1983. METHODS AND MATERIALS Rhagoletis completa is a univoltine species in western Texas that attacks ripening fruit of so-called “little walnut,” Juglans micro- carpa Berlandier, following late-summer rains. Larvae can only be obtained in fruit during a short period during mid August and early September (S. H. Berlocher, pers. commun.; E. F. Legner, unpub. data). Most fruit drop from trees by the second week of September and the larvae immediately exit from the fallen fruit and enter the ground to pupariate. Surveys were conducted on J. microcarpa from near Carlsbad, New Mexico through- out the Davis Mountains of Texas and south to the northern boundary of Big Bend Na- tional Park (Table 1). This region lies within the Chihuahuan Desert between the Rio Grande and Pecos Rivers and has many prominent mountains which usually sup- port a grassland climax vegetation (War- nock, 1970). Juglans microcarpa 1s a com- mon tree along arroyos at elevations between 1200 and 1600 m. Samples of whole, blackened infested walnuts were taken from trees during the final week of August in 1978, 1980, 1981 and 1983. However, all sites could not be sampled each year because the fruit had fal- 740 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Sample sites of Rhagoletis completa Cresson on Juglans microcarpa in western Texas and south- eastern New Mexico during 1978-1983. No. Trees Km & Direction from Site No. Sampled Ft. Davis, TX Location l 20 4 NE Rt. 17, NE of Ft. Davis, TX 2 10 4N Along Hwy. 118, N of Ft. Davis (Kent Rd.), TX 3 10 7N Indian Lodge (adjacent to buildings), TX 4 15 5 N Indian Lodge (camping area) 5 9 13 NE Route 17, NE of Ft. Davis, TX 6 10 248 North of Alpine, TX in canyon on Rt. 118 7 1 28 N Rt. 118, 2.5 km N of McDonald Observatory Rd., TX 8 6 38 NE Along Rt. 17, 13 km S of Toyahvale, TX 9 10 40 NW Rt. 118, Madera canyon picnic area, TX 10 5 48 W Rt. 166, W slope of Davis Mts., TX 11 6 53 N Hwy. 118, N of Ft. Davis, TX 12 5 70 N Hwy. 118, N of Ft. Davis, TX 13 5 748 North of Big Bend Natl. Park, Hwy. 118, TX 14 8 243 NW Rattlesnake Springs, Guadalupe Mts., NM 15 10 275 NW Carlsbad Natl. Park, NM len prior to our arrival. Fruit fall depended on variable weather conditions (wind and rainfall). An estimate of larval density per walnut was made in 1980 by sampling 300 walnuts at random per tree. Walnuts were placed on 4 cm of local soil in polyethylene buckets for 5 days to allow larvae to exit the nuts and pupariate in the soil. Puparia were then carefully and with minimum abrasion sifted from the soil and placed in 12 dram screened polystyrene containers. The caged puparia were transported to Riv- erside and placed in quarantine at the Uni- versity of California, where they were stored in refrigerators at 3° + 1°C and 55% RH for 6 months. They then were incubated at Zoe eC yao Ra anda? 1421/0) he LD photoperiod to allow emergence of adult flies and parasitoids. Unemerged puparia were refrigerated for another 6 months beginning the following September, followed by another period of incubation to promote ad- ditional emergence. A third such refrigera- tion/incubation cycle also was performed. Identifications.— The identity of Rhago- letis completa was verified by comparing adult specimens with those identified for the Department of Entomology by F. L. Blanc and R. H. Foote, and by referring to the descriptions in Boyce (1934) and Michel- bacher and Ortega (1958). The 7rybliogra- pha sp. was identified by G. Gordh.* Bio- steres sublaevis Wharton was identified by R. A. Wharton from material we gave K. Hagen, some of which probably were in- cluded in Wharton’s type series. We iden- tified subsequent collections of sublaevis by reference to Wharton and Marsh (1978). Statistical analyses.—Larval walnut huskfly densities per walnut at the time of initial field sampling were compared to the final densities after parasitization and other mortality factors had acted. These density differences measured parasitoid response to varying host densities in the field and de- termined whether such response was regu- lative, i.e. an increasing proportion of hosts were parasitized at higher host densities. First, the initial larval density in 300 sam- pled walnut fruit per tree was compared to the parasitized host density using a bivariate correlation analysis. Secondly, host regu- * Voucher specimens are deposited in the Depart- ment of Entomology, University of California, Riv- erside. VOLUME 89, NUMBER 4 741 Table 2. Emergence of Biosteres sublaevis and Tribliographa sp. parasitoids from Rhagoletis completa puparia collected as larvae in the Davis and Guadalupe Mountains of western Texas and southeastern New Mexico during 1978-1983. % Puparia No. Biosteres emerged! No. Trybliographa Emerged' IN US ye/eCG VU sme 8 OY NNN) 0 No. Puparia Flies Nothing % % Site Year Obtained Emerged Emerged 22 38 Parasitism 92 33 Parasitism 1 1978 7812 3953 28.4 Ta2 891 29.4 0 0 0 1980 — — — _ - = = 1981 = _ _ — — — = — = 1983 1358 251 Teel 0 20 6.4 22. 18 12.9 2 1978 2855 654 45.3 323 584 58.1 0) 0 0 1980 700 310 42.9 60 30 2255 0 0 0 1981 823 03 81.3 21 40 39.6 0 0 0 1983 480 10 93.8 10 10 66.7 0 0 0 3 1978 B27 740 30.8 803 22 67.3 0 0 0 1980 1320 680 35.6 100 70 20.0 0 0 0 1981 474 44 87.3 8 8 PX II 0 0 0 1983 1047 513 S97 39 66 24.9 2 2 0.9 4 1978 2961 410 30.1 7 910 80.2 0 0 0 1980 1233 692 28.9 102 83 DALE 0 0 0 1981 2391 2 91.4 31 62 45.4 0 0 0 1983 3250 1080 55.4 300 50 24.2 113 8 eS 5) 1978 -- = - — = — — — 1980 361 162 46.8 20 10 15.6 0 0) 0 1981 419 21 90.5 10 9 47.5 0 0 0 1983 290 180 37.9 0 0 0 0 0 0 6 1978 44] 322 27.0 0 0 0 0 0 0 1980 383 222 33.9 31 0 223 0 0 0 1981 611 7) 94.9 0 10 32:3 0 0 0 1983 1661 148 89.2 11 0 6.1 12 8 1 2 dl 1978 1658 693 19.5 291 351 48.1 0 0 0 1980 1930 1440 20"2 90 10 6.5 0 0 0 1981 681 23 96.6 0 0 0 0 0 0 1983 Sil 55 90.4 0 0 0 0 0 0 8 1978 580 50 27.6 250 120 88.1 0 0 0 1980 273 174 3653 0 0 0 0 0 0 1981 349 50 82:5 0 11 18.0 0 0 0 1983 369 0 100.0 0 0 0 0 0 0 9 1978 Sli 85 30.0 139 138 76.5 0 0 0 1980 1072 822 23.4 0 0 0) 0 0 0 1981 763 42 93.1 11 0 20.8 0 0 0 1983 = a = = = _ _ — _ 10 1978 ~ _ — = = — — = 1980 161 62 61.5 0 0 0 0 0 0 1981 378 35 90.2 0 2 5.4 0 0 0 1983 — = = = = = = 11 1978 us = = = — _ _ _ ~ 1980 a 2 = = ES — = _ _ 1981 2821 279 89.4 20 0) 6.7 — _ _ 1983 — — - = — = = = 12 1978 720 80 30.6 240 180 84.0 = = =A 1980 323 182 43.7 0 0 0) 0 0 0 1981 = — _ = = a = = = Na) oo Ww | | | | | | | | | 742 Table 2. Continued. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON i % Puparia No. Biosteres emerged! No. Tribliographa sp. Emerged' No. Adult from which No. Puparia Flies Nothing % % Site Year Obtained Emerged Emerged 92 38 Parasitism 99 36 Parasitism a 13 1978 576 387 29.2 21 0) > 0 0 0) 1980 270 160 40.7 0 0 0 0 0 0 1981 _ _ - — - - _ _ = 1983 349 181 48.1 0 0) 0 0 0 0 14 1978 _ — — - _ - — - — 1980 2930 750 65.5 110 140 24.8 6 4 1.0 1981 343 0 100.0 0 0 0 0 0 0 1983 _ — — _ _ _ _ —- _ 15 1978 203 37 28.6 58 50 74.5 0 0 0 1980 1322 923 30.3 0 0 0 0 0 0 1981 = = — — — — - — 1983 - — — _ _ — — - = | % parasitism = no. emerged parasitoids/total emerged parasitoids + hosts x 100. lative response was analyzed by correlating the log,, (initial density + 1.0) with the dif- ference between log,, (initial density + 1.0) and the log,, (final density + 1.0), 1.e. the “killing power” or ‘“‘k-value”’ of Varley et al. (1974). Correlation coefficients were all tested at P = 0.05. Determination of a parasitoid’s activity from incubation and emergence data in the laboratory, however, may underestimate its actual impact. Some hosts may be killed by the probing and oviposition of parasitoids, and die without giving rise to adult para- sitoids as suspected previously for other in- sects (Legner, 1979; Legner and Silveira- Guido, 1983). Also, although considerable care was taken to provide a natural situation for pupariation, and handling was done as little as possible, some developmental anomalies may have occurred during the pupal stage. These may result in adult emer- gence failures. RESULTS AND DISCUSSION Adults of only R. completa emerged from the nuts collected, although Rhagoletis ju- glandis (Muesebeck) is known to occur in the northwestern portion of the study area (sites 14 and 15). The first refrigeration/incubation cycle stimulated >95% of the total emergence of host flies and parasitoids in every sample (Table 2). Biosteres sublaevis was the most prominent parasitoid species reared. Try- bliographa sp. occurred at much lower fre- quencies, and always in conjunction with the former species. Parasitism was wide- spread throughout the sampled area and varied considerably from year to year at any given site (Table 2). There may have been a trend toward higher parasitism in areas protected from the full impact of storms from the north by rises of the Davis and Guadalupe Mountain ranges, whereas in the more-open, northernly exposed and wind- swept areas, e.g. 10, 11 and 13 (Tables 1 and 2), parasitism was comparatively lower. Site 12 was sheltered by the northernmost foothills of the Davis Mountains and showed relatively high parasitism. High mortality in puparia also was re- corded at all collection sites (Table 2). This mortality was not correlated with intensity of parasitoid emergence (r = —0.186, 41 df), and probably was caused by combinations of handling, parasitoid probing and aborted parasitism. There was a significant correlation be- tween the initial within walnut larval den- sity and the final adult fly emergence density VOLUME 89, NUMBER 4 in 1980 (r = 0.777, 14 df). A subsequent k-value analysis (Varley et al., 1974) also showed a significant correlation (0.494, 14 df). This indicated that fly mortality from all natural factors combined occurred in greater proportions at relatively higher ini- tial larval densities. However, it cannot be ascertained whether parasitism was the main regulative factor, because there was no sig- nificant correlation between the initial host larval density and the Biosteres density (r = 0.308, 14 df). Data pertaining to inter-tree and -walnut density might give further clues to the regulative ability of the parasitoids. However, problems associated with mea- suring the full impact of any parasitoid on its host in the wild reviewed recently (Leg- ner, 1983; Legner and Silveira-Guido, 1983) obviously also contributed to our inability to access this natural parasitism more fully. The wide distribution and high intensity of walnut husk fly larval parasitization by B. sublaevis in the surveyed areas has prompted an effort to introduce this species into California from Texas for biological control. LITERATURE CITED Berlocher, S. H. 1976. The genetics of speciation in Rhagoletis (Diptera: Tephritidae). Ph.D. Thesis, Univ. of Texas, Austin. 203 pp. 743 Boyce, A. M. 1934. Bionomics of the walnut husk fly, Rhagoletis completa. Hilgardia 8: 363-579. Legner, E. F. 1979. The relationship between host destruction and parasite reproductive potential in Muscidifurax raptor, M. zaraptor, and Spalangia endius [Chalcidoidea: Pteromalidae]. Entomopha- ga 24: 145-152. . 1983. Requirements for appraisal of the role of parasitic insects in the natural control of syn- anthropic Diptera. Proc. Calif. Mosq. and Vect. Contr. Assoc., Inc. 51: 97-98. Legner, E. F. and A. Silveira-Guido. 1983. Estab- lishment of Goniozus emigratus and Goniozus leg- neri [Hym.: Bethylidae] on navel orangeworm, Amyelois transitella [Lep.: Phycitidae] in Califor- nia and biological control potential. Entomophaga 28: 97-106. Michelbacher, A. E. and J. C. Ortega. 1958. A tech- nical study of insects and related pests attacking walnuts. Calif. Agric. Exp. Stn. Bull. 764. 86 pp. Varley, G. C., G. R. Gradwell, and M. P. Hassell. 1974. Insect Population Ecology, an Analytical Ap- proach. Univ. of Calif. Press, Berkeley and Los Angeles. 212 pp. Warnock, B. H. 1970. Wildflowers of the Big Bend Country, Texas. Sul Ross State University, Alpine Texas: 157, pp: Wharton, R. A. and P. M. Marsh. 1978. New world Opiinae (Hymenoptera: Braconidae) parasitic on Tephritidae (Diptera). J. Wash. Acad. Sci. 68: 147- 167. PROC. ENTOMOL. SOC. WASH. 89(4), 1987, pp. 744-758 REDESCRIPTION OF AN ERGOT BEETLE, ACYLOMUS PUGETANUS CASEY, WITH IMMATURE STAGES AND BIOLOGY (COLEOPTERA: PHALACRIDAE) WARREN E. STEINER, JR. AND BHISHAM P. SINGH (WES) Department of Entomology, University of Maryland, College Park, Maryland 20742; (Present address: Department of Entomology, NHB-169, Smithsonian Institution, Washington, D.C. 20560); (BPS) Plant Protection and Quarantine, Animal and Plant Health Inspection Service, USDA, Federal Building, Hyattsville, Maryland 20782. Abstract.—Acylomus pugetanus Casey (Coleoptera: Phalacridae) feeds on the toxic sclerotia of ergot (Claviceps spp.) on grains in North America. The biology and potential economic importance of the beetle as a control agent and vector of the fungus disease are discussed. Specimen data and distribution records are given. The adult, larval, and pupal stages are described and illustrated. A species of Acylomus Sharp (Coleoptera: Phalacridae) has been associated with ergot fungi of grains in Michigan (Singh, 1981) and was thought to be undescribed. An ear- lier study, apparently concerning the same species, but unidentified by the authors, at- tempted to determine the effects of the bee- tles’ feeding on sclerotial germination (Lambert and Mcllveen, 1976). The species has now been identified as Acylomus pu- getanus Casey. This species was described from a single specimen from Washington, but is now known to have a wide distri- bution across the northern United States, south to Arkansas and Virginia. The beetle feeds on the sclerotia of ergot fungi (Claviceps spp.) which are known to contain a number of toxic alkaloids, some of which are related to the drug LSD (Bar- gar, 1931; Bove, 1970). The implication of this and other species of Phalacridae in the spread or biological control of ergot and smut diseases of grasses has created an in- terest in the group, but identification of the species with the existing literature is not possible. Studies to determine if the beetles have detoxification mechanisms and coevo- lutionary relationships with the fungi and grasses, also of great interest, could be car- ried out if dcylomus species can be recog- nized. The North American species of Acylomus were studied by Casey (1916), who de- scribed all of the species presently known from the United States. The genus is pri- marily Neotropical, but some species occur northward to southern Canada. The group has not recently been studied systematical- ly, and, as with most phalacrid beetles, the biology and larval stages of most members are unknown. Acylomus pugetanus, which has been shown to be of potential economic importance, is redescribed and illustrated in this paper. Its larva and pupa are also described, with information on life history. The larval stages and ways of life of A. puge- tanus are thus far unique among the Phal- acridae, and even aberrant among the mem- bers of the genus, being different from those recently associated with a few other adult Acylomus (Steiner, 1984). A key to the species of Acylomus will be presented in a VOLUME 89, NUMBER 4 future review of North American Acylomus members. Acylomus pugetanus Casey Description. — Length |.6—2.1 mm; great- est width (at basal 4 of elytra) 1.0-1.2 mm; greatest thickness (at metasternum) 0.6-0.7 mm. Body form oval, strongly convex dor- sally, with head, pronotum and elytra form- ing a continuous oval outline in dorsal view; dorsal surface strongly shining, polished; coloration very dark brown to black dor- sally, brown ventrally, with more darkly pigmented area across metasternum; ap- pendages brown, femora darkened. Head rounded, 2 as wide as pronotum, with eyes large; frons with surface finely, randomly punctate, black, becoming brownish very narrowly along clypeus; la- brum light brown, setose; mandibles with sharp, bidentate apices; antennae setose, with 11 segments and a 3-segmented, elon- gate club with apical segment nearly as long as 2 preceding segments combined. Pronotum 2 x wider than long, widest ba- sally, narrowly margined laterally, with fine, shallow, random punctures (each bearing a minute seta) and extremely fine, transverse- ly strigulate microsculpture dorsally; pros- ternal process (Fig. 1A) abruptly truncate between front coxae, not extending poste- rior to them, with a sharp, shelflike edge bearing 5 small, posteriorly directed setae. Elytron 2x longer than wide, with mi- crostrigulate surface (Fig. 1C, D) as on pronotum; punctures in regular, alternating, longitudinal rows of larger and smaller size, shallow, crescentiform, each with a fine, de- cumbent seta; sutural stria well developed, reaching basal % of elytron. Mesosternum well developed behind middle coxae, forming sharply angulate lobe behind each coxa (Fig. 1B) with mesal side slightly curved, distal side nearly straight. Metasternum, abdominal sterna and legs se- tose. Hind tibiae without any apparent sex- ual dimorphism, slender, gradually widen- ing from base to middle, nearly parallel from 745 middle to apex; truncation of apex slightly oblique. Hind tarsi with basal segment '/ as long as second; 3rd segment of intermediate length, narrow, with apex bilobed; 4th seg- ment very small, appearing as basal part of 5th segment. Tarsal segments 1-3 with dense pubescence ventrally. Claws with apices slender, sharp; basal tooth broad, not point- ed. Male genitalia (Fig. 2a—d) with basal piece short, broad, arched dorsally, with 2 sym- metrical “false parameres”’ at apex of each side of true parameres (which are fused into a single pointed apex); basal ring asymmet- rical; dorsolateral struts slender, very weak- ly sclerotized, each with a heavily sclero- tized protuberance near base; median dorsal strut short, pointed. Inner lining sclerite lightly sclerotized, membranous basally ex- cept lateral struts; apex broadly rounded, dorsoventrally flattened, with a slightly up- turned edge in lateral view. Parameres entirely fused into a small, conical sclerite hinged and movable be- tween false parameres, with 2 pairs of long setae apicolaterally and one pair basoven- trally. Median lobe not heavily sclerotized; main sclerite with a broadly rounded, spoon shaped apex, narrow lateral rods, and a sim- ple, flattened, somewhat asymmetrical base; internal sac with 2 apical, external, dorso- lateral sclerites forming convergent, tongue- like lobes; system of internal sclerites well developed, with hook and crescent sclerites strongly sclerotized, pigmented; small toothlike spicules lining sac and sperm duct above hook sclerites, forming 2 dense lat- eral rows of longer, fine spicules immedi- ately before internal sac sclerites; sperm duct with a globular brush of fine, hairlike spic- ules where duct narrows (usually adjacent to basal tip of main sclerite). Variation. — There are occasionally 6 or 7 setae across the apex of the prosternal pro- cess rather than the usual number of 5. Also, the setae are easily broken off and in some specimens may appear to be entirely lack- 746 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Acylomus pugetanus, adult structures, scanning electron micrographs. A, Prosternal process and front coxae. 235 x. B, Left mesocoxa and mesosternal plate. 200 x. C, Dorsal surface of right elytron showing sutural edge, stria and microsculpture. 220 x. D, Detail of dorsal elytral microsculpture and setae. 1100. ing. The presence and number of setae ap- nearly so, but teneral specimens or thor- parently are important in separating species oughly degreased, dry specimens may ap- of Acylomus. pear more brownish. The unique type spec- The dorsal coloration is usually black or imen (data below) has a much paler VOLUME 89, NUMBER 4 0.2mm Fig. 2. Acylomus pugetanus, male genitalia. a—b, Basal piece and parameres, dorsal and lateral view. c-d, Median lobe, dorsal and lateral view. brownish coloration than is normal; it ap- parently has faded with age since Casey (1916) described the color as “blackish-pi- ceous.” Diagnostic characters. —In addition to the distinctive form of the male genitalia, the following combination of external features will distinguish 4. pugetanus from other de- scribed U.S. species: Blackish dorsally and very dark ventrally, with areas of metaster- num and femora darker than other parts of the venter; form of the prosternal process; sharply angulate mesosternal side plates; lack of sexual dimorphism in hind tibiae (males in some other species have tibiae broadly expanded at apex and enlarged api- cal spurs). Type data.—The unique type specimen has only the label ““W. T.”’ as a locality, with Casey’s dot code. With the description Ca- sey gave ‘“‘Washington State (Tacoma), Wickham” for the locality and collector, but Tacoma is not the locality given for the label code on the specimen. The pin partially ob- scures one of the code dots which indicates “N. Yakima,” not Tacoma, as the locality, according to Casey’s code list. The true type locality is probably North Yakima, Wash- ington. The specimen is in the Casey collection in the U.S. National Museum of Natural History, Smithsonian Institution, Washing- ton, D.C., and is identified with the type number 48994 on a red label, and Casey’s handwritten label “‘pugetanus.” Remarks.—In his redefinition of the ge- nus, Casey (1916) stated that the enlarged hind tibiae and spurs in male Acylomus 748 Fig. 3. Acylomus pugetanus, known distribution. members were typical of the genus; genitalia were not studied by Casey. Because of this, he identified the type of A. pugetanus as a female. Upon dissection, this specimen was found to be a male, with genitalic structures identical to those of specimens found throughout the known range (Fig. 3; listed below). Prior to this discovery, and because the type specimen of A. pugetanus is brown, specimens recently collected were thought to belong to a separate, undescribed species. Casey must have been aware of some kind of association of Acylomus spp. with ergot, having named another common and wide- spread species A. ergoti, but he wrote noth- ing on phalacrid biology. At present, it 1s uncertain that A. ergoti is associated with ergot. Some specimens earlier identified as A. ergoti, with labels referring to ergot as- sociation, were found to be A. pugetanus, while no specimens of A. ergoti examined bear biological data referring to ergot. Past references to 4. ergoti (e.g. Lewis, 1945) may PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON have been based on misidentifications and actually involved 4. pugetanus. Specimens examined and data.—Num- bers in parentheses refer to the number of specimens from each locality and date. Any additional label data regarding habitat or method of collection are quoted verbatim, following the name of collector. CANADA: MANITOBA: Husavick, 5 July 1915 (1), H. C. Fall. ONTARIO: Carle- ton Co.: Ottawa, 24 May 1955 (3), 21 Aug. 1956 (1), 13 Aug. 1958 (3), M. H. Hatch; Aug. 1980 (5), C. Young. QUEBEC: Ber- ther Co. Berth; 6 Sept, 19212) eae Scott; Brome Co.: Knowlton, 21 Aug. 1956 (1), M. H. Hatch; Deux Montagnes Co.: St. Placide; 10 July 1931 (1); H.C: Fall Rae mouski Co.: Rimouski, Aug. 1936 (3), H. C. Fall. U.S.A.: ARKANSAS: Hempstead Co.: Hope, 24 Nov. 1921 (1), H. C. Fall; Polk>Co:: 21 Aug: 192870) -JsGe Shaw: Washington Co.: 7 Sept. 1961 (2), W. H. Whitcomb, “‘on ergot on rye”; state label VOLUME 89, NUMBER 4 only: H. C. Fall (1). CONNECTICUT: New London Co.: Barn Island, 30 Jan. 1975 (1), L. E. Watrous, “‘leaf litter”, DELAWARE: Newcastle Co.: Newark, 15 Sept. 1953 (2), C. A. Triplehorn. DISTRICT OF COLUM- BIA: Washington, 30 May (1). ILLINOIS: Cook Co.: Chicago Heights, July 1969 (1), “light trap’; Du Page Co.: Wheaton, 6 June 1959 (1); Henry Co.: Annawan, Sauk Tr. St. Pk., 17 June 1975 (1), “at light’’; Jo Daviess Co.: 6 mi. S of Stockton, 29 July—3 Aug. 1972 (3), Reaves & Hollander, “‘at light’’; Ogle Co.: White Pines St. Pk., 24-25 June 1966 (2), R. Chevalier, “sweeping”; Pike Co.: Pittsfield, 27 June 1947 (2), 7 June 1948 (1), B. Cadwell; Union Co.: 22 July 1938 (1), W. F. Turner; Williamson Co.: Carter- ville, 6 Sept. 1958 (1), 22 Mar. 1959 (1), V. Cole. INDIANA: Benton Co.: Oxford (5); Kosciusko Co.: Winona L. (1), E. B. Wil- liamson; La Porte Co.: Michigan City, | June 1972 (5), M. Prokop, ‘“‘on beach.”” IOWA: Rage ‘Co: Norwich; 227Aug. (1); Az) 1: McClay; Pottawattamie Co.: 1 mi. W. Cres- cent, 21 Aug, 1970"(1))G. Godfrey ‘& J. Ameel; Scott Co.: Davenport, May 1964 (1), S. Peck, “‘light.””» KANSAS: Douglas Co.: 21 Mar. 1929 (1), R. H. Beamer; no date (3), F. H. Snow; Lone Star Lake, 30 Sept. 1951 (1), P. J. Spangler; Sumner Co.: Wellington (6), ED). Kelly sKENTUCKY: Hardin Co: Fort Knox, 3 July 1952 (1), R. D. Alex- ander, “Light trap.” MAINE: Oxford Co.: Paris, 9 July 1914 (1), C. A. Frost. MARY- LAND: Baltimore Co.: Baltimore, 17 June 1909 (1), F. E. Blaisdell; Calvert Co.: Ches- apeake Beach, 23 May 1970 (1), M. Druck- enbrod, “UV Light’; Carroll Co.: Elders- burg, 9 June 1985 (8), W. E. Steiner & J. E. Lowry; Dorchester Co.: Crapo, 8 Sept. 1984 (35), W. E. Steiner & D. S. Bogar, ““Reared from ergot sclerotia on Spartina patens”’; Garrett Co.: 2 mi. E Keysers Ridge, 2500’, 18 June 1968 (2), S. Peck; Montgomery Co.: Gaithersburg, 6 June 1979 (1), W. E. Stei- ner; Potomac, Aug. 1972 (1), W. E. Steiner; Prince Georges Co.: Bowie, 1-7 July 1981 749 (2), R. F. Denno, “Malaise trap’; 4 July 198411) We ES Stemerc.2E.cownyc: G: L. Williams, ““Reared from ergot sclerotia on Festuca pratensis’; College Park, 17 July 1970 (1), R. A. Belmont; 5 km SE Croom, 30 May 1986 (1), W. Steiner, S. Larcher, J. Swearingen; Talbot Co.: 3 km SE Easton (Seth Forest), 3 Aug. 1986 (1), W. E. Steiner, J. M. Hill, J. M. Swearingen; McDaniel (Wades Pt.), 1 June 1979 (2), 14 June 1980 (1), W. E. Steiner; Wittman, 28 July 1973 (@2)725 Aug: 1973.() eSepta 19 73°@).23 Sept. 1974 (1), 24 May 1975 (1), 25 July 1975 (5), 5 July 1976 (3), 28 Aug. 1976 (1), 25 June 1977 (1); 26 June 197716); 1 July 1977 (8), 5 July 1977 (1), 11 May 1979 (1), 19 July 1980 (2), 6 Sept. 1980 (1), 1 Aug. 1981 (2), 8 Aug. 1981 (5), 9 Aug. 1981 (2), 13 Sept. 1981 (1), W. E. Steiner, “At black- light near mixed forest, open fields and tidal creek”; 5 July 1982 (91), 12 July 1982 (40), W.E. Steiner, “In sclerotia of ergot on Fes- tuca pratensis”; 28 July 1985 (13), W. E. Steer, J.) Es Hownyicé& Ge jks) Walhamis: “Reared from ergot sclerotia on Festuca pratensis.”” MASSACHUSETTS: Berkshire Co.: Monterey, 9 July 1919 (2), C. A. Frost; Dukes Co.: Cuttyhunk Is., Elizabeth Is., 21 Aug. 1971 (3), C. T. Parsons, “‘Flowers of Achillea’; 17 Sept. 1972 (4), C. T. Parsons, “Sweeping Solidago”; Nashawena Is., Eliz- abethuls al Julys19 72.42) ..Cs Tz Parsons: Penekise Is., Elizabeth Is., 30 Aug. 1971 (1), C. T. Parsons, “Beating Acer platanoides’’; Hampshire Co.: Ware, 23 Aug. 1957 (1), C. A. Frost, “‘“Flowers Solidago”; Middlesex Co.: Pickman area, Bedford, 7 Aug. 1974 (10), J. F. Lawrence, ‘“‘Claviceps purpurea’; Belmont, 10 Aug. 1974 (11), J. F. Lawrence, “Claviceps purpurea’; Billerica, 7 Sept. 1944 (1), A. I. Bourne, “in grass heads”; 15 Sept. 1944 (6), A. I. Bourne, “‘In smutted seeds of grass’; Concord, June (4), A. Fenyes; Fra- mingham, 20 Nov. 1943 (1), C. A. Frost, “sifting humus”; 24 Sept. 1958 (1), C. A. Frost; Holliston, 11 Aug. (1), 6-13 Sept. (3), N. Banks; Lowell, 20 July 1893 (2), H. C. 750 Fall; Natick, 17 Apr. 1937 (1), C. A. Frost, ‘sifting’; 19 Aug. 1945 (1), C. A. Frost; Sherborn, 15 Mar. 1925 (1), P. J. Darling- ton; Tyngsboro, 30-31 Aug. 1904 (17), F. Blanchard; Norfolk Co.: Milton, 26 July 1903 (1), P. G. Bolster; Suffolk Co.: W. Rox- bury, 14 June 1908 (1), P. G. Bolster; Worcester Co.: Ashburnham, 17 July 1952 (8), M. H. Hatch. MICHIGAN: Allegan Co.: Aug. 1980 (22), B. P. Singh, “‘In sclerotia of ergot on rye”; Cheboygan Co.: 30 July 1929 (1), 28 July 1953 (1), H. B. Hungerford; 5 July 1935 (1), M. Sanderson, “Collected at light’; 26 July 1935 (1), L. R. Penner; Ing- ham Co.: East Lansing, 2 Aug. 1979 (2), B. P. Singh, “In sclerotia of ergot on wheat”; 1 Aug. 1980 (24), B. P. Singh, “In sclerotia of ergot on rye”; Livingston Co.: E. S. George Reserve, 20 June 1939 (11), I. J. Cantrall; 8 July 1955 (1), M. L. Cantrall, ““On flowers Daucus carota”, Whitmore Lake, 10 Aug. 1956 (2), G. H. Nelson; Macomb Co.: Sel- fridge Field, Mt. Clemens, 15-21 June 1944 (3), B. Malkin; Menominee Co.: Lake Mary, 8 July 1940 (1), I. J. Cantrall; Oakland Co.: Birmingham, 6 Aug. 1936 (1), M. H. Hatch; Milford, 21 May 1922 (1), T. H. Hubbell; Rochester, 10 July 1936 (1), M. H. Hatch; St. Clair Co.: Memphis, 16 Aug. 1979 (13), B. P. Singh, “In sclerotia of ergot on wheat”’; 9 Oct. 1979 (46), B. P. Singh, “‘In sclerotia of ergot on quack grass, Agropyron repens”’; 9Oct: 1979"(9); By P-Singh;, “Intsclerotia of ergot on rye”; Washtenaw Co.: 20 Aug. 1921(4)" 1Aug« 1922-(1),. MH Hatch: Locality and date unknown: “Mich. E.P.A.,” F. Blanchard (1). MINNESOTA: Aitkin Co.: Tamarack, 3 July 1936 (1), H. R. Dodge; 24 June & 12 July 1973 (4), D. E. Rau; Anoka Co.: 23 June 1936 (1), H. R. Dodge; Chisago Co.: (1), O. W. Oestlund; Clay Co.: Buffalo River St. Pk., 11 June—20 Aug. 1972 (6), E. F. Cook, ““N. J. Mosquito trap’; 26 July—9 Oct. 1973 (7), ‘““Malaise Trap’’; Crow Wing Co.: Crow Wing St. Pk., 20 July 1972 (1), E. F. Cook, “N. J. Mosquito Trap”; Douglas Co.: Brandon, 22 Aug. 1922 (1), PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Wm. E. Hoffmann; Freeborn Co.: Helmer Myre St. Pk., 30 May-14 Aug. 1972 (19), E. F. Cook, “N. J. Mosquito trap’’; Houston Co.: Beaver Creek Valley St. Pk., 2-8 July 1972 (1), “N. J. Mosquito trap”; 16-22 July 1972 (1), E. F. Cook, ““N. J. Mosquito trap’’; 23 May-2 Oct. 1973 (58), “Malaise trap”; Houston Co.: La Crescent, 2 July 1956 (6), C. S. Li; SE tip of Houston Co., 23 May 1936 (1), H. R. Dodge; 31 Aug. 1936 (1), H. R. Dodge; Hubbard Co.: Lower La Salle L., 2-24 July 1971 (4), E. F. Cook, “N. J. Mosquito trap’; Itasca Co.: Grand Rapids N. Central Exp. Sta., 10 July-1 Aug. 1973 (19), ‘“Malaise trap’’; Itasca St. Pk., 22 June— 8 July 1954 (10), C. S. Li; Itasca Park, 3 July 1939 (1), C. E. Mickel, “Light trap’’; 13 July 1941 (3), H. P. Nicholson; Itasca Park, La Salle Valley, 9 July 1940 (3), C. E. Mickel, ‘“‘at light’’; Jackson Co.: Kilen Woods St. Pk., 13 June—11 Sept. 1972 (3), E. F. Cook, ““N. J. Mosquito trap”; Kan- diyohi Co.: Sibley St. Pk., 31 May—7 June 1974 (5), E. F. Cook, “‘Malaise trap’’; Kitt- son Co.: Lake Bronson St. Pk., 7 June 1972 (1), ““N. J. Mosquito trap’; Lac Qui Parle Co.: Lac Qui Parle St. Pk., 7 June—10 Sept. 1974 (15), E. F. Cook, “‘Malaise trap’’; Mar- ietta, 27 June 1921 (1), H. H. Knight; 5 mi. S Milan, 5 July 1974 (1), B. Tollefson; Lake Co.: Basswood Lake, Washington Id., 14—- 18 Aug. 1950 (1), J. W. Barnes & R. Namba; Lyon Co.: Camden St. Pk., 9 May—25 Sept. 1973 (30), “Malaise trap’; Olmsted Co.: 9 May 1905 (1), C. N. Ainslie; Pine Co.: Little Sand Ck., 1 mi. E Cloverdale, 4 July—-11 Sept. 1973 (14), “Malaise trap’’; Pipestone Co.: Pipestone Nat. Mon., 28 May 1973-9 Oct. 1974 (93), ““Malaise trap’; Polk Co.: Red Lake River nr. Crookston, 6 June—27 Sept. 1973 (7), ‘““Malaise trap’; Pope Co.: 30 July 1939 (1), C. E. Pederson; Glacial Lakes St. Pk:, 12:Aug. 1972 (2); EuxkeG@ook: “N. J. Mosquito trap’; 31 May—-19 Sept. 1973 (27), ‘““Malaise trap’; Ramsey Co.: 9 Aug. 1922 (4), A. A. Nichol, “‘Ergot-rye”’; 23 June 1936 (1), H. R. Dodge; Lake Vad- VOLUME 89, NUMBER 4 nais, 18 Aug. 1956 (1); White Bear, 10 July 1921 (1), Wm. E. Hoffmann; Rock Co.: Blue Mounds St. Pk., 21 June 1973 (1), 23 Aug. LOTS 7) S0FSept. 1973 (8).-E. F. Cook: Roseau Co.: Roosevelt, 1 July 1964 (6), A. G. Peterson, ““Bluegrass’’; 8 mi. SE Roseau, 29 June 1961 (1), “ex Blue grass”; Stearns Co.: St. Anthony Pk., 5 June 1921 (1), Wm. E. Hoffmann; Swift Co.: 3 mi. SW Apple- ton, 4 July 1974 (1), B. Tollefson; Traverse Co.: 1 mi. NW Browns Valley, 12 June 1974 (1), B. Tollefson; 6 mi. NE Browns Valley, 19 June 1974 (3), B. Tollefson; Washington Co.: Valley Ck. nr. Afton, 18 June-21 July 1971 (4), ““N. J. Mosquito trap”; Wright Co.: Buffalo, 30 July 1947 (2), S. I. Parfin. MISSOURI: Boone Co.: Columbia, 7 June 1970 (1), F. D. Parker, ‘““Malaise trap 7AM-— 7AM”; Phelps Co.: Rolla, 24 June 1952 (4), M. H. Hatch; Rolla, Dry Fork Cr., 26 June 1950) (3), 27 May 1951 (4), H. Frizzell; 25 June 1952 (8), M. H. Hatch. MONTANA: Lake Co.: Moiese Nat. Bison Range, 17 June 1952 (1), M. H. Hatch. NEBRASKA: Lan- caster Co.: Lincoln (4), F. M. Webster. NEW JERSEY: Hudson Co.: Arlington, 4 June (1); Middlesex Co.: Morgan, 4 May (1), Weiss & West; Monmouth Co.: Horners- town, 14 May 1910 (1). NEW HAMP- SHIRE: Grafton Co.: Rumney, 16 June 1924 (1), P. J. Darlington. NEW YORK: Essex Co.: Elizabethtown, July 1981 (1), A. E. Lacy; Tompkins Co.: Ithaca, Savage Farm, 5 Aug. 1968 (1), A. G. Wheeler, “‘Vernal Alfalfa”; McLean Bogs, 30 May 1919 (1), Inlet Brook, McLean Res., 22 May 1925 (1); Ulster Co.: Oliverea, 12 July 1919 (1), E. Shoemaker; Slide Mt., Catskills, 19 June 1918 (1); Wayne Co.: 14-17 Sept. 1948 (3), E. Shoemaker. OHIO: Ashtabula Co.: An- dover (2), F. M. Webster; Clinton Co.: 18 May 1963 (1), F. J. Moore; Delaware Co.: 2 yunerGh)s3 Sept 1957 (2), WD: J. & J. N: Knull; Franklin Co.: 25 May 1963 (1), F. J. Moore; Columbus, 1957 (1), L. Schramm; Greene Co.: 12 May (2), 2 June (1), 18 May 1959 (1), D. J. & J. N. Knull; Hocking Co.: 751 20-May (1), 15 Sept. 1958 (1), D. J. & J. N. Knulls Mercer Co:: 35-July (@); &.-H. Parks; “from ergot”; Ross Co.: Tar Hollow St. For., 16 Feb. 1976 (1), L. E. Watrous, ““Berlese pine duff’; Trumbull Co.: Hubbard, 16-17 June 1975 (1), A. Newton & M. Thayer, “‘on flowers & vegetation.”” OREGON: Benton Co.: 12 mi. S Corvallis, 13 March 1949 (2), V. Roth, “‘in moss’’; Jackson Co.: Prospect, 5 May 1939 (1), A. T. McClay; Multnomah Co.: Portland, 6 June 1915 (1), A. K. Fisher; Tillamook Co.: Sandlake, 5 July 1939 (1), K. M. & D. M. Fender. PENNSYLVANIA: Delaware Co.: 20 June 1894 (2), Geo. M. Greene; Huntingdon Co.: Furnace, 11 Aug. 1974 (7), D. Lambert, “‘ergot on rye’’; Mon- roe Co.: Effort, 6 Aug. 1930 (7), 2 July 1939 (2), J. W. Green; Northampton Co.: Easton, 4 July 1926 (3), 30 June 1947 (1), J. W. Green. SOUTH DAKOTA: Roberts Co.: 12 mi. SE Sisseton, Lk. Traverse, 24 June 1974 (1), 19 July 1974 (3), B. Tollefson. UTAH: Box Elder Co.: Mouth Bear River, 17 July 1914 (2), A. Wetmore. VERMONT: Ben- nington Co.: East Dorset, 22 July 1935 (1), C. T. Parsons; Manchester, 25 June 1970 (1), C. T. Parsons, “black light’’; Lamoille Co.: Stowe, 22 July 1981 (1). VIRGINIA: Accomack Co.: Wachapreague (Parramore I.) 19° May 197351); "W. Es stemer, “in sweep of dune grasses”; Bath Co.: Blowing Springs Rec. Area, 7 July 1973 (1), M. Druckenbrod, “‘Beatg.”’; Warren Co.: 3 July 1973 (2), M. Druckenbrod, “Beating.” WASHINGTON: King Co.: North Bend, | Sept. 1940 (2), M. H. Hatch; Snohomish Co.: Silver Lake, 20 July 1939 (1). WEST VIRGINIA: Hampshire Co.: 8 km NW Ca- pon Bridge, Buffalo Gap Camp, | 2-14 Sept. 1986 (1), W. E. Steiner & J. M. Swearingen, **At black light, edge of mixed forest, sandy soil slope’; Marion Co.: Fairmont, 1929 (5), Musgrave; Randolph Co.: Spruce Knob Mtn., 6 July 1973 (1), M. Druckenbrod, “Beatg.”; Summers Co.: 5 km NE Talcott at Greenbrier River, 30 May 1982 (8), Stei- ner & Collins, “At light.”” WISCONSIN: Tz PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Acylomus pugetanus, mature larva. a, Dorsal habitus. b, Ventral view of head. c, Abdominal spiracle. d, Right mandible, ventral view. e-f, Abdominal apex, dorsal and lateral view. Length of larva 3.5 mm. Kenosha Co.: Chiwaukee Prairie, 3 Sept. 1967 (1), W. Suter, “sweep”; Waukesha Co.: Dousman, 11 June 1959 (2). Acylomus pugetanus, Mature Larva Description. — Length 3.0-3.6 mm; width of head capsule 0.4—-0.45 mm; body elon- gate, somewhat flattened dorsoventrally; head prognathous. Color whitish, with sclerotized parts (head, thoracic and ab- dominal sclerites, urogomphi, legs, spira- cles) brownish; dorsal sclerites, especially on thorax, with lighter pigmented areas lat- erally; sclerites with setae of varying size, arranged as illustrated (Fig. 4a); some sur- faces finely granular. Head robust, wider than long, with dorsal outline nearly semicircular; large dorsolat- VOLUME 89, NUMBER 4 T33 Fig. 5. Acylomus pugetanus, mature larva, scanning electron micrographs. A, Mouthparts, anterior view, with right mandible partially broken out. 266 x. B, Head and prothorax, dorsal view. 92x. C, right mandible, dorsal view. 550. D, Mola of right mandible. 2100~. eral, lateral and ventrolateral setae as illus- trated (Figs. 4a, 4b, 5b); a cluster of six ocelli on each side of head immediately posterior to antennae, with dark pigment spots be- neath them; epicranial suture not pigment- ed, lyriform, with stem absent. Antennae very short, 3-segmented; Ist segment wider than long, resting loosely on a membranous base; 2nd segment cylindrical, slightly long- er than wide, with 2 small subapical setae and an apical, conical accessory process; 3rd segment very small, as long as accessory process, cylindrical, with setae and sensillae apically. Mandibles (Figs. 4d, SA—D) short, robust, heavily sclerotized, palmate, with 3 apical teeth, the middle one most prom1i- nent; smaller accessory teeth on inner edges of dorsal and ventral teeth; pseudomola prominent, well developed, with a concave, toothed, grinding surface; basal molar area smooth; 2 setae on lateral surface of man- dible. Labrum transverse, broadly rounded anteriorly, with a few small setae along api- cal edge. Maxillae with cardo, stipes and mala fused, stout, setose as illustrated (Fig. 4b); palpi 3-segmented, nearly as long as 754 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON antennae. Labium with mentum small, bi- setose; prementum broad, with 2 setae ven- trolaterally; ligula short, rounded, with 2 apical setae; palpi short, 2-segmented. Thoracic segments with dorsal sclerites not pigmented along midline; with small ac- cessory sclerites anterior to these on meso- and metathorax; legs robust, short, with a few scattered, small setae; tarsunguli sharp- ly, finely pointed. Abdominal segments 1- 7 with dorsal sclerites broad, trapezoidal to oblong; lateral sclerites small; ventral scler- ites of basal segments lightly sclerotized, becoming much more hardened and pig- mented on posterior segment; 8th abdom- inal segment heavily sclerotized except for narrow lateral division between tergum and sternum; 9th segment completely, darkly sclerotized, with roughly granular areas me- dially and laterally; urogomphi strongly up- turned, with sharply pointed apices (Fig. 4e, f); dorsal articulation between abdominal segments 8 and 9 internally with 4 ante- riorly directed, darkly sclerotized, pointed spicules (visible beneath the integument). Spiracles on mesothorax and abdominal segments 1-8; openings annular, with a curved lateral duct (Fig. 4c). Variation. — The extent of darkness of the sclerotized parts of the body is variable; this may be because of the age of the larva since the last molt. Setae vary in size and exact placement among individuals, and the smaller setae (e.g. on dorsum of abdomen) may vary in number. In larvae in the pre- pupal stage, the head assumes a hypogna- thous position because the thoracic seg- ments become arched dorsally. Acylomus pugetanus, Pupa Description. — Length including urogom- phi 2.4-2.5 mm; width 1.2-1.3 mm; body white, eyes often darker (reddish to black, depending on age); larger setae light brown; body form oval, convex and arched dor- sally, with head hidden in dorsal view; body with many long setae, as illustrated (Fig. 6a— Cc); setae stiff, erect, finely tapered. Head with 6 setae on each side of frons and | above (on) each eye; a posterolateral seta usually much larger than others; clyp- eus with 4 small, inconspicuous setae along edge; 2 larger setae on labrum, and 1 on each mandible; palpi directed posteriorly, parallel with body midline; antennae par- tially hidden, lying beneath front femora, with apices between elytra and hind angles of pronotum. Pronotum heavily setose dorsally except on midline; 1 seta on each side above front angle of pronotum much larger than others; 2 ventral setae arising medial to lateral mar- gin, directed mesally. Mesothorax dorsally with 3 small setae in a triangular arrange- ment on each side; metathorax with 5 setae on each side, as illustrated (Fig. 6c). Elytra nearly meeting each other ventrally; sur- faces smooth, with shallow longitudinal striae; a single seta near middle of basal ' of each elytron; hind wings completely hid- den beneath elytra. Front and middle legs in a similar posi- tion, with femora and tibiae oriented obliquely to midline; femora each with 3 setae on apex and a row of 3 setae along anterior face. Hind legs not setose, hidden beneath elytra except for tips of tarsi. Abdomen with setae small, inconspic- uous dorsally, larger and more prominent laterally, with a larger seta at posterior cor- ners of each of segments 1—6; each side of segment 8 with an extremely long, promi- nent seta laterally; segment 9 without setae, urogomphi not sclerotized, with finely ta- pered, slightly upturned and divergent api- ces. Genital segment in male (Fig. 6d) trans- verse, twice as wide as long, with a central transverse groove; in female (Fig. 6e) with 2 rounded papillae with short tips directed posterolaterally. Spiracles on abdominal segments 1-7 annular, inconspicuous; spi- racle on segment 8 very small, vestigial. Variation. — Number, size and placement of setae, particularly on the dorsum, varied slightly among individuals of the series ex- amined; the single elytral seta is often ab- VOLUME 89, NUMBER 4 ul, / \i \ MW, Heer \\s x a Fig. 6. Acylomus pugetanus, pupa. a, b and c, Ventral, lateral and dorsal views respectively. d, Ventral abdominal apex of male and, e, Female. Length of pupa 2.5 mm. sent. Smaller setae apparently vary in num- ber more often than the larger “key” setae. The largest setae on the head, pronotum, and sides of the abdomen, and those of the femora, are apparently constant in number and placement. Remarks.—Pupal specimens have the larval exuvium attached to the apex of the abdomen; this conceals the urogomphi, gen- ital segment, and the long setae of seg- ment 8. Larval and pupal material examined.— MARYLAND: Talbot Co.: Wittman, 5 July LOS2i Gio EX6P) 2 July 1982 3: L); WaE- Steiner, ‘In sclerotia of ergot on Festuca pratensis”; 28 July 1985 (13), W. E. Steiner, J.E. Lowry & G. L. Williams, ““Reared from ergot sclerotia on Festuca pratensis.” MICHIGAN: Allegan Co.: Aug. 1980 (14 L, 13 P), B. P. Singh, “‘in sclerotia of ergot on rye”; Ingham Co.: East Lansing, 2 Aug. 1979: (GL; WP); BoP: Singh; imesclerotia of ergot on wheat”; 8 Aug. 1980 (21 L, 25 P), B. P. Singh, “‘in sclerotia of ergot on rye”; St. Clair Co.: Memphis, 16 Aug. 1979 (6 L, 6 P), B. P. Singh, “in sclerotia of ergot on wheat”; 9 Oct. 1979 (1 L, 10 P), B. P. Singh, “in sclerotia of ergot on quack grass, Agro- 756 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON pyron repens.’ PENNSYLVANIA: Hun- tingdon Co.: Furnace, 11 Aug. 1974 (4 L, 6 IP), Dykambert:is erzouoniye.. Larvae and pupae were found in associ- ation with adult A. pugetanus in all of the above collections, and in several cases lar- vae and pupae were reared to the adult stage in the laboratory to confirm their identity. Specimens of larvae and pupae are pre- served in alcohol; a few larvae were cleared and kept in glycerine. The material is in the USNMNH collection, Smithsonian Insti- tution, Washington, D.C. BIOLOGY AND DISCUSSION Life cycle and feeding habits. —Acylomus pugetanus has been found feeding during the summer months on the sclerotia of ergot fungi (Claviceps spp.) on grains (wheat and rye), quack grass (Agropyron repens (L.) Beauv.), Festuca pratensis Huds., and Spar- tina patens (Ait.) Muhl. The sclerotia are eaten by both adults and larvae; gut con- tents of dissected adults and larvae were made up of only the dark fungal tissue. Lar- vae apparently consume more of the fungal mass than adults because they develop in- side the sclerotia and hollow them out as they feed. Adult beetles lay their eggs on the surface of the sclerotium. Larval development time has not been closely studied, but is fairly rapid (1-2 weeks) and there are apparently 3 instars. Several larvae (up to 7 but usually 2-3) can develop in a single sclerotium. Pu- pal cells are formed inside the sclerotium and adults emerge in 2—3 weeks. The entire life cycle, then, takes roughly | month and occurs during late summer. In early July 1982 near Wittman, Talbot County, Maryland, A. pugetanus adults, lar- vae and pupae were collected from sclerotia of ergot on grasses growing along the edge of corn and soybean fields and mixed for- ests. Most sclerotia were on Festuca praten- sis and a few were found on Agropyron re- pens. Nearly all sclerotia examined were infested by 4. pugetanus; adult beetles were emerging when sclerotia were collected and continued emerging until the end of July. Sclerotia were dissected to determine the number of beetles per sclerotium and extent of feeding. A total of 308 sclerotia from in- florescences of F. pratensis collected on 5 and 12 July were examined. Only 2 were intact (without feeding damage); 306 were at least partially eaten and usually contained a larva, pupa, newly eclosed adult or empty pupal cell of A. pugetanus. Ten of the sclero- tia contained 2 individuals; all others in- fested (296) had produced only a single bee- tle. The small size of these sclerotia (2-5 mm long) probably is the reason for the lower number of beetles per sclerotium as compared to that in rye and wheat ergot. Most sclerotia were thoroughly hollowed out by the larval feeding but often the exposed tip of the sclerotium was left uneaten. Period of activity.—In Pennsylvania, Lambert and Mcllveen (1976) first noted adult A. pugetanus in mid June on rye and other grasses infested with ergot in the hon- eydew (infective conidial) stage; similarly in Michigan, beetles appeared on infested grasses at this stage. Adults were collected or seen from then until frost in mid Sep- tember. In Maryland (Dorchester County, Crapo) beetles emerged until late September from ergot sclerotia collected 8 September 1984 on Spartina patens. At Wittman, Tal- bot County, Maryland, 4. pugetanus spec- imens were collected at blacklight from 11 May to 13 September. The early appearance of adults in May and their presence until frost suggest that they overwinter as adults; a few other specimens labeled as being col- lected by sifting leaf litter and moss in the winter months further confirm this. The number of generations per year has not been determined. If their development is dependent on the ergot disease cycle, it is likely that they are univoltine, but alternate hosts of the beetle (e.g. ergot on wild grasses) may also be involved and allow for multiple generations each summer. Singh (1981) has demonstrated the lack of host specificity in VOLUME 89, NUMBER 4 A. pugetanus, but the total ergot and grass host range is unknown. Disease relationships.—If A. pugetanus adults are in fact attracted to the conidial stage of ergot fungi, they could be an im- portant vector of the disease. Beetles could easily pick up the sticky honeydew bearing the conidia and fly long distances to other susceptible flowering grasses, as well as in- fect other ovaries on the same inflorescence or adjacent plants. The lack of host speci- ficity in the beetle would also facilitate the spread of the fungus from wild reservoir hosts to grain crops. The active feeding on the sclerotia by adults and larvae, on the other hand, is po- tentially beneficial in reducing disease in- oculum for the following year. In sclerotia heavily infested by A. pugetanus, percent germination and number of stromata pro- duced were greatly reduced (Lambert and Mcllveen, 1976). The value of A. pugetanus as a biological control agent has not been determined. The vector role of the beetle may outweigh the beneficial aspects of its feeding. A systematic study to determine the num- ber of Acylomus species, their distributions, host ranges, and biologies, is in progress. This will allow further studies on the rela- tionships among species of Acylomus, Clav- iceps, and host grasses, which will aid in evaluating the agricultural importance of these beetles. ACKNOWLEDGMENTS This work was supported in part by a co- operative agreement between the Univer- sity of Maryland and the Biosystematics and Beneficial Insects Institute, Agricultural Re- search Service, USDA. We thank the U.S. National Museum of Natural History, Smithsonian Institution, for providing work space and equipment. John M. Kingsolver, Systematic Entomology Laboratory, Agri- cultural Research Service, called our atten- tion to specimens and literature. We also thank John M. Kingsolver, Lloyd Knutson, 757 and Theodore J. Spilman for their encour- agement and critical reviews of the manu- script. We are indebted to the following insti- tutions and curatorial staff for their gener- ous loan of material used in this study: American Museum of Natural History, New York, New York (L. H. Herman); California Academy of Sciences, San Francisco, Cali- fornia (D. H. Kavanaugh); Division of Entomology and Parasitology, University of California, Berkeley, California (J. A. Chemsak); Department of Entomology, University of California, Davis, California (R. O. Schuster); Department of Entomol- ogy, Cornell University, Ithaca, New York (L. L. Pechuman, J. A. Schafrik); Division of Insects, Field Museum of Natural His- tory, Chicago, Illinois (L. E. Watrous); Snow Entomological Museum, University of Kansas, Lawrence, Kansas (P. D. Ashlock); Los Angeles County Museum of Natural History, Los Angeles, California (C. L. Hogue); Museum of Zoology, University of Michigan, Ann Arbor, Michigan (L. Blu- mer, T. E. Moore); Department of Ento- mology, Fisheries and Wildlife, University of Minnesota, St. Paul, Minnesota (P. J. Clausen); Museum of Comparative Zoolo- gy, Harvard University, Cambridge, Mas- sachusetts (A. F. Newton); Department of Entomology, Ohio State University, Co- lumbus, Ohio (C. A. Triplehorn); Depart- ment of Entomology, Oregon State Univer- sity, Corvallis, Oregon (G. L. Peters); U.S. National Museum of Natural History, Smithsonian Institution, Washington, D.C. (D. M. Anderson, J. M. Kingsolver). LITERATURE CITED Bargar, G. 1932. Ergot and Ergotism. Gurney and Jackson, London, Edinburgh, pp. 279-297. Bove, F. J. 1970. The Story of Ergot. S. Karger, Basel (Switzerland), New York. Casey, T. L. 1916. Some random studies among the Clavicornia. Memoirs on the Coleoptera 7(2): 35— 292. Lambert, D. H.and W. D. MclIlveen. 1976. Acylomus 758 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON sp. infesting ergot sclerotia. Annals of the Ento- Steiner, W. E., Jr. 1984. A review of the biology of mological Society of America 69: 34. phalacrid beetles, pp. 424-445. Jn Wheeler, Q. Lewis, R. 1945. The field inoculation of rye with and M. Blackwell, eds. Fungus-Insect Relation- Claviceps purpurea. Phytopathology 35: 353-360. ships, Perspectives in Ecology and Evolution. Co- Singh, B. P. 1981. A new species of insect that feeds lumbia University Press, New York. on ergot sclerotia. Phytopathology 71: 255 (ab- stract). PROC. ENTOMOL. SOC. WASH. 89(4), 1987, pp. 759-769 MATING BEHAVIOR AT FLORAL RESOURCES IN TWO SPECIES OF PSEUDOMASARIS (HYMENOPTERA: VESPIDAE: MASARINAE) ROBERT W. LONGAIR Department of Zoology & Entomology, Colorado State University, Fort Collins, Col- orado 80523; Current address: Dept. of Zoology, Erindale College, University of Toronto, Mississauga, Ontario LSL 1C6, Canada. Abstract.—Males of Pseudomasaris vespoides and P. zonalis (Hymenoptera: Vespidae) patrolled patches of flowers where females obtained pollen and nectar for provisioning nests. Females of each wasp species foraged only at a single plant species in the study locality, and males restricted searching to areas and times when those plants were flowering. In P. vespoides, both males and females spent more time in larger patches, but females spent less time in patches when males were present, probably because males harassed them. Males patrolled several patches, but frequently remained within one patch for extended periods, perching and investigating insects which entered the patch. Absolute numbers of males were low, and while interactions between males were thus rare, they were sometimes intense. No size difference could be distinguished between males that copulated and males not observed to copulate. Available data on mating behavior of Pseudomasaris and other masarine wasps indicates that males search for females at lo- cations where females are concentrated in time and space, whether this occurs at nest sites, water or floral resources. Males are predicted to search for females at locations where receptive females are highly concentrated. In insects, if large numbers of emerging females are easily lo- cated, for example in aggregated nest sites (refs. in Thornhill and Alcock, 1983), mat- ing may occur there. If nests are difficult to locate, or nest densities are low, males are predicted to search for females at requisite resources which are spatially and tempo- rally restricted in availability (Emlen and Oring, 1977; Thornhill and Alcock, 1983). Patterns of male searching are expected to be influenced by both resource distribution and by female distribution at resources. The Masarinae (Hymenoptera: Vespidae) are an unusual group of solitary wasps that closely parallel the behavior of bees in pro- visioning their nests. Food for larvae con- sists of a pollen-nectar mixture, the source of which is often a limited number of plant species. North American Masarinae (sensu Carpenter, 1982) belong to the genus Pseu- domasaris and are restricted to areas of moderate elevation in the western part of the continent (Richards, 1962, 1963). De- spite some confusion in earlier reports con- cerning foraging specificity, most species probably use just one or a few host plants, particularly in a given locality (Cooper, 1952; Cooper and Bequaert, 1955). Pseudomasaris vespoides specializes on flowers of Penstemon spp. (Scrophularia- ceae), and males mate with females there (Hicks, 1929) but details of resource distri- bution have not been closely examined. 760 Similarly, males and females of P. zonalis have been collected at Phacelia spp. (Hy- drophyllaceae), but little is known of spe- cific details of behavior there. Males of sol- itary Hymenoptera often display one of two major behavior patterns: 1) patrolling of areas where females are present and 2) def- ence of areas in which females are concen- trated. Males of bees that specialize on one forage plant species often defend patches of that plant (e.g. Eickwort, 1977; Alcock et al., 1981), and I predicted that where Pen- stemon or Phacelia was patchily distribut- ed, males of P. vespoides and P. zonalis would show similar behavior. Specifically, males of Pseudomasaris spp. are expected to concentrate mate searching at forage plants where females come for larval pro- visions and will defend discrete patches where females are most common. The dis- tribution and behavior of males within and among patches should be influenced by the distribution of plants and patterns of re- source use by females. In Hymenoptera, the influence of male body size on mating success is well docu- mented for some species (Thornhill and Al- cock, 1983) and, where male-male aggres- sive behavior occurs, larger males often experience greater mating success. Female Pseudomasaris construct mud nests, attaching them to trees and rocks (but see Dorr and Neff, 1984 for one exception of a cavity nesting female of P. marginalis). These nests contain several cells, up to 13 for P. vespoides (Hicks, 1929) which are mass provisioned. Females scrape soil particles, which are mixed with liquid (nectar regur- gitated from the crop in at least one case (Torchio, 1968)) for nest construction. In- dividual nests are usually dispersed and very difficult to locate (Torchio, 1968; Richards, 1963), though they may be common in some areas (Torchio, pers. comm.). This study examines the behavior of males and females of P. vespoides and P. zonalis at flowers with respect to male mate-locat- ing behavior, male-male and male-female PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON interactions, and resource distribution and availability, and the effect of male size on mating success. Stupby AREA AND METHODS Observations were made from 22 June to 7 July 1981, 7 July to 21 July 1982, and 20 June to 18 July 1983, at Hewlett Gulch (el- evation 2000 m), about 30 km northwest of Fort Collins, Colorado (Larimer Co.). A small stream flows through the gulch, which contains several open meadows. Forage plants for Pseudomasaris spp. were patchily distributed throughout the gulch. The main study site for P. vespoides was a large open meadow marked by many large, discrete clumps of Penstemon unilateralis Rydb. Thirteen major clumps and numer- ous isolated plants were present (Fig. 1). Clumps were categorized by number of stalks at the beginning of 1982 and 1983, and in 1983 periodic determinations of flower numbers were made. Pseudomasaris activity was censused at specific clumps, and sequential checks of clumps were made several times each day to determine the presence or absence of males and females and marked individuals. In 1983, an additional large, dense clump at an isolated site further up the gulch was also studied. In addition, observations were made on each study day along a transect of several km leading to the study site. This allowed examination of other plants, sec- tions of dirt roadway, stream crossings and other areas in the gulch for presence or ab- sence of masarine wasps. Observations on male activity are taken from 1982 and 1983 data. Individual males were captured at flowers and marked with dots of colored enamel paint on the thorax. Head width measurements were made with Mitutoyo dial calipers. Times are given in Mountain Daylight Time. RESULTS At least three species of Pseudomasaris occur in Hewlett Gulch. P. vespoides, one VOLUME 89, NUMBER 4 761 : a hill b 11 July hill 12 July YO—-as bs ©) oD) YG Se) SS 6) mre se se Se road S<> Foch meadow ee re) meadow “o>. S 91 0 SS stream 25m 55m C d hill a 15 July 13 July 1300 Re. > D ee) Gps 80 as—'2 = * 1 ye YG GED CS —_YO GED UI (o-RB FA fa lor" } Ss No — \ road 2 oad Sa etiam belie Ase ers @) 7G ‘@) meadow a meadow ? BY J }? Y of Yy oy pi; Yj 25m 25m e f hill ? 15 July 15 July 1400 1500 ex 25S oe vomigesy. C80 ae coe aes 5 . SS Ne} WC THe ee as O)-3P Sie] 5} roed SS s (GD) m W eado meadow ie) ene ee str Sey stream 25m 25m Fig. 1. Main study site indicating clumps of Penstemon (stippled shapes) visited by males of Pseudomasaris vespoides in 1983. Dotted and dashed lines enclose clumps in which males patrolled or perched. Cross-hatching shows wooded area. YG, YO, 2F1, Ol, RB, FIR, O, G, 2S, BO, 3Y, BY, SB and 3P are male wasps. Maps a-— d indicate changes in which clumps were patrolled or occupied over five days, d—f indicate changes in which clumps were patrolled over three hours on one day. 762 of the larger species in the genus, appears to be a specialist on several species of Pen- stemon (Richards, 1963), although other species may sometimes be used for forage (Tepedino, 1979). P. zonalis visits flowers of Phacelia spp. (Richards, 1963). Despite repeated searches for nests of these species, none were found, although numerous suit- able nesting sites appeared to be present. Pseudomasaris vespoides This species was observed only at Pen- stemon unilateralis, which flowers from late June/early July to late July/early August at Hewlett Gulch. While systematic sampling of other areas was not carried out, numerous other flowering plant species were observed each day, as were stream crossings and the dirt road, but individuals of P. vespoides were never seen, except flying between patches of P. wnilateralis. Plants were patch- ily distributed throughout the gulch, but the largest concentration was in a large open meadow. In 1983, the size of clumps of plants ranged from 2 to 158 stalks/clump in the gulch (%¥ = 61.3 + 52.5, N = 19), and in the main study site from 6 to 158 stalks/ clump (X = 61.6 + 56.5, N = 14). The max- imum mean number of open flowers/stalk was 15.6 on 12 July. Peak populations of P. vespoides occurred at different times in the three years, presum- ably due to differences in weather among years. In 1981, wasps were observed from 26 June to 6 July; in 1982 from 7 July to 21 July and in 1983 from 1 July to 18 July. During inclement weather (rain or cloudy or windy conditions), wasps were either ab- sent or, under less severe conditions, males spent more time on the ground or on perch- es (see also Alcock, pers. obs. in Alcock et al., 1978). During rains, males occasionally sought refuge in the corolla tube of P. uni- lateralis flowers, resuming normal behavior when the sun came out (see Hicks, 1929; Cooper, 1952 for similar observations). Males and females may also spend the night in flowers (Gwynne, pers. comm.). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON All clumps, and isolated single plants, were visited by males at some time. Few wasps were observed prior to 1000. Between about 1000 and 1500, when most wasps were active, the proportion of censuses dur- ing which clumps were occupied or visited by at least one male ranged from 0.5 to 1.0 in 1982 (x = 0.70 + 0.15; 138 censuses of 10 clumps) and from 0.25 to 1.0 in 1983 (X= 0:63. + (0:20; 22) censuses ony clumps). For females, the equivalent figures are’ 0:33 to 0:67 in 1982+ =10.56 On: 43 censuses of 6 clumps) and 0.33 to 1.0 in 1983 (x = 0.66 + 0.31; 100 censuses on 13 clumps). Females. — Females foraged for pollen and nectar in a manner similar to that of many bees (Kevan and Baker, 1984), moving up a stalk in a spiral, entering most open flow- ers, occasionally visiting some blossoms more than once, and departing upon reach- ing the unopened immature flowers at the top of the stalk. Females foraged rapidly (Xx = 3.4 + 2.7 s/flower; N = 95 flowers; 17 observations of 14 females). Numerous stalks in each clump were visited and fe- males often visited more than one clump on a single foraging trip, before departing from the study site. The duration of female foraging visits within a clump varied widely, from 20 s to over 480 s. All but one (the exception being an unusually long visit) were from 20 s to 160 s (¥ = 47.7 + 42.25, N = 20). The amount of time a female spent foraging in a clump on one visit was posi- tively correlated with the number of flow- ers in the clump (Kendall’s Tau: S = 56, 0.025 < P < 0.05). Numbers of females showed a typical pattern of increase and de- crease over the course of the season (Fig. 2). Males.—In three years of study, all 60 males seen were marked and measured (11 in 1981, 21 in 1982, 28 in 1983), indicating the small numbers of males present at these sites. A large proportion of marked males was not observed after the day of marking (42% in 1981, 67% in 1982, 54% in 1983). Males present on days subsequent to mark- VOLUME 89, NUMBER 4 Mean No. of Females ieee oe 4S. 1G. 7. 819 MO) Ti 12 13514545) 16: 37 18 July Date Fig. 2. Mean number (+SE) of females of Pseu- domasaris vespoides (based on instantaneous samples) present in clump PH4 during 1983. ing were present from 2 to 9 days (1981 * = Sa see Tide wl 982 xX = 420 == 1.4.:ds 1983 X= 4.9 + 2.8 d). The maximum number of individually identifiable males (newly ob- served males and males previously marked) observed on different days was 8 in 1981, 12 in 1982 and 8 in 1983, although late in the season the number declined to a single male (Fig. 3). The date of the maximum number of males varied from year to year (1981—3 July; 1982—13 July; 1983—11 July). Head capsule measurements of males were not significantly different in the three years of the study (Kruskal-Wallis test: x7, = 3.32, 0.10 < P < 0.20). Head capsule widths ranged from 3.55 mm to 4.0 mm (X= 3.81 + 0.11, N = 60) (Fig. 4b). One female was measured in 1983, and had a head capsule width of 4.05 mm. Males displayed two relatively distinct mate searching behaviors, and individuals sometimes alternated between them. Males flew between numerous clumps in sequence (patrolling), perched and flew within a single clump, or both. Males patrolled a subset of available clumps for extended periods, reiterating a particular route. Usually, clumps were rap- idly examined by hovering, but occasion- ally, flight was interrupted by feeding at flowers, or by approaching other insects. Pa- trolled areas varied during the study. Males 763 Number Oo _ ny wo oS uo [o>] N @ Ie) (Sees Oe On va Saco: 1Omil 2) 19) 14 45.169 17 A8 July Date Fig. 3. Seasonal distribution in 1983 of Pseudo- masaris vespoides. a) Total number of males observed (A); b) number of males newly marked (@). Frequency histogram indicates number of copulations observed. Asterisks indicate days with no observations. usually patrolled some of the same clumps throughout a day, and occasionally on the following days as well. Patrolled areas ex- panded or contracted, but sometimes males shifted to an entirely different set of clumps (Fig. 1). In the three years, the number of clumps in which a given male was observed ranged from one to five over the season (X = Ose TINE = 56): Patrolling flights included both large and small clumps. The mean length of visits to clumps for individual males varied from | s to 48 s (x = 4.73 + 7.9 s, N = 79), and the number of visits to a specific clump by one male during 10 min censuses varied from 2 to 10. Occasionally visits included feeding or interactions with females, thus lengthening the visitation period. For ex- ample, the duration of visits of one male ranged from 12 s to 47 s (* = 23.6 + 12.2, N = 11); if those visits involving feeding or interactions with conspecifics are excluded (i.e. only surveys for females are consid- ered), the range narrows to 12 to 19 s (x¥ = 15.3 + 2.7 s, N = 6). When only survey flights are considered, the length of time males spent examining a clump was posi- tively correlated with the number of flowers in a clump (r = 0.71, P < 0.0001, N = 58). Since males patrol numerous clumps se- quentially, clump size may affect the num- ber of clumps patrolled. The relation be- 764 roportion Oo Oo ye) ine) Oo ao P oO an s a 3.55 3.6 3.65 3.7 3.75 3.8 Head Capsule Width (mm) 3.85 3.9 3.95 4 oe N=60 Proportion ° a 3/55 1S Ome SGD 3 :/ue OOo OM SSO) 610-0), Go.0o 4 Head Capsule Width (mm) Fig.4. Size frequency distribution of males of Pseu- domasaris vespoides (1981-1983). Graph a, copulating males; b, all males. tween mean time away from a clump and the number of stalks in a clump was ex- amined, but found to be non-significant (Kendall’s Tau: S = —9, P < 0.10, N = 13). Similarly, number of male visits was not linearly related to mean time away from the clump. In contrast to patrolling flights between clumps, males sometimes spent extended periods within a single clump. Males perched constantly on, or beside, a given plant for long periods of time, taking flight periodi- cally to investigate flowers and responding to the presence of other insects, and then returning to the perch. Perch sites were usu- ally located on the periphery of clumps, es- pecially at small clumps or near isolated plants on patrolled routes. During obser- vation periods, the time spent within a patch ranged from 300 s to 485 s (x = 416.2 + 77.0 s, N = 5). Time males spent in the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON clump under these circumstances was di- vided between perching and investigative flights. The percentage of time in flight var- ied from 8.5 to 59.6 (excluding one 6 min 16 s survey during which the male never took flight) (¥ = 39.1 + 23.3, N = 5 surveys of 5 males for 36.6 min). For six males ob- served for a total of 60 min (mean obser- vation time/male = 8 min 56 s) the mean rates of various activities were as follows: male approaches to females 0.1 1/min (SD = 0.11); feeding episodes 0.43/min (SD = 0.39); approaches to other insects 0.27/min (SD = 0.31). Each male foraged on one to nine flowers per feeding episode (X = 4.73 + 2.6, N = 57 flowers: 12 bouts of 9 males). The percentage of time males were re- corded in clumps during censuses varied from 22.7 to 87.3% (X = 53.4 + 28.9%, N = 5 censuses of 4 males for 52.9 min total). Absences usually consisted of a series of short departures (e.g. BBY male, time away from clump X = 23.9 + 17.5 s, N = 14, 3Y male < = 128.4 + 99.6, N = 5). A number of males clearly switched from perching be- havior to patrolling behavior during the course of observation, and it appears that all males spend some time in both forms of mate searching. Males did not switch from patrolling to perching behavior in any ob- servable pattern consistent among individ- uals. Some males patrolled at the same time as other males perched. Male/male interactions. — Interactions be- tween males usually commenced when a patrolling male passed a perched male, but also when two patrolling males met. Only 18 male/male interactions were observed 1n 1982 and 1983 combined, and one in 1981. As has been described for some bees (Eick- wort, 1977), interactions could involve three levels of aggression: chasing, face-to-face hovering and grappling. Chasing and face- to-face hovering were sometimes charac- terized by one male striking another. In the course of 19 interactions, on five occasions a male oriented toward a second male and flew towards it, striking it from behind. One VOLUME 89, NUMBER 4 male chased the other from the patch on five occasions. Swirling flights, with the two males hovering and facing each other, while moving in a circle, occurred four times, twice after an initial strike, and twice when two males observed each other at approximately the same time. At this point, males might depart the patch, with one returning. On three occasions, males contacted each other with their heads, twice while in the swirling flight. In one case, two males grappled and landed on the ground, separating immedi- ately afterwards. One highly aggressive strike resulted in the attacked male being knocked to the ground where it remained immobile, on its back, for 3-4 s. After an interaction, one male departed the patch immediately, while the other male remained for at least a short time, before moving on to another patch, or returning to a perch. The “‘losing”’ male sometimes returned to the patch, with similar results if it was seen by the resident male. The possible function of aggressive inter- actions to displace competing males can best be shown by a description of two males which concurrently occupied the upper (male marked Orange-Yellow (OY)) and lower halves (Blue-Silver (BS)) of one clump on 15 July 1982. At one point, when OY left the clump, BS began flying through the whole clump. OY returned to the upper half, and shortly thereafter, BS encroached upon the upper half of the clump. OY struck BS to the ground, the latter then flying back to the lower half, while OY flew a larger por- tion of the entire clump. Another confron- tation occurred with a series of facing strikes, with BS retreating. This occurred a second time when OY flew further into the lower half, but BS avoided the strike and departed from the clump. Subsequently, OY pa- trolled much of the clump, and BS did not return. The size of males involved in male/male interactions was known for 11 cases. Three times, the outcome was uncertain. In five cases, the larger male “won” (remained in 765 the clump), and three times, the smaller male won. While sample size is too small to allow statistical analysis, there is no indication that escalated confrontations (head butting, swirling flights, grappling) occur more fre- quently between males of roughly equal size. Males involved in swirling flights differed by at least 0.15 mm in head capsule width, and in one case by 0.35 mm. Similarly, fac- ing strikes, as opposed to those where one male struck another from behind, occurred between males differing by the same amount. In one case where a strike knocked one male to the ground, males differed by only 0.05 mm. Resident status of males is often difficult to determine, since two males may simul- taneously patrol the same route for some time before encountering one another. Nevertheless, in 14 interactions where one male was present before a second male, the first male remained on 11 occasions. When the first male present was displaced, the sec- ond male had previously been in the patch several times, in two cases out of three. Male/female interactions.—Interactions with females involved three levels of reac- tion prior to copulation. Upon sighting a female in flight or on a flower, a male usually approached her slowly, hovering behind un- til she settled in a flower. Sometimes a single contact (strike or touch) was all that oc- curred, but other times the male grasped the female from behind and began to antennate the ventral-lateral portion of the female’s head capsule. The antennae of Pseudoma- saris spp. are distinctive structures, unlike those of other vespids. The terminal seg- ments form a club which, in P. vespoides, is large and distinctly concave beneath. The placement of these clubs during copulation, possibly over the female palps, strongly im- plies that they play a role in copulatory be- havior. A series of strokes with the antennae fully extended, moving slightly farther from the female’s head with each subsequent stroke, was followed by the male extending his abdomen toward the female’s genital 766 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON opening. Coupling was accompanied by vi- olent buzzing, at which time the female often released her grip and the pair fell to the grass below, separating shortly thereafter. During 68 observed interactions, males oriented to, but did not move toward, fe- males on seven occasions (10%). Most fre- quently, a male touched or lightly struck a female (44 of 68 —65%). Males grappled with females without copulating on 17 occasions (25%). Males seemed to be able to distin- guish females with which they had recently attempted copulation. Males ignored a fe- male that they had unsuccessfully mounted, even when such a female contacted the male during the course of her foraging. Twenty-one copulations by 13 marked males were observed in the three years. In 1983 one male copulated six times, but most males were observed to copulate only once. Of a total of 71 copulation attempts ob- served in 1983, 58 (82%) were unsuccessful. Combining data from all years, males that copulated (i.e. the size of individual males which copulated, whether once or more than once) and size of copulating males (i.e. the size of the male for each copulation) do not differ significantly from the population size distribution (x?,, goodness-of-fit for males = 0.221, P = 0.66; for copulations x7, = 3.26, P = 0.77) (Fig. 4b). Males appear to interfere with female for- aging. The proportion of time females were in a clump was negatively correlated with both number of male visits, and the pro- portion of time males were in the clump (Kendall’s Tau: S = — 54, P < .005, N = 15 and S = —43, .010 < P< .025, N = 15). The mean duration of female visits was also negatively correlated with these two vari- ables (Kendall’s Tau: S = —37, P < .005, N= 11; S = —23, P= .05, N = 11). Pseudomasaris zonalis Females of P. zonalis foraged at clumps of Phacelia sp. (probably P. hastata Dougl. ex Lehm. Stirp.), which are typically re- stricted to dry, sandy patches of soil, or well- drained hillsides in the gulch and flowered somewhat earlier than Penstemon unilater- alis. Plant distribution varied from year to year, as did the use of plants by wasps. For example, an area with ihe largest concen- tration of plants in 1983 was never observed to be visited by wasps. Individuals of P. zonalis were observed on 25 and 26 June 1981, between 16 and 28 June 1982 and between 23 June and 1 July 1983. Thirteen males were marked and measured (3 in 1981; 3 in 1982; 7 in 1983); in 1982, nine females were marked and measured. Individually marked males and females were observed returning to the same patches of P. hastata for up to 8 days (for individuals observed after marking, ¥ = 6.7 + 2.3 d, N = 6). Females used more than one plant/patch on a single foraging trip, and while in one case, a female inter- acted aggressively with other females, most encounters between females were not ag- gressive. Females collect soil for use in nest construction, and were observed at bare patches of soil, scraping the surface. Males. — Males flew from clump to clump, investigating flowering plants, and they ap- proached females from behind, attempting to copulate. Males flew a circuit of the plants in the patch, and approached different species of insects, especially leafcutting bees present on the flowers. Following a circuit, males perched on the sand and remained there, occasionally orienting to, and some- times investigating, passing insects. Interactions between males were rare, and usually occurred when a patrolling male passed a perched male, the latter flying up to investigate. On one occasion, a male pounced ona second male which was resting on the sand. The former immediately de- parted. Interactions between males and fe- males were also infrequent. One male was observed to strike foraging females on two separate occasions, but no grapples, mounts or copulations were observed. Some males were observed at bare patches of soil, distant from any observed host plants VOLUME 89, NUMBER 4 but similar to sites at which females col- lected soil. Males perched on the sand or on plants at these sites. They made periodic short flights and occasionally departed from the site to return later. At one such site, observed over a 730 s period, a male was present for 54% of the observation period. While present, it took apparently sponta- neous short flights (N = 28), changing perches frequently. In addition, longer flights out of the immediate area occurred, in- volving an irregular, weaving flight follow- ing bombyliid and tachinid flies. No contact with the flies was observed. DISCUSSION The scattered distribution of nests of Pseudomasaris spp., combined with the concentration of females at resources re- quired for nesting and rearing of brood, has resulted in males searching for females at sites other than nests. Some males may search at nests, or remain on their natal nests. Richards (1963) reports a male of P. co- quilletti resting on a nest containing two unemerged females (though this may simply be a freshly-emerged male prior to depar- ture), and sib-mating on natal nests is known from other vespids (Cowan, 1979; Jayakar and Spurway, 1966). Mating with females in or near the natal nest requires little in- vestment in energy searching for nests. Nevertheless, searching at other sites is a major component of mating behavior in some species. If forage plants are not scarce and numbers of females at these locations are large, as is the case when females forage at only a single, or at most a few, species of plants, male searching behavior is expected to evolve to concentrate on these areas. This is the case for P. vespoides and for some males of P. zonalis and emphasizes the con- vergences between these wasps and many bees (Alcock et al., 1978). Some males of P. zonalis and an uniden- tified Pseudomasaris sp. in the gulch may search for females at patches of soil where 767 females scrape material for nest construc- tion. Details of the mating system of P. ves- poides raise a number of questions. The movement of females at Penstemon is prob- ably influenced by the limited nature of pol- len release and nectar secretion, which forces females to visit numerous flowers on each provisioning trip (Torchio, 1974, pers. comm.). This results in a high turnover of individuals within a given clump. Addi- tionally, evidence suggests that not all fe- males are receptive (or acceptable) and males which remain in one large patch will not encounter “new” females as frequently as if they move. Torchio (unpub.) has observed that females of P. edwardsii in the green- house mate more than once, but are recep- tive only during the periods of cell construc- tion. All males may spend some time patrol- ling a number of clumps and some time remaining within a single clump, perched and making short flights within the clump. Why do males engage in both perching and patrolling behavior? This may be an evo- lutionary compromise between the time and energy costs of travel between clumps, as- sociated with patrolling, which would yield the highest encounter rate with newly for- aging, virgin females, compared to less en- ergetically expensive perching, which is coupled with a greater probability of ob- serving and contacting all females in one area. The proportion of time spent patrol- ling or perching will depend on the inter- action of a number of factors, possibly in- cluding air temperature, rate of female visitation at patches, and length of time within a clump of a given quality. Perch sites appear to be located to in- crease observation of new females arriving at clumps. Males normally perch on the pe- riphery of clumps. The use of perch sites beside small clumps and isolated plants may be due to these plants having a greater turn- over of females, since a female cannot ob- tain all necessary provisions for a forage trip 768 from a single plant or a few plants. In these cases perching, interspersed with occasional patrolling, may provide the most energy- efficient means of encountering virgin fe- males. While males do not defend discrete ter- ritories, aggressive interactions occasionally occur and may serve to displace competing males from patrolling the same route. Why do males not defend a clump or several ad- jacent clumps, as is the case for males of some bees, such as Hoplitis anthocopoides (Eickwort, 1977)? Females of this species are also highly specific to forage plants used for provisioning offspring. Many of the dif- ferences between the behavior of these two species may be explained by differences in the number of males present in the area. In H. anthocopoides, particularly during the mid-part of the season, 60 males emerged and numerous males competed for territo- ries around forage plants. In contrast, the largest number of males of P. vespoides present in the area was 10 in 1982. This difference could explain the lack of defence since there is little requirement to spend considerable time excluding other males and the costs of patrolling territorial boundaries rather than a larger number of plants in a given area would not result in a sufficiently large increase in number of matings. Pa- trolling males also rarely encounter other males at any patch of plants, reducing the chance of lost time or physical damage in fights, and are likely to have access to any females encountered. Thus, there appears to be a reduced requirement to defend areas used by females against conspecific males. The contrasting behavior of other ma- sarine wasps indicates the importance of fe- male distribution to male searching behav- ior. In contrast to examples of searching at resources, males of P. maculifrons perch in open areas on peaktops and ridges in central Arizona. Foraging females are scarce and widely scattered, like those of other hilltop- ping insects, and this probably led to the evolution of a landmark-based mating sys- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tem (Alcock, 1985). Some genera of Ma- sarinae (Jugurtia, Celonites, Masaris) nor- mally visit members of at least two families of plants (Cooper, 1952; Gess and Gess, 1980). Under such circumstances, searching for females at flowers would be much less profitable than where specific forage plant species are used. Jugurtia confusa, a South African masarine, nests in large aggrega- tions of ground burrows, and males are re- ported flying in the nesting area and chasing females and other males, though no matings were observed (Gess and Gess, 1980). In three species of Ceramius (capicola, linearis and lichtensteinii) both females and males are present at temporary water sources. Fe- males land on the surface of the water, pre- sumably to collect water for nest construc- tion, and males which patrol these water sources mount females on the water, and the pair fly off. Ceramius also nest in ag- gregations. In contrast, females of Jugurtia were observed at water, while males were not (Gess and Gess, 1980). These differ- ences emphasize the importance of the dis- tribution of females relative to emergence sites and resource availability, in the evo- lution of male searching behavior for mates. Further studies on the mating systems and host choice of species which appear to be polylectic, in contrast to the oligolectic North American Pseudomasaris, would be partic- ularly illuminating. ACKNOWLEDGMENTS I thank K. M. O’Neill and H. E. Evans for help in the field, support and manuscript review. J. Alcock, V. Tepedino and P. Tor- chio also offered comments on the manu- script and unpublished information. C. R. Tracy reviewed an earlier draft of the paper. D. Gwynne kindly provided his observa- tions of P. vespoides at Great Sand Dunes National Monument. J. Hare assisted with the figures. Funding was provided by grants from Sigma Xi and NSF grant 79-26655 to H. E. Evans. This study was done in partial VOLUME 89, NUMBER 4 fulfillment of requirements for a Ph.D. de- gree at Colorado State University. LITERATURE CITED Alcock, J. 1985. Hiulltopping behavior in the wasp Pseudomasaris maculifrons (Fox) (Hymenoptera: Masaridae). J. Kans. Entomol. Soc. 58: 162-166. Alcock, J., G. C. Eickwort, and K. R. Eickwort. 1977. The reproductive behavior of Anthidium macu- losum (Hymenoptera: Megachilidae) and the evo- lutionary significance of multiple copulations by females. Behav. Ecol. Sociobiol. 2: 385-396. Alcock, J., E. M. Barrows, G. Gordh, L. J. Hubbard, L. Kirkendall, D. W. Pyle, T. L. Ponder, and F. G. Zalom. 1978. The ecology and evolution of male reproductive behavior in the bees and wasps. Zool. J. Linn. Soc. 64: 293-326. Carpenter, J. M. 1982. The phylogenetic relation- ships and natural classification of the Vespoidea (Hymenoptera). Syst. Entomol. 7: 1-38. Cooper, K. W. 1952. Records and flower preferences of masarid wasps. II. Polytrophy or oligotrophy in Pseudomasaris (Hymenoptera: Vespidae). Amer. Midl. Nat. 48: 103-110. Cooper, K. W. and J. Bequaert. 1950. Records and flower preferences of masarid wasps (Hymenop- tera: Vespidae). Psyche 57: 137-142. Cowan, D. P. 1979. Sibling matings in a hunting wasp: Adaptive inbreeding? Science 205: 1403- 1405. Dorr, J. E. and L. J. Neff. 1983. Pseudomasaris mar- ginalis nesting in logs in Colorado. Pan-Pac. Ento- mol. 58: 124-128. Eickwort, G. C. 1977. Male territorial behaviour in the mason bee, Hoplitis anthocopoides (Hyme- noptera: Megachilidae). Anim. Behav. 25: 542- 554. Eickwort, G. C. and H. S. Ginsberg. 1980. Foraging and mating behavior in Apoidea. Ann. Rev. Ento- mol. 25: 421-446. Emlen, S. T. and L. W. Oring. 1977. Ecology, sexual 769 selection and the evolution of mating systems. Sci- ence 198: 215-223. Gess, F. W. and S. K. Gess. 1980. Ethological studies of Jugurtia confusa Richards, Ceramius capicola Brauns, C. /inearis Klug and C. lichtensteinii (Klug) (Hymenoptera: Masaridae) in the eastern Cape Province of South Africa. Ann. Cape Prov. Mus. (Nat. Hist.) 13: 64-83. Hicks, C. H. 1929. Pseudomasaris edwardsii Cr., another pollen provisioning wasp, with further notes on P. vespoides. Can. Entomol. 61: 121-125. Jayakar, S. D. and H. Spurway. 1966. Re-use of cells and brother-sister mating in the Indian species Stenodynerus miniatus (Saussure) (Vespidae: Eu- meninae). J. Bombay Nat. Hist. Soc. 63: 368-378. Kevan, P. G. and H. G. Baker. 1984. Insects on flow- ers, pp. 607-631. Jn Huffaker, C. B. and R. L. Rabb, eds., Ecological Entomology. Wiley and Sons, Ltd., New York. Richards, O. W. 1962. A revisional study of the ma- sarid wasps (Hymenoptera: Vespidae), British Museum (Natural History). William Clowes and Sons, Ltd., London. vii + 294 pp. 1963. The species of Pseudomasaris Ash- mead (Hymenoptera: Masaridae). Univ. Calif. Publ. Entomol. 27: 283-310. Tepedino, V. J. 1979. Notes on the flower-visiting habits of Pseudomasaris vespoides (Hymenoptera: Masaridae). Southwest. Nat. 24: 380-381. Thornhill, R. and J. Alcock. 1983. The evolution of insect mating systems. Harvard Univ. Press, Cam- bridge, Mass. 1x + 547 pp. Torchio, P. F. 1970. The ethology of the wasp, Pseu- domasaris edwardsii (Cresson), and a description of its immature forms (Hymenoptera: Vespoidea: Masaridae). L.A. Co. Mus. Contrib. in Sci. 202: 32 pp. 1974. Mechanisms involved in the pollina- tion of Penstemon visited by the masarid wasp, Pseudomasaris vespoides Cresson. Pan-Pac. Ento- mol. 50: 226-234. PROC. ENTOMOL. SOC. WASH. 89(4), 1987, pp. 770-775 A NEW SPECIES OF PARACRIAS (HYMENOPTERA: EULOPHIDAE) PARASITIC ON ANTHONOMUS SPP. (COLEOPTERA: CURCULIONIDAE) JAMES B. WOOLLEY AND MICHAEL E. SCHAUFF (JBW) Department of Entomology, Texas A&M University, College Station, Texas 77843; (MES) Systematic Entomology Laboratory, BBII, Agricultural Research Service, % U.S. National Museum NHB 168, Washington, D.C. 20560. Abstract.—Paracrias anthonomi Woolley and Schauff, new species, is described and illustrated. This species was reared from Anthonomus hunteri in southern Mexico and 1s being investigated as a possible biological control agent for the boll weevil (Anthonomus grandis). The relationship of the new Paracrias to others in the genus is discussed and a revision to the key to the species 1s given. During foreign exploration for parasites of the boll weevil, Anthonomus grandis Boheman (Curculionidae), J. Cate and P. Krauter of Texas A&M University discov- ered a new species of Paracrias (Eulophidae: Entedoninae) attacking Anthonomus hun- teri Burke and Cate. In order to provide a name for this species, we take this oppor- tunity to describe it and discuss its rela- tionship to other Paracrias species. Very little is known of the biology and host associations of the previously de- scribed species of Paracrias. Only two species have been reared and both of these are also associated with curculionids (in the genera Lignyodes and Conotrachelus (Schauff, 1985)). In addition, one of us (MES) has recently seen specimens of another un- described species reared from Anthonomus aeneolus Dietz in the state of Nuevo Leon, Mexico. The concept of Paracrias given by Schauff (1985) is altered considerably by the inclu- sion of the species that we describe here. In spite of the numerous character differences between this species and the others cur- rently placed in this group, naming a new genus for this species would likely render Paracrias paraphyletic. We believe that the character differences in this species are aut- apomorphic (e.g. the elongated median pro- podeum and greatly reduced metanotum) and do not justify a separate genus. Since one of us (MES) is currently analyzing the relationships of this genus and its relatives, we will defer additional comment on ge- neric relationships until that work can be completed. Species of Paracrias can be separated from other genera of Eulophidae by the following characters: scutellum with a single pair of setae and submarginal vein with two setae (subfamily Entedoninae); head and meso- soma well sclerotized and not collapsed when dry; notauli incomplete, usually only present as small indentations at posterior margin of mesoscutum; mesoscutum and scutellum alveolate (Figs. 3, 5); scrobal grooves united below Y-shaped facial groove (Fig. 1); pronotum without transverse ca- rina; metapleural protuberance with antero- lateral carina (Fig. 6); scutellum without median groove; propodeum with a median raised area (Fig. 5); metasoma petiolate; mid VOLUME 89, NUMBER 4 and hind coxal insertions widely separated; stigmal vein sessile, only about as long as wide (Fig. 9). This species will not key through the cou- plets given in Schauff (1985), because it has light yellow fore and mid femora, but a dis- tinctly ridged occiput. The key should be modified as follows: l(a). Female antennae with 5 flagellomeres (without a differentiated club) (Fig. 10); pronotum reduced, barely visible from above (Figs. 3, 4); propodeum medially about equal in length to scutellum, an- terior edge advanced under the scutellum (Fig. 5) and metanotum greatly reduced Beta £9 fie RAE cat Wiehe, or CD P. anthonomi n. sp. - Female antennae with 2 or 3-segmented funicle (club 2 or 3-segmented); pronotum visible from above as a narrow band; propodeum medially much shorter than scutellum, anterior edge sep- arated from scutellum by metanotum and meta- notum clearly visible as a narrow band .... 1 METHODS Terminology for surface sculpturing fol- lows Harris (1979). Morphological termi- nology largely follows that of Graham (1959) or Gibson (1985). Ratios are presented in the form of “‘scape/pedicel”’ (length of scape divided by length of pedicel) followed by the observed ranges. Measurements were taken from the holotype @, allotype 6, and 10 2 paratypes and 5 ¢ paratypes using a Zeiss stereomicroscope and an eyepiece mi- crometer. All measurements were made at the widest or longest point of the structure. Because this species does not possess a dif- ferentiated antennal club in either sex, we use the term flagellomeres to refer to anten- nal segments distal to the annellus, and the abbreviations, FL1, FL2 refer to the first flagellomere, second flagellomere, etc. We use mesosoma and metasoma rather than thorax and gaster. Metasoma refers to the abdomen posterior to the propodeum, in- cluding the petiole, and tergal numbering (e.g. T2) and sternal numbering (e.g. S2) re- fer to metasomal terga and sterna. The metapleural protuberance refers to the cone shaped enlargement of the metapleuron (Fig. 4) that is common in these species and re- 771 lated genera. Accession numbers preceded by a “T” in Types and Other Material Ex- amined were assigned in the Quarantine Laboratory, Biological Control Center, De- partment of Entomology, Texas A&M Uni- versity. Paracrias anthonomi Woolley and Schauff, New SPECIES Diagnosis. — Paracrias anthonomi can be separated from the other described species in this genus by the following: mandibles with 2 teeth of equal size (Fig. 2) (other species with one large tooth and one small tooth); female antennae without club (Fig. 10) (other species have a 2- or 3-segmented club); pronotum reduced dorsad, barely vis- ible from above (Fig. 3) (pronotum visible as a narrow band dorsad in other species); anterior edge of propodeum produced for- ward under scutellum (Fig. 5) (scutellum clearly separated from propodeum by meta- notum in other species) and propodeum (Fig. 6) medially about equal in length to length of scutellum (propodeum much shorter than scutellum in other species). The elongate, neck-like, posterior medial extension of the propodeum of this species is distinctive and immediately distinguishes it from the other described species of Par- acrias. Although there generally is some posterior elongation of the propodeum in other Paracrias spp., the length of the ex- tension in P. anthonomi far exceeds that of the others. However, like some of the other characters in which P. anthonomi differs from other species of this genus (e.g. anten- na without differentiated club), an elongate propodeum is found in other Entedoninae and it commonly occurs in Neotropical species of related genera such as Horismen- US. Female.—Length 2.1-3.2 mm. Color as follows: scape, fore, middle, and hind legs except coxae pale tan; apical spur on hind tibia white in proximal half and black in distal half; frons, vertex, mesoscutum, scu- tellum, axillae, propodeum, and all coxae M2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VOLUME 89, NUMBER 4 Figs. 9-11. black with metallic green, bronze, or occa- sionally blue or purple reflections; propo- deum occasionally with reddish reflections, particularly in medial area; metasoma black with metallic blue reflections. Fore and hind wings hyaline. Head.— Head height 0.70-0.80 the length of head width. Inner margins of eyes sub- parallel, diverging slightly medially (Fig. 1); occipital ridge distinct behind ocelli, con- tinuing about halfway to eye margin; occi- se Paracrias anthonomi, female paratype. 9, Fore and hind wings. 10, Antenna. 11, Hind leg. put reticulate; vertex smooth and shining; facial grooves deep and well-defined, ex- tending laterad to eye margin; frons above facial grooves alveolate, below facial grooves minutely alveolate; area between toruli raised, shining, faintly coriaceous, raised triangular area defined by lateral carinae ex- tending down face to mouth margin; clypeus undifferentiated; malar sulcus present; scape/pedicel 2.27—2.85, one transverse an- nellus present, FL1/pedicel 1.12—1.50, FL2/ Figs. 1-8. SEM’s of Paracrias anthonomi. 1, Head, frontal view. 2, Mandibles. 3, Thorax, dorsal view. 4, Thorax, lateral view (mp = metapleural protuberance). 5, Scutellum and propodeum. 6, Propodeum, postero- lateral view. 7, Metasoma, lateral view. 8, Male scape. 774 pedicel 1.0-1.25, FL3/pedicel 0.80-1.10, FL4/pedicel 0.86-1.17, FLS/pedicel 1.12- 150: Mesosoma.—Prepectus minutely alveo- late, smooth along dorsal and posterior margins; metapleural protuberance smooth (Fig. 4), coriaceous below antero-lateral ca- rina, alveolate to coriaceous posteriad; mesoscutum, scutellum and axillae alveo- late; propodeum (Figs. 5, 6) with antero- medial area only slightly convex and cori- aceous-reticulate with weak longitudinal striations, smooth shining lateral areas set off by sharp carinae, posterior “neck” with alveolate dorsal and minutely alveolate ventral areas divided by ventro-longitudi- nal carinae, sunken spiracular area coria- ceous, callus smooth and bearing one seta. Fore femur and fore tibia subequal in length, fore basitarsus/fore tibia 0.20-0.27, middle femur/middle tibia 0.88—0.97, mid- tibial spur/middle basitarsus 0.75-0.91, hind femur/hind tibia 0.98—1.14, hind tibia (Fig. 11) with apical spur subequal in length to or slightly shorter than the basal two tar- someres; hind coxae elongate (Fig. 11), length/width 1.73-2.14, rounded and smooth in dorsal view, medial surface re- ticulate, lateral surface with medial patch of coriaceous sculpture; length of fore wing (Fig. 9) 2.23-2.47 times its width, marginal vein/ submarginal vein 0.92-1.11, stigmal vein/ marginal vein 0.05-0.10, postmarginal vein about as long as wide, membrane under the proximal portion of marginal vein evenly setose, setae shorter and more dense distal to marginal vein. Metasoma.—Ovate-acuminate, tapering strongly posteriad; petiole (Fig. 7) subcon- ical, smooth, shining; T2 slightly longer than all remaining terga combined, smooth, shining, without setae; T3-T4 or T3-T5 often hidden beneath T2, exposed portions of T3-TS transverse, T6 slightly so, T7 subquadrate, syntergum long, acuminate; combined lengths at midline of exposed portions of terga in following proportions: T2 slightly longer than T3-syntergum, T3- 5 subequal to T6, T3—6 subequal to T7, T3- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 7 subequal to syntergum; S2 (Fig. 7) exposed anterior to T2, other sterna concealed by terga; ovipositor sheaths concealed under syntergum in basal half, exposed parts about ’; length of syntergum in lateral aspect. Male. — Length 2.0-2.4 mm. As described for the female except: anterior margin of scape (Fig. 8) darkened in dorsal 2—*4, with 3 small pits set in an elongate indentation; frons and face with brilliant metallic blue or occasionally green reflections, mesoscu- tum and propodeum with purple or occa- sionally green or gold reflections, pleural re- gions and coxae metallic blue, scape/pedicel 2.50-2.83, lst annellus transverse, 2nd an- nellus reduced to a small semicircular ridge on ventral surface, FL1/pedicel 1.33-1.50, FL2/pedicel 1.17-1.33, FL3/pedicel 1.0- 1.25, FL4/pedicel 1.0-1.25, FL5/pedicel 1.67-2.0; medial surface of hind coxae weakly coriaceous-reticulate; petiole longer and more cylindrical than female, widening apically, about *4 length of medial propo- deum; T2 ovate, strongly truncate apically, with large antero-medial depression, cov- ering remaining terga or occasionally with syntergum exposed (air-dried specimens), with 3-5 scattered, lateral setae; fore wing 2.16—2.28 times longer than wide, marginal vein/submarginal vein 0.87-1.11, stigmal vein/marginal vein 0.05-0.06. Biology and distribution.— Known hosts of P. anthonomi are Anthonomus hunteri and an undescribed Anthonomus sp. (not closely related to hunteri, det. H. R. Burke). Material was reared primarily from buds of Hampea trilobata Standley (Malvaceae). In one case specimens were reared from fruit of Lycianthes synanthera Bitt. (Solanaceae), which were infested with weevil larvae. Adult parasites emerge from 2nd or 3rd in- star weevil larvae. The species is apparently a primary parasite, as no evidence of hy- perparasitism has been noted (J. R. Cate and P. C. Krauter, personal communica- tion). The species is known only from the states of Yucatan, Quintana Roo, Cam- peche, and Chiapas in southern Mexico. Types.— Holotype 2 on point with data: VOLUME 89, NUMBER 4 ‘““Mexico, Quintana Roo, Akumal-Tulum, 18-20 November 1985, ex Anthonomus hunteri, on: Hampea trilobata, colls. P. Krauter & D. Hutchinson, T85708 & T85709” (deposited in USNM). Six 2 para- types on points and one slide-mounted 2 paratype (USNM) with same data. Allotype 6 (USNM), 10 2 paratypes on points, 6 4 paratypes on points and one slide-mounted 6 paratype (USNM) with data: “Mexico, Yucatan, East of Valladolid, 2-X-1984, colls. J. R. Cate and P. C. Krauter, ex. A. hunteri on: H. trilobata, T84073 & T84074.” One 2 and 2 6 paratypes on points with data ‘““Mexico, Yucatan, Merida, 2-X-1984, coll. R. Fisher, ex. 4. Aunteri on: H. trilobata, T84075 & T84076.” 16 2 and 6 6 paratypes on points with data: ““Quintana Roo (S. Tul- um to N. Coba) and Yucatan (Xalau & Yal- coba to Nueva X-Can), 28-X-1-XI-1986, collseb aC. Krautern, & 1)... Cate, ex: A: hunteri, on: H. trilobata, T86084 & T86085.” Paratypes have been deposited in the USNM (10 2, 3 4), CNC (4 9, 2 6), BMNH (1 2, 1 8), and Texas A&M University (TAMU) (19 ¢, 9 8). Other material examined.—AIll from Mexico (deposited in TAMU): CAM- PECHE,.Escarcega,, 9-X-=1976,.colls.. J. R. R. Cate & W. E. Clark, ex: A. hunteri, on: H. trilobata, 5 ?. CHIAPAS, Belisario Do- minguez, Km. 27 & 32, Hwy 200, 10-IX- 1985, ex: Anthonomus sp. (not hunteri), on: Lycianthes synanthera, colls. J. R. Cate & P. C. Krauter, T85056, 1 6 and host re- mains. QUINTANA ROO, Coba, 20-21-X- 1985, ex: A. hunteri, on: H. trilobata, colls. joReCate& PC iKrauter 185069.) 9.11 6; Tulum S. to Felipe Carrillo Puerto, 22- X-1985, ex: A. hunteri, on: H. trilobata, colls. J. R. Cate & P. C. Krauter, T85070, 1 4. YUCATAN, 9-I-1984, coll. R. Fisher, ex: A. hunteri, on: H. trilobata, T84001, 1 9; 15 Km. E. Celestun, 15-X-1981, coll. D. W. Williams, ex: A. hunteri, on: Hampea bud, 1 6; Dzitas, 8-VIII-1984, ex: A. hunteri, on: H. trilobata, coll. J. R. Cate, T84044; 4 Km. SE Tunkas, 12-X-1983, coll. P. Stansley, reared from H. trilobata bud, T83021, 1 2; 775 Valladolid, 14-X-1976, colls. J. R. Cate & W.E. Clark, ex. A. hunteri, on: H. trilobata, 3 2, and 2 6; Valladolid (Km 185/164 -Xe- lon/Tezold), 29-X-1984, ex. A. hunteri, on: H. trilobata, coll. R. Fisher, T84084, 1 4; Yalcoba, 19-X-1985, ex. A. hunteri on: H. trilobata, colls. J. R. Cate & P. C. Krauter, T85068, 2 2, 1 2. Etymology.—The specific name is the genetive form of Anthonomus, the generic name of the host. ACKNOWLEDGMENTS We thank J. Cate and P. Krauter for pro- viding the unpublished biological infor- mation and all of the specimens from their research program on biological control of boll weevil. Horace Burke assisted with taxonomic determinations of host species. Jim Cate, Pete Krauter (Department of Entomology, Texas A&M University), John LaSalle (Department of Entomology, Uni- versity of California, Riverside), Gary Gib- son (Biosystematics Research Center, Ag- riculture Canada, Ottawa), Robert Smiley and Eric Grissell (Systematic Entomology Laboratory, Agricultural Research Service, USDA) reviewed the manuscript and each made valuable suggestions and comments. This paper is Technical Article No. 22703 from the Texas Agricultural Experiment Station. LITERATURE CITED Gibson, G. A. P. 1985. Some pro- and mesothoracic structures important for phylogenetic analysis of Hymenoptera, with a review of terms used for the structures. Can. Entomol. 117: 1395-1443. Graham, M. W. R. deV. 1959. Keys to the British genera and species of Elachertinae, Eulophinae, Entedontinae, and Euderinae (Hym., Chalcidoi- dea). Trans. Soc. Br. Entomol. 13: 169-204. Harris, R.A. 1979. A glossary of surface sculpturing. Occasional Papers in Entomology, no. 28. Cali- fornia State Department of Food and Agriculture, Sacramento, CA. 31 pp. Schauff, M.E. 1985. The New World genus Paracrias Ashmead (Hymenoptera: Eulophidae). Proc. Entomol. Soc. Wash. 87: 98-109. PROC. ENTOMOL. SOC. WASH. 89(4), 1987, pp. 776-789 BIONOMICS OF MICROMUS POSTICUS (WALKER) (NEUROPTERA: HEMEROBIIDAE) WITH DESCRIPTIONS OF THE IMMATURE STAGES GARY L. MILLER AND RONALD D. CAVE Department of Entomology, Auburn University, Alabama 36849. Abstract. —The egg and larva of Micromus posticus (Walker) are redescribed and illus- trated. While setal patterns appear similar for all instars, head capsule dimensions can be used to distinguish between instars. A key to the instars is provided. The life history of M. posticus was studied using Aphis gossypii Glover as prey. Laboratory-reared females survived an average of 61 days and males an average of 74 days. Adult female survivorship was 100% up to 25 days. Mean total egg production was 933.2 eggs/female, with a maximum of 1484. Net reproductive rate was 461.3 females/female/generation, mean generation time was 56.3 days, intrinsic rate of increase was 0.153 females/female/day, doubling time was 4.5 days, and finite rate of increase was |.17/individual/day. Laboratory and field observations in east-central Alabama indicated that females oviposited most often on fibrous material such as cotton fibers and spider mite webbing. Pupation in the field on cotton occurred within bracts on bolls and in folds of leaves. Charitopes mellicornis (Ashmead) and Anacharis melanoneura Ashmead together parasitized 6% of the M. pos- ticus larvae in an unsprayed cotton field. The Hemerobiidae continue to be over- looked despite the widely held belief that they are important predators. Withycombe (1924) ranked them as one of the three most economically important Neuroptera fami- lies in Britain. Balduf (1939) considered the hemerobiids the most important Neurop- tera next to the Chrysopidae in controlling soft-bodied agricultural pests. Although all hemerobiids are predatory, many are lim- ited in habitat range or exhibit a prey spec- ificity that limits their effectiveness (New, 1975). However, some species do frequent low vegetation, and could be considered for use In many agroecosystems. Among the species that New (1975) con- sidered to be of agronomic importance were species of the genus Micromus. In the Nearctic region, Micromus posticus (Walk- er) has been collected in the eastern United States from Massachusetts west to Minne- sota and south to Florida and Texas (Car- penter, 1940) and as far north as Manitoba, Canada (Batulla and Robinson, 1983). The most common species of the genus (Car- penter, 1940), it is associated with a variety of different habitats including mixed forest (Batulla and Robinson, 1983), citrus groves (Muma et al., 1961), pecan orchards (Edel- son, 1982), vineyards (Jubb and Masteller, 1977), cotton fields (Whitcomb and Bell, 1964), potato fields (Mack and Smilowitz, 1980) and alfalfa (Smith, 1923). The pest prey of M. posticus include the cotton aphid, Aphis gossypii Glover, Aphis spiraecola Patch, the green peach aphid, Myzus per- sicae (Sulzer), the black citrus aphid, Tox- optera aurantii (Fonscolombe), and Macro- VOLUME 89, NUMBER 4 siphum sp. (Selhime and Kanavel, 1968), the cabbage aphid, Brevicoryne brassicae (L.), and eggs of the imported cabbage- worm, Pieris rapae (L.) (Cutright, 1923), yellow pecan aphids, Monellia caryella (Fitch) and Monelliopsis nigropunctata (Granovsky) (Edelson, 1982) and Uroleucon ambrosiae (Thomas) (Batulla and Robin- son, 1983). Smith (1922, 1923) and Cutright (1923) provided the first life history accounts of M. posticus. Smith (1922) observed hatching and (1923) briefly described the larval in- stars. Cutright (1923) determined consump- tion rates of the larvae and included notes on preoviposition time, fecundity, and adult longevity. Selhime and Kanavel (1968) reared M. posticus on a variety of aphids at 26.67°C and recorded developmental time. They determined a fecundity of 509 eggs for a single field-collected female. Identification of the Hemerobiidae con- tinues to be based primarily on adult mor- phology (e.g. MacLeod and Stange, 1981), while other stages are frequently over- looked. The potential of many hemerobiids as biological control agents is also over- looked. The purpose of this study was to provide morphological descriptions of the egg and larval stages of /. posticus and to better qualify and quantify various aspects of its life history. MATERIALS AND METHODS Larval and adult M. posticus were ob- tained by rearing eggs and larvae collected from a cotton field near Tallassee, Alabama. Larvae were caged in 50 x 9 mm plastic petri dishes and provided daily with cotton aphids obtained from the field or green- house. Voucher material of adults and lar- vae of M. posticus is deposited in the Au- burn University Entomological Museum. Egg morphology.—Eggs were measured with an ocular micrometer using a Wild ste- reoscope to determine length and width. For examination with the scanning electron mi- croscope, other eggs were removed from 10% Heel formalin preservative, rinsed in distilled water, air dried, mounted on aluminum stubs with double-coated cellophane tape, and coated with gold-palladium alloy in a sputter coater. Egg ultrastructure and ultra- structure measurements of these eggs were determined using an ISI-SS40 scanning electron microscope (SEM) with an accel- erating voltage of 5 kV. Photographs were taken with Polaroid 55 film. Measurements were recorded in millimeters (mm) with means followed by ranges in parentheses. Larval morphology.—Larvae were ex- amined under a Zeiss compound micro- scope, and measurements were made with an ocular micrometer. Larvae were ob- tained from the laboratory colony and were preserved and mounted on slides as de- scribed by Wilkey (1962). Setal numbers are given for each half of the body. Occasion- ally, setal pattern is expressed as a formula. For example, the setal pattern 2-1-2 indi- cates one long and one short seta together, a single long seta, and one long and one short seta together from the midline to the lateral margin. Length of the setae (long or short) is relative to other setae of the same segment being described. Short setae are ap- proximately one-half or less the length of long setae. Measurements were recorded in millimeters (mm) with means followed by ranges in parentheses. Life table.— Larvae reared from eggs were checked daily for molt to the next instar, cocoon formation, and pupation. Upon eclosion, adult females were transferred in- dividually to 60 x 20 mm petri dishes. A single male accompanied each female. Each mating pair was supplied daily with fresh cotton aphids and a ball of cotton saturated with 10% honey water. A portion of loose cotton fibers was supplied as an oviposition site. Females were checked daily for ovi- position and survivorship. Eggs were re- moved and counted and fresh cotton fibers were placed in the cage. Occasional samples of eggs were saved for rearing to determine time to egg hatch. Dead males were replaced 778 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-3. substrate) (97 x). 2, Anterior pole (469 x). 3, Micropyle (939 x). with live ones when necessary. All cages were housed in a rearing room at 28°C and 70- 80% RH with a 14L:10D photoperiod. A survivorship and fertility (= number of offspring produced) table was constructed for adult females by determining for each day, or age interval (x), the proportion of surviving individuals (1,) and the mean number of female progeny per surviving fe- male (m,). The sum of the mean develop- mental times of eggs, larvae, and pupae was used to estimate the age at which adult eclo- sion occurred. Egg survival was recorded as 100% since rearing of eggs for develop- mental time, colony maintenance, and vi- ability checks indicated complete egg hatch. Due to exclusion of cannibalism, disease and natural enemies, survivorship was also 100% for larvae and pupae. Sex ratio was SEM photographs of Micromus posticus eggs. 1, Habitus (arrow indicates side of attachment to assumed to be 50:50. This assumption 1s supported by our determination of the sex ratio of field-collected individuals and the findings of Deyrup and Deyrup (1978), who reported rearing nearly equal numbers of both sexes from field-collected pupae of five species of hemerobuids. Life table statistics were calculated as follows: R,, the net re- productive rate, is the sum of the products 1.«m, calculated for each age interval; G, the generation time, is the sum of the products ].«m,*x divided by R,; r, the innate capacity for increase, was calculated by substituting values of r into the equation: > (1,*«m,*e™) = 1 until equilibrium was reached; and X, the finite rate of increase, was calculated as e'. Field observations. — From July—Septem- ber 1984, field observations of the life his- tory of M. posticus were recorded from an VOLUME 89, NUMBER 4 unsprayed cotton field 3.2 km S of Tallas- see, Elmore Co., Albama, which was heavily infested with cotton aphids. The field was sampled at the end of the growing season (1 September) to determine density of co- coons per plant. Density was estimated by visually sampling 50 plants chosen at ran- dom, 10 in each of the four corners and in the center of the plot. RESULTS Descriptions of immature stages Egg.—(n = 15): Length .91 mm (.88-.96), width .42 mm (.38-.46). Ellipsoidal, slightly flattened on surface attached to substrate (Fig. 1); pale yellow to cream colored after deposition, darkening to pinkish brown prior to hatching. Eyespots and abdominal stria- tions from developing embryo visible through chorion before eclosion. Chorion primarily smooth but exhibits high concen- tration of irregular projections near micro- pylar pole (Fig. 2); projections occasionally separated by fissures to form irregularly shaped polygons (Fig. 3) but are usually in- terconnected by narrow bridges basally. Prominent, buttonlike micropyle, height .005 mm, width .045 mm, present on center of anterior pole. Anterior pole slightly dark- er than remainder of chorial surface, color of micropyle remaining chalk-white throughout embryonic development. Third instar. —(Fig. 4a—d) (n = 10): Length 7.96 mm (6.28—9.50), width at metathorax 1.35 mm (.98-1.75). Head Capsule: Length .59 mm (.55-.61), width .59 mm (.57-.62). Head prognathous, Y-shaped ecdysial line terminating at base of mandibles and run- ning medially to posterior margin of head; frons deltoid with two long setae on fron- toanterior margin and | on lateral margin. Gena with 5 long setae in center, 3 short setae on posterior margin of sclerite, 2 long setae on lateral margin near 3 corneal swell- ings, and 3 long setae and | short seta ven- trally. Antenna three segmented, extending well beyond tips of jaws; scape .10 mm (.09- TUS, .12) long, .09 mm (.08-.11) wide; pedicel .99 mm (.93-1.02) long, .043 mm (.037- .049) wide; flagellum .32 mm (.30-.34) long gradually tapering to a long apical bristle. Pedicel and flagellum with irregular annular sclerotizations. Cervix: Venter with 3 small anterolateral setae, 3 long median setae in vertical row, 3 long median setae in oblique row, 2-4 long lateral setae; dorsum with posterior row of 3—4 long setae, 3-4 sub- median setae in vertical row, 2-3 long an- teromedial setae. Prothorax: Anterior sub- segment: Venter with 3 short setae near anteromedial margin, 2 short median setae, 3 long setae in anterior transverse row, 2 long median setae and | short posterior seta; dorsum with 3 short setae on anterior me- diolateral margin, 3 long setae in anterior transverse row, | long seta posterior to pre- ceding row of setae, 2-3 long median setae, median sclerite with 2 long setae and | short seta, and a transverse row of 3 long setae near posterior margin, 2 long setae laterally. Posterior subsegment: Venter covered with microspines only; dorsum with | short seta on anterior submedian margin, | short sub- median seta, median transverse row of 4-6 long setae, a spiracle and long seta laterally. Mesothorax: Anterior subsegment: Venter with 2 short setae on anterior submedian margin, | short seta near anteromedial mar- gin, circulet of 4-5 long anterolateral setae, 2 long median setae; dorsum with 2 anterior submedial short setae and transverse row of 4—5 long setae, median transverse row of 3 long setae, posterior row with 4 long and 1- 2 short setae, 2-3 long setae near lateral margins. Posterior subsegment: Venter with microspines only; dorsum with 2 short setae anterosubmedially and a transverse row of 6-7 long median setae. Metathorax: Venter with 2 short setae on anterior sublateral margin, | short seta on anterior submedial margin, 5—6 long setae in anterolateral clus- ter, 2 long submedian setae; dorsum with | anterior submedian short seta and trans- verse row of 4 long setae, median transverse row with 3 long setae, 2—3 lateral setae and 780 a short seta on small sclerotized area, pos- terior transverse row of 4 long setae, 3-4 long lateral setae. Abdomen: Segment I: Venter with 2 long median setae; dorsum with | short seta on anterior sublateral mar- gin, 1 short seta near anterior submedian margin, anterior transverse row of 3 long setae, posteriorly with a transverse row of setae with either a single long seta or a long and short seta on small sclerotized areas arranged 2-1-2, from midline to lateral mar- gin, spiracle and 2 long setae laterally. Seg- ment II: Venter with 2 short setae and 2 long setae near anterior margin, posterior row of long and short setae arranged 1-1-2 from midline to lateral margin; dorsum with anterior subdivision with 1-2 short antero- submarginal setae, posterior subdivision with anterior transverse row of 3-4 long setae, posterior transverse row of long and short setae arranged 2-1-2 from midline to lateral margin, spiracle and 2 long setae lat- erally. Segment III: Venter with 2 short se- tae and 2 long setae near anterior margin, posterior transverse row 1-1-2 midline to lateral margin; dorsum with anterior sub- division with 1-2 short anterosubmarginal setae, posterior subdivision with anterior transverse row of 3 long setae and | short seta between inner 2 setae, posterior trans- verse row of long and short setae arranged 2-1-2 from midline to lateral margin, spi- racle and 2 long setae laterally. Segment IV: Venter with 2 setae anteriorly and 3 short setae approximately midway between these setae, posterior row of long and short setae arranged 1-1-2 from midline to lateral mar- gin; dorsum same as abdominal segment III. Segment V: Venter with 2 long setae anteriorly and 2 short setae slightly above innermost seta, posterior transverse row of long and short setae arranged 1-1-2 from midline to lateral margin; dorsum same as abdominal segment III. Segment VI: Venter with 1-2 short setae above innermost an- terior long setae, posterior transverse row of long and short setae arranged 1-1-2 from midline to lateral margin; dorsum same as PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON segment III. Segment VII: Venter same as segment VI; dorsum same as segment III. Segment VIII: Venter with 2 long setae an- teriorly and 2 small pores and a short seta above the innermost long seta, posteriorly with a transverse row of long and short setae arranged 1-2 from midline toward lateral margin; dorsum same as segment III except lateral setae lacking. Segment IX: Venter with | long seta medially, 2 pores and a short seta adjacent to long seta, posterior transverse row of long and short setae ar- ranged 1-2 from midline to lateral margin; dorsum with anterior subsegment with mi- crospines only, posterior subsegment with anterior transverse row of 2 long setae and 1 short seta anteromedially to these setae, posterior row of long and short setae ar- ranged 2-1-2 from midline to lateral mar- gin, 1-2 small pores above innermost setae, spiracle absent, | long anterolateral seta. Segment X: sclerotized with dorsal deltoid plate, lateral plates, and ventral deltoid plate. Venter with 2 long setae and 2 pores on inner anterior margin of lateral plates, 1 long lateral seta on lateral plate, 3 short setae on posterior margin of ventral deltoid plate; dorsum with 3 long setae on outer margin, 1 median pore and 3—4 shorter setae on bas- al margin of deltoid sclerite, 2 long setae on inner margin of lateral plate. Second instar.—(n = 10): Differing from third instar as follows: Length 5.48 mm (4.73-6.03); width at metathorax .86 mm (.59-1.0). Head Capsule: Length .42 mm (.40-.46), width .42 mm (.40-.43). Anten- nal scape .073 mm (.062-.093) long, .060 mm (.056-.093) wide; pedicel .65 mm (.58- .71) long, .038 mm (.037-.043) wide; fla- gellum .29 mm (.27-.32) long. Cervix: Ven- ter with 4-6 setae arranged randomly from midline to lateral margin; dorsum with 4— 5 setae in posterior row, 6-8 setae arranged randomly from midline to lateral margin. Prothorax: Anterior subsegment: Venter with 5 short anterolateral setae; dorsum with 0-3 short setae near anteromedial margin, 10-14 long and short setae arranged ran- Fig.4. Micromus posticus third instar. A, Dorsal habitus. B, Dorsal setal pattern. C, Ninth and 10th abdominal segments (dorsum and venter). D, Head capsule (dorsum and venter). 782 domly from midline to lateral margin (setal pattern may resemble that of third instar). Posterior subsegment: Dorsum with 2 short submedian setae, median transverse row of 4 long setae. Mesothorax: Anterior subseg- ment: Venter with circulet of 4-6 long setae, 1 short seta posterolateral to posteromedial long seta; dorsum with 3 long setae in an- terior transverse row, median transverse row of 2 long setae and | short seta near lateral margin. Posterior subsegment: Dorsum with 1 short seta near anterior margin, 2 short setae posterior to anterior marginal seta and median transverse row of 6 long setae. Metathorax: Dorsum with ill-defined me- dian transverse row of setae but 2 long me- diolateral setae and | long medial seta, pos- terior transverse row with | short and 3 long setae. Abdomen: Segment II: Venter with | short seta near anterior margin; dorsum oc- casionally with | additional long seta in pos- terior transverse row. Segment IV: Venter with | short seta near anterior margin; dor- sal anterior subdivision more noticeable than in previous segments and with only | small median seta. Segment V: Venter with 2 long setae anteriorly and | short seta slightly above innermost long seta; dorsum with anterior subsegment with | short me- dian seta. Segment VI: Venter with anterior oblique row of 2 setae and | short seta slightly above innermost long seta. Segment VII: Venter with anterior oblique row of 2 setae, sometimes with 2 short setae between innermost long seta. Segment VIII: Venter with posterior transverse row arranged 1-2-2 from midline to lateral margin; dorsal pos- terior subsegment without lateral setae, in- tegument of posterior transverse row of se- tae more heavily sclerotized. Segment IX: Venter with 2 long setae in anterior trans- verse row and | short seta slightly above; innermost, posterior transverse row of long and short setae arranged 2-1-2 from midline to lateral margin; posterior submargin dor- sally with 2 long setae in anterior transverse row. First instar.—(n = 10): Differing from PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON third instar as follows: Length 3.04 mm (2.43-3.55), width at metathorax .51 mm (.34-.64). Head Capsule: Length .30 mm (.26-.32), width .33 mm (.30-.37), 4 long setae and | short seta on gena and 4 long setae and | short seta on venter. Antennal scape .055 mm (.037-.062) long, .055 mm (.043-.062) wide; pedicel .46 mm (.43-.48) long, .036 mm (.031-.043); flagellum .29 mm (.23-.34) long. Palpi more robust. Cer- vix: Venter with only 3 small anterolateral setae; dorsum with minute spinules only. Prothorax: Anterior subsegment: Venter without anterior transverse row of long se- tae, occasionally 2 short median setae; dor- sum with 2 long and | short anterior seta, 1 long mediolateral seta, and a posterior transverse row of 3—4 long setae. Posterior subsegment: Dorsum without transverse row of long setae and long lateral setae. Meso- thorax: Anterior subsegment: Venter with 2-3 short setae on anterior submedian mar- gin, 1 short median seta, and a group of 5- 6 short setae laterally; dorsum variable but usually with 2 short setae near anterior sub- margin, anterior transverse row of 3-5 long setae, posterior transverse row of 3-4 long setae and 1-2 long lateral setae. Posterior subsegment: Dorsum with 2 short antero- lateral setae only. Metathorax: Venter vari- able but usually with 2 short setae on an- terior sublateral margin, 1 short seta on anterior submedial margin, anterior trans- verse row of 2-3 long setae, 1-2 long pos- terior setae; dorsum without noticeable me- dian transverse row of setae, posterior transverse row of 3 long setae, lateral setae variable. Abdomen: Segment I: Dorsum with 2 long setae in anterior transverse row. Legs: Tarsi with trumpet-shaped empodia. For quick identification, first instars are distinguished by the presence of tarsal em- podia. Second instars can be separated from third instars by head capsule dimensions. BIOLOGY Table 1 summarizes the developmental times for the immature stages of M. posticus VOLUME 89, NUMBER 4 Table 1. Developmental time (days) of Micromus posticus immature stages at 28°C. Quies Active cent First Second Third Third Egg Instar Instar Instar Instar Pupa Mean SiS) Ps) 1.4 1.9 pS) 3.9 Mode 4 3 l 2 2 4 n 115 29 18 24 61 65 at 28°C. Eggs hatched between 3 and 4 days after deposition. Newly emerged larvae readily fed on small aphids, drank from water droplets, or cannibalized other larvae and eggs if no other food source was avail- able. Duration of the first instar was 1-5 days. The shortest instar developmental time was that of the second, which lasted 1-2 days. Third instars were active and fed for 1-4 days, then formed a loosely spun two-layered cocoon (Fig. 11). A non-feed- ing, quiescent period of 2—5 days was spent within the cocoon prior to pupation. The mean total time larvae were active and feed- ing on cotton aphids was 5.8 days. Duration of the pupal stage was 2-5 days. Mean longevity of adult males exceeded that of adult females by nearly two weeks (Table 2). Student’s t-test revealed these means are significantly different (¢ = 2.149, df = 38, P < 0.05). One female lived 86 days and two lived 85 days. Seven males lived longer than 85 days, the maximum being 100 days. The age-specific survivorship curve for adult female M. posticus in the laboratory is shown in Fig. SA. Initial survivorship was very high; the percentage of females surviv- ing to age 25 days was 100%, and 96% for Table 2. Longevity (days) of adult male and female Micromus posticus at 28°C. Sex X(+SD) Range N Male T3602 16.2) 45-100 17 Female 61:0) 20.2 25-86 23 ' Means are significantly different (t-test; P < 0.05). 783 100 auegsoe eo 8.9 9 9 —- DO OO Of C2 Ono ADULT FEMALE SURVIVORSHIP (%) a =} NO. OF EGGS / FEMALE 0 15 30 45 60 79 90 ADULT AGE (DAYS) Fig. 5. Age-specific survivorship (A) and age-spe- cific egg production (B) of adult female Micromus pos- ticus at 28°C. Shaded area 1n (B) encompasses | stan- dard error above and below the daily mean. eight days thereafter. Following this, there was a steady decline in survival except for a few short plateaus which are probably sample artifacts. Mean preoviposition period was 7 days (range 4-12) and mean oviposition period was 54.3 days (range 21-79). The mean number of eggs produced by the 26 females studied was 933.2 (range 298-1484). The most frequent daily clutch sizes were 14 and 17 eggs per day (Fig. 6). Sixty-seven percent of the clutch sizes were between 10 and 29 eggs. Ten percent were greater than 30 eggs. The greatest number of eggs laid by a female in our study in one 24h period was 58. This female laid 44 eggs the day before laying 58 and oviposited another 43 the day after for a total of 145 eggs in 72 h. The pattern of age-specific egg production per female had two peaks (Fig. 5B). Mean daily egg production rose to an initial peak of 23.4 eggs/female on day 41. A second FREQUENCY wo > J eae % caoS nee z om SERIES BSE am 5 bs Fi NUMBER OF EGGS Fig.6. Observed frequencies of daily egg clutch sizes. higher peak of 28.2 eggs/female occurred on day 76. Due to reduction in adult survi- vorship through time (Fig. 5A), the second peak was based on an average of only eight females, but all of these females displayed this second peak in each of their individual Oviposition patterns. This suggests that, al- though 31% of the initial cohort survived for 76 days, females that survived this long were still capable of producing large num- bers of eggs similar to or greater than youn- ger females. Plotting the product of age-specific sur- vivorship and fertility (1,*m,) against age (x) shows that the cohort as a group is most productive from 18-43 days (Fig. 7). The egg production peak displayed by individ- ual females around age 76 days was damp- ened by the reduced survival of females, but was still evident due to the large number of eggs laid by the remaining females at this age. Using the data obtained in this study, a survivorship and fertility table were con- structed and statistics describing the pop- ulation growth potential of M. posticus were calculated. The net reproductive rate (R,), or the number of times a population mul- tiplies per generation, was 461.3 females/ female/generation. Mean generation time (G) was calculated as 56.3 days. The intrin- sic rate of increase (r) was 0.153 females/ female/day with a resulting doubling time (DT = (In 2)/r) of 4.5 days. The intrinsic PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON -_ xt ADULT AGE-SPECIFIC lym, 0 15 30. 45 60 75 90 ADULT AGE (DAYS) Fig. 7. Age-specific survivorship-fertility (1,+m,) of Micromus posticus. rate of increase was converted into a finite rate of increase of 1.17 per individual per day. FIELD OBSERVATIONS We have observed M. posticus eggs and larvae in flowering crimson clover, Trifoli- um incarnatum L., fields containing large infestations of the pea aphid, Acyrthosiphon pisum (Harris), in April. Adults were col- lected at incandescent lights in residential areas and near wooded areas in April and May. Although an occasional egg was found during June and July, eggs were not seen on most cotton plants until the first week of August. By 26 July, nearly all plants were heavily infested with large colonies of the cotton aphid. By mid-August, nearly every plant harbored numerous eggs and larvae of M. posticus. About one week later, the aphid population began to subside; possibly due to parasitism, fungal disease, and predation by M. posticus and coccinellids. Nearly all aphid colonies on the cotton terminals were eliminated, while a few remained on the lower leaves. At this time, eggs and larvae of M. posticus became less abundant as pu- pae became more abundant. By late August and early September, adults were more common in the field than at any other time and relatively few eggs, but no larvae, were observed. No adults or larvae were ob- VOLUME 89, NUMBER 4 served in the unsprayed cotton field after mid-September. Females usually laid several eggs sepa- rately on the undersides of leaves, in leaf folds, or on leaf petioles. Females appar- ently preferred to oviposit on fibrous sub- strates. On leaf petioles, eggs were often at- tached to plant trichomes (Fig. 8). Eggs on leaves were most often deposited on the webbing of the two-spotted spider mite, 7e- tranychus urticae Koch (Fig. 9), although neither M. posticus larvae nor adults fed on mites in the laboratory. Eggs were also once observed on an abandoned spider web in the field. Pupation sites were mostly within the bracts of large green bolls on the lower por- tions of plants (Fig. 10). Cocoons were found less frequently within tight, twisted folds of senescing green leaves and dead, dried leaves hanging on the plant or lying on the ground. The mean number of cocoons per plant was 5.0 + 0.7 (SE). Half the plants sampled had 4 or more cocoons, with 18 the maximum. There were often 2 or more cocoons on a single boll (Fig. 10). One boll had 12 co- coons within its bracts. Upon adult emergence, the sex of 52 in- dividuals collected either as eggs, larvae, or pupae was determined. The sex ratio (30:22 females : males) was not significantly differ- ent (x? = 1.33; df = 1; P < 0.05) from a 1:1 ratio. On 1 September, two species of Hyme- noptera were found parasitizing M. posticus pupae: Charitopes mellicornis (Ashmead), an ectoparasitic ichneumonid; and Anach- aris melanoneura Ashmead, an endopar- asitic figitid. Of 252 cocoons examined, 15 (6%) were parasitized by these parasitoids. These 15 parasitized hosts were distributed among 11 plants, none of which contained fewer than five M. posticus cocoons. DISCUSSION Our egg measurements were greater than Smith’s (1923) findings of 0.66 mm long and 0.40 mm wide at the largest diameter. 785 Smith (1922, 1923) described the egg cho- rion as unsculptured and without reticu- lations. Our investigation with the SEM revealed a sculptured chorion of intercon- nected projections around the anterior pole. These projections are similiar to those found on the eggs of Chrysoperla carnea (Ste- phens) (Mazzini, 1976) and Hemerobius stigma Stephens (Miller and Lambdin, 1982). The micropyle of M. posticus is not as pronounced as that of H. stigma (Miller and Lambdin, 1982), but the tubercles and fissures described on the micropylar surface of H. stigma appear similar in M. posticus. Smith (1922) described the hatching process and illustrated the egg burster of M. posti- CUS. Although body and head capsule sizes were not considered reliable characters for determining instars of H. stigma (Miller and Lambdin, 1984), we found that these char- acters are more uniform in M. posticus. Ex- cept for some individual variability, chae- totaxy is relatively consistent throughout the three instars and is therefore not a good basis for separating the instars of M. pos- ticus. This agrees with Withycombe’s (1923) belief that chaetotaxy probably does not provide useful taxonomic characters. Our egg development results are com- parable to the 4-day period reported by Smith (1923) and Cutnght (1923), who did not report their temperature conditions. Selhime and Kanavel (1968) obtained egg hatch in 5-6 days at 26.67°C. The active larval developmental time in our study is one day shorter than that ob- served by Selhime and Kanavel (1968), who reared their specimens at a similar temper- ature to ours, but provided their larvae with a mixture of five aphid species, including the cotton aphid. Cutright (1923) obtained an average active larval developmental time of 5.7 days with B. brassicae serving as prey. Cutright (1923) and Selhime and Kanavel (1968) reported a mean pupal period of 4.0 and 4.1 days, respectively, which compares favorably with our results. 786 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VOLUME 89, NUMBER 4 Cutright (1923) observed that female lon- gevity ranged from 6-36 days (n = 13). Moreover, he recorded a preoviposition pe- riod of 3-4 days for 3 females and noted the longest oviposition period was only two weeks. These data are much lower than our observations, but direct comparisonsare dif- ficult to make since Cutright did not reveal the temperature regime of his study and used a different prey species. The mean fecundity of W/. posticus de- termined in our study is 6.5-fold higher than the mean of 144 eggs/female reported by Cutright (1923). Our finding of 933 eggs/ female is the highest average known for any hemerobiid species. Selhime and Kanavel (1968) obtained a mean egg production of 898 eggs/female for Micromus subanticus (Walker). Neuenschwander (1976) ob- served a mean total egg production of 714.8 eggs/female for Hemerobius pacificus Banks; one female produced a maximum of 2554 eggs. Our observed maximum daily clutch size of 58 eggs is equal to that reported by Smith (1923) for M. posticus. Cutright (1923), however, reported a maximum of 68 eggs laid by one female in one day. The attachment of eggs to plant trichomes superficially resembles the stalked eggs of chrysopids and berothids, except in M. pos- ticus the point of attachment is on the side. This behavior could represent an interme- diate evolutionary step between oviposition in crevices or directly on the substrate and the behavior of forming a stalk for the egg. Other hemerobiid species need to be inves- tigated for this intermediate behavior. Ob- servations of females laying eggs most often on cotton lint in the laboratory also indicate that they have a proclivity to oviposit on something fibrous. Similar to our observations of M. posticus 787 pupation sites in cotton leaf folds, Smith (1923) found cocoons in curled apple leaves. Cutright (1923) suspected M. posticus pu- pated under stones or in the soil, but we found no evidence of this. The life table data and 1984 field obser- vations indicate that M. posticus popula- tions can increase rapidly as long as favor- able food and climatic conditions are present. Adults are long-lived and females can produce a large number of eggs which are deposited on a substrate (cotton) that is easily removed. Therefore, this insect may have mass-rearing potential for use in bio- logical control programs against aphids. In- undative or inoculative releases in green- houses would be particularly inviting because climatic conditions are usually fa- vorable and constant in these structures and the confines of the building prevent disper- sion from the target area. The wide range in aphid species fed upon by M. posticus gives it an advantage over host-specific parasitoids. In addition, both larval and adult M. posticus are predaceous, unlike some chrysopids which are not predaceous in the adult stage. Moreover, Edelson (1982) showed that aphid consumption by ™. pos- ticus equalled or exceeded that of four other aphidophagous predators. Cutright (1923) reported that first, second, and third instar M. posticus larvae consumed an average of 10, 11, and 20 cabbage aphids, respectively. Dunn (1954) found that temperature did not affect the consumption rate of Micro- mus variegatus (F.) a predator of Acyrtho- siphon pisum (Harris). Micromus posticus adults and_ larvae, however, prey on other beneficial species. Cutright (1923) found larvae feeding on coccinellid eggs and eggs of their own species. On a few occasions, we observed adult M. ee ss ne —- Figs. 8-10. 8, Egg of Micromus posticus attached to cotton trichome. 9, Micromus posticus egg attached to spider mite webbing on underside of cotton leaf. 10, Micromus posticus cocoons at base of cotton boll within the removed calyx. 788 posticus biting holes into the mummies of parasitized aphids, and then extracting and devouring the parasitoid larvae. Therefore, M. posticus may not be compatible with other aphidophagous biological control agents. The idea of using lacewings for aphid con- trol in greenhouses was suggested by Reau- mur in the 1 700’s (Flint and van den Bosch, 1981). However, little information has been gathered on the manipulation of hemero- biids as biological control agents. Adequate control of the citrus mealybug, Planococcus citri (Risso), was obtained in Palestine by mass releasing Sympherobius amicus Navas (Bodenheimer, 1928). Larvae from released H. pacificus eggs consistently reduced the aphid population throughout the artichoke growing season in California (Neuen- schwander and Hagen, 1980). Hussein (1984) developed a spray technique for the mass release of eggs of Micromus tasmaniae Walker in greenhouses and proposed the use of this species in a pest management pro- gram for control of pests on potatoes in Aus- tralia. Conversely, in the 1930's, eggs of Hemerobius nitidulus F. and H. stigma were released for control of Adelges piceae (Ratzeburg) without apparent success (Gar- land, 1978). We hope the information in our study of M. posticus will stimulate more investiga- tion into this and other hemerobiid species for use in aphid control. ACKNOWLEDGMENTS We thank Wayne E. Clark, Michael J. Gaylor, Gary R. Mullen, Michael L. Wil- liams, Department of Entomology, Auburn University, and Alfred G. Wheeler, Jr., Pennsylvania Department of Agriculture, Harrisburg, for their suggestions and critical reviews of the manuscript. LITERATURE CITED Balduf, W. V. 1939. The Bionomics of Entomopha- gous Insects, Part II. (Reprinted 1974 by E. W. Classey Ltd.) 384 pp. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Batulla, B. A. and A. G. Robinson. 1983. A list of predators of aphids (Homoptera: Aphididae) col- lected in Manitoba, 1980-1981. Proc. Entomol. Soc. Manitoba 39: 25-45. Bodenheimer, F.S. 1928. Contributions towards the knowledge of the citrus insects of Palestine. I. Pre- liminary report on the work of the Palestine breed- ing laboratory at Petah-Tikwa, 1924-1927. Pal- estine Citrol. 1: 5-6, 16 pp. Carpenter, F. M. 1940. A revision of the Nearctic Hemerobiidae, Berothidae, Sisyridae, Polystoe- chotidae, and Dilaridae (Neuroptera). Proc. Am. Acad. Arts Sci. 74: 193-280. Cutright, C. R. 1923. Life history of Micromus pos- ticus Walker. J. Econ. Entomol. 16: 448-456. Deyrup, M. and N. Deyrup. 1978. Pupation of Hem- erobius in Douglas-fir cones. Pan-Pac. Entomol. 54: 143-146. Dunn, J. A. 1954. Micromus variegatus Fabricius (Neuroptera) as a predator of the pea aphid. Proc. R. Entomol. Soc. Lond. Ser. A, 29: 76-81. Edelson, J. V. 1982. Seasonal abundance, distribu- tion and factors affecting population dynamics of yellow pecan aphids (Monellia caryella and Mo- nelliopsis nigropunctata). Ph.D. Dissertation, Au- burn Univ., Auburn. 85 pp. Flint, M. L. and R. van den Bosch. 1981. Introduc- tion to Integrated Pest Management. Plenum Press, New York. 240 pp. Garland, J. A. 1978. Reinterpretation of information on exotic brown lacewings (Neuroptera: Heme- robiidae) used in a biocontrol programme in Can- ada. Manitoba Entomol. 12: 25-29. Hussein, M. Y. 1984. A spray technique for mass release of eggs of Micromus tasmaniae Walker (Neuroptera: Hemerobiidae). Crop Protection 3: 369-378. Jubb, G. L., Jr. and E. C. Masteller. 1977. Survey of arthropods in grape vineyards of Erie County, Pennsylvania: Neuroptera. Environ. Entomol. 6: 419-428. Mack, T. P. and Z. Smilowitz. 1980. The develop- ment of a green peach aphid natural enemy sam- pling procedure. Environ. Entomol. 9: 440-445. MacLeod, E. G. and L. H. Stange. 1981. The brown lacewings of Florida (Neuroptera: Hemerobiidae). Fla. Dept. Agric. Consum. Serv. Div. Plant Ind. Entomol. Circ. 227: 1-4. Mazzini, M. 1976. Fine structure of the insect mi- cropyle—III. Ultrastructure of the egg of Chrysopa carnea Steph. (Neuroptera: Chrysopidae). Int. J. Insect Morph. Embryol. 5: 273-278. Miller, G. L. and P. L. Lambdin. 1982. Hemerobius stigma Stephens (Neuroptera: Hemerobiidae): ex- ternal morphology of the egg. Proc. Entomol. Soc. Wash. 84: 204-207. 1984. Redescriptions of the larval stages of VOLUME 89, NUMBER 4 Hemerobius stigma Stephens (Neuroptera: Hem- erobiidae). Fla. Entomol. 37: 377-382. Muma, M. H., A. G. Selhime, and H. A. Denmark. 1961. Anannotated list of predators and parasites associated with insects and mites of Florida citrus. Fla. Agric. Exp. Stn. Tech. Bull. 634. 39 pp. Neuenschwander, P. 1976. Biology of the adult Hem- erobius pacificus. Environ. Entomol. 5: 96-100. Neuenschwander, P. and K. S. Hagen. 1980. Role of the predator Hemerobius pacificus in a non-insec- ticide treated artichoke field. Environ. Entomol. 9: 492-495. New, T. R. 1975. The biology of Chrysopidae and Hemerobiidae (Neuroptera), with reference to their usage as biocontrol agents: a review. Trans. R. Entomol. Soc. Lond. 127: 115-140. Selhime, A. G. and R. F. Kanavel. 1968. Life cycle and parasitism of Micromus posticus and M. su- banticus in Florida. Ann. Entomol. Soc. Am. 61: 1212-1215. 789 Smith, R.C. 1922. Hatching in three species of Neu- roptera. Ann. Entomol. Soc. Am. 15: 169-176. 1923. The life histories and stages of some hemerobiids and allied species. Ann. Entomol. Soc. Am. 16: 129-151. Whitcomb, W. H. and K. Bell. 1964. Predaceous insects, spiders, and mites of Arkansas cotton fields. Ark. Agric. Exp. Stn. Bull. 690. 84 pp. Wilkey, R. F. 1962. A simplified technique for clear- ing, staining and permanently mounting small ar- thropods. Ann. Entomol. Soc. Am. 55: 606. Withycombe, C. L. 1923. Notes on the biology of some British Neuroptera (Planipennia). Trans. Entomol. Soc. Lond. 70: 501-594. 1924. Notes on the economic value of the Neuroptera with special reference to the Coniop- terygidae. Ann. Appl. Biol. 11: 112-125. PROC. ENTOMOL. SOC. WASH. 89(4), 1987, pp. 790-793 UROPODID MITES PHORETIC ON FLEAS OF GROUND SQUIRRELS IN CALIFORNIA Tom G. SCHWAN AND DAN CorRWIN Department of Health and Human Services, Public Health Service, National Institutes of Health, National Institute of Allergy and Infectious Diseases, Laboratory of Patho- biology, Rocky Mountain Laboratories, Hamilton, Montana 59840. Abstract.—Uropodid mites were found attached to two species of fleas, Diamanus montanus and Opisocrostis oregonensis, associated with ground squirrels in California. This is the first record of such mites attached to fleas in the New World. The phoretic attachment of deutonymphs of uropodid mites to insects is well known (Krantz, 1978). However, observations of such associations involving fleas as pho- rionts are uncommon. Previous records of fleas as phoretic hosts of uropodid mites are restricted to Poland (Oudemans, 1913), Russia (off, 1926; Zasukhin et al., 1936; Popov, 1957), China (Zasukhin et al., 1936), and Sweden (Brinck-Lindroth and Smit, 1971). From September 1980 to June 1983, approximately 15,000 fleas were examined for phoretic mites during surveillance ac- tivities for plague in California. Herein we document the association of uropodid mites with two species of ground squirrel fleas, the first such records of this relationship involving fleas in the New World. METHODS AND MATERIALS Fleas were collected in saline by staff of the Vector Biology and Control Branch, California Department of Health Services, and county health departments. Fleas were identified and those with mites attached were preserved in 70% ethanol. For scanning electron microscopy (SEM), fleas were at- tached to a stub with double stick tape, crit- ical point dried, coated with a 15 nm layer of a gold-palladium alloy, and viewed with a J.E.O.L. 35-CF scanning microscope. RESULTS Of the approximately 15,000 fleas ex- amined, only 2 specimens of Diamanus montanus (Baker) and 6 of Opisocrostis or- egonensis (Good and Prince) had uropodid mites attached. Records are: 23 February 1981, 1 mite ex D. montanus 2 ex Sper- mophilus beecheyi, Santa Barbara Co., CA; 7 May 1982, 1 mite ex D. montanus 2 ex S. beecheyi, Los Angeles Co., CA; 20 April 1982, 5 mites ex 2 O. oregonensis 2? ex 2 S. beldingi, Modoc Co., CA; 13 May 1982, 6 mites ex 4 O. oregonensis (2 6, 2 2) ex S. beldingi, Modoc Co., CA. Six fleas had only 1 mite attached while 2 fleas had 3 and 4 mites attached. Phoresy provides the means for many mites, which have only limited movement of their own, to disperse greater distances by attaching in various ways to insect hosts (Binns, 1982). Uropodid deutonymphs at- tach by means of an anal pedicel (Figs. 1, 2) formed by a glandular secretion that hardens on contact with the air. The broad base of a pedicel is shown in Fig. 3. Nu- merous examples of fleas as phoretic hosts for acaridid hypopodes are known (e.g. Fain and Baker, 1983; Fain and Schwan, 1976, 1984). However, such associations between uropodid mites and fleas are rare. When > a Y mt ee 2) Figs. 1, 2. Uropodid deutonymphs attached to the abdomen of a female Opisocrotis oregonensis. Bar equals 100 um. 792 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 3s they occur, the mites probably represent a very small proportion of the total popula- tion, the majority of which are dispersing from rodent burrows on beetles and flies. ACKNOWLEDGMENTS We thank B. Nelson, R. Doty, E. Lusk, C. Smith, K. Townzen, J. Clover, M. Ma- don, R. Yescott, C. Meyers, and T. Smith, Vector Biology and Control Branch, Cali- fornia Department of Health Services, for their help in collecting fleas throughout Cal- ifornia. C. Welbourn, Acarology Labora- tory, Ohio State University, Columbus, provided some of the literature. We also thank Betty Kester and Gary Hettrick for their assistance in preparing the manu- script. Basal attachment of caudal stalk to the flea’s cuticle. Bar equals 10 um. LITERATURE CITED Binns, E.S. 1982. Phoresy as migration—some func- tional aspects of phoresy in mites. Biol. Rev. 57: 57-620. Brinck-Lindroth, G. and F.G. A.M. Smit. 1971. The Kemner collection of Siphonaptera in the Ento- mological Museum, Lund, with a check-list of the fleas of Sweden. Entomol. Scand. 2: 269-286. Fain, A. and G. T. Baker. 1983. Trichopsyllus ore- gonensis g.n., sp.n. (Acari, Acaridae), a new hyp- opus phoretic on a flea Trichopsylloides oregon- ensis, parasitic on the rodent Aplodontia rufa in the United States. Can. J. Zool. 61: 928-929. Fain, A. and T. G. Schwan. 1976. Paraceroglyphus xenopsylla sp.n., a new hypopus phoretic on Xenopsylla cheopis in Kenya. Rev. Zool. Afr. 90: 634-639. 1984. Three new hypopial nymphs (Acari: Acaridae) phoretic on fleas parasitic on rodents in California, USA. Bull. Ann. Soc. R. Belge Ento- mol. 120: 91-97. VOLUME 89, NUMBER 4 loff, I. G. 1926. Arbeiten zum Studium der Ekto- parasitenfauna in sud-ostlichen Steppen U.S.S.R. Trudy 5 protivochumn. Konfer. Saratov. 200-204 (in Russian). Krantz,G. W. 1978. A Manual of Acarology. Oregon State University Book Stores, Inc., Corvallis. 509 pp. Oudemans, A. C. 1913. Acarologische Aanteeken- ingen XLVII. Entomol. Ber. Amst. No. 71: 372- 376. 193 Popov, P. P. 1957. Onectoparasites of soft Ornithod- oros ticks. Dokl. Akad. Nauk Azerb. S.S.R. 13: 701-703 (in Russian). Zasukhin, D. N., I. G. Ioff, and V. E. Tiflov. 1936. Materials for the study of the parasites and ene- mies of fleas. Vestn. Mikrobiol. Epidem. Parazit. 15: 27-44 (in Russian). PROC. ENTOMOL. SOC. WASH. 89(4), 1987, pp. 794-802 BEHAVIORAL CHARACTERS WITH SYSTEMATIC POTENTIAL IN STONEFLIES (PLECOPTERA) DAvipD D. ZEIGLER AND KENNETH W. STEWART (DDZ) Department of Biology, Southwest Texas State University, San Marcos, Texas 78666; (KWS) Department of Biological Sciences, North Texas State University, Denton, Texas 76203. Abstract. — Based on rapidly accumulating knowledge of drumming, a complex inter- sexual vibrational communication system in stoneflies, we propose and discuss several behavioral characters that may provide input for phenetic and/or cladistic analysis, in- cluding: (1) ability to drum (non-drumming, drumming), (2) signal type (sequential, grouped), (3) phasic structure of sequential signals (monophasy, diphasy), and (4) mode of vibration production (tapping, rubbing, tremulation). The more complex variations of signal structure (diphasy, grouping, rubbing, tremulation) appear to be derived when known species in various taxa of the suborder Arctoperlaria are tested by out-group comparison. Male-call diphasy and signal production by rubbing are derived among genera of Peltoperlidae and Perlidae, as are grouped signals in the family Perlodidae, and trem- ulation in the Chloroperlidae. Other finer aspects of drumming exchange or signal structure such as complex exchange structure, a large number of beats/signal, and non-constant beat intervals likewise appear to be apomorphic in contrast to extant simpler states in each category. The potential for innate behavior to pro- vide lines of evidence for systematic and phylogenetic analysis has long been recog- nized (Mayr, 1958; Ross, 1974). Behavioral systems suitable for such use have been identified and quantified in several insect groups including the Gryllidae (Alexander, 1962; Otte and Alexander, 1983) and the Chrysopidae (Henry, 1984). Recent studies have shown that the drumming behavior of stoneflies (Plecoptera) seems to offer a sim- ilarly useful behavioral system since it is relatively stereotyped at the species level, is most likely a homologous category of be- havior, and has evolved into considerably diverse and complex patterns within the or- der (Stewart and Zeigler, 1984a; Maketon and Stewart, 1984; Zeigler and Stewart, 1985). Zwick (1973) suggested non-drum- ming and drumming as ancestral and de- rived characters, respectively, separating the suborders Antarctoperlaria and Arctoper- laria. Nelson (1984) concluded, after a reas- sessment of 113 mostly morphological characters (but including drumming vs. non- drumming), using a computerized version of the Wagner Parsimony Method, that the available characters were insufficient for re- solving phylogenetic relationships in the Plecoptera. He strongly suggested that the acquisition of additional characters would significantly increase our understanding of Plecoptera phylogeny. We have concentrated over the past ten years on elucidating the nature of drum- ming in North American Arctoperlaria species in part to establish a behavioral data base for phylogenetic applications. A pre- VOLUME 89, NUMBER 4 liminary assignment of polarity was made for certain drumming characters by Stewart and Zeigler (1984b). It is our objective in this paper to further define and better de- lineate pair-forming characters that appear to be potentially useful for phylogenetic analysis, and to give additional examples of preliminary character polarity testing using out-group comparison (Watrous and Wheeler, 1981). THE NATURE OF DRUMMING AND SOME PROBLEMS OF RECENT QUANTITATIVE STUDIES Stonefly drumming 1s an intersexual com- munication behavior for mate identification and location. The principal mode of drum- ming exchange involves substrate-borne vi- brations produced when the insects strike or rub the substratum with the posteroven- tral portion of the abdomen (Rupprecht, 1967; Maketon and Stewart, 1984), al- though some species have been found to tremulate without abdomen/substratum contact (Rupprecht, 1981). The number and spacing of the resultant pulses provide the necessary information for species and sex recognition. Both sexes typically drum, and one or both partners search between drum- ming exchanges until tactile contact is es- tablished. Mating follows immediately. These generalizations are based on the descriptions and quantitative data collected over the last twenty years, before which only a few qualitative descriptions were avail- able. Some of these works have stressed characters such as signal duration (Zeigler and Stewart, 1977; Rupprecht, 1982) and beat frequency in beats/sec (Rupprecht, 1967; Szczytko and Stewart, 1979) which have limited usefulness, even as species level descriptors. Signal duration is a continuous variable which is affected by temperature (Zeigler and Stewart, 1977) and also by the number of beats/signal which can vary even at the individual level. Beat frequency may be useful (at noted temperatures) where the signal consists of a series of evenly spaced 795 beats, but the signals of many species have increasing or decreasing beat intervals or varying phasic structure (see diphasic calls below) which render a beat frequency value essentially useless in describing signal struc- ture. Similar problems are inherent in the “Xx beat interval” which is an average of all beat intervals in a signal. What is needed are drumming characters with few character states which will be useful in categorizing genera and families and for which character polarity can be tentatively determined. Fol- lowing are several characters which look promising in these respects. DRUMMING CHARACTERS THAT MAY PROVE USEFUL IN SYSTEMATIC STUDIES Drumming vs. non-drumming.—This character has been used by Zwick (1973) and Nelson (1984), and indeed it should be very important except for two problems. First is the question of non-drumming. Most often drumming is observed and recorded under laboratory conditions that facilitate uniformity, increase efficiency of research time, and increase recording quality, all of which are difficult to accomplish in the field. If individuals fail to drum under laboratory conditions, we cannot necessarily assume that they are non-drummers in nature. Physical or temporal requirements or cues may limit or prevent drumming behavior under laboratory conditions. Hydroperla crosbyi (Needham and Claassen) was be- lieved to be a non-drummer until more thorough observations proved otherwise (Zeigler and Stewart, 1985). A species must be observed in large numbers and at various ages under “natural conditions” over a 24 h period(s) in order to realistically deter- mine its ability and/or tendency to drum. Consequently, Zwick (1973) and Nelson (1984) actually misused this character since Zwick’s non-drumming Antarctoperlaria and Nelson’s non-drumming families have either been inadequately investigated or not studied at all in terms of drumming capa- bility. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VOLUME 89, NUMBER 4 Secondly, it is possible that some species may have secondarily lost drumming in fa- vor of some other mate-location mecha- nism such as pheromones or specialized searching behavior so that, although origi- nally considered primitive (Stewart and Zeigler, 1984b), in some cases non-drum- ming might be viewed as a derived trait. This will be a difficult question to resolve until a broader spectrum of data is available for the various families in both suborders. Signal phase structure.— Males of many species produce what have been termed di- phasic calls. Originally these calls were con- sidered to have two “phases” or beat group- ings with distinctively different temporal structure (beat spacing) in the two phases (Fig. 1). The category could also be extended to include calls of two beat groupings with similar beat spacing in the two beat group- ings as in H. crosbyi (Fig. 2). It is certainly possible that as the calls of more species are recorded and described, the distinction be- tween these two types will fade. However, both types of diphasic calls must logically be considered as derived in contrast to the simpler monophasic call (Fig. 3) typical of most species (Stewart and Zeigler, 1984b). To date, diphasic signal structure has not been found in the male calls of the Tae- niopterygidae, Capniidae, or Pteronarcyi- dae, nor in the female signals of any family. The origin of diphasic calls can be hy- pothesized from observations on available data. In the genus Soliperla (Peltoperlidae), males of S. quadrispinula (Jewett) and S. thyra (Needham & Smith) have simple monophasic calls while S. fenderi (Jewett) males have a derived diphasic call with dis- tinctively different beat spacing in the two — 797 phases. It could be hypothesized, judging from the similarity of S. thyra’s call and the second phase of S. fenderi’s call (Figs. 1, 3), that the fenderi call has been produced by adding a few widely spaced beats (apo- morphic phase) before the original mono- phasic call sequence (plesiomorphic phase). Stark (1983) found that S. guadrispinula and S. thyra share many key morphological characters which distinguish their common recent lineage from that of S. fenderi’s. Another possible origin of diphasic calls could be from simply repeating the original monophasic call sequence twice in rapid succession. The diphasic calls of Kathro- perla perdita (Banks) are suggestive of this hypothesis (Fig. 4) as are those of Jsoperla gortzil Illies (Rupprecht, 1969). Finally, a diphasic call could originate by the dropping of one or more beats from the middle of a long monophasic call. Some calls from “fatigued” males of Taeniopteryx burksi Ricker & Ross (Fig. 5) showed miss- ing beats resulting in a call of two distinct beat groupings (personal observation), and the typical call of H. crosbyi (Fig. 1) is suggestive of a monophasic call missing two internal beats. However, if interspecific call differences are in part the result of sexual selection, as is currently suggested (Thorn- hill and Alcock, 1983; West-Eberhard, 1983), this latter possibility 1s perhaps least likely since such calls would initially be re- ceived as “deficient” aberrations of the typ- ical call. Grouped calls.—Male calls of three or more beat groupings have so far been ob- served only in the family Perlodidae, and they appear to be derived (Fig. 9) when tested by out-group comparison (Watrous and Figs. 1-8. Oscillographs of assorted plecopteran drumming signals. 1, Diphasic male call of Soliperla fenderi, 22°C. 2, Diphasic male call of Hydroperla crosbyi, 24°C. 3, Monophasic male call of Soliperla thyra, 24°C. 4, Diphasic male call of Kathroperla perdita, 23°C. 5, A male call of Taeniopteryx burksi showing a deleted 4th beat (after 25 min of male-female drumming contact). 6, Grouped male call of Isogenoides zionensis, 22°C. 7, Rubbing male call of Doroneuria baumanni, 22°C. 8, Overlapping male-female drumming exchange of Prer- onarcys princeps, 23°C. 798 < ue Q Ne Vy, cd (tp L — NG Q Tt a &y © Q Qa ONS ae) Vln ae) a we TOG PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON a i) ~w Zz yy TZ Sa eae eg RS) Q Ly ©) yy ~N g (e) Q — Na Q a Ly a SS oS = Zz Oo SE LG ONO Omer 1 1 1 Ue" a? A oe ---PERLODIDAE TIG 1 = Male calls of one or two beat groupings 1’ = Male calls of three or more beat groupings Fig. 9. Out-group comparison for male calls of three or more beat groupings (TOG = taxonomic out-group, TIG = taxonomic 1n-group). Wheeler, 1981). These calls often appear to be closely spaced repetitions of an ancestral monophasic call. Evidence to support this view is best illustrated by [sogenoides zio- nensis Hanson in which the first few calls in a male-female exchange are grouped calls of three or four groupings (Fig. 6), but as exchanges continue, the male shifts to a sim- pler monophasic call which is very similar to a single beat grouping within the initial grouped calls (Stewart and Zeigler, 1984a). Beating, rubbing, and tremulation.— There are at least three methods of signal production in the Plecoptera. Most com- monly a pulse is produced as the abdomen strikes (beats) the substratum. Maketon and Stewart (1984) found that several perlids rub or scrape the substratum with their ab- domen producing a distinctly different type of vibrational signal (Fig. 7). This manner of signal production seems to be a derived character state in the family Perlidae VOLUME 89, NUMBER 4 (Maketon and Stewart, 1984), although rub- bing has apparently arisen separately in some of the peltoperlids as well (Maketon and Stewart, unpublished). Rupprecht (1981) has shown that the males of some species of Siphonoperla (family Chloroperlidae) pro- duce their signals by repeatedly jerking their bodies without abdomen/substratum con- tact. This method, termed tremulation (Busnel et al., 1956; Morris, 1980), also ap- pears to be derived because other chloro- perlid species produce the more typical beating type signals (Stewart and Zeigler, 1984a). Exchange structure.— Exchange structure refers to the number of male and female signals in a typical drumming exchange and their arrangement. The simplest observed exchanges consist of a male call followed by a female answer. More complex is a three- way (or three-part since only two individ- uals are involved) exchange where a male response signal is added to the basic call- answer sequence. The presence of a re- sponse signal is most likely a derived character even though it is found in most families where drumming is known. Per- haps this character originated before many of the present families had diverged. We believe the male response signal must func- tion to acknowledge receipt of the female’s answer, thus establishing the certainty of communication contact for both partners and affecting proper behavior patterns (often including a cessation of locomotor activity in the female) for the duration of drumming contact. Rarer four- and five-part exchanges have also been observed (6-92-3-? and 3-9-é- 9-6), but these often occur only in the first few exchanges of a longer exchange se- quence (Zeigler and Stewart, 1985). Still, their occurrence could be important if cer- tain families or genera uniformly lack them, as the Pteronarcyidae apparently do (Stew- art et al., 1982a). The non-synchronous overlap of male and female signals within an exchange has been observed in some families (Fig. 8) and might also prove useful 799 ifit is found to be common or totally lacking in certain groups. Beat number/signal.—As mentioned above, this character can be highly variable both within and between species. However, there are instances where creating categories of number of beats/signal may be useful. For instance, male pteronarcyids produce calls ranging between 4-8 beats (Stewart et al., 1982a; Zeigler and Stewart, 1985), whereas males of the family Leuctridae pro- duce calls of more than 20 beats (Zeigler and Stewart, 1977; Rupprecht, 1977). A large number of beats/signal has been sug- gested as a derived character state (Stewart and Zeigler, 1984b). In the perlids, this seems to be the case for the female answers of Phasganophora capitata (Pictet) (Maketon and Stewart, 1984) and Paragnetina kan- sensis (Banks) (Stewart et al., 1982b) which exceed 100 beats per signal (Fig. 10). To date, answers of other species rarely exceed 20 beats. Beat intervals.—In the family Pteronar- cyidae, at 22—25°C, Pteronarcella species produce signals with beat intervals of less than 130 msec, whereas Preronarcys species produce signals with intervals greater than 200 msec (Stewart et al., 1982; Zeigler and Stewart, 1985). In most cases, the actual beat interval means (in msec) are probably too variable to be useful above the family level, and for diphasic calls, two such values must be reported (one/phase) making com- parisons more difficult. However, there are other possible characters to be derived from beat interval measurements. As pointed out earlier, beat intervals can be constant throughout the signal, or they can increase (lengthen) or decrease (shorten) as the signal progresses. To date, the male call and fe- male answer of all taeniopterygid species have increasing beat intervals, while pel- toperlid signals have decreasing or constant beat intervals. Constant beat intervals, being the simpler situation, can be tentatively considered as the plesiomorphic condition. Female calling.—In most stonefly species 800 TOG PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ---- PERLIDAE TIG 1 = Female answers of less than 100 beats 1’ = Female answers of more than 100 beats Fig. 10. Out-group comparison for female answers of more than 100 beats (TOG = taxonomic out-group, TIG = taxonomic in-group). studied to date, the male initiates drum- ming communication with a calling signal. However, in some species of Soliperla (Stewart and Zeigler, 1984a) and Doroneu- ria (Maketon and Stewart, 1984) females will produce signals (calls?) in the absence of male drumming activity (though males do call in these species). Since this trait is apparently rare, it may prove to be apo- morphic and to have some systematic val- ue, Or 1t may prove to be convergent in sev- eral genera due to some common set of environmental conditions. For example, calling in females would seem to be bene- ficial to a species which typically exists at low population densities, since calling by both sexes would effectively increase the chances for male-female contact where VOLUME 89, NUMBER 4 drumming is a major mechanism for mate location. CONCLUSIONS Currently, drumming has been described for approximately 5% of the world Plecop- tera fauna, currently estimated at around 1600 species (Arnett, 1983), and for some of these, only male signals have been ob- tained. Consequently, most of the ideas pre- sented herein must be considered tentative. As the drumming of more species is eluci- dated, the systematic utility of these char- acters will be reinforced, weakened, or elim- inated accordingly. Also, questions concerning the evolutionary rate in behav- ioral vs. morphological trends, as well as possible correlations between plesiomor- phic/apomorphic character states in both areas, may be resolved. Interestingly, the pteronarcyids, which are the most completely studied family (drum- ming descriptions for over 50% of the species) show several primitive drumming traits: monophasic calls, signal production exclusively by beating, few beats/signal, and lack of four- and five-part exchanges. This family exhibits many primitive morpholog- ical characteristics and is considered the most primitive extant group within its sub- order (Nelson and Hanson, 1971). In con- trast, the perlids, which are a diverse and relatively “derived group,” exhibit many of the proposed apomorphic drumming char- acters discussed herein. LITERATURE CITED Alexander, R. D. 1962. The role of behavioral study in cricket classification. Syst. Zool. 11: 53-72. Arnett, R. H. 1983. Status of the taxonomy of the insects of America north of Mexico, a preliminary report prepared for the subcommittee for the In- sect Fauna of North America Project. Committee on Systematics Resources of the Entomological Society of America. R. H. Arnett, privately pub- lished. 49 pp. Busnel, R. G., B. Dumortier, and M. C. Busnel. 1956. Recherches sur le comportement acoustique des ephippigéres (Orthoptéres, Tettigoniidae). Bull. Biol. Fr. Belg. 3: 219-286. 801 Henry, C.S. 1984. The sexual behavior of green lace- wings, pp. 101-110. 7m Canard, M., Y. Semeria, and T. R. New, eds., Biology of Chrysopidae. W. Junk, The Hague. Maketon, M. and K. W. Stewart. 1984. Further stud- ies of the drumming behavior of North American Perlidae (Plecoptera). Ann. Entomol. Soc. Am. 77: 770-778. Mayr, E. 1958. Behavior and systematics, pp. 341- 362. In Roe, A., and G. G. Simpson, eds., Behav- ior and Evolution. Yale Univ. Press, New Haven. Morris, G. K. 1980. Calling display and mating be- haviour of Copiphora rhinoceros Pictet (Orthop- tera: Tettigoniidae). Anim. Behav. 28: 42-51. Nelson, C. H. 1984. Numerical cladistic analysis of phylogenetic relationships in Plecoptera. Ann. Entomol. Soc. Am. 77: 466-473. Nelson, C. H. and J. F. Hanson. 1971. Contribution to the anatomy and phylogeny of the family Ptero- narcidae (Plecoptera). Trans. Am. Entomol. Soc. 97: 123-200. Otte, D. and R. D. Alexander. 1983. The Australian crickets (Orthoptera: Gryllidae). Acad. Nat. Sci. Phila. Mon. 22: 1-477. Ross, H. H. 1974. Biological Systematics. Addison- Wesley Publishing Co., Inc., Reading, Mass. 345 pp. Rupprecht, R. 1967. Das Trommeln der Plecopteren. Z. Vergl. Physiol. 59: 38-71. . 1969. Zur Artspezificitat der Trommelsignale der Plecopteren (Insecta). Oikos 20: 26-33. 1977. Nachweis von Trommelsignalen bei einem europaischen Vertreter der Steinfliegen- Familie Leuctridae (Plecoptera). Entomol. Ger. 3: 333-336. . 1981. Anew system of communication with- in Plecoptera and a signal with a new significance. Biol. Inland Waters 2: 30-35. 1982. Drumming signals of Danish Plecop- tera. Aquat. Insects 4: 93-103. Stark, B. P. 1983. A review of the genus Soliperla (Plecoptera: Peltoperlidae). Great Basin Nat. 43: 30-44. Stewart, K. W.,S. W. Szczytko, and B. P. Stark. 1982a. Drumming behavior of four species of North American Pteronarcyidae (Plecoptera): Dialects in Colorado and Alaska Pteronarcella badia. Ann. Entomol. Soc. Am. 75: 530-533. Stewart, K. W., S. W. Sczcytko, B. P. Stark, and D. D. Zeigler. 1982b. Drumming behavior of six North American Perlidae (Plecoptera) species. Ann. Entomol. Soc. Am. 75: 549-554. Stewart, K. W. and D. D. Zeigler. 1984a. Drumming behavior of twelve North American stonefly (Ple- coptera) species: First descriptions in Peltoperli- dae, Taeniopterygidae and Chloroperlidae. Aquat. Insects 6: 49-61. 802 1984b. The use of larval morphology and drumming in Plecoptera systematics, and further studies of drumming behavior. Ann. Limnol. 20: 105-114. Szczytko, S. W. and K. W. Stewart. 1979. Drumming behavior of four western Nearctic /soperla (Ple- coptera) species. Ann. Entomol. Soc. Am. 72: 781- 786. Thornhill, R. and J. Alcock. 1983. The Evolution of Insect Mating Systems. Harvard Univ. Press, Cambridge, Mass. 576 pp. Watrous, L. E. and O. D. Wheeler. 1981. The out- group comparison method of character analysis. Syst. Zool. 30: 1-11. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON West-Eberhard, M. J. 1983. Sexual selection, social competition, and speciation. Quart. Rev. Biol. 58: 155-183. Zeigler, D. D. and K. W. Stewart. 1977. Drumming behavior of eleven Nearctic stonefly (Plecoptera) species. Ann. Entomol. Soc. Am. 70: 495-505. 1985. Drumming behavior of five stonefly (Plecoptera) species from central and western North America. Ann. Entomol. Soc. Am. 78: 717-722. Zwick, P. 1973. Insect: Plecoptera. Phylogenetisches System und Katalog. Das Tierreich, 94. Walter de Gruyter and Co., Berlin. 465 pp. PROC. ENTOMOL. SOC. WASH. 89(4), 1987, pp. 803-810 RESOURCE PARTITIONING IN CHLOROPIDAE (DIPTERA) OF A FRESHWATER MARSH J. L. Topp AND B. A. Foote Department of Biological Sciences, Kent State University, Kent, Ohio 44242; JLT current address: Department of Entomology, Ohio Agricultural Research and Develop- ment Center, The Ohio State University, Wooster, Ohio 44691. Abstract. —Information is presented on the spatial distribution, temporal occurrence, and larval trophic habits of Chloropidae found in a northeastern Ohio freshwater marsh. Differential utilization of marsh vegetation was apparent, with the majority of species occurring in a stand of the spike-rush, Eleocharis smallii Britt. Within this vegetation type, chloropid species exhibited trophic partitioning by utilizing different larval foods. The Chloropidae consist of 53 genera and some 287 species in the Nearctic Region (Sabrosky, 1987). Many chloropids are phy- tophagous, feeding on crop species (Cun- liffe, 1925; Empson, 1957) as well as eco- nomically unimportant monocots (Wendt, 1968; Wearsch, 1968; Rogers, 1970), whereas other species are saprophagous. Valley et al. (1969) presented information concerning the larval foods of 26 Nearctic species, reporting that several species were associated with freshwater monocots such as sedges, rushes, and grasses. A recent study of a freshwater marsh in northeastern Ohio (Todd, 1985) also indicates that chloropids are well represented in stands of wetland monocots, with a particularly large array of species occurring in a plant association dominated by spike-rush, Eleocharis smal- lii Britt. This paper presents information concerning the spatial distribution, tem- poral occurrence, and larval food utilization of chloropid species occurring in a fresh- water marsh. We then propose possible re- source partitioning mechanisms affecting the Chloropidae encountered in a stand of E. smallii. MATERIALS AND METHODS The freshwater marsh utilized in this study was located near Kent, Ohio, 0.8 km east of the Kent State University campus. The marsh measured approximately 4978 m? and consisted of a spatial mosaic of 8 vege- tation types, most of which existed in nearly monoculture condition (Todd and Foote, 1987): (1) Phalaris arundinacea L. (reed ca- nary grass), (2) Typha-Phalaris (equally di- vided between cattail and grass), (3) Carex lacustris Willd. (sedge), (4) Carex stricta Lam. (sedge), (5) Eleocharis smallii Britt. (spike-rush), (6) Typha latifolia L. (broad- leaved cattail), (7) Nuphar lutea Sibth. and Smith (yellow water lily), and (8) Sparga- nium eurycarpum Engelm. (bur-reed). Eleocharis smallii formed a single stand of 70 m? in the marsh, and during spring and early summer consisted of scattered patches growing in approximately 12-15 cm of standing water. Eleocharis did not be- come a noticeable component of the marsh vegetation until mid-July, at which time the stems grew above the water surface. Once established, the patches measured 40—45 cm 804 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Chloropidae collected in marsh vegetation. Vegetation types are listed in decreasing order of chlo- ropid occurrence. PA = Phalaris arundinacea; TP = Typha-Phalaris; CL = Carex lacustris; CS = Carex stricta; ES = Eleocharis smallit,; NL = Nuphar lutea, TL = Typha latifolia; SE = Sparganium eurycarpum. SY Species Vegetation Types —_—————————— Apallates neocoxendix (Sabrosky) Chlorops certimus Adams Chlorops obscuricornis Loew Conioscinella sp. Diplotoxa inclinata Becker Diplotoxa nigripes (Coquillett) Diplotoxa sp. nr. nigripes Diplotoxa sp. Elachiptera nigriceps (Loew) Elachiptera sp. Epichlorops exilis (Coquillett) Eribolus longulus (Loew) Eribolus nanus (Zetterstedt) Liohippelates bishoppi (Sabrosky) Oscinella frit L. Pseudopachychaeta approximatonervis (Zetterstedt) Rhopalopterum atriceps (Loew) Rhopalopterum carbonarium (Loew) Thaumatomyia bistriata (Walker) Thaumatomyia glabra (Meigen) Thaumatomyia grata (Loew) ESMRAS'CS ES; CL, SE CL CEYSENES, BA ESS CIE NL PACE SE ES; SES TE,CS PA CS, CLA PA; NED TLy TPAESZSE PA PA; CL;iGS SE in height and were interspersed among areas of open water. As summer progressed, the water level dropped, and the Eleocharis stand existed on a highly organic mud sub- strate. A few individuals of Sagittaria la- tifolia Willd. (broad-leaved arrowhead) in- vaded Eleocharis in early August, and a noticeable algal flora usually was present on the mud substrate. Decaying plant material accumulated throughout the summer. For collecting purposes, a 5.0 x 10.0 m quadrat was established within each vege- tation type. Sampling for adult chloropids was conducted between June and October, 1984, using a simplified version of the pan trap described by Grigarick (1959). A 38.5 x 14.0 x 4.5 cm yellow plastic container was filled with a detergent-water solution to a depth of approximately 2.0 cm. Disney et al. (1982) found yellow to be most effective in collecting Diptera from grass-like areas. One pan was placed in the center of each quadrat biweekly and left in place for 24 h. Trapped insects were removed and stored in 70% ethyl alcohol. Additional specimens were obtained by sweeping vegetation with a standard 30.5 cm aerial insect net. Sweep- ing was conducted biweekly before place- ment of the pan traps and consisted of 10 back and forth sweeps along the 10 m center line of a quadrat. All sampling was done in the early afternoon hours. Material obtained from the sweeps and pan traps was combined to compile species lists for each vegetation type. Relative abundance and percent presence values were calculated for all species collected from each vegetation type. Relative abundance values were further categorized according to a method developed by Scheiring and Foote (1973) and are defined as follows: 1—2% rare (r); 3-8% occasional (occ); 9-14% common (c); 15-25% abundant (a); and 26-—100% very abundant (va). To quantify utilization of spatial and tem- poral resources, niche breadth and overlap VOLUME 89, NUMBER 4 805 16 14 12 ” 2 So 10 © Q. 2 8 je re) é 6 Zz. 4 2 ) PAS [PoC =4CSVVES “FTE ANI Se Vegetation Type Fig. 1. Number of species of Chloropidae collected in 8 vegetation types of the freshwater marsh. PA = P. arundinacea, TP = Typha-Phalaris, CL = C. lacustris, CS = C. stricta, ES = E. smallii, TL = T. latifolia, NL = N. lutea, SE = S. eurycarpum. values (Colwell and Futuyma, 1971) were calculated for all chloropids determined to species. Breadth values ranged from 0.00 to 1.00. The minimum value (0.00) indicates extreme specialization, and the maximum value (1.00) indicates generalization. Over- lap values also ranged from 0.00 to 1.00. The minimum value (0.00) was obtained when two species shared no resource in common, whereas the maximum value (1.00) was obtained when the proportional distribution of two species among a resource was the same. Information on larval feeding habits was taken from the literature. RESULTS A total of 225 chloropids of 21 species and 12 genera was collected from the 8 quadrats established within the marsh (Ta- ble 1). Chloropid species and individuals were most commonly collected in E. smal- lii, C. lacustris, and P. arundinacea (Figs. 1, 2). Temporal data indicate that species rich- ness was highest in the marsh between mid- July and mid-August (Fig. 3). A secondary peak was observed in early June, followed by a period in which no species were col- lected again until early July. From late Au- gust until the completion of the study, species richness remained low (X = 3) but fairly constant. The majority of chloropid individuals (>70%) was also collected be- tween mid-July and mid-August (Fig. 4). Field collections indicate that the stand of E. smallii contained 44% of all chloropid species and 23% of all chloropid individuals taken during the study (Figs. 1, 2). Ninety- nine individuals were collected, comprising 7 genera and 11 species (Table 2). Relative abundance values indicate that 4 species can be considered very abundant (va), abundant (a) or common (c) in Eleocharis: Apallates neocoxendix (Sabrosky) (c), Chlorops ob- scuricornis Loew (va), Diplotoxa inclinata 806 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Relative Abundance % NO on PA TP 52 3 30 CL CS 11 99 6 10 ES G7 TL AGNL. [Se 14 Vegetation Type Fig. 2. Relative abundance of species of chloropidae collected in 8 vegetation types of the freshwater marsh. PA = P. arundinacea, TP = Typha-Phalaris, CL = C. lacustris, CS = C. stricta, ES = E. smallii, TL = T. latifolia, NL = N. lutea, SE = S. eurycarpum. Actual number of specimens collected in each vegetation type is given below the x-axis. Becker (c), and Diplotoxa nigripes (Coquil- lett) (a). These and other species are dis- cussed below. Apallates neocoxendix was collected only between early and mid-August (Table 2), and its percent presence (Table 2) and tem- poral breadth values (Table 3) were low. This species was most similar in temporal occurrence to C. obscuricornis (Table 3). Little information has been published con- cerning the trophic habits of A. neocoxen- dix, although larvae probably feed on de- caying plant material (Starks and Thurston, 1962; Wheeler, 1973). Chlorops obscuricornis was collected from early August to late September, with peak emergence in mid-August (Table 2). This species exhibited a moderately high tem- poral breadth value, and was most similar in temporal occurrence to A. neocoxendix (Table 3). Larvae of this species feed as pri- mary invaders of Eleocharis stems (Valley et al., 1969). Diplotoxa inclinata was collected only be- tween early and mid-August (Table 2), and was most similar in temporal occurrence to Elachiptera nigriceps (Table 3). Larvae at- tack rhizomes of Eleocharis (Wearsch, 1968). Females deposit eggs on stems just above the water surface, and newly hatched larvae penetrate the stem and do some feed- ing before moving to the rhizomes to com- plete development. Diplotoxa nigripes was collected between early August and mid-October (Table 2), with the majority of individuals occurring in late September. Wearsch (1968) collected numerous adults of this species from mid- June through early October in stands of E. smallii. Therefore, the temporal breadth VOLUME 89, NUMBER 4 807 16 14 12 10 8 No. of species oOo NM FS OD 6—1 6-6 6-21 7-9 7-19 8-6 8-17 8-31 9-13 9-30 10-11 10—27 Time Fig. 3. Number of species of Chloropidae occurring at 12 time intervals in the freshwater marsh. 90 45 40 35 50 25 Relative Abundance % 6-1 6-6 6-21 7-9 7-19 8-6 8-17 8-31 9-15 9-30 10-11 10-27 17 0 0 8 48 64 49 6 5 17 5 6 Time Fig. 4. Relative abundance of species of Chloropidae occurring at 12 time intervals in the freshwater marsh. Actual number of specimens collected on each date is given below the x-axis. 808 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 2. Abundance data, flight periods and larval foods of Chloropidae collected in E. smallii. Number Col- Relative Percent Species lected Abundance Presence Flight Period Larval Food A, neocoxendix 10 10.10 16.67 Aug. Decaying vegetation C. obscuricornis 44 44.44 33.33 Aug.—Sept. Eleocharis stems D. inclinata 12 AN? 8.33 Aug. Eleocharis rhizomes D. nigripes IS SIMS) 33.33 Aug.—Oct. Eleocharis stems D. sp. nr. nigripes 2 2.02 8.33 Aug. Unknown Diplotoxa sp. 2 2.02 16.67 Aug. Unknown E. nigriceps 4 4.04 25.00 Aug. Decaying vegetation E. longulus 3 3.03 8.33 Oct. Damaged monocot stems E. nanus 2 2.02 8.33 Oct. Damaged monocot stems P. approximatonervis 3 3.03 8.33 Aug. Eleocharis seeds R. carbonarium 2 2:02 16.67 Aug. Damaged monocot stems value attained for D. nigripes in the present study (0.47) (Table 3) probably would have been considerably higher had this vegeta- tion existed in the marsh in June. Diplotoxa nigripes was most similar in temporal oc- currence to C. obscuricornis (Table 3). Lar- vae of D. nigripes feed as primary invaders of Eleocharis stems (Valley et al., 1969). Elachiptera nigriceps was collected be- tween early and late August (Table 2), and was most similar in temporal occurrence to D. inclinata (Table 3). Larvae of E. nigriceps feed on decaying vegetation (Valley et al., 1969). Eribolus longulus was collected only in early October (Table 2), and thus exhibited extreme temporal specialization (0.00) (Ta- ble 3). This species was most similar in tem- poral occurrence to D. nigripes (Table 3). Larvae feed as secondary invaders of a va- riety of wetland monocots (Valley et al., 1969), although we did not encounter larvae in stems of Eleocharis. Eribolus nanus was collected in late Oc- tober (Table 2), and exhibited extreme tem- poral specialization (0.00) (Table 3). The flight period of this species did not overlap with that of any other chloropid collected in Eleocharis (Table 3). Larvae of E. nanus feed as secondary invaders of various wet- land monocots (Valley et al., 1969). Pseudopachychaeta approximatonervis was collected in mid-August (Table 2), and was most similar in temporal occurrence to A. neocoxendix (Table 3). Larvae are seed predators in Eleocharis inflorescences (Val- ley et al., 1969). Rhopalopterum carbonarium was collect- ed from mid- to late August (Table 2), and was most similar in temporal occurrence to C. obscuricornis (Table 3). Larvae are known to be secondary invaders of damaged stems of herbaceous wetland plants (Valley et al., 1969). We did not encounter larvae in stems of Eleocharis. DISCUSSION Prior to the 1970’s, most of the research on insects in wetlands focused on taxonom- ic composition or the biology of selected species (Judd, 1949; Davis and Gray, 1966). More recently, research has concentrated on determining the mechanisms affecting in- sect community organization (Tschirnhaus, 1981: Price, 1983). Several authors have suggested that resource partitioning oper- ates to segregate similar species that could be potential competitors for a limited re- source. Partitioning may involve utilization of different plant species (Blair and Foote, 1984) or plant parts (Root and Chaplin, 1976), microhabitat distribution (Hicks and VOLUME 89, NUMBER 4 809 Table 3. Temporal niche breadth ( ) and overlap values for 9 chloropid species collected in E. smallii. i EEE EE EE ee eee a eee eee Species l 2 3 A. neocoxendix (.29) =— 0.84 . obscuricornis (.47) = . nigripes (.47) E. nigriceps (.45) E. longulus (.00) E. nanus (.00) P. approximatonervis (.00) R. carbonarium (.30) CAHN DARWN = A 0.65 (Ei O52 D. inclinata (.24) = D 4 5 6 i 8 9 0.13 0.65 0:00'F 0.009) 10:60! +1050 0:27 0.61 0:00) ), 0:00;)) (0:57 0.59 0.13 0.75 0.00 0.00 0.25 0.25 — 0:20. O13 0.00 0.07 0.13 - 0.00 0.00 0.25 0.50 — 0.00 0.00 0.00 - 0.00 0.00 _ 0.50 Tahvanainen, 1974), or temporal occur- rence (Thornhill, 1980; Stave and Shiff, 1981). The results of this study indicate that re- source partitioning does occur among the Chloropidae inhabiting the marsh. This is revealed by examining the spatial distri- bution, temporal occurrence, and larval trophic habits of the component species (Tables 1, 2). In agreement with previous studies (Schoener, 1974), the spatial re- source axis (vegetation type) is of primary importance in segregating species, followed by the trophic and temporal axes, respec- tively. For example, collection data indicate that A. neocoxendix, C. obscuricornis, D. nigripes, D. inclinata, and P. approxima- tonervis were largely or exclusively collected from E. smallii, whereas Oscinella frit and Thaumatomyia glabra were collected most abundantly in P. arundinacea (Table 1). Be- cause the majority of chloropid species (11) and individuals (99) was collected from E. smallii, this discussion focuses on possible partitioning mechanisms among its com- ponent species. Several authors have sug- gested that competition may not be impor- tant in dictating the niches of phytophagous insects (Strong et al., 1984). Similarly, we have no evidence that resources were lim- iting or that competition was operating to structure chloropid community organiza- tion in the marsh. Trophic habits appear to be important in segregating at least 5 of the species occurring in Eleocharis (Table 2). Larvae of A. neo- coxendix feed on decaying vegetation, a trophic resource which was utilized by only one other chloropid species, E/achiptera ni- griceps. Larvae of D. inclinata feed in the rhizomes of Eleocharis, and are thereby trophically isolated from other species. In contrast, C. obscuricornis and D. nigripes are both primary invaders of Eleocharis stems. However, they differ in their peak emergence times (Table 2). Chlorops ob- scuricornis 1s abundant much earlier in the season than is D. nigripes, and temporal seg- regation appears to be important in pre- venting potential competition between lar- vae of these species should their populations attain levels that could result in Eleocharis stems becoming a limiting resource. Three species, E. Jongulus, E. nanus, and R. car- bonarium are known to be secondary in- vaders of wetland monocots damaged by other insect species. Adult numbers in the Eleocharis stand were relatively low, and it is probable that the larvae do not utilize this particular plant species. Finally, P. approx- imatonervis 1S in a unique trophic niche in that its larvae are seed predators within E/- eocharis inflorescences. In summary, data indicate that the Chlo- ropidae of a northeastern Ohio freshwater marsh exhibit resource partitioning along the spatial resource axis by differential uti- lization of available vegetation types. The 810 majority of species and individuals was ob- tained from the spike-rush, EF. smallii. Within this vegetation type, chloropid species primarily exhibited trophic parti- tioning by utilizing different plant parts as larval food (seeds, stem, rhizomes, decaying tissue). Temporal partitioning was of minor importance in segregating species, with the flight period of most species being between early and mid-August. ACKNOWLEDGMENTS This research was supported by a grant (DEB-79 12242) from the National Science Foundation. We thank Karl Valley, Penn- sylvania Department of Agriculture at Har- risburg, for assistance with chloropid species identifications, and Barbara Andreas, Cuy- ahoga Community College in Cleveland, for aid in determining plant species. LITERATURE CITED Blair, J. M. and B. A. Foote. 1984. Resource parti- tioning in five sympatric species of Scatella (Dip- tera: Ephydridae). Environ. Entomol. 13: 1336- 1339. Colwell, R. K. and D. Futuyma. 1971. On the mea- surement of niche breadth and overlap. Ecology 52: 567-576. Cunliffe, N. 1925. Studies on Oscinella frit (L.). A preliminary investigation of the extent of recovery power of oats when subjected to injury. Ann. Appl. Biol. 12: 276. Davis, L. V.and I.E. Gray. 1966. 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Hypotheses on the organization and evolution of herbivorous insect communities, pp. 559-596. Jn Price, P. W., R. F. Denno, and M. S. McClure, eds., Variable Plants and Herbi- vores in Natural and Managed Systems. Acad. Press, N.Y. Rogers, T. J. 1970. Biology and immature stages of certain phytophagous Chloropidae (Diptera). M.S. Thesis, Kent State University, Kent, Ohio. Root, R. B. and S. J. Chaplin. 1976. The lifestyles of tropical milkweed bugs, Oncopeltus (Hemiptera: Lygaeidae) utilizing the same hosts. Ecology 57: 132-140. Sabrosky, C. W. 1987. Chloropidae, pp. 1049-1067. In McAlpine, J. F. et al., eds., Manual of Nearctic Diptera. Vol. 2. Res. Br. Agric. Can. Monogr. 28. Scheiring, J. F. and B. A. Foote. 1973. Habitat dis- tribution of the shore flies of northeastern Ohio (Diptera: Ephydridae). Ohio J. Sci. 73: 152-166. Schoener, T. W. 1974. Resource partitioning in eco- logical communities. Science 185: 27-39. Starks, K. J. and R. Thurston. 1962. Silver top of bluegrass. J. Econ. Entomol. 55: 865-867. Stave, M. E. and C. J. Shiff. 1981. Temporal segre- gation in North American Membracidae. Oeco- logia 5: 408-411. Strong, D. R., J. H. Lawton, and T. R. E. Southwood. 1984. Insects on Plants. Harvard Univ. Press, Cambridge, Massachusetts. 313 pp. Thornhill, R. 1980. Competition and coexistence among Panorpa scorpionflies (Mecoptera: Pa- norpidae). Ecol. Monogr. 50: 179-197. Todd, J. L. 1985. The community organization of acalyptrate Diptera in a freshwater marsh. M.S. Thesis, Kent State University, Kent, Ohio. Todd, J. L. and B. A. Foote. 1987. Spatial and tem- poral distribution of shore flies in a freshwater marsh (Diptera: Ephydridae). Proc. Entomol. Soc. Wash. 89: 448-457. Tschirnhaus, M., von. 1981. Die Halm- und Min- ierfliegen im Grenzbereich Land-Meer der Nord- see. Spixiana (Suppl. 6): 1-405. Valley, K., T. Wearsch, and B. A. Foote. 1969. Larval feeding habits of certain Chloropidae (Diptera). Proc. Entomol. Soc. Wash. 71: 29-34. Wearsch, T. 1968. Biology and immature stages of certain Chloropidae (Diptera). M.S. Thesis, Kent State University, Kent, Ohio. Wendt, H. 1968. Faunistisch-okologische Unter- suchungen an Halmfliegen der Berliner Umgebung (Dipt. Chloropidae). Dtsch. Entomol. Z. (N.F.) 15: 49-105. Wheeler, A. G., Jr. 1973. Studies on the arthropod fauna of alfalfa. IV. Species associated with the crown. Can. Entomol. 105: 353-366. PROC. ENTOMOL. SOC. WASH. 89(4), 1987, pp. 811-817 THE GENUS POL YPLAX (ANOPLURA: POLYPLACIDAE) IN SULAWESI, INDONESIA, WITH THE DESCRIPTION OF A NEW SPECIES LANCE A. DURDEN 134 Wesley Hall, Vanderbilt University, Nashville, Tennessee 37240. Abstract. — Three species of sucking lice assignable to the genus Polyplax are documented from Sulawesi, Indonesia. Polyplax wallacei, new species, is described from a series of females collected from three species of sympatric murid rodents from primary lowland rain forest in North Sulawesi. Po/yplax eropepli (Ewing) is redescribed from the original type material from the Celebes soft-furred rat, Eropeplus canus, from montane forest in Central Sulawesi; adults of both sexes are illustrated. Polyplax spinulosa (Burmeister) is recorded from two species of commensal rats in North Sulawesi. Sulawesi (Celebes) is one of the principal islands in the Indonesian archipelago and is situated between Wallace’s and Lydek- ker’s zoogeographical lines. Excluding bats, 97% of the mammal species that occur nat- urally on Sulawesi are endemic (Musser, 1987). Few of these mammal species have been processed for ectoparasites and, con- sequently, knowledge of their sucking lice (Anoplura) is poor. At present count, 36 species of endemic murids are known from Sulawesi (Musser, 1987) and only Polyplax eropepli (Ewing), redescribed here, and P. wallacei, new species, are known from this potential host group. At least some of these murid species surely must be parasitized by additional undescribed species of Polyplax (and Hoplopleura) sucking lice. With regard to the commensal murids of Sulawesi, none of which are native to the island (Durden, 1986; Musser, 1987), Po- lyplax spinulosa (Burmeister) is reported here from two species of Rattus. Further collecting from this commensal host group probably will show P. spinulosa to infest also one or more of the other three com- mensal Rattus species now on Sulawesi, and, possibly, Polyplax serrata (Burmeister) will be recorded from the house mouse, Mus musculus castaneus Waterhouse, there. Another commensal mammal now inhab- iting Sulawesi, the house shrew, Suncus mu- rinus (L.), supports an assemblage of Poly- plax spp. lice over its range in southern Asia and it seems likely that one or more of these also parasitize it in Sulawesi. For more than 50 years, P. eropepli has been the only sucking louse documented from any endemic mammal in Sulawesi. Two additional Polyplax species were iden- tified from a collection of mammal ecto- parasites made by the author in North Su- lawesi in 1985. Ectoparasites were collected mainly by intensive visual pelage searches of anesthetized animals but a few hosts were subjected to ectoparasite extractions by skin digestion techniques (Cook, 1954). One of these lice was P. spinulosa, which was taken only from commensal rats, a host group that this species parasitizes on an almost global basis. The second Polyplax species collected was taken only from sympatric, endemic, murids; it represents a new species and a description of it follows. Morphological ter- 812 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON minology designated by Kim (1966) and by Kim and Ludwig (1978) has been followed for the descriptions. The three species of Polyplax reported here are easily separated. Polyplax wallacei, new species, possesses a distinct thoracic sternal plate with central and lateral anterior projections. It has very stout hindleg claws. Although P. spinulosa and P. eropepli both have similarly shield-shaped thoracic ster- nal plates, they are easily separated by the longer paratergal setae and shorter DPHS (dorsal principal head setae) of P. eropepli. Also, P. spinulosa has hindleg claws of in- termediate size, while those of P. eropepli are narrow. Polyplax wallacei Durden, NEw SPECIES Type data.—Holotype @ from adult @ Bunomys chrysocomus (Hoffman) (LAD No. 69), Dumoga-Bone National Park, North Sulawesi (Sulawesi Utara), INDONESIA (0°34'N, 123°53’E) at c. 230 m in primary forest, 26 February 1985, L. A. Durden. Twelve 2 paratypes all from the same lo- cality and by the same collector, but with various dates in February 1985 and host identities as follows: 3 additional specimens from the same host individual as for holo- type; 2 from adult 6 B. chrysocomus (LAD No. 24); 2 from adult 4 B. chrysocomus (LAD No. 32) (there is also a single third-instar Polyplax sp. nymph from this host that is very likely assignable to P. wallacei); 2 from adult 2° B. chrysocomus (LAD No. 51), 1 from adult 2 Bunomys fratrorum (Thomas) (LAD No. 43); 1 from adult 2° B. fratrorum (LAD No. 53); and | from adult ? Taeromys sp. (LAD No. 73). The holotype and one paratype will be deposited in the U.S. Na- tional Museum of Natural History, Wash- ington, D.C. (USNM); two paratypes will be deposited in both the British Museum (Natural History), London, England, and the Museum Zoologicum Bogoriense, Indone- sia. The remaining seven paratypes (and the Polyplax sp. nymph), most of which are not in perfect condition, are in the author’s ref- erence collection. Diagnosis.—Similar to P. spinulosa but distinguished from it most easily by the shape of the thoracic sternal plate which has anterior projections in P. wallacei and by the hindleg claws which are very stout in P. wallacei. In life, P. wallacei is more barrel- shaped than P. spinulosa (dorsoventral flat- tening was accentuated with a blunt probe to prevent skewing of specimens during slide-mounting). The thoracic sternal plate of 2 P. cutchicus Mishra and Kaul from In- dia is partially similar to that of 2 P. wallacei but in the former species the lateral anterior projections are less developed and the plate does not taper posteriorly to a rounded point; there are numerous other differences be- tween these two species. Description.—Total body lengths: Fe- male holotype 1.13 mm, x = 1.19 mm (n = 13, range 1.13-1.31 mm). Female (Figs. 1, 2, 3, 6).—Head, thorax and abdomen well sclerotized. Head: About as long as wide with anterior apex broadly rounded; 2 DMHS (dorsal marginal head setae) on each side; DPHS just extending to thoracic spiracle; DAcHS (dorsal accessory head setae) much larger than DMHS num- bering 2 on each side, both medial to DPHS; DPoCHS (dorsal posterior central head se- tae) and VPHS (ventral principal head se- tae) distinct; antennae 5-segmented with the basal segment much larger than the second segment, slightly wider than long. Thorax: Slightly broader than long; thoracic sternal plate (Fig. 6) very distinct with central an- terior apex prolonged and with less well de- veloped lateral anterior projections, taper- ing to an evenly rounded posterior apex; mesothoracic spiracle fairly small (0.017 mm in diameter); DPTS (dorsal principal thoracic setae) moderate in length (0.105 mm) just extending beyond the insertions for the first TeAS (tergal abdominal setae); DMssS (dorsal mesothoracic setae) well de- veloped. Legs: With subtriangular coxae; forelegs small with acuminate claws; hind- VOLUME 89, NUMBER 4 Figs. 1-7. 7 1, Polyplax wallacei, female. 2, P. wallacei, female genitalia. 3, P. wallacei, female paratergites. 4, Polyplax eropepli, male paratergites. 5, P. eropepli, female paratergites. 6, P. wallacei, female thoracic sternal plates showing variation (the plate to the left represents the most commonly encountered shape). 7, P. eropepli, thoracic sternal plates. Females on left, males on right. legs larger than midlegs, both with strong pointed claws and those on the hindlegs being particularly robust. Abdomen: Wider than thorax with two plates per segment dorsally and ventrally; 12 rows of TeAS, each with 4-8 setae; segments one and two with one seta on each side and tenth row with two setae on each side; one DLAS (dor- sal lateral abdominal seta) on each side of segments 3-6; abdominal sternites equally well sclerotized as tergites; 12 rows of StAS (sternal abdominal setae) each with 4-8 se- tae; first two rows with a total of five (four in three specimens) and four (three in one specimen) setae, respectively; one VLAS (ventral lateral abdominal seta) on each side of segments 3-7. Paratergites (Fig. 3) pres- ent on segments 2-8; plates II-VI subtrian- gular; plates I-V each with two short apical setae; plates VI and VII each with two long- er apical setae; dorsal angles of paratergites I-III produced into points; paratergites II- Vil each with spiracles; paratergite I slightly displaced medially. Genitalia (Fig. 2): With subgenital plate triangular and with its mid- dle apex extended posteriorly; four small 814 setae immediately anterior to the center of this plate; vulvar fimbriae distinct; gono- pods VIII distinct with three setae each, consisting of one long distal seta, one short medial seta and a central seta of interme- diate length; gonopods IX with one short stout seta each. Male.— Unknown. Immatures. — Unknown. Etymology.—The species is named for Alfred Russel Wallace who worked exten- sively in Wallacea, the zoogeographical transition zone named after him that lies between Asia and Australasia. Project Wal- lace, during which this new species was col- lected, was also named for this eminent zo- ologist. Host associations.—Curiously, all 13 known specimens of the new species are fe- males. Also, infestation levels were very low when compared to those for the four Hoplo- pleura species sucking lice also collected from Sulawesi-endemic murids in this sur- vey (Durden, 1986). Bunomys chrysocomus appears to be the principal host of P. wal- lacei; single specimens taken from sympat- ric B. fratrorum and Taeromys sp. rats could represent accidental infestations through ecological associations of these hosts. All three of these mammal species are endemic to Sulawesi. Polyplax eropepli (Ewing) Eremophthirius eropepli Ewing, 1935: 209. Polyplax eriopepli Ewing [sic]: Ferris, 1951: 207. Polyplax eropepli (Ewing): Johnson, 1958: 47. Lectotype designated. Type data.—Lectotype ¢ and 3 paralec- totypes (1 6, 2 2) on one microscope slide (USNM type no. 44906) from young adult 2 Eropeplus canus Miller and Hollister (Ce- lebes soft-furred rat) pelt (USNM 219711) presumably removed by H. E. Ewing. The host specimen was collected from tropical montane rain forest (mossy, primary forest) at an elevation of c. 1800 m at Rano Rano, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Central Sulawesi (Sulawesi Tengah), IN- DONESIA (1°30’S, 120°28’E) on 15 De- cember 1917 by H. C. Raven (Musser, 1969, 1970, and Musser, American Museum of Natural History, New York, pers. comm.). Diagnosis.—Similar to P. spinulosa but distinguished from it by a number of char- acters and particularly by the longer para- tergal setae of P. eropepli. Whereas both paratergal setae on the same plate are ap- proximately equal in length in P. spinulosa, the dorsal seta is longer than the ventral seta on plates II-VI of males and plate VI of females in P. eropepli. Other notable differ- ences between both sexes of these two species are the much longer DPHS and the stouter tarsal claws of P. spinulosa. Polyplax ero- pepli and P. wallacei, new species, are sep- arable from numerous characters and most easily by the shapes of the thoracic sternal plates and the more robust tarsal claws of P. wallacei. Description.— Total body lengths: Male lectotype 1.03 mm, male paralectotype 0.98 mm, female paralectotypes 1.42 and 1.39 mm. Male (Figs. 4, 7, 10, 11).—Head, thorax and abdomen well sclerotized. Head: About as long as broad, anterior apex separable into four low festoons with a small frontal tubercle on the dorsal, central festoon; DPHS fairly short, not quite reaching sec- ond leg coxa; 2 DAcHS on each side with the medial seta short and the other seta about half the length of the DPHS; DPoCHS and VPHS distinct; antennae 5-segmented with strong sexual dimorphism, with the basal segment much larger than the second seg- ment, about as wide as long; third segment highly modified with a long straight apical process situated at its posterodistal corner and about twice the length of the segment proper—see Ewing (1935) for further de- scription including a detailed drawing of this segment. Thorax: Broader than long; tho- racic sternal plate (Fig. 7) similar to that of P. spinulosa with posterolateral angles pro- duced into a blunt point and lateral margins VOLUME 89, NUMBER 4 Figs. 8-11. almost parallel but with the anterior edge less rounded than in P. spinulosa; mesotho- racic spiracle fairly small (0.019 mm in di- ameter); DPTS moderate in length (0.124 mm), not quite extending to the first ab- dominal tergite; DMsS distinct. Legs: With subtriangular coxae; forelegs smaller than other legs, each with a narrow acuminate claw. Abdomen: Wider than thorax, with one broad plate per segment dorsally and ventrally except for the venter of segment three which has two narrower plates; 7 rows of TeAS and StAS, each with 6-8 setae. Paratergites (Fig. 4) present on segments 2 815 Polyplax eropepli. 8, Female. 9, Female genitalia. 10, Male. 11, Male genitalia. to 8; plates I-VI subtriangular; plate I with 2 short setae of equal length; plates II-VI with dorsal seta longer than ventral seta, particularly so in plates V and VI; setae on plate VII broken; paratergites showing dif- ferential sclerotization especially marked on plate III; dorsal angles of plates H-V show- ing some degree of development into points; plates II-VII each with spiracles. Genitalia (Fig. 11): With basal apodeme broadening anteriorly and extending to the sixth ab- dominal segment; parameres very short and curved, each articulating distally with base of pseudopenis; pseudopenis broadly 816 rounded apically and extending beyond the apices of the parameres. Female (Figs. 5, 7, 8, 9).—Head, thorax and abdomen as in male unless indicated otherwise. Head: Longer than broad, with- out frontal tubercle; first antennal segment longer than broad; third antennal segment not highly modified but instead similar to the other adjacent segments. Legs: Missing except for coxae which are similar to those of the male. Abdomen: With two plates per segment dorsally (except for segments three and four which have one) and ventrally (ex- cept for segment three which has one); ster- nites of segment four narrow; 12 rows of TeAS, each with 4—7 setae (except for the first row which has two); 13 rows of StAS, each with 4-7 setae; 6 rows of DLAS and VLAS each with one seta per side. Parater- gites (Fig. 5) all showing clear differential sclerotization and all subtriangular; plates I-V each with both setae of about equal length (the dorsal setae of plates I and II are very slightly longer than their correspond- ing ventral setae); dorsal seta of plate VI clearly longer than corresponding ventral seta; setae of plate VII much longer than other paratergal setae and about equal in length. Genitalia (Fig. 9): With subgenital plate broadly triangular and with the medial apex extended posteriorly; four small setae immediately anterior to this plate; vulvar fimbriae distinct; gonopods VIII with three setae each, with broken lateral setae, long central setae and shorter medial setae; gon- opods IX with two short setae each, with the lateral setae longer than the medial se- tae. Immatures.— Unknown. Remarks.—In his original description of P. eropepli, Ewing (1935) relied heavily on characters of the third antennal segment of the male, and the only drawing he gave was of this structure. Although this segment is highly modified and probably characteris- tic, by current standards (Kim, 1966; Kim and Ludwig, 1978; Weisser and Kim, 1973) the original description is inadequate. To PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON date, no additional specimens of P. eropepli have been collected to supplement the type series. Collection data are given here for the type host for the first time. One problem with this redescription has been the poor quality of the type specimens, all of which are mounted on the same microscope slide. Appendages are missing, setae are broken, and specimens are not mounted in good dorsoventral aspect. The illustrations given are composites. Both female lice lack all leg segments except coxae. Both sexes are de- void of some other minor features. Johnson (1958) designated the male specimen that is second from the right on the slide as the lectotype for P. eropepli (and circled this specimen by etching the back of the slide with a diamond marker); this is the most complete specimen and therefore a good choice as such. Johnson (1958) had intend- ed to redescribe P. eropepli some years ago but was unable to arrange this (Johnson, National Marine Fisheries Service, Oxford, MD, pers. comm.). Host associations. —Polyplax eropepli is rare, having been collected just once, but the host, E. canus, is also rare with fewer than ten documented museum specimens. This murid has been taken only from the mountainous areas of central Sulawesi in primary forest at or above 1800 m elevation (Musser, 1970, 1987). Ferris (1951) mistak- enly listed the type host of P. eropepli as Eriopeplus incanus, which he stated to be Cricetulus longicaudatus (Johnson, 1958). Ewing (1935) presented the type host infor- mation correctly in his original description. Polyplax spinulosa (Burmeister) Sulawesi collecting records.—Two 2 from adult 2 Rattus argentiventer (Robinson and Kloss) (the ricefield rat) on 8 February 1985, L. A. Durden, Toraut Base Camp (0°34’N, 123°54’E), Sulawesi Utara, INDONESIA; 1 2 from adult ¢ Rattus exulans (Peale) (the Polynesian rat) on 12 February 1985 by the same collector at the same locality; 1 2, 1 4 from adult ¢ R. argentiventer on 13 Feb- VOLUME 89, NUMBER 4 ruary 1985 by the same collector at the same locality. Remarks. — Polyplax spinulosa, the spiny rat louse, is almost cosmopolitan in distri- bution and probably invaded Sulawesi rel- atively recently with its commensal rat col- onizers (Durden, 1986). Further collecting will probably show that P. spinulosa also infests one or more of the other three com- mensal rats now on Sulawesi [R. rattus (L.), R. norvegicus (Berkenhout) and R. nitidus (Hodgson)]. ACKNOWLEDGMENTS I am indebted to Phyllis Johnson for for- warding much useful information including copies of her notes on P. eropepliand related Anoplura and to Ke Chung Kim for his helpful communications and comments on earlier drafts of this paper. Guy Musser identified voucher mammal specimens and provided collecting data on the type host for P. eropepli including distribution par- ticulars for this and other murid species in Sulawesi. Gary Hevel of the U.S. National Museum arranged the loan of the P. eropepli syntypes. Chris Lyal and Jenny Palmer made arrangements for me to examine the Ano- plura collections in the British Museum (Natural History). Nixon Wilson provided a large collection of reprints on the Ano- plura. This paper is based partly on material collected while the author was a participant on Project Wallace, sponsored by the Royal Entomological Society of London and the Indonesian Institute of Sciences (Results of Project Wallace No. 33). My participation in Project Wallace was funded by the Com- 817 mittee for Research and Exploration of the National Geographic Society. LITERATURE CITED Cook, E. F. 1954. A modification of Hopkin’s tech- nique for collecting ectoparasites from mamma- lian skins. Entomol. News 65: 35-37. Durden, L. A. 1986. Ectoparasites and other arthro- pod associates of tropical rain forest mammals in Sulawesi Utara, Indonesia. Nat. Geog. Res. 2: 320- 331. Ewing, H. E. 1935. The taxonomy of the anopluran genera Polyplax and Eremophthirius, including the description of new species. Proc. Biol. Soc. Wash. 48: 201-210. Ferris, G. F. 1951. The sucking lice. Mem. Pac. Coast Entomol. Soc. 1: 1-320. Johnson, P. T. 1958. Type specimens of lice (Order Anoplura) in the United States National Museum. Proc. U.S. Nat. Mus. 108: 39-49. Kim, K. C. 1966. The species of Enderleinellus (An- oplura, Hoplopleuridae) parasitic on the Sciurini and Tamiasciurini. J. Parasitol. 52: 988-1024. Kim, K. C. and H. W. Ludwig. 1978. The family classification of the Anoplura. Syst. Entomol. 3: 249-284. Musser, G. G. 1969. Results of the Archbold expe- ditions. No. 89. Notes on the taxonomic status of Rattus aspinatus Tate and Archbold and Mus cal- litrichus Jentink (Rodentia, Muridae). Am. Mus. Novit. 2365: 1-9. 1970. Results of the Archbold expeditions. No. 93. Reidentification and reallocation of Mus callitrichus and allocations of Rattus maculipilis, R. m. jentinki, and R. microbullatus (Rodentia, Muridae). Am. Mus. Novit. 2440: 1-35. . 1987. The mammals of Sulawesi, pp. 73-93. In Whitmore, T. C., ed., Biogeographic Evolution of the Malay Archipelago. Oxford University Press, Oxford. Weisser, C. F. and K. C. Kim. 1973. Rediscovery of Solenopotes tarandi (Mjoberg, 1915) (Linognath- idae: Anoplura), with ectoparasites of the barren ground caribou. Parasitology 66: 123-132. PROC. ENTOMOL. SOC. WASH. 89(4), 1987, pp. 818-820 HOST PLANTS AND SEASONAL ABUNDANCE OF ADULT CAPRAITA SUBVITTATA (COLEOPTERA: CHRYSOMELIDAE: ALTICINAE) OwEN D. V. SHOLES Division of Natural Sciences, Assumption College, Worcester, Massachusetts 01609. Abstract. —In central New York, adults of Capraita subvittata fed nocturnally on leaves of Aster divaricatus, A. macrophyllus and Veronica officinalis. Adults were active at least from early May to mid August, but no immature stages were found on or near the adult host plants at any time during this study. North American species of Capraita feed on hosts from at least 17 plant families. Adults of the flea beetle Capraita subvit- tata (Horn) (formerly in the genus Oediony- chis; Blake, 1927) are 3-5 mm long, and tan or yellow brown with brown or black dorsal markings (Blake, 1927). During an herbi- vore removal experiment with Aster divar- icatus L. (Asteraceae), I observed C. sub- vittata adults at night on plants in the forest understory. METHODS On 62 nights from 6 May to 18 August 1985, I counted all herbivorous insects and gastropods that I found between 2200 and 2400 h on 30 0.65-m? plots. The plots were established in early May on the first 30 clumps of Aster divaricatus I encountered along a transect running parallel to and 150 m upslope from the southwestern shore of Lake Myosotis in the E. N. Huyck Preserve in Rensselaerville, New York. On half of the plots, I removed the herbivores by hand (mammals were excluded by cages of 1-inch- mesh chicken wire). Within an hour after removal, representative individuals were placed singly into vials with undamaged leaves of A. divaricatus. Some individuals of C. subvittata were put into vials with un- damaged leaves of Veronica officinalis (Scrophulariaceae). The vials were checked the following morning for damage to the leaves and for accumulation of frass. RESULTS AND DISCUSSION Adult C. subvittata chewed rectangular or arc-shaped holes in the leaves of A. divari- catus and V. officinalis. Single beetles ate 10-30 mm*? of leaf area during 8-10 h in a vial, probably less than what an undisturbed individual could eat in one night in the field. Of the 118 C. subvittata adults seen at night, 114 were on Aster divaricatus, two were on Veronica officinalis L. (Scrophular- iaceae), and two were on Aster macrophyllus L. The holes in A. macrophyllus leaves were indistinguishable from those produced by C. subvittata in the feeding trials. Clearly, Aster divaricatus is the main host plant for adult C. subvittata, but not the only one. Capraita subvittata feeds on other as- ters, on Scrophulariaceae, and perhaps on Rosaceae (Table 1). In North America, the genus has been found on at least four su- perorders, 13 orders, 17 families and 22 genera of dicots (Table 1). Individual species of Capraita have 1-6 species of host plants used by adult beetles (Table 1). Larvae of C. gibbitarsa have been collected from mint, VOLUME 89, NUMBER 4 819 Table 1. Plant hosts for adults of North American species of Capraita. USNM: United States National Museum: MCZ: Museum of Comparative Zoology, Harvard University. Plant superorders are from Stebbins (1974). RS Host plant taxa Capraita Source i I. Asteridae— Asteraceae Aster divaricatus subvittata this study Aster macrophyllus subvittata this study Caprifoliaceae Lonicera sp. tenuilingata USNM Oleaceae Chionanthus virginicus sexmaculata Wilcox, 1979 Fraxinus sp. sexmaculata Wilcox, 1979 Lamiaceae Teucrium canadense thyamoides Wilcox, 1979 Verbenaceae Verbena urticifolia circumdata Wilcox, 1979 circumdata eggs Blake, 1927 Plantaginaceae Plantago lanceolata circumdata Wilcox, 1979 Convolvulaceae jacobiana USNM Solanaceae petaurista USNM Scrophulariaceae jacobiana USNM Dasistoma macrophylla hornii USNM Mimulus ringens subyittata Wilcox, 1979 Veronica officinalis subvittata this study Bignoniaceae Chilopsis linearis durangoensis USNM II. Rosidae— Rosaceae Rubus sp. circumdata Wilcox, 1979 Amelanchier flowers subvittata MCZ Aquifoliaceae Tlex glabra suturalis Wilcox, 1979 Ilex opaca obsidiana Wilcox, 1979 Tlex verticillata obsidiana Wilcox, 1979 Celastraceae Euonymus americanus obsidiana Wilcox, 1979 Fabaceae several spp. USNM Juglandaceae Juglans cinerea circumdata Wilcox, 1979 III. Hamamelidae—Fagaceae Fagus grandifolia young leaves circumdata Blake, 1927 Quercus spp. obsidiana Wilcox, 1979 quercata Blake, 1927 IV. Dilleniidae—Ericaceae suturalis Wilcox, 1979 Florida spp. Blake, 1927 Vaccinium sp. obsidiana Wilcox, 1979 circumdata Wilcox, 1979 Vaccinium virgatum obsidiana Wilcox, 1979 820 o NUMBER OF BEETLES £ Je Le oO MAY JUNE JULY AUG Fig. 1. Numbers of adult Capraita subvittata ob- served at night (2200-2400 h) on 30 0.65-m? plots between 6 May and 18 August, 1985, at the E. N. Huyck Preserve in Rensselaerville, New York. but are otherwise poorly known (Blake, 1927). The numbers of adult C. subvittata fluc- tuated widely between nights, but the range of variation was fairly steady through the sampling period until the numbers of adults declined in August (Fig. 1). I observed mat- ing pairs of beetles on 10 and 11 June. Both observations were in the same plot, so it may have been the same pair both nights. I saw no eggs, larvae or pupae of C. subvit- tata on any plants at any time of day in these plots. The duration of adult presence was longer for C. subvittata in this study than it was for univoltine chrysomelid species on Sol- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON idago host plants in old fields (Messina and Root, 1980). Even so, no firm conclusion on voltinism of C. subvittata can be reached until more is known about its immature stages. ACKNOWLEDGMENTS Deborah Gordon, Susan Beatty, George Bryce, and Tim McCabe provided help at various stages of this study, E. R. Hoebeke identified the beetles, and two anonymous reviewers made helpful comments on the manuscript and provided the host records from specimens in the U.S. National Mu- seum. This research was supported by a Summer Fellowship from the Edmund Niles Huyck Preserve, and by Faculty Develop- ment Funds from Assumption College. LITERATURE CITED Blake, D. H. 1927. Revision of the beetles of the genus Oedionychis occurring in America north of Mexico. Proc. U.S. Nat. Mus. 70: 1-44. Messina, F. J. and R. B. Root. 1980. Association between leaf beetles and meadow goldenrods (Sol- idago spp.) in central New York. Ann. Entomol. Soc. Am. 73: 641-646. Stebbins, G. L. 1974. Flowering Plants: Evolution above the Species Level. Harvard University Press, Cambridge. 399 pp. Wilcox, J. A. 1979. Leaf Beetle Host Plants in North- eastern North America (Coleoptera: Chrysomeli- dae). E. J. Brill Publishing, New York; Biological Research Institute of America. 30 pp. PROC. ENTOMOL. SOC. WASH. 89(4), 1987, pp. 821-824 THE ESTABLISHMENT OF CHILOCORUS KUWANAE (COLEOPTERA: COCCINELLIDAE) IN EASTERN UNITED STATES J. J. DREA AND R. W. CARLSON (JJD) Beneficial Insects Laboratory, BBII, Agricultural Research Service, USDA, BARC-E, Beltsville, Maryland 20705; (RWC) Biosystematic and Beneficial Insects Insti- tute, Agricultural Research Service, USDA, BARC-W, Beltsville, Maryland 20705 (for- merly at the Asian Parasite Laboratory, % American Embassy, Seoul, Republic of Korea). Abstract. —The coccinellid Chilocorus kuwanae (Silvestri) was introduced from the Re- public of Korea into the Washington, D.C. area in 1984 as part of an Agricultural Research Service project for the biological control of diaspine scales. The beetle was released on euonymus trees and shrubs infested with the euonymus scale, Unaspis euonymi (Com- stock). By the third year after release, the beetle had greatly increased in number and had drastically reduced the population of the scale on the test plants. A brief review of the biology of the beetle is given, and a morphological character is figured to differentiate adults of C. kuwanae from the similar native species, C. stigma (Say). Coccinellids are the most important in- sect predators of diaspine scale insects throughout the world. Species of this family are credited with complete, substantial, or partial control of many species of armored scales (DeBach and Rosen, 1976). Conse- quently, when the Agricultural Research Service, USDA established a Small Farms Research Project for the control of scale pests, lady beetles were among the prime candidates considered for introduction as part of the biological control effort. One of the main target insect pests of the project was the euonymus scale, Unaspis eu- onymi (Comstock) (Homoptera: Diaspidi- dae), a very serious pest of many ornamen- tal trees and shrubs throughout much of the temperate region, including the United States (Gill et al., 1982). Because this scale appears to be of Asian origin, the Asian Par- asite Laboratory, Seoul, Republic of Korea (ROK), was assigned the task of surveying for, studying, and collecting natural enemies of this scale in Asia. The coccinellid Chilocorus kuwanae (Sil- vestri) was among the 10 or more species of biotic agents associated with this scale in Korea. Eventually the predator was shipped to the United States and released at various locations in eastern United States. This re- port documents the establishment of the coccinellid in the eastern states and fur- nishes biological data relating to the species. DISTINGUISHING FEATURES The exotic Chilocorus kuwanae (Fig. 1) is very similar in appearance to C. stigma (Say), a native species commonly found on other diaspine scales in North America (Gordon, 1985). Both species are black ex- cept for the abdomen and the two elytral spots. These spots are reddish and some- what laterally rectangular in C. kuwanae and are more orange-yellow and round in C. stigma. However, for the untrained, the liv- ing adults are indistinguishable especially if specimens of only one species are on hand. For recovery and evaluation purposes, 822 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Adult of Chilocorus kuwanae. Fig. 2. Reticulate surface of pronotum of Chilocorus stigma. Fig. 3. Smooth surface of pronotum of Chilocorus kuwanae. Fig. 4. Third larval instar of Chilocorus kuwanae. Fig. 5. Pupa of Chilocorus kuwanae. Bars of Figs. 1, 4, 5 represent 2 mm. there was need for a differentiating character that would require neither dissection nor a direct comparison of specimens of both species. The presence or absence of sculp- turing on the pronotum of the adult beetle provided just such a character. Under the high power (100) of a dissecting micro- scope the intersetal area of the pronotum of C. stigma is distinctly reticulate (Fig. 2), whereas the same area of C. kuwanae is smooth and has an oil-like film on living specimens (Fig. 3). BIOLOGY Several references in the literature de- scribe various aspects of the biology of C. kuwanae: Ishii (1937), Nohara (1962), and Xia et al. (1986) give detailed information on the life cycle; Kato (1968) and Xia et al. (1986) report on predation and host range of the species; Tanaka and Kobayashi (1970) review the development and food relation- ship with the prey; Tanaka and Wada (1973) consider insectary production; and the ef- fectiveness of the species as a natural enemy of scales is given by Zhang (1983). Several species of coccids and aphids are recorded as prey of C. kuwanae, but diaspine scales are the most common food of this cocci- nellid (Kamiya, 1966). Chilocorus kuwanae 1s essentially Far Eastern in origin and is widely distributed in China (Xia et al., 1986) and Japan (Ka- miya, 1966). According to Clausen (1956) the species was introduced into the United States in Georgia in 1901-1902 under the VOLUME 89, NUMBER 4 name C. similis, but the beetle was not re- covered after 1905. Smith (1965) reported the same species (i.e. C. similis) established in the Santa Barbara area of California from an introduction made in 1923. However, Gordon (1985) considered that the C. sim- ilis of California is actually C. kuwanae and the other records are misidentifications. As with other species of Chilocorus the eggs are deposited singly or in small num- bers under empty scale coverings or in cracks and other protected places of the substrate. At room temperature eggs hatch in about 8 days and larvae feed for 2 to 4 weeks, de- pending upon the availability of food. The larva (Fig. 4) is voracious and will consume several hundred scales during its develop- ment (Nohara, 1962). The pupa (Fig. 5) is formed on the plant where the larva developed. Pupation often occurs in congregations or small groups. This stage lasts for 1-2 weeks, depending on am- bient temperatures. The species overwinters as an adult in protected places on or about the host plant. According to Nohara (1962), activity begins in spring with the onset of warm weather, over 10°C. At our release site, however, bee- tles were active even on January days if the temperature rose above 10°C. There are three generations per year, but there is an extensive overlapping of all stages during the growing season. Adults are found on plants throughout the year. RELEASES AND RECOVERIES All insects released were from laboratory cultures established from adult beetles col- lected in Sacheon, Kyeong Sangnam Prov- ince, Republic of Korea, on Evonymus ja- ponica infested with U. euonymi. The shipments from Korea were received through the quarantine facility of the Ag- ricultural Research Service, Beneficial In- sects Research Laboratory, Newark, Dela- ware. In late September and October, 1984, 171 823 larvae and 9 adults were released on a eu- onymus tree, Evonymus europaeus L., heavily infested with U. euonymi at the United States National Arboretum in Washington, D.C. In October of 1985 an additional 25 adults were released on the same tree. By mid-1986 all stages of the beetle were evident on all the trees of the small grove of euonymus, including E. hamiltonianus Wallich var. nikoensis (Na- kai) Blakelock and E. kiautschovicus, about 15 trees on less than 0.5 hectares. From July to October of 1986 more than 400 adults and larvae of C. kuwanae were collected from the release site for distribution to other regions in eastern United States, but no ap- parent reduction of the field population of the coccinellid was observed at the collec- tion site. Therefore, the potential for pop- ulation growth appears to be great. Although both adults and larvae were re- leased, the adults tended to disperse and leave the tree. Consequently, the establish- ment of C. kuwanae at the location appears to be the result of the use of larvae for release purposes. Samways (1984), in the use of Chilocorus nigritus F. for control of Cali- fornia red scale, Aonidiella aurantii Mas- kell, proposed the release of eggs of the coc- cinellid to avoid rapid dispersion. Eggs of C. kuwanae from laboratory culture were placed on the trees at the Arboretum but no larvae were observed. Failure of this tech- nique is attributed to predation and desic- cation. Very few specimens of the native C. stig- ma were ever recovered from any of the euonymus scale infestations that were stud- ied, although the predator commonly is as- sociated with many other species of dias- pine scales in the same general area. CONCLUSION The level of control affected by C. ku- wanae has not been established, but the vis- ible level of the scale population at the re- lease site is considerably reduced. This 824 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON colony of beetles will continue to serve as a source of specimens for redistribution to other areas of the United States. ACKNOWLEDGMENTS We thank Ho-Yeon Han and Jang-Hoon Lee, personnel of the Asian Parasite Lab- oratory who assisted in the original collec- tions in the Republic of Korea; S. G. March of the U.S. National Arboretum, Washing- ton, D.C. for assistance in locating scale populations and support during the project at the arboretum; R. D. Gordon, Systematic Entomology Laboratory, Agricultural Re- search Service, USDA, Washington, D.C. for determination of the coccinellid speci- mens; L. R. Ertle and R. M. Hendrickson, Beneficial Insects Research Laboratory, Ag- ricultural Research Service, USDA, New- ark, Delaware for assistance in quarantine clearance and release of the beetle in the northern areas; and E. McClunin, Depart- ment of Entomology, University of Mary- land, College Park, who took the photo- graphs. LITERATURE CITED Clausen, C. P. 1956. Biological control of insect pests in the continental United States. U.S. Dep. Agric. Tech. Bull. 1139: 1-151. DeBach, P. and D. Rosen. 1976. Armoured scale in- sects, pp. 139-178. In Delucchi, V. L., ed., Studies in Biological Control. International Biological Programme 9. Cambridge Univ., London. 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. Univ. Md. Agric. Exp. Stn. Misc. Pub. 969. 36 pp. Gordon, R. D. 1985. The Coccinellidae (Coleoptera) of America north of Mexico. J. N.Y. Entomol. Soc. 93: 1-912. Ishii, T. 1937. On the natural enemies of arrowhead scale, Prontaspis yanonensis Kuw. Agric. Hort. (Tokyo) 12: 60-70. (Rev. Appl. Entomol. Ser. A, 25: 303-304. 1937). Kamiya, H. 1966. On the Coccinellidae attacking the scale insects and mites in Japan and the Ryukyus. Mushi 39: 65-93. Kato, T. 1968. Predaceous behavior of coccidopha- gous coccinellid Chilocorus kuwanae on the hedge of Euonymus japonicus Thunberg. Kontyt 36: 29- 38. Nohara, K. 1962. Studies on the life history and pred- atory behavior of Chilocorus kuwanae Silvestri (Coleoptera, Coccinellidae). Sci. Bull. Fac. Agric. Kyushu Univ. 20: 29-32. Samways, M. 1984. Biology and economic value of the scale predator Chilocorus nigritus (F.) (Coc- cinellidae). Commonw. Inst. Biol. Control, Bio- control News and Information 5: 91-105. Smith, S. G. 1965. Chilocorus similis Rossi: Disin- terment and case history. Science 148: 1614-1616. Tanaka, M. and M. Kobayashi. 1970. Food con- sumption of Chilocorus kuwanae Silvestri (Coc- cinellidae). Proc. Assoc. Plant Prot. Kyushu 16: 56-59. Tanaka, M. and K. Wada. 1973. Factors affecting the rate of emergence of Chilocorus kuwanae Silvestri (Coccinellidae) in an insectary. Proc. Assoc. Plant Prot. Kyushu 19: 71-73. Xia, B., Y. Zhang, and B. Shen. 1986. Biology of Chilocorus kuwanae and its control of insects in the field. Chin. J. Biol. Control 2: 70-74. Zhang, S. Y. 1983. Observations on the effectiveness of the natural control of Chilocorus kuwanae Sil- vestri (Col.: Coccinellidae) against two species of scales in the orchard. Natural Enemies of Insects 5: 86-88. (Hort. Abstr. 1984, 54: 3109). PROC. ENTOMOL. SOC. WASH. 89(4), 1987, pp. 825-830 SALMOPERLA, A NEW STONEFLY GENUS FROM NORTHERN CALIFORNIA (PLECOPTERA: PERLODIDAE) RICHARD W. BAUMANN AND DaAvID R. LAUCK (RWB) Department of Zoology and Monte L. Bean Life Science Museum, Brigham Young University, Provo, Utah 84602; (DRL) Department of Biology, Humboldt State University, Arcata, California 95521. Abstract. — Salmoperla is designated as a new genus in the subfamily Perlodinae, tribe Arcynopterygini, with the new species Salmoperla sylvanica as the type species. A brief description of the genus is given that includes characters that delineate it from the other genera in Perlodinae. Salmoperla sylvanica is described for the adult male and mature male and female nymphs. Illustrations are provided, along with a diagnosis that compares it with Oroperla barbara and Perlinodes aureus. During the spring of 1984 Peter Wilkin- son did extensive collecting of stonefly nymphs and adults in the Willow Creek drainage in Humboldt County, California (Wilkinson, 1986). His study was restricted to Willow Creek and several tributaries above the confluence with the Trinity River. He obtained several nymphs and a single male adult of a peculiar perlodid species that he had trouble identifying correctly. This interesting large stonefly could not be classified even to the generic level by using the recent key to North American Perlo- dinae genera (Stewart and Stark, 1984). In addition, another population of nymphs was discovered in a 1981 collection from a trib- utary of the Klamath River north of the Hoopa Indian Reservation. These events led to the organization of a special field trip in April 1987 for the ex- press purpose of collecting additional spec- imens of this interesting stonefly. Three re- searchers from Brigham Young University: R. W. Baumann, C. R. Nelson, and S. A. Wells; B. P. Stark from Mississippi College, and D. A. Lauck from Humboldt State Uni- versity joined together to learn more about this stonefly and attempted to collect an adult female. However, we only succeeded in collecting an additional male adult and mature nymphs from several localities. Be- cause nymphs of this species were studied by K. W. Stewart and B. P. Stark as part of their survey of the nymphs of North Amer- ican Plecoptera, it was decided that a de- scription of this previously undescribed ge- nus and species should be done now so it can be included in their forthcoming book on stonefly nymphs. Salmoperla Baumann and Lauck, New GENUS Type species.—Salmoperla sylvanica Baumann and Lauck, new species. Size.— Large: adult male 22-25 mm; fe- male unknown; mature nymphs 25-37 mm. Gills. —Submental (SM) small and tri- angular; anterior thoracic (AT,) tiny; pro- thoracic (ASC, ) small and single; meso (AT;) and meta thoracic (AT,;) double, large and finger-like (Fig. 2). Mesosternum of nymph.—Arms of Y-ridge meet posterior corners of furcal pits (Fig. 2). 826 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Te tld Fig. 1. Salmoperla sylvanica, mature nymphal habitus. VOLUME 89, NUMBER 4 Genitalia.—Epiproct large, darkly scler- otized and covered with short, stout spi- nules; lateral stylets present and well de- veloped. Nymphal mouthparts.—Lacinia_ biden- tate, with fringe of thin spinules on inner margin; right mandible moderately cleft with small denticles on ventral teeth (Figs. 5, 6). Distribution. — Northern Carolina. Salmoperla sylvanica Baumann and Lauck, NEw SPECIES Figs. 1-11 Male.—Slightly brachypterous. Length of forewings 15-17 mm; length of body 22-25 mm. General color dark brown but with yellow markings; legs yellow with brown areas at joints; antennae dark brown. Head wider than prothorax, mostly yellow but with dark brown areas laterally behind eyes. Pronotum wider than long, brown, with broad median yellow stripe, which extends onto mesonotum. Wings clear, veins brown, numerous crossveins and small cells pres- ent. Abdominal terga dark brown, with paired light colored patches on first four seg- ments; sterna yellow. Cerci light brown. Ninth sternum enlarged, extending poste- riorly over base of paraprocts; ninth tergum with enlarged lateral humps, covered with small stout setae and long fine hairs. Tenth tergum with moderate-sized genital lobes, narrowly rounded, tips of lobes darkly scler- otized and bearing a few short, stout setae (Fig. 11); paragenital plates darkly sclero- tized, basal sclerotizations as flattened three- pronged fork, lateral sclerotized bars large and expanded at base. Epiproct large and almost completely sclerotized, broadest at base, tapering to narrower truncate tip, which is membranous, dorsal anterior half of epiproct covered with small dark spi- nules. Lateral stylets, short broad and dark- ly sclerotized, apex shaped as stout outward directed hook. Female.— Unknown. Nymph.—Length: mature males 25-30 mm; mature females 32-37 mm. Dorsal color brown with light paisley patterns dor- 827 sally (Fig. 1). Head with light colored areas on median of labrum and anterior portion of head beyond ocelli, ovoid light area in ocellar triangle, large light areas on poster- iorlateral margins behind compound eyes; antennae light brown. Pronotum lighter me- dially and darker near lateral margins. Mesonotum, metanotum and abdominal terga basically brown, with faint median stripe. Ventral color pattern much lighter with distinctly sclerotized areas on thoracic sterna. Cerci light brown. Abdominal seg- ments one to seven with enlarged membra- nous areas on lateral margins, best devel- oped on anterior segments and gradually becoming smaller posteriorly. Mandibles and maxillae greatly enlarged laterally and visible in dorsal view. Lacinia bidentate, with only thin spinules on inner margin (Fig. 6). Right mandible moderately cleft, with small denticles on ventral teeth and small patch of tiny denticles at base of dorsal teeth, fringes of hairs present on inner margins of both teeth below cleft (Fig. 5). Legs covered by numerous short spinules, bearing short but heavy lateral fringe of hairs (Fig. 7). Abdominal segments covered with short in- tercalary spinules, terminal sterna bearing fringe of posterior spinules (Fig. 4), fringe interrupted on females where truncate lobe occurs along posterior median margin of eight sternum (Fig. 3). Cerci with lateral fringe of long fine hairs, basal segments very short and tightly compacted, bearing few short stout spinules on posterior margin, oc- casionally larger lateral spines present (Fig. 8), middle segments with intercalary spi- nules and small setae, lateral spines larger and forming whorls along posterior margin (Fig. 9), terminal segments with intercalary setae and well developed whorls of spines at apex of each segment (Fig. 10). Diagnosis.—Salmoperla sylvanica be- longs to the perlodine tribe Arcynopterygini (Stark and Szczytko, 1984). Its sister species is Oroperla barbara Needham, which was described from the nymphal state because of its distinctive abdominal gills (Needham, 1933). Salmoperla has the same pattern of 828 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 2-11. Salmoperla sylvanica: 2, Nymphal thoracic sterna. 3, Female nymph, terminal abdominal sterna. 4, Male nymph, terminal abdominal sterna. 5, Right nymphal mandible, ventral. 6, Right nymphal lacinia, ventral. 7, Right front leg, nymph, dorsal. 8, 9, 10, Basal, middle and apical cercal segments, nymph. 11, Male terminalia, adult, dorsal. VOLUME 89, NUMBER 4 nymphal gills on the submentum and tho- racic sterna (Shepard and Stewart, 1983) and it has large, expanded membranous areas on abdominal segments one to seven, where Oroperla has paired gills. These genera have very similar mandibles and maxillae, which are also greatly expanded laterally and are easily visible in dorsal view. The males are very similar in size and basic morphology, including their wings in both size and ve- nation. Their genitalia are also rather sim- ilar and they both have short, stout lateral stylets. Salmoperla has a much larger epi- proct but it is structurally similar to that found in Oroperla. Even though these gen- era are relatively similar, the differences, es- pecially in the gills, are large enough to war- rant the naming of a new genus. The only other species that is at all similar to S. sy/- vanica is Perlinodes aureus (Smith). Perli- nodes, however, has the thoracic gills single and some peculiar, derived male characters that are absent in Sa/moperla (Stark and Stewart, 1982). Types.—Holotype 4, stream one-third mile west of Ruby Creek, Hwy 299, above junction Willow Creek, Humboldt County, California, 25 April 1987, R. W. Baumann, C. R. Nelson, B. P. Stark, and S. A. Wells. Paratype 4, stream one-half mile east of Ce- dar Creek, Hwy 299, above junction Willow Creek, Humboldt Co., California, 20 April 1984, P. Wilkinson (BYU). Nymphs ex- amined: California: Humboldt Co.: stream north of Fish Lake above junction Bluff Greek, Jul April 1981, D, A. Lauck,, 10 nymphs (BYU, HSU, NTSU); same local- ity, 24 April 1987, Baumann, Nelson, Stark and Wells, 11 nymphs (BPS, BYU); stream one-half mile east of Cedar Creek, Hwy 299, above junction Willow Creek, 25 April 1987, Baumann, Nelson, Stark and Wells, 5 nymphs (BPS, BYU); stream one-third mile west of Ruby Creek, Hwy 299, above junc- tion Willow Creek, 25 April 1987, Bau- mann, Nelson, Stark, and Wells, 7 nymphs (BYU, USNM). Nymphs are recorded in Wilkinson (1986) from some additional lo- calities in the Willow Creek drainage. Ho- 829 lotype deposited at the National Museum of Natural History, Smithsonian Institu- tion, Washington, D.C. Collection abbre- viations: BPS, Bill P. Stark, Mississippi Col- lege, Clinton, Mississippi; BYU, Brigham Young University, Provo, Utah; HSU, Humboldt State University, Arcata, Cali- fornia; NTSU, North Texas State Univer- sity, Denton, Texas; USNM, National Mu- seum of Natural History, Smithsonian Institution, Washington, D.C. Etymology.—The generic name Salmo- perla is based on the observation that in- dividuals of this genus seem to occur only in pristine trout streams. Sylvan is the stem for the specific name and it refers to the fact that all the streams that contain Salmoperla sylvanica are located in mesic, heavily wooded areas in the Trinity Alps area of Six Rivers National Forest. Ecological notes.—Salmoperla has only been collected in small tributary streams of larger creeks that soon join large rivers. In every case the stream above the collecting site 1s very steep and then levels out for a relatively short distance before becoming torrenticolous again. The nymphs are found only in loose gravel-rubble areas with a moderate rate of flow. ACKNOWLEDGMENTS We are grateful to Peter Wilkinson for collecting the first adult specimen of Sal- moperla sylvanica and several nymphs, which started us on this exciting study. Spe- cial thanks are given to our colleagues C. Riley Nelson, Bill P. Stark, and Samuel A. Wells, who were members of the collecting team that searched for the elusive Salmo- perla in April 1987. Finally we appreciate the excellent illustrations that were made by Jean A. Stanger, except for the habitus nymph, which was done by the junior au- thor. LITERATURE CITED Needham, J.G. 1933. A stonefly nymph with paired lateral abdominal appendages. J. Entomol. Zool. 25: 17-19. 830 Shepard, W. D. and K. W. Stewart. 1983. Compar- ative study of nymphal gills in North American stonefly (Plecoptera) genera and a new, proposed paradigm of Plecoptera gill evolution. Misc. Publ. Entomol. Soc. Am. 55: 1-57. Stark, B. P. and K. W. Stewart. 1982. Notes on Per- linodes aurea (Plecoptera: Perlodidae). Ann. Ento- mol. Soc. Am. 75: 84-88. Stark, B. P. and S. W. Szczytko. 1984. Egg mor- phology and classification of Perlodinae (Plecop- tera: Perlodidae). Ann. Limnol. 20: 99-104. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Stewart, K. W. and B. P. Stark. 1984. Nymphs of North American Perlodinae genera (Plecoptera: Perlodidae). Great Basin Nat. 44: 373-415. Wilkinson, P. 1986. The spring emergence of Ple- coptera (stoneflies) in the Willow Creek drainage, Humboldt Co., CA, April—July, 1984. Unpub- lished Masters thesis, Humboldt State University, Arcata, California. 100 pp. NOTE Accuracy in reporting types I have recently made a card file for several hundred new holotypes described by about twenty authors during the past few years. The cards record the references to the orig- inal descriptions and the data on the ho- lotypes. Comparing data on the pin labels with the data reported in the original de- scriptions reveals discrepancies in about three per cent of the cases, not just free in- terpretations of pin label data but actual disagreements. When data on the holotypes and those in the publications do not agree there 1s a question whether the specimen labeled holotype is the correct one. Did the author make a careless mistake in copying the data, or did he label the wrong specimen as holotype? If some paratype bears the data reported for the holotype and the specimen labeled holotype does not, is that paratype the true type? When the labeled holotype has data different from the data published, some later investigator could correctly (or incorrectly?) conclude that the true type is lost. This note is to show the need for care when reporting data on types. A common mistake is to report an incorrect date of col- lection. H. Townes, American Entomological In- stitute, 3005 SW 56th Avenue, Gainesville, Florida 32608. PROC. ENTOMOL. SOC. WASH. 89(4), 1987, pp. 831-833 NOTE Two Cynipid Wasp Acorn Galls Preserved in the La Brea Tar Pits (Early Holocene) Published accounts of fossil plant galls have been derived almost exclusively from studies of leaf impressions (Larew. 1986. Proc. Entomol. Soc. Wash. 88: 358-388), and gall structure is rarely preserved. This report describes pieces of two acorns taken from the Rancho La Brea Tar Pits, Los An- geles, California, USA that have been galled and preserved intact. While studying the fossil seeds of the Rancho La Brea Tar Pits, Templeton (Un- published Ph.D. dissertation. 1964. Oregon St. Univ., Corvallis, 224 pp.) identified sev- eral acorns removed from the tar matrix in excavated sabre-toothed cat (Smilodon) skulls. The specimens were excavated be- tween 1913 and 1929. One specimen is de- scribed as a “fossil acorn of Quercus agri- folia showing insect infestation” (ibid., Fig. 34). This acorn (specimen number 1481, Templeton Collection, G. C. Page Museum, Los Angeles, California) was excavated from Pit A at Rancho La Brea. Based on carbon dating of wood found in Pit 3, Templeton estimated the age of the plant remains at La Brea as 13,000-15,000 years (early Holo- cene). The specimen label states “LACM Paleobotany Plesiotype No. PB 1481.” I found another similarly damaged acorn in the Templeton Collection (specimen num- ber 1418B) that was removed from a sabre- toothed cat skull in Pit 3 and is from an undetermined species of oak. Both speci- mens had been cleaned of tar with kerosene and xylol so that external features were ev- ident. To study affinities with extant spec- imens, comparisons were made with galled acorns from the L. H. Weld cynipid collec- tions (California Academy of Sciences, San Francisco, California and the Museum of Natural History, Smithsonian Institution, Washington, D.C.). Specimen 1481 consists of cotyledon tis- sue (1.6 cm long, 1.0 cm wide) (Fig. 1) and detached remnants of the inner surface of the seed coat. The cotyledon tissue is dark brown and woody in texture. An acorn cup is not present. The outer surface of the cot- yledons bears 10 blister-like swellings on average 4.5 mm in diameter. The outer sur- face of many of the swellings has been chewed through or worn away so that an empty internal cavity is exposed (1.5 mm deep). Specimen 1418B (Fig. 2) is smaller (0.9 cm long, 0.6 cm wide) and consists of a dark brown remnant of core cotyledon tissue capped by what is most likely the smooth, cup-shaped remains of cotyledon tissue preserved under the acorn cup. The cup 1s not present. The cotyledons bear two swellings (3 mm in diameter) each of which contains an empty central cavity. Comparison with extant acorn galls caused by several species of cynipids reveals that the La Brea galls resemble those caused by Callirhytis milleri Weld (Fig. 3) which galls acorns of Quercus agrifolia Nee, Q. wisli- zenil A. de Cand., and Q. californica Cooper (= Q. kelloggii Newberry) in California (Weld. 1922. Proc. U.S. Nat. Mus., No. 2440. 61(19): 1-32). Weld (ibid.) described the damage caused by this wasp as a ““com- pact stony-hard mass containing four to a score or more confluent cells (chambers). . . more or less filling the interior of the acorn, which is frequently reduced in size. The woody mass thus occupies the center of the acorn ..., extends its whole length, and when the acorn is cut open can be lifted out intact.” It is this woody mass that has been preserved in the La Brea specimens. The swellings are cells in which the larval and later, pupal cynipid lived (one/cell). Adults emerging from the cells most likely chewed 832 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-3. Acorn galls. Scale line = 5 mm. 1, Specimen 1481; preserved galled acorn of Quercus agrifolia. Opened swellings (arrows) on the surface of disrupted cotyledonous tissue are larval/pupal cells. 2, Specimen 1418B; preserved galled acorn, unknown oak. Larval cell (arrow) showing presumed exit hole is evident. 3, Modern-day acorns of Q. wislizenii galled by Callirhytis milleri. Cut-away view of clumped larval cells (arrows) forming woody mass inside seedcoat. exit holes in the outer cell wall thus exposing the internal cavity as seen in the preserved specimens. One swelling on specimen 1481 shows a very small, circular hole in the outer wall suggestive of a parasite’s exit hole. Emergence of adult C. milleri occurs in March and April (Weld, ibid.). Thus the La Brea galls appear to be of a mature stage and were most likely preserved after ab- scission and after the adult gall formers emerged. Very few Quaternary fossil galls have been described (Larew, ibid.). All available Ho- locene specimens are cynipid-caused oak galls such as leaf galls collected at a pre- Roman Iron Age hill fort in Sussex, England (Cunliffe. 1976. Council for British Archae- ology, Res. Rept. 16, London), oak root galls at Chew Valley Lake, probably from a Roman well, Somerset, England (Eady. 1977. Dept. Environment, Archaeological Reports, London. p. 373), or leaf galls in moss between stones lining a 4th century Roman well in Oxon, England (Robinson. 1980. J. Archaeological Sci. 7: 93-95). Oak galls preserved at Herculaneum by the erup- VOLUME 89, NUMBER 4 tion of Mount Vesuvius (A.D. 79) were most likely items of commerce (Larew. 1987. Econ. Bot. 41: 33-40). The La Brea speci- mens enrich the fossil gall record by virtue of their being the first Holocene galls col- lected outside of Europe, and the first fossil acorn galls ever collected. They provide the first tangible evidence that recognizable galls of seeds were being produced several thou- sand years ago, possibly by an extant wasp species. I thank the staffs of the George C. Page Museum, Los Angeles, the California Acad- emy of Sciences, San Francisco, and the 833 Museum of Natural History, Smithsonian Institution, Washington, D.C. for the loan of specimens. H. K. Phinney, Department of Botany and Plant Pathology, Oregon State University, called my attention to B. C. Templeton’s dissertation. Wayne Brewer and Scott Wing made helpful comments on the manuscript. Hiram G. Larew, Florist and Nursery Crops Laboratory, B-470, Agricultural Re- search Service, USDA, Beltsville, Maryland 20705: PROC. ENTOMOL. SOC. WASH. 89(4), 1987, pp. 834-835 NOTE Urocerus sah (Mocsary) (Hymenoptera: Siricidae) New to North America and Key to North American Species of Urocerus Siricids are commonly dispersed by com- merce. The secretive wood-boring habits of the larvae and several-year life cycle of many species, sometimes not emerging until after wood has been used for construction, com- bine to allow easy access into alien territory. Numerous interceptions have been made by U.S. Plant Protection and Quarantine in wood and wood products coming into the United States. The most recent introduction of a siricid to the United States was Eri- otremex formosanus (Matsumura) (Smith, D. R. 1975. Coop. Econ. Insect Rep. 25(44): 851-854) from southeastern Asia. In a collection of Symphyta loaned to me by D. S. Chandler of the University of New Hampshire, I discovered three specimens of Urocerus sah (Mocsary) collected in that state. This is anew North American record. Urocerus sah occurs in North Africa, Asia Minor to Afghanistan, and southern U:S-S.R. (Smith, D. R. 1978. Hym. Cat., pars 14, 193 pp.); hosts have not been re- corded. In the literature, U. sah is some- times regarded as a subspecies of Urocerus augur (Klug). Urocerus augur is more wide- spread, occurring in most of Europe and Asia Minor; its hosts include Abies spp., Picea spp., and Pinus spp. Benson (1943. Bull. Entomol. Res. 34: 27-51) separated females of the two by the coloration of the abdomen and wings: ‘“‘abdominal tergites 7 entirely and 9 below black; wings rich am- ber in colour with clearly defined infuscate margins” in augur sah, and “abdominal ter- gites 7 with lateral pale band and 9 pale below; wings yellowish-hyaline without clearly defined infuscate margins” in augur augur. Benson did not separate the males. The New Hampshire specimens agree with sah, and, inasmuch as sah has more recently been regarded as a separate species, I am treating it as much. Urocerus sah is separated from other North American species of Urocerus by the following description and key to North American species: Female.— Antenna orange; head mostly orange with interocular area from posterior ocelli to antennae black; sometimes narrow black line at center of postocellar area sep- arating yellow on each side of head. Thorax blackish with pronotum (except lower an- gles), upper 3 mesepisternum, and most of mesonotum yellow orange; mesonotum may be suffused with black anteriorly and on scu- tellum. Abdomen black with terga 1, 2, 8 (except posterior margin), 9 (except ante- riorly and laterally), and cornus yellow. Legs yellow with coxae, trochanters, basal '2 midfemur, apical '2 midtibia, hindfemur, and apical *4 hindtibia black. Wings yellow- ish with apical margins slightly blackish in- fuscated. Male.—Nearly all yellowish, apical ab- dominal segment may be blackish, and legs usually with black as in female. Hindbasi- tarsus 5.0-5.8 x longer than broad. North American records. —_ NEW HAMP- SHIRE: Rockingham Co., New Market, IX- 7-1981, D. Chandler (1 2); Strafford Co.., Durham, VIII-22-1958, W. J. Morse (1 9); Strafford Co., Durham, Aug. 22, 1974, L. J. & M. Milne (1 9). In the collection of the University of New Hampshire and National Museum of Natural History. KEY TO NoRTH AMERICAN SPECIES OF UROCERUS 1. Female — Male 2. Wings black; antennal flagellum partly white VOLUME 89, NUMBER 4 with some basal and/or apical segments HYG pew lhe icuisll pa eat Ssaliriie 7+ ike Att ter ia) 6) mri~w 1 | i ce i} 7 e: , ' no*) bow eehiitey or iaites? | D fe Pparys i uc bain melt) ~~ ' HBT ‘ "T> og witht (RL 1 } d ae) 2 ore Th j a aay Thy bh) ina rt > eew ate ae ie rt © oat oat = ; wi a aot ol fo + ai ee eee ne tp : 7 A ee - ct Ie = 1. ae ogi 4 — So. ets. | “2 (LC + OLS ‘) MOE att le 4 AN Cynipid Galls of the Eastern United States, by Lewis H. Weld Cynipid Galls of the Southwest, by Lewis H. Weld Identification of Alaskan Black Fly Larvae, by Kathryn M. Sommerman Both papers on cynipid galls PUBLICATIONS FOR SALE BY THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MISCELLANEOUS PUBLICATIONS Unusual Scalp Dermatitis in Humans Caused by the Mite Dermatophagoides, by Jay R. Waa Nge ewer or eels A ML ie, pe RL A He RT RT A Short History of the Entomological Society of Washington, by Ashley B. Gumey Pictorial Key to Species of the Genus Anastrepha (Diptera: Tephritidae), by George C. SES Rea nee ie Sy Lah Aa BN Noe NM ee ee RE Me RN ee EE RE BN Bg ea al ia Taxonomic Studies on Fruit Flies of the Genus Urophora (Diptera: Tephritidae), by George C. BS LEN Scale eea wh Pela ir SN GU bah Ol EE BEE he Ae oI ie a EER CL i LOU MU Wee ABE 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. SEND PITY s Waa eee eae Oe BT Ee ANE Ed PB Oe ST Oe SB PES No. 3. The Nearctic Leafhoppers, a Generic Classification and Check List, by Paul Wilson Oman. PAS Ppl ao ese eee Behe eC CLE Ny Bore bb a ee OVA EP Bea ae Be ee No. 4. A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 1952... No.5. A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp. TOS Gem hee wie ke DEE eve eb Pel bE PE Ub Leigh bl ty MLL CCE Biot RS JUL RAR AIS Cg La BA oh le No.6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi MakahasiecoOlppin sage. fret MS ek Od bE EEA eS OI ee Ee eas No.7. Ant Larvae: Review and Synthesis, by George C. Wheeler and Jeanette Wheeler. 108 pp. US aE a 11 eae BSE I See Sa ead Pe Re eA IPs Ta PE to a 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. cee No. 9. The Flower Flies of the West Indies (Diptera: Syrphidae), by F. Christian Thompson. 200 peyote CRSP Se CE TAN Noh ae la DAE LA Red Le SY DNS A OLE Be Thy Tae LL) a 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. OPES! DY EL Ce MOY LCD NO OL OES IAMONS Mink G0 aE OO Dee EN SM Nd DP No. 12. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff. TE Le Sn OD SL$ EN a SARL A) le a ee EEN SA a TO Le 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 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) McCABE, T. L.—Phylogenetic placement of two genera of Hadeninae from southwest Russia (Lepidoptera: Noctuidae) MARSHALL, S. A.—Herniosina voluminosa: a new sphaerocerid of isolated phylogenetic position described from northeastern North America (Diptera: Sphaeroceridae) MILLER, G. L. and R. D. CAVE—Bionomics of Micronus posticus (Walker) (Neuroptera: Hemerobiidae) with descriptions of the immature stages PLATT, A. P.— Banded admirals from western Maryland: analysis of the Limenitis (Basilarchia) arthemis-astyanax (Lepidoptera: Nymphalidae) at Green Ridge State Forest SCHAUFF, M. E. and Z. BOUCEK — Alachua floridensis, a new genus and species of Entedoninae (Hymenoptera: Eulophidae) parasitic on the Florida carpenter ant, Camponotus abdominalis (Formicidae) SCHWAN, T. G. and D. CORWIN—Uropodid mites phoretic on fleas of ground squirrels in California SHOLES, O. D. V.—Host plants and seasonal abundance of adult Capraita subvittata (Coleop- tera: Chrysomelidae: Alticinae) STEINER, W. E. and B. P. SINGH—Redescription of an ergot beetle, Acylomus pugetanus Casey, with immature stages and biology (Coleoptera: Phalacridae) TODD, J. L. and B. A. FOOTE—Resource partitioning in Chloropidae (Diptera) of a freshwater WOOLLEY, J. B. and M. E. SCHAUFF—A new species of Paracrias (Hymenoptera: Eulophidae) parasitic on Anthonomus spp. (Coleoptera: Curculionidae) ZEIGLER, D. D. and K. W. STEWART—Behavioral characters with systematic potential in stoneflies (Plecoptera) NOTES CARROLL, J. F.—Larvae of soldier beetle, Chauliognathus pennsylvanicus (De Geer) (Coleop- tera: Cantharidae): predators of engorged tick larvae and nymphs in the laboratory LAREW, H. G.—Two Cynipid wasp acorngalls preserved in the La Brea Tar Pits (Early Holo- McCAFFERTY, W. P. and R. D. WALTZ—Baetis caelestis Allen and Murvosh, an available name for Baetis sp. A of Morihara and McCafferty (Ephemeroptera: Baetidae) MAIER, C. T.—First Connecticut records of Williamsonia lintneri (Hagen) (Odonata: Cordu- liidae) and Mitoura hesseli Rawson and Ziegler (Lepidoptera: Lycaenidae) SMITH, D. R.— Urocerus sah (Mocsary) (Hymenoptera: Siricidae) new to North America and key to North American species of Urocerus TOWNES, H. V.—Accuracy in reporting types ZACK, R. S.—New records of Brachydeutera (Diptera: Ephydridae) in Malaysia BOOK REVIEWS GERBERG, E. J.— Immature Insects GERSON, U.— Biological Control of Pests by Mites GORDH, G. and R. E. ORTH—IJnsects and Mites: Techniques for Collection and Preserva- OBITUARY JOHNSON, P. 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