oF tye wit Re Tete aie te eRe aE vate Stee Linea sien Hy hi . i ol a a mie | BA o ew) ‘ i { Lay ‘ : My PP Bee: ‘4 in ul ¢ ¥ Ave. id ( iy } ' y i : ayy 1 } 44 | y ‘a xt wy Wi ; } * 7 s ’ , 2 meh) DOME eked Po La Ray bop ally ' s ag rf ve si } : Mie el pon \ , ry: he Age ie eer ere HG | VOL. 104 JANUARY 2002 NO. 1 G EO iK (ISSN 0013-8797) "PROCEEDINGS of the ENTOMOLOGICAL SOCIETY ADAMSKI, DAVID—A new species of Glyphidocera Walsingham (Lepidoptera: Gelechioidea: Gilyphidacerdac) bison CostarRacay ! by. oko Sates ota tee A Raa eee erste ee 119 BRAILOVSKY, HARRY—A new species of Maevius Stal from Australia and some notes on the family akoyocephalidae:(Hemipteras Heteroptera) ete ncilsaoe de dane ee ae eee eben a sioner 41 BRAILOVSKY, HARRY—A new genus and a new species of Daladerini (Hemiptera: Heteroptera. Coreidac) wromiviad ag ascary wae a). pense cet Nee cee Ole eet eres ees ise BOTS: 111 BURKS, ROGER A. and JOHN D. PINTO—Reproductive and electrophoretic comparisons of Trichogramma californicum Nagaraja and Nagarkatti with the 7) minutum complex (Hyme- Hoptera: Uric hOpramiatatidae yy em aah oe te ner creat NONE eect aR tal ct tel ea aa Pr Pa 33 CARROLL, J. F—Notes on the responses of host-seeking nymphs and adults of the ticks Ixodes scapularis and Amblyomma americanum (Acari: Ixodidae) to canine, avian, and deer-produced SUROSTAMGE S357, co alate ais Vara a ree OL ae Raa REE RL SRY So SE CCG eal A Lee RE or) a qe) DELLAPE, P. M., M. pet C. COSCARON, and B. F AMARAL FILHO—Immature stages of Montina confusa (Stal) (Heteroptera: Reduviidae: Harpactorinae) ........................005- 168 DEWALT, R. EDWARD, DONALD W. WEBB, and AMY M. SOLI—The Neoperla clymene (Newman) complex (Plecoptera: Perlidae) in Illinois, new state records, distributions, and an LASN tit CA OMI C Ye 4e02-... lsh! cio J ae nate baie Se marae pte Rae ee neat: CRM Seas ARR an aa am 126 FERREIRA, PAULO SERGIO FIUZA and THOMAS J. HENRY—Descriptions of two new species of Fulvius Stal (Heteroptera: Miridae: Cylapinae) from Brazil, with biological and bioscorraphie notes on thes Senus {kee eres tas acs isle Mopars ese eee ears ee Meenas eee 56 GOEDEN, RICHARD D.—Descriptions of TJephritis footei and T. headricki, new species (Dip- tera: Tephritidae), with notes on their life histories in southern California .................... 142 HALL, JASON P. W.—A review of the new riodinid butterfly genus Panaropsis (Lepidoptera: ING CorOHeRCG Ver Mtsy arava ntzted SVU) AT) Weesamtyde iat smerNey crate PeeKe A UN ents doves era ny en} Can uate ne OS 63 HARRIS, STEVEN C. and OLIVER S. FLINT, JR.—New Alisotrichia (Trichoptera: Hydroptili- dae) from Central and South America and the Greater Antilles ................0. 0... eee eee 195 (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON OFFICERS FOR 2002 GABRIELA CHAVARRIA, President MICHAEL G. PoGug, 7reasurer JONATHAN R. Mawps_Ley, President-Elect RonaLp A. Ocuoa, Program Chair Sruart H. McKamey, Recording Secretary STEVEN W. LINGAFELTER, Membership Chair Hotuis B. WILLIAMS, Corresponding Secretary JOHN W. Brown, Past President Jon A. Lewis, Custodian Davip R. Smitn, Editor . Publications Committee RAYMOND J. GAGNE THOMAS J. HENRY Wayne N. Martuis Honorary President Louise M. RUSSELL Honorary Members KARL V. KROMBEIN RONALD W. HopGEs DoNnALD M. ANDERSON WILLIAM E. BICKLEY All correspondence concerning Society business should be mailed to the appropriate officer at the following address: Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Wash- ington, D.C. 20560-0168. MEETINGS.— Regular meetings of the Society are held in the Natural History Building, Smithsonian Institu- tion, on the first Thursday of each month from October to June, inclusive, at 7:00 P.M. Minutes of meetings are published regularly in the Proceedings. 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This issue was mailed 24 January 2002 Periodicals Postage Paid at Washington, D.C. and additional mailing office. PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PROC. ENTOMOL. SOC. WASH. 104(1), 2002, pp. 1-23 REDESCRIPTION OF PARACOCCUS MARGINATUS WILLIAMS AND GRANARA DE WILLINK (HEMIPTERA: COCCOIDEA: PSEUDOCOCCIDAE), INCLUDING DESCRIPTIONS OF THE IMMATURE STAGES AND ADULT MALE DOUGLASS R. MILLER AND GARY L. MILLER Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart- ment of Agriculture, Bldg. 005, Room 137, BARC-W, Beltsville, MD 20705, U.S.A. (e-mail: dmiller@sel.barc.usda.gov; gmiller@sel.barc.usda.gov) Abstract.—Paracoccus marginatus Williams and Granara de Willink, commonly known as the papaya mealybug was first discovered in the Caribbean in 1994 and was collected in Florida in 1998. To facilitate implementation of control measures we have prepared descriptions, illustrations, and keys for all stages of this species including: First instar, second-instar male and female, third-instar male (prepupa) and female, fourth-instar male (pupa) and female (adult), and fifth-instar male (adult). Comparisons are given between P. marginatus and other species of Paracoccus as well as other commonly encountered mealybug species in Florida and the Caribbean. Key Words: mealybugs, Coccoidea, Pseudococcidae, ontogeny, pest, papaya, Carica pa- paya, hibiscus, invasive species, pest, Florida, Caribbean The genus Paracoccus includes some 79 species of varied distribution from _ the ‘*‘Austro-Oriental, Ethiopian, Madagasian, Nearctic, Neotropical, New Zealand, Pacif- ic, Palaearctic, and Oriental regions”’ (Ben- Dov 1994). Although most assigned species have not been recognized as major econom- ic pests, there are two notable exceptions. Paracoccus burnerae (Brain) is considered a serious pest of citrus in South Africa (Hattingh 1993) and Paracoccus margina- tus Williams and Granara de Willink has recently received attention as a pest of pa- paya and other economically important crops in the Caribbean and Florida. Paracoccus marginatus or the papaya mealybug was originally reported from the Neotropical Region in Belize, Costa Rica, Guatemala, and Mexico (Williams and Granara de Willink 1992) and is believed to be native to at least part of Mexico and/ or Central America (Miller et al. 1999). In 1993 or 1994, it was apparently introduced into the Caribbean Islands where it spread rapidly and is considered a pest of papaya in some areas (Miller et al. 1999). It has subsequently been reported from St. Martin (Matile-Ferrero and Etienne 1996) and Guadeloupe and St-Barthelemy Islands (Matile-Ferrero and Etienne 1998). Other Caribbean distribution records include: An- tigua (Watson and Chandler 1999); Baha- mas, San Salvador Island and New Provi- dence (Watson, personal communication), Paradise Island (Meyerdirk, personal com- munication); British Virgin Islands (Watson and Chandler 1999); Cuba (Meyerdirk, per- sonal communication); Dominican Repub- lic (Meyerdirk, personal communication); Haiti (Meyerdirk, personal communica- tion); Montserrat (Watson and Chandler 1999): Nevis (Watson and Chandler 1999); >~ROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 2 Table | lo eferences for Paracoccus marginatus. #9 Reference Acacia Miller et al. (1999) Acalypha s} Miller et al. (1999) Acalypha wilkesiana Muell.-Arg. Ambrosia cumanensis auct. non Kunth Annona squamosa L. Bauhinia sp. Carica papaya L. Carica sp. Cestrum nocturnum L. Citrux x paradisi Macfad. (pro sp.) Clerodendrum paniculatum L. Coccoloba sp. Fistulosa sp. Guazuma ulmifolia Lam. Hamelia patens Jacq. Hamelia sp. Hibiscus sp. Hibiscus rosa-sinensis L. Ipomoea carnea Jacq. Ipomoea sp. Jatropha integerrima Jacq. Jatropha sp. Malvaviscus penduliflorus DC. Hamon (personal communication) Williams and Granara de Willink (1992) Miller et al. (1999) Hamon (personal communication) Hamon (personal communication) Williams and Granara de Willink (1992) Hamon (personal communication) Hamon (personal communication) Hamon (personal communication) Miller et al. (1999) Hamon (personal communication) Miller et al. (1999) Hamon (personal communication) Hamon (personal communication) Hamon (personal communication) Hamon (personal communication) Hamon (personal communication) Miller et al. (1999) Hamon (personal communication) Hamon (personal communication) Hamon (personal communication) Manihot chlorosticta Standl. & Goldman Manihot esculenta Crantz Mimosa pigra L. Williams and Granara de Willink (1992), Miller et al. (1999) Williams and Granara de Willink (1992), Miller et al. (1999) Williams and Granara de Willink (1992) Williams and Granara de Willink (1992), Miller et al. (1999) Parthenium hysterophorus L. Persea americana P. Mill. Plumeria rubra L. Plumeria sp. Rhaphiolepis umbellata (Thunb.) Makino Sida sp. Solanum melongena L. Uniola paniculata L. Zea mays L. Miller et al. (1999) Hamon (personal communication) Hamon (personal communication) Hamon (personal communication) Williams and Granara de Willink (1992), Miller et al. (1999) Miller et al. (1999) Hamon (personal communication) Miller et al. (1999) Puerto Rico (Meyerdirk, personal commu- nication); St. Barthelemy (Meyerdirk, per- sonal communication); St. Kitts (Watson and Chandler 1999); and U.S. Virgin Is- lands (Watson and Chandler 1999). Addi- tionally, P. marginatus has recently been reported from French Guiana, South Amer- ica (Matile-Ferrero et al. 2000). Paracoccus marginatus was first discovered in the Unit- ed States in Manatee and Palm Beach coun- ties, Florida in 1998 and has since been col- lected in Alachua, Brevard, Broward, Col- lier, Dade, Hillsborough, Manatee, Martin, Monroe, Palm Beach, Pinellas, Polk, Sara- sota, and Volusia counties (Hamon, person- al communication). Since its introduction into Florida, it has been collected 80 times in 30 different cities on 18 species of hosts. The species has apparently spread rapidly in the state. Paracoccus marginatus has been report- ed from more than 25 genera of host plants including economically important crops such as papaya, citrus, yams, cassava, and hibiscus. Papaya mealybug hosts along with associated references are included in Table 1. Although this mealybug has also been recorded from many other hosts, these have yet to be confirmed and are not included in Wable ws VOLUME 104, NUMBER 1 Williams and Granara de Willink (1992) published the first taxonomic paper on Par- acoccus marginatus and included a descrip- tion and illustration of the adult female. They also provided descriptions and a key for the 21 species of Paracoccus occurring in Central and South America. The purpose of this research is to describe and illustrate all instars (including the adult male and fe- male) of this potentially serious invasive species in order to facilitate its recognition as part of any control initiatives that may be undertaken. METHODS Terminology in the descriptions follows that of Williams and Granara de Willink (1992) and Gimpel and Miller (1996) for adult females and immatures and that of Af- ifi (1968) for adult males. Measurements were made with an ocular micrometer using a Leica DMRB compound microscope. Numbers and measurements are from 10 specimens and are given as an average fol- lowed by the range in parentheses. Asso- ciated enlargements of various structures on the illustrations are not proportional. All specimens studied are from the National Museum of Natural History, Coccoidea Collection, Beltsville, MD. Information list- ed in the SPECIMENS EXAMINED section is verbatim from information recorded on the microscope slides. The abbreviation ‘‘ad.”’ refers to adult specimens. Tables 3—6 contain 12 of the most com- mon mealybugs in the Caribbean region and were selected for comparison to P. marginatus. Included species are: Dysmi- coccus brevipes (Cockerell); Ferrisia vir- gata (Cockerell); Maconellicoccus hirsutus (Green); Nipaecoccus nipae (Maskell); Phenacoccus gossypii Townsend and Cock- erell; Phenacoccus madeirensis Green; Phenacoccus solenopsis Tinsley; Planococ- cus citri (Risso); Planococcus minor (Mas- kell); Pseudococcus jackbeardsleyi Gimpel and Miller; Pseudococcus longispinus (TYar- gioni Tozzetti); and Pseudococcus viburni (Signoret). Phenacoccus gossypii and P. madeirensis are listed together as are Plan- ococcus citri and P. minor because of mor- phological similarities. There are other spe- cies of Phenacoccus, Pseudococcus, and Nipaecoccus that could be included in this list, but they are also so similar in general appearance to other species in the genus that characters of one species in the genus are sufficient to distinguish all species in the immature instars. Host plant names included in Table 1 were verified in Integrated Taxonomic In- formation System (Anonymous 2001). RESULTS Paracoccus marginatus Williams and Granara de Willink Suggested Common Name: Papaya mealybug (Figs. 1-8) KEY TO INSTARS ile Without wings or wing buds on thorax ... 4 - With wings or wing buds on thorax ..... 2 2(1). Genitalia weakly sclerotized, aedeagus ab- sent; wings represented by buds less than 2 length of body, without a vein ......... 3 ~ Genitalia heavily sclerotized, aedeagus ap- parent; wings approximately as long as body, with small basal vein (Fig. 8) ..... Os PRES el! Pie Od at S| Ac. Oe ERR ME Se OREM C adult male . Antenna unsegmented; wing buds small, less than length of thorax; head without sclerotization (Fig. 6) prepupal male - Antenna with 10 segments; wing buds large, about same length as thorax; head with weak sclerotization (Fig. 7) ..... pupal male . Antennae 6-segmented Antennae with 7 or 8 segments . Without tubular ducts or with less than 5 SUCHIGUGISE Sin ces o = ces = Sr etecre.s 6 With more than 5 tubular ducts particularly onidorsalfabdomens(Bi1e7s5)) tan tee second-instar male Third antennal segment with more than 4 setae; anal-lobe cerarius with at least 1 aux- iliary seta - Third antennal segment with 4 setae; anal- lobe cerarius without auxiliary setae (Fig. A) ccoreg cy. s Venchep acy Pye Rags Gycatebye Paes first instar . Third antennal segment with 5 setae; tibia divided by tarsus 0.9(0.8—1.0) (Fig. 3) easter suse ee Rees eee ee second-instar female - Third antennal segment with 9 setae; tibia ey) NM ay 6(5). 4 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON divided by ta 1.2-1.4) (Fig. 2) ... Te third-instar female (in part) 8(4). Antenna nted; with less than 5 mul- tilocular pores; without vulva (Fig. 2) . third-instar female (in part) Antenna 8-segmented; with more than 5 multilocular pores; with vulva (Fig. 1) ... adult female Type data.—We have not examined type material of this species, but have studied many of the specimens that were used in the original description but were not in- cluded in the type series. Etymology.—The species epithet is de- rived from a Latin word meaning enclosed within a border and refers to the border of oral rim tubular ducts (Williams and Gran- ara de Willink 1992). ADULT FEMALE (Fig. 1) Field features.—Body yellow, dusted with mealy wax not thick enough to hide body color, without discrete bare areas on dorsum, with many short waxy filaments around body margin. Ovisac developed be- neath and behind adult female. Slide-mounted characters—Body 2.2(1.5— 2.7) mm long, 1.4(0.9—1.7) mm wide. Dorsum with 16(14—17) pairs of cerarii; ceramml. 2.4.5. 7.-and) 9 wath 2 conical setae (Fig. 1M); cerarii 3, 6, and 16 with 3(2—3) conical setae; cerarii 8, 11, and 17 with 2(0—2); cerarii 10 and 14 with 1(0—2) conical setae; cerarii 12, 13, and 15 with 2(0—3) conical setae. Cerarius 12 without auxiliary setae, with 2(0—3) conical setae, 5(0—8) trilocular pores, 1(0—3) discoidal pores. Anal-lobe cerarius (Fig. 11) with I(1—3) auxiliary setae (Fig. 1J), 2 conical setae, 13(10—18) trilocular pores Gigs i€); 2(0—3) discoidal pores (Fig. 1K). Dorsal body setae (Fig. 1L) more slender than cer- arian setae. Multilocular pores absent; tri- locular pores scattered over surface, most abundant near setae; discoidal pores rare, about 2 diameter of trilocular pore. Oral- rim tubular ducts (Fig. 1B) usually restrict- ed to marginal areas associated with cerarii, | specimen examined with | mediolateral duct on segment I and | in medial area of mesothorax; of 21 specimens examined cer- arius | without associated oral rim, cerarius 2 with associated oral rim in 20 of 21 spec- imens, cerarius 3 with 4 on 21 specimens, cerarius 4 with 12 on 21, cerarius 5 with 11 on 21, cerarius 6 with 17 on 21, cerarius 7 with 18 on 21, cerarius 8 with 20 on 21, cerarii 9 and 10 without associated oral rims, cerarius 11 with 15 on 21, cerarius 12 with 4 on 21, cerarii 13, 15, and 16 without associated oral rim, cerarius 14 with 2 on 21, and cerarius 17 with 15 oral rims on 21 specimens. Oral-collar tubular ducts absent. Longest submedial seta on segment VII 10(8—14) p long; 1(0—2) submedial setae on segment VIII, when present longest seta 13(8—18) p long. Anal-ring seta. 136(120—150) wp long; 1.4(1.2—1.7) times as long as width of anal ring. Venter with multilocular pores (Fig. 1G) usually in posterior and anterior bands on segments VI—-VIII and restricted to poste- rior band on segments IV and V, | or 2 specimens with | or 2 pores on segment III or with a few pores on anterior margins of segments IV and V; 3 of 10 specimens with 1 multilocular pore near base of front or hind leg. Trilocular pores concentrated near setal bases. Discoidal pores uncommon, of same size as on dorsum. Oral-rim tubular ducts in mediolateral areas from prothorax to segment I, with 4(3—6) ducts on each side of body. Oral-collar tubular ducts of 1 size, IN Conspicuous marginal clusters along body margin from segments II-VI, often with 2 or 3 pores on segment I, also present in medial and mediolateral areas of abdom- inal segments HII—VIII, present on thorax in seta clusters near hind 2 pairs of legs, oc- casionally with | or 2 along body margin of thorax especially in area laterad of an- terior spiracle and front legs, absent from head. Setae as follows: 4 cisanal (Fig. 1H), longest 52(45—68) pw long; longest anal-lobe seta 170(155—200) w long; longest seta on trochanter 104(95—110) p long. Anal-lobe VOLUME 104, NUMBER | Fig. 1. Adult female of Paracoccus marginatus. A, Detail of front leg. B, Oral-rim tubular duct. C, Trilocular pore. D, Translucent pores. E, Oral-collar tubular duct. E Detail of hind leg. G, Multilocular pore. H, Cisanal seta. I, Anal-lobe cerarius. J, Auxiliary seta. K, Discoidal pore. L, Dorsal body seta. M, Cerarian seta. 6 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON bar conspicuoi ler than base of anal bar set Circulu 80) wp wide, generally di- vided b segmental line. Labium 137(125—i162) » long. Antenna 8-segment- ed, 372(335—400) pw long. Legs with trans- lucent pores restricted to hind coxa, ventral surface (when leg is lying flat as shown in illustration) with 40(14—62) pores, dorsal surface (Fig. 1D) with 79(54—108) pores. Hind femur 209(195—225) w long; hind tib- ia 211(185—228) pw long; hind tarsus 94(91— 100) » long. Hind tibia/tarsus 2.2(2.0—2.3); hind femur/tarsus 1.0(0.9—1.1). Length of hind femur divided by greatest width of hind femur 3.7(3.0—4.2). Hind tibia with 15(14—19) setae. Claw digitules on all legs clubbed, approximately same size. Tarsal digitules (Fig. 1F) on hind 2 pairs of legs clubbed, each tarsus with 1| digitule notice- ably longer and with club slightly larger than other; tarsal digitules (Fig. 1A) on front pair of legs of 2 different sizes and shapes, | digitule on each tarsus clubbed and robust, other digitule without club, slender. Notes.—The above description is based on 240 specimens from 41 localities. The adult female can be distinguished from all other instars by having multilocular pores, translucent pores on hind coxa, and a vulva. THIRD-INSTAR FEMALE (Fig. 2) Slide-mounted characters—Body 1.1(0.7— 1.8) mm long, 0.7(0.3—1.1) mm wide. Dorsum with 6(1—10) pairs of cerarii; cerarii indefinite, when present, with 2 con- ical setae and | trilocular pore between con- ical setae. Cerarius 12 absent. Anal-lobe cerarius with 1(1—2) auxiliary setae, 2 con- ical setae, 5(4—7) trilocular pores, 0(O0—1) discoidal pores. Dorsal body setae more slender than setae. Multilocular pores absent; trilocular pores scattered over surface, most abundant near setae; discoidal pores rare, about % diameter of trilocular pore. Oral-rim tubular duct rarely present near position of cerarius 8 (of 10 specimens cerarian examined, 4 had | oral rim or large oral collar on at least one side of body). Oral- collar tubular ducts absent. Longest sub- medial seta on segment VII 7(5—10) p long; 1(0—2) submedial setae on segment VIII, when present longest seta 7(5—9) p long. Anal-ring seta 87(78-92) wp long; 1.6(1.3—1.8) times as long as width of anal ring. Venter with multilocular pores absent (present on | of 10 specimens examined, restricted to segment VIII). Trilocular pores concentrated near setal bases. Discoidal pores uncommon, of same size as on dor- sum. Oral-rim tubular ducts sometimes pre- sent near body margin on abdominal seg- ment II, III, or on metathorax (of 10 spec- imens examined, 4 had 1 or 2 oral rims or large oral collar on at least one side of body). Oral-collar tubular ducts absent. Se- tae as follows: 4 cisanal, longest 33(22—50) w long; longest anal-lobe seta 132(112— 158) pw long; longest seta on trochanter 63(50—-85) ww long. Anal-lobe bar slightly narrower than on adult female. Circulus 47(22—80) w wide, generally di- vided by intersegmental line. Labium 85(70—-100) w long. Antenna 6- or 7-seg- mented (of 10 specimens examined 5 had 6-segments with weak indication of partial division between segments 3 and 4), when 6-segmented, segment 3 with 9 setae, 233(205—262) pw long. Legs without trans- lucent pores. Hind femur 114(100—138) p long; tibia 108(98—-127) w long; tarsus 84(80-90) pw long. Hind tibia/tarsus L2C1.2=14): hind femiur/tarsus 121 G@e0= 1.2). Length of hind femur divided by greatest width of hind femur 2.5(2.0—3.1). Hind tibia with 9(7—10) setae. Claw and tar- sal digitules same as on adult female except sometimes | claw digitule slightly smaller that other. Notes.—The above description is based on 42 specimens from 12 localities. The third-instar female can be distinguished from all other instars by having 6- or 7-seg- mented antennae, when 6-segmented with VOLUME 104, NUMBER 1 Fig. 2. Third-instar female of Paracoccus marginatus. 8 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON hind tibia divided by hind tarsus length usu- ally 1.2. and about 9 setae on the hind tibia. SECOND-INSTAR FEMALE (Fig. 3) Field features.—Body color yellow. Slide-mounted characters ——Body 0.7(0.5— 0.8) mm long, 0.4(0.3—0.5) mm wide. Dorsum with 6(4—11) pairs of cerarii; cerarii indefinite, when present, with 2 con- ical setae and | trilocular pore between con- ical setae. Cerarius 12 absent. Anal-lobe cerarius with | auxiliary setae, 2 conical se- tae, 2(2—3) trilocular pores, sometimes with 1 discoidal pore. Dorsal body setae more slender than cerarian setae. Multilocular pores absent; trilocular pores scattered over surface, most abundant near setae; discoidal pores rare, about % diameter of trilocular pore. Oral-rim tubular ducts absent. Oral- collar tubular ducts absent. Longest sub- medial seta on segment VII 6(5—6) pw long; without submedial setae on segment VIII. Anal-ring seta 60(52-70) w long; 1.5(1.3-1.8) times as long as width of anal ring. Venter with multilocular pores absent. Trilocular pores concentrated near setal ba- ses, absent in medial area of abdomen. Dis- coidal pores rare. Oral-rim tubular ducts ab- sent. Oral-collar tubular ducts absent. Setae as follows: 4 cisanal, longest 20(18—25) long; longest anal-lobe seta 95(88—102) w long; longest seta on trochanter 49(42—55) w long. Anal-lobe bar narrower than on adult female. Circulus 40(20—75) j wide, generally di- vided by intersegmental line. Labium 69(62—75) w long. Antenna 6-segmented, 173(152—185) pw long; antennal segment 3 with 5 setae. Legs without translucent pores. Hind femur 78(72—82) w long; hind tibia 66(58—72) ww long; hind tarsus 69(65— 72) w long. Hind tibia/tarsus 0.9(0.8—1.0); hind femur/tarsus 1.2(1.1—-1.3). Length of hind femur divided by greatest width of hind femur 2.3(2.1—2.5). Hind tibia with 9(8—9) setae. Claw and tarsal digitules same as on adult female except | claw digitule conspicuously smaller on all legs. Notes.—The above description is based on 64 specimens from 9 localities. The sec- ond-instar female can be distinguished from all other instars by lacking oral-collar tu- bular ducts and multilocular pores, and by having 5 setae on the third antennal seg- ment. First INSTAR (GENDER NOT DETERMINED) (Fig. 4) Slide-mounted characters.—Body 0.4(0.3— 0.5) mm long, 0.2(0.2—0.3) mm wide. Dorsum with 9(7—10) pairs of cerarii; cerarii indefinite, when present, with 2 con- ical setae and | trilocular pore between con- ical setae. Cerarius 12 absent. Anal-lobe cerarius without auxiliary setae, 2 conical setae, | trilocular pore, without discoidal pores. Dorsal body setae more slender than cerarian setae. Multilocular pores absent; trilocular pores scattered over surface, forming 2 longitudinal lines on each side of abdomen, excluding cerariian setae. Discoi- dal pores absent. Oral-rim tubular ducts ab- sent. Oral-collar tubular ducts absent. Lon- gest submedial seta on segment VII 5(4—8) w long; without submedial setae on segment VUl. Anal-ring seta 37(30—42) w long; 1.4(1.2—1.6) times as long as width of anal ring. Venter without multilocular pores. Tri- locular pores restricted to 10 positions on each side of body on head, thorax, and an- terior abdomen (see Fig. 4 for position la- bels); triloculars in positions | to 7 and 10 rarely absent, pores in positions 8 and 9 of- ten absent. Discoidal pores in submarginal line on each side of abdomen, usually with | pore posterior of each spiracle. Oral-rim and oral-collar tubular ducts absent. Setae as follows: 4 cisanal, longest 17(12—22) w long; longest anal-lobe seta 58(45—69) w long; longest seta on trochanter 38(35—42) w long. Anal-lobe bar narrower than on adult female. Circulus 37(20—55) wide, generally di- VOLUME 104, NUMBER 1 Fig. 3. Second-instar female of Paracoccus marginatus. 9 10 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON , 1ret—i Pi, aa) > Paes . : : = San = Fig. 4. First-instar of Paracoccus marginatus, gender not determined. Numbers refer to positions of ventral trilocular pores. VOLUME 104, NUMBER 1 vided by intersegmental line. Labium 48(44—-52) ww long. Antenna 6-segmented, 132(120—148) ww long; antennal segment 3 with 4 setae. Legs without translucent pores. Hind femur 59(55—62) pw long; hind tibia 50(45—55) pw long; hind tarsus 57(53— 62) w long. Hind tibia/tarsus 0.9(0.8—0.9); hind femur/tarsus 1.2(1.1—1.3). Length of hind femur divided by greatest width of hind femur 2.3(1.8—2.5). Hind tibia with 9 setae. Claw and tarsal digitules same as on adult female except | claw digitule con- spicuously smaller on all legs. Notes.—The above description is based on 86 specimens from 6 localities. The first- instar can be distinguished from all other instars by having no auxiliary setae in the anal-lobe cerarius, 2 longitudinal lines of trilocular pores on dorsal abdomen on each side of body (excluding cerariian setae), and antennal segment 3 with 4 setae. SECOND-INSTAR MALE (Fig. 5) Field features.—Body color usually pink, occasionally yellow. Slide-mounted characters.—Body 0.6(0.5— 1.0) mm long, 0.3(0.2—0.6) mm wide. Dorsum with 4(2—5) pairs of cerarii; cer- arii indefinite, when present, with 2 conical setae and | trilocular pore between conical setae. Cerarius 12 absent. Anal-lobe cerar- ius with 1(1—2) auxiliary setae, 2 conical setae, 2(2—3) trilocular pores, without dis- coidal pores. Dorsal body setae more slen- der than cerarian setae. With 1(0—2) multi- locular pores in medial areas of thorax and/ or head, present on 6 of 10 specimens ex- amined; trilocular pores scattered over surface, most abundant near setae; discoidal pores rare, about % diameter of trilocular pore. Oral-rim tubular ducts absent. Oral- collar tubular ducts abundant over surface, of 1 size. Longest submedial seta on seg- ment VII 6(5—8) p» long; without submedial setae on segment VIII. Anal-ring seta 54(48—-58) w_ long; 1.3(1.2-1.5) times as long as width of anal ring. 11 Venter with multilocular pores mesad of each pair of legs, for hind pair of legs lo- cated on segment III, with 4(2—5) pores on body. Trilocular pores concentrated near se- tal bases. Discoidal pores rare. Oral-rim tu- bular ducts absent. Oral-collar tubular ducts of 2 sizes: larger size same as on dorsum, located marginally; smaller size present in longitudinal line along submargin of abdo- men. Setae as follows: 4 cisanal, longest 21(17—26) pw long; longest anal-lobe seta 91(83-108) ww long; longest seta on tro- chanter 47(40—55) pw long. Anal-lobe bar narrower than on adult female. Circulus 50(40—75) w wide, generally di- vided by intersegmental line. Labium 67(62—72) w long. Antenna 6-segmented, 171(160—188) ww long; antennal segment 3 with 5 setae. Legs without translucent pores. Hind femur 82(75—92) long; hind tibia 68(61—80) pw long; hind tarsus 62(52— 70) w long. Hind tibia/tarsus 1.1(1.0—1.2); hind femur/tarsus 1.2(1.1—1.4). Length of hind femur divided by greatest width of hind femur 2.6(2.2—3.0). Hind tibia with 9(8—9) setae. Claw and tarsal digitules same as on adult female except | claw digitule conspicuously smaller on all legs. Notes.—The above description is based on 86 specimens from 7 localities. The sec- ond-instar male can be distinguished from all other instars by having dorsal oral-collar tubular ducts and multilocular pores near | or more pairs of legs. In addition, live and alcohol preserved specimens tend to appear more narrowly elongate than other associ- ated second-instar females, third-instar fe- males, or first instars. Both field collected and laboratory reared early instars of P. marginatus exhibit pink and yellow forms. We hypothesize that the pink forms are males. Pink forms were col- lected in the field on St. Kitts, W.I. In ad- dition, Richard Warkentin (USDA, APHIS- PPQ, Biological Control Technical Special- ist) collected and sorted a vial of pink spec- imens and a vial of yellow ones from the Papaya mealybug rearing facilities in St. Thomas, U.S.V.I. However, the results from PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON leis 2) Second-instar male of Paracoccus marginatus. VOLUME 104, NUMBER 1 both collection sites were inconclusive. Al- though field collected pink forms were mostly second instar males, there were a few second instar females present. Results of the laboratory reared specimens were also inconclusive. While specimens of the pink form yielded only second instar males, specimens of the yellow form yielded both second instar males and second instar fe- males although there were many more fe- males than males. Contamination, especial- ly for field collected specimens may be a possibility for the discrepancies but, this is probably unlikely for the specimens col- lected at the rearing facilities. THIRD-INSTAR MALE (PREPUPA) (Fig. 6) Slide-mounted characters.—Body 0.9(0.8— 1.1) mm long, 0.4(0.3—0.4) mm wide. Dorsum without cerarii; posterolateral margins of segments V, or VI, VII, and VIII each with 2 setae conspicuously longer than remaining setae on abdominal segments. Multilocular pores in medial areas of head, forming row on prothorax and metathorax, usually without pores on mesothorax, oc- casionally with | or 2 medially, in rows on most abdominal segments, fewer in medial area, absent from segments VIII and IX; tri- locular pores absent; discoidal pores rare. Oral-rim tubular ducts absent. Oral-collar tubular ducts present around body margin, medial and submedial ducts sometimes pre- sent on prothorax, metathorax and 1 or 2 abdominal segments. Longest submedial seta on segment VII 18(15—20) wp long; without submedial setae on segment VIII. Anal-ring setae absent; anal ring 25(20— 28) ww wide. Venter with multilocular pores near an- terior margin on head, near spiracles, legs, and in medial areas of pro- and mesothorax on thorax, in rows on abdominal segments, sparse medially, absent from segments VIII and IX. Trilocular pores absent. Discoidal pore located near each pair of legs. Oral- rim tubular ducts absent. Oral-collar tubular 13 ducts restricted to margin. Longest anal- lobe seta 67(50—78) wp long. Circulus appearing collapsed, 62(45—98) ut wide, resting on intersegmental line. La- bium absent. Antennal segments indistinct, 226(215—248) pw long. Hind femur 90(82— 95) » long; division between hind tibia and tarsus indistinct, hind tibia + tarsus 134(130—140) p long. Length of hind fe- mur divided by greatest width of hind fe- mur 2.7(2.2—3.0). Wing buds of mesothorax protruding from lateral margin, 67(50—78) w long. Hamulohalterae represented by small swelling on lateral margin of meta- thorax. Notes.—The above description is based on 16 specimens from 3 localities. The pre- pupa can be distinguished from all other in- stars by having multilocular pores, oral-col- lar tubular ducts, antennae without definite segmentation, tibia + tarsus fused, no la- bium, no aedeagus, and no definite constric- tion for the head. FOURTH-INSTAR MALE (PUPA) (Fig. 7) Slide-mounted characters.—Body 1.0(0.9— 1.0) mm long, 0.3(0.3—0.4) mm wide. Dorsum without cerarii; posterolateral margins of segments III, IV, or V, to seg- ment VIII each with 2 setae conspicuously longer than remaining setae on abdominal segments. Multilocular pores absent from head, forming conspicuous row on protho- rax, mediolateral cluster on metathorax, without pores on mesothorax, in mediolat- eral clusters on each side of abdominal seg- ments I-VI or VII; trilocular pores absent; discoidal pores associated with multilocu- lars and oral collars. Oral-rim tubular ducts absent. Oral-collar tubular ducts present near body margin of prothorax and abdom- inal segments I or II to VII or VII, forming clusters of 2(1—5) ducts. Longest submedial seta on segment VII 20(16—28) wp long; without submedial setae on segment VIII. Anal-ring setae absent; anal ring 27(25— 30) » wide. Venter with multilocular pores absent PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Third-instar male or prepupa of Paracoccus marginatus. VOLUME 104, NUMBER 1 Fig. 7. Fourth-instar male or pupa of Paracoccus marginatus. 16 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON from head. row of pores between front cox- mainder of thorax near spi- ae, present on rr racles and legs, in mediolateral clusters of 2(1—4) pores on each side of segments H— VI. VII, or VIL. Trilocular pores absent. Discoidal pores associated with oral-collars and multiloculars. Oral-rim tubular ducts absent. Oral-collar tubular ducts present near body margin of prothorax usually forming cluster of several ducts, sometimes absent from abdomen or with | duct near body margin on each of abdominal seg- ments II-VII or VIII. Longest anal-lobe seta 60(48—72) pw long. Circulus ill-defined. Labium absent. An- tenna 10-segmented, 357(345—375) w long. Hind femur 109(105—115) p long; hind tib- ia 111(105—112) p long; hind tarsus 80(75— 88) » long. Hind tibia/tarsus 1.4(1.3—1.5); hind femur/tarsus 1.0. Length of hind femur divided by greatest width of hind femur 3.3(3.2—3.4). Wing buds of mesothorax pro- truding from lateral margin, 336(250—385) w long. Hamulohalterae 36(22—42) w long. Notes.—The above description is based on 8 specimens from | locality. The pupa can be distinguished from all other instars by having multilocular pores, oral-collar tu- bular ducts, 10 segmented antennae, no la- bium, no aedeagus, and a slight constriction between the thorax and head. ADULT MALE (Fig. 8) Slide-mounted characters.—Body elon- gate oval, 1.0(0.9—1.1) mm long; greatest width at thorax 0.3(0.2—0.3) mm. Dorsum with | pair of tail-forming pore clusters; each cluster with 2 elongate setae approximately 250 wp long, | sometimes 2 additional shorter setae, 38(34—42) multi- locular pores, and | or 2 discoidal pores. Multilocular pores in marginal areas of pro- thorax and each abdominal segment, with 5(3-7) on each side of segment I, 2(1—3) on each side of segment II, 2(1—2) on III, 2(1—3) on IV, 2(1—3) on V, 2(1-3) on VI, 1(1—2) on each side of segment VII, mul- tilocular pores with 4 or 5 loculi, quinquel- oculars predominate (Fig. 8G); normally without pores on head (1 of 10 specimens with | pore near lateral arm of midcranial ridge). Discoidal pores (Fig. 8F) associated with lateral abdominal multilocular pores. Body setae bristle shaped. Small abdominal tergites present on mid-dorsum of segments I-III and dorsum of segment VIII. Dorsal abdominal tergites usually without associ- ated setae. Metapostnotal ridge conspicu- ous. Scutellum rectangular, with several medial setae. Scutum sclerotized through- out except for a median longitudinal clear area which bears several setae. Prescutum rectangular with well defined prescutal ridge, weakly defined prescutal suture and several setae. Pronotal ridge heavily scler- otized. Hamulohalterae 75(67—82) w long, with 1 apical hooked seta. Mesothoracic wings 932(889—988) p long, each with 2—3 basal setae. Head width 180(148—193) p; dorsal eye 34(30—40) p in diameter, lateral ocellus 17(12—20) wp in diameter and locat- ed at junction of preocular and postocular ridges. Dorsal arm of midcranial ridge (Fig. 8A) extending beyond posterior margin of dorsal eye. Median crest weakly sclerotized with several setae. Ocular sclerite weakly sclerotized. Penial sheath (Fig. 8C and 8E) 95(86— 106) pw long, 70(62—74) pw wide with distinct ventral lobes; length/width ratio 1.4. Ae- deagus (Fig. 8D) 68(54—79) w long, broad and apically truncate. Venter with hair-like setae only, present medially, submedially and laterally of most abdominal segments as well as few scat- tered prosternal and basisternal setae. Ab- dominal sclerotization confined to segment VIII. Prosternal ridge well developed, ster- nite weakly sclerotized. Preoral ridge weak- ly developed. Ocular sclerite weakly scler- otized near ventral eye. Ventral midcranial ridge well developed, with lateral arms. Ventral eye 39(32—44) w in diameter. Antenna 10-segmented with bristle- shaped and fleshy setae, capitate setae pre- sent on apical segment; segments I 37(30— 40) w long (Fig. 8H); I 55(49—-62) pw long VOLUME 104, NUMBER 1 17 Fig. 8. Adult male of Paracoccus marginatus. A, Frontal view of midcranial ridge. B, Detail of front leg. C, Ventral view of penial sheath. D, Aedeagus. E, Lateral view of penial sheath. F Discoidal pore. G, Quin- quelocular pore. H, Detail of scape and pedicel. I, Detail of apical segment. 18 ROCEREDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 2 omparisons of New World Paracoccus. Bold indicates characters that differ from Paraco fare fee Marginal Cluster Oral Collars Number of Near Anterior Pores on Par Circulus Cerarii Oral Rim Distribution Spiracle Hind Tibiae marginatus yes 16-17 margin only no no alazanensis yes 17 margin and abdomen no yes ascius yes IL7/ scattered yes yes baccharidicola yes a scattered yes yes circuliprivis no 7 margin and thorax no yes decorus no 7-9 throughout except head no yes ferrisi variable 12-17 usually margin and abdomen yes yes hamoni yes 12 scattered no yes herreni yes 16 scattered yes yes juniperi yes 5-17 scattered yes yes lycopersici yes 4-7 scattered yes yes mexicanus yes 15-17 scattered no yes myrtacearum no 14 scattered no yes oneratus yes 5 many, scattered no yes ordinis yes 15-16 scattered yes yes reductus yes 0 absent no yes salviacola yes 16 scattered yes yes solani variable 6-7 scattered no yes townsendi yes 13 absent yes no turrialbensis no wT abdomen no yes villanuevai yes 7 scattered yes no (Fig. 8H); III 68(62—74) wp long; VI 52(45— 57) w long; VII 53(47-59) pw long; VIII 53(47—-59) w long; IX 46(37—49) pw long; X 46(49-69) w long (Fig. 81); total length 534(469-563) pw long. Hind femur 145(133—148) p long; hind tibia 199(163- 222) » long; hind tarsus 83 (69-86) yp long; hind tarsal claw 27(22—32) w long; hind fe- mur/tibia ratio 0.7; hind tibia/tarsus ratio 2.4(2.3—2.5); leg setae bristle shaped. Tarsal digitules capitate; claw digitules acute (Fig. 8B). Notes.—The above description is based on 64 specimens from 8 localities. The adult male can be distinguished from all other instars by having a distinct aedeagus, lateral pore clusters, a heavily sclerotized thorax and head, and by having well-de- veloped wings. SPECIMENS EXAMINED UNITED STATES: Florida: Broward County, Ft. Lauderdale, on Hibiscus sp., 5-XI-1998, by W. Thiel. Manatee County, Bradenton, on H. rosa-sinensis, 24-VII- 1998, by W. Clifton (2 ad. 2). Palm Beach County, Boca Raton, on Hibiscus rosa-si- nensis, 1998; by J. Lofquist (l\;adeeg): BRITISH VIRGIN ISLANDS: Great Ca- manoe Island, on /pomoea sp., 16-X-1996, by R. E Denno (7 ad. 2°). DOMINICAN REPUBLIC: Locality unknown, on Persea americana, 19-VII-1994, by J. Sanchez (1 ad. 2). MEXICO: Baja California—La Paz, on Carica papaya, 13-1X-1978, by G. Bux- ton (16 ad. 2, 3 third-instar 2, 3 second- instar d). Colima—Cofradia de Juarez, on Carica papaya, 8-V1-1999, by H. Gonza- lez, J. Villaneuva,; D; «R. Miller (iadare): Crucero de Perquillos, on Carica papaya and Manihot esculenta, 8-V1-1999, by H. Gonzalez, J. Villaneuva, D. R. Miller (3 ad. 2); Rinxon de Lopez near Ameria, on Car- ica papaya, 8-V1-1999, by H. Gonzalez, J. Villaneuva, D. R. Miller (1 ad. 2); near Te- coman, Boca de Pazcuales, on Carica pa- VOLUME 104, Morphological comparisons of selected first instars, second-instar females, third-instar females, and adult females of Paracoccus marginatus and other economically important mealybugs. Bold indicates characters that differ from Paracoccus marginatus. Table 3. Adult Female Second-Instar Female Third-Instar Female First Instar NUMBER 1 # Cerarli Oral Rims Anal Bar # Cerarii Oral Rims Anal Bar # Cerarii Oral Rims Anal Bar # Cerarli Anal Bar yes no yes yes no 17 17 yes no no no no yes yes no 7-10 yes 4-1] no yes 17 no / 17 yes no Paracoccus marginatus no yes yes no 17 no yes Dysmicoccus brevipes no no Ferrisia virgata 4-7 10+ 18 18 yes no 3-4 10+ 18 18 yes yes yes yes 34 1-2 17 15-18 15-18 yes yes yes Maconellicoccus hirsutus Nipaecoccus nipae no no yes yes no no no no 15-18 15-18 es yes yes no yes yes no s solenopis zacoccus gossypii/madeirensis 1acoccu ~ oS Phe Phe rare yes yes yes 18 17 17 iN7/ yes no no 18 17 17 17 yes no no 18 17 17 17 18 17 17 17 citri/minor yes yes yes no yes no is jackbeardsleyi yes no yes no yes no yes is longispinus no ts viburni y~ re 2 = ococcl = Plar dococc = Psei dococc = Psei = 19 paya, 8-VI-1999, by H. Gonzalez, J. Vil- laneuva, D. R. Miller (2 ad. @, 1 third-in- star 6); Laguna de Chanchopan, on Carica papaya, 10-II-2000, by H. Gonzalez, J. Vil- laneuva, M. E. Schauff, D. R. Miller (8 first instar); Puerta de Caberar, on Carica pa- paya, 7-l-2000, by H. Gonzalez, J. Villa- neuva, M. E. Schauff, D. R. Miller (1 ad. 2, 4 first instar); Plan de Zapotes, on Car- ica papaya, 10-II-2000, H. Gonzalez, J. Villaneuva, M. E. Schauff, D. R. Miller (4 ad. 2, 2 ad. 6, 3 third-instar 2, 2 second- instar 6); Colima City, 7-H-2000, on Car- ica papaya, H. Gonzalez, J. Villaneuva, M. E. Schauff, D. R. Miller (2 ad. 9, 1 third- instar 2, | second-instar 2, 1 second-instar 3d). Guerrero—Acapulco Airport, on Sida sp., 15-IV-1984, by J. Gillet and H. Miran- da (1 ad. 2); El Carrizal, on Mimosa pigra, 7-VII-1986, by J. Gillett (1 ad. @). Jalis- co—San Marcos, on Carica papaya, 6-II- 2000, by H. Gonzalez, J. Villaneuva, M. E. Schauff, D. R. Miller (1 ad. 2). Michoa- can—near Nueva Italia, on Carica papaya, 5-II-2000, by H. Gonzalez, J. Villaneuva, M._ Es; Schaufiz) Ds R: (Miller (2iads"o5 5 third-instar 2°, 4 second-instar ¢); Santa Casilda, on Carica papaya, 9-VI-1999, by H. Gonzalez, J. Villaneuva, D. R. Miller (20 ad. 2, 3 ad. 6, 2 third-instar 2, 1 third- instar ¢, 3 second-instar 2, 15 first instar). Tobasco—Ejido Lopez Portillo, on Hibis- cus sp., 4-VI-1999, by H. Gonzalez, D. R. Miller (1 ad. 2); Rancherias Barrancar y Amate, on Carica papaya and Manihot es- culenta, 4-VI-1999, H. Gonzalez, D. R. Miller (2 ad. 2); Villahermosa, on Mimosa pigra, 18-II-1985, by J. Gillet (11 ad. &); Villahermosa, on Hibiscus sp., 4-V1-1999, by H. Gonzdlez, D. R. Miller (1 ad. @). Vera Cruz—Campo Cotaxtla, on Carica papaya, 2-V1-1999, by H. Gonzalez, J. Vil- laneuva, D. R. Miller (4 ad. 2); Curva del Pato Santa Fe, on Hibiscus sp. and Manihot esculenta, 2-V1-1999, by H. Gonzalez, J. Villaneuva, D. R. Miller (8 ad. 2); El Man- gal, on Carica papaya, 2-VI-1999, by H. Gonzalez, J. Villaneuva, D. R. Miller (7 ad. 2); Huexotla, on Carica papaya, 2-VI- 20 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1999, by H Villaneuva, D. R. Miller a ). Manilo Fabio Altamirano, on Hibis VI-1999, by H. Gonza- lez. J. Villanei D:175 CAYK Yakima, WA >55 T. minutum MCVA Chula Vista, San Diego Co., CA >675 T. platneri PRV1I Riverside (UCR Campus), CA >475 T. exiguum EXSL Selma, AL >500 EXHN. Hendersonville, NC >60 a platneri in the west. Although reproductive crosses of T. californicum showed complete incompatibility with several similar species including 7. platneri (Pinto 1999), these re- sults were based on single crosses, each in- volving no more than 20 pairs in each di- rection. This paper investigates the distinctness of T. californicum and members of the 7. min- utum complex with further reproductive compatibility tests and allozymic electro- phoresis. A large number of crosses were performed between three lines of 7. cali- fornicum and T. platneri to determine if cases of morphological intermediacy could be explained by a degree of reproductive compatibility. We also include the eastern North American 7. exiguum Pinto and Plat- ner in this study because of its similarity to the 7. minutum complex and, in particular, to T. californicum (Pinto 1999). Tricho- gramma exiguum also is known from cod- ling moth and other fruit tree tortricid pests. It has frequently been taken on these hosts at localities that also harbor T. minutum (Pinto et al., in prep.). The interspecific studies presented here are similar to those performed between members of the T. min- utum complex (Pinto et al. 1991, 1992), and on the closely related species pairs 7. deion/ T. pretiosum (Pinto et al. 1993) and T. deion/T. kaykai (Pinto et al. 1997). MATERIALS AND METHODS Cultures.—Cultures from three graphically distant populations of 7. fornicum were available for study. geo- cali- They were initially identified using morphology, and this was supported by their nearly iden- tical ITS2 ribosomal transcript sequences (R. Stouthamer, pers. comm.). All are as- signable to Form A of this species as de- fined by Pinto (1999). We utilized our stan- dard laboratory cultures of 7. minutum and T. platneri (Pinto et al. 1991), and two cul- tures of T. exiguum. The origin of all cul- tures used is given in Table 1. Cultures were collected and maintained as detailed in Pinto et al. (1991). Each originated from a single mated female that emerged from a field-collected host egg, and was main- tained in the laboratory at 21—27° C on ir- radiated Trichoplusia ni (Hiibner) eggs. Slide-mounted vouchers of all cultures studied are on deposit in the collection of the University of California, Riverside, De- partment of Entomology Research Muse- um, and are labelled with the voucher code RBI and numbers UCRC ENT 43850— 43984. Crosses.—The three cultures of 7. cali- fornicum were crossed with each other, with standard cultures of 7. platneri (PRV1) and T. minutum (MCVA) that were used in pre- vious studies (Pinto et al. 1992, 1993), and with the Hendersonville (EXHN) culture of T. exiguum for a total of ten crosses. The PRV1 and MCVA standard cultures have each been shown to be reproductively com- patible with numerous other conspecifics (Pinto et al. 1992, in prep.). Procedures used for crossing experiments are detailed in Pinto et al. (1991). A single cross be- tween two cultures consisted of an equal VOLUME 104, NUMBER 1 number of heterogamic (males from the other culture) and homogamic (males from the same culture) replicates. Each replicate consisted of a single virgin male and a sin- gle virgin female in a 29.6 cc (8 dram) glass vial with many (40 or more) host eggs. The male and female progeny from each repli- cate were counted, and the mean sex ratio (msr) for the cross calculated as the per- centage of female progeny. The number of heterogamic and homogamic replicates of each cross were designed to number from 12 to 20 each, but fewer were performed in some cases due to extremely poor viability of certain T. californicum cultures. A total of 296 pairs of T. californicum and T. plat- neri were crossed to increase the chance of detecting rare hybridization. This included an expanded number of heterogamic cross- es conducted in both directions between PRV1 X CAAD and PRV1I X CASB, and 39 between PRV1 X CAYK (Table 2). For statistical analyses, however, only the first 20 replicates were compared with the ho- mogamic replicates. For each cross, an ad- ditional 10 virgin females were placed in- dividually into separate vials with host eggs but without males to confirm that cultures were arrhenotokous. Reproductive compatibility of a cross in each direction is expressed as a percentage: 100% X msr (heterogamic combination)/ msr (homogamic combination) (Fig. 1). In arrhenotokous Trichogramma, females hatch only from fertilized eggs, while males hatch from unfertilized eggs. The absence of female progeny indicates complete in- compatibility. Relative degrees of compat- ibility were measured using the non-para- metric Mann-Whitney U test to compare the mean sex ratio of the heterogamic cross- es with that of the homogamic crosses (Sor- ati et al. 1996). Electrophoresis.—A total of 14 enzyme systems were examined in the three cultures of 7. californicum, the two reference cul- tures of T. minutum and T. platneri, and one of the two cultures of 7. exiguum (Table 3). The enzyme systems, their Enzyme Com- 35 mission numbers, and the abbreviations representing them in this paper are: aconi- tase (4.2.1.3) Acon, acid phosphatase (3153.2) Acp-2\ esterase (Gal I) est marase (4.2.1.2) Fum, glyceraldehyde-3- phosphate dehydrogenase (1.2.1.12) Gapd, a-glycerol-phosphate dehydrogenase (1.1.1.8) aGpd-1 and aGpd-2, glucose- phosphate isomerase (5.3.1.9) Pgi, glucose- 6-phosphate dehydrogenase (1.1.1.49) G6pd, B-hydroxybutyrate dehydrogenase (1:1..1.30), Hbdh, hexokinase (2.7.1.1) Ak isocitrate dehydrogenase (1.1.1.42) Idh, malate dehydrogenase (1.1.1.37) Mdh-2, malic enzyme (1.1.1.40) Me, phosphoglu- comutase (2.7.5.1) Pgm. The same culture (EXHN) of 7. exiguum used for crosses could not be used for all loci because of a shortage of available specimens. The scores of another 7. exiguum culture from Selma, AL (EXSL) were substituted for Est-/, Me, and Pegi. The electrophoretic analysis followed procedures reported in Pinto et al. (1992), originally detailed in Kazmer (1991). Whole females, four from each culture per run, were individually analyzed at each lo- cus by isoelectric focusing in one or two layers of cellulose acetate membranes using a single blend of carrier ampholytes (8% pH 4—6.5 and 2% pH 3-10 Pharmalytes), and an effective gel length of 4.5 cm. BIOSYS-1 (Swofford and Selander 1989, release 1.7) was used to analyze the data. Nei’s (1972) genetic distances (D) were calculated with individual allozyme profiles as input. All specimens were ho- mozygous at the loci examined, probably due to the fact that each culture was estab- lished from a single mated female and be- cause of the large number of generations that each culture had undergone prior to study (Table 1). RESULTS Crosses.—Results of the crossing studies are summarized in Fig. 1 and Table 2. Of the three crosses conducted among cultures of T. californicum, only that between 36 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 2. Results of 7richogramma crosses conducted in this study.! ——————_ Heterogam Homogamic Cross Yo. Replicates msr Crosses No. Replicates msr p-value nnn ne UE EEE DENI ED UpSEI NEU REE EE REE ESUESRERR EERE! CASB 16 0.49 CAAD 16 0.62 0.2333 CAAD CAAD 8 0.67 CASB 8 0.43 0.0306* CASB CASB« 20 0 CAYK 20 0.59 _ CAYK? CAYK¢ 17 0 CASB 17 0.48 = CASB? CAADS i 0 CAYK 20 0.68 ase CAYK? CAYK¢ 20 0.19 CAAD 20 0.59 0.0001* CAAD? oe 20 (50) 0 PRVI 20 0.66 at PRVI° PRVId 0 (80 0 CASB 20 0.66 = CASB? 2050) CAADS 20 (4 0 PRVI 20 0.66 ats PRV19 DAG) PRVId 2 0 CAAD 20 0.33 oe CAAD? a) CAYKS PRVI 19 0.66 ia PRV1° te e PRVId AYK 20 0.59 ei CAYK? ou o e CASBS MCVA 12 0.65 = MCVA@ ie. 0 MCVAS 2 12 0.29 = mah | 0 CASB CAYK¢ 2 12 0.70 = ee | 0 MCVA MCVAS 2 0.34 = pee i 0 CAYK 12 CASBS aaa 12 0 BXHN 12 0.53 EXHN6 nage 12 0 CASB 12 0.49 eid CAYKé i) — Saigo 12 0 EXHN 12 0.72 EXHN2 9 ==. Zee 12 0 CAYK 12 0.48 ' Numbers in parentheses indicate the actual number of replicates conducted in expanded crosses of T. cali- fornicum and T. platneri, with the accompanying number indicating the number of replicates used in the Mann- Whitney test. The p-value is that of the Mann-Whitney test, with those values significant at an a value of 0.05 indicated by an asterisk (*). Mean sex ratio (msr) is the average proportion of females in all replicates. See Table 1 for explanation of culture acronyms. VOLUME 104, NUMBER 1 0 een ee VO [) [ose Se S lv #, CAAD <2 SCASBS —” ibe > MCVA > 1.0 me Oly Fig. 1. Crossing results among cultures of Tricho- gramma. Solid arrows represent complete reproductive compatibility according to a Mann-Whitney U test. Dashed arrows represent complete incompatibility. Hatched arrows represent partial compatibility. Num- bers along arrows represent level of reproductive com- patibility for heterogamic cross relative to appropriate homogamic check cross. Arrows point to the parental female. See Table | for explanation of acronyms. CAAD and CASB showed full bidirectional compatibility. Although the Mann-Whitney test indicated a significant difference be- tween the homogamic check and the cross involving CAAD males and CASB females, the direction of significance was that of more females in the heterogamic cross. This difference is likely due to the generally poor viability of 7. californicum. There was incomplete unidirectional compatibility be- tween CAYK males and CAAD females as indicated by the Mann-Whitney test. Only 10 of the 20 CAAD females produced fe- male progeny in this heterogamic cross, whereas 19 of the 20 CAAD females pro- duced daughters in the homogamic check. The cross between CASB and CAYK was incompatible in both directions. 37 Table 4. Nei genetic distances (D) among cultures of Trichogramma examined electrophoretically. EXHN/ Culture CASB CAYK MCVA _ PRVI EXSL CAAD} 10470) /10:693; 0!693. 0:6937 5101981 CASB — 0.470 0.693 0.470 0.981 CAYK — OSI OlG93 eles 86 MCVA _ 0.470 0.693 PRV1 — 0.625 None of the interspecific crosses yielded female progeny, including the relatively large number of replications between T. californicum and T. platneri. This is con- sistent with an earlier cross between CAAD and a collection of 7. platneri from Cow Head Lake (Modoc Co.), CA (Pinto 1999). It is not known whether interspecific mat- ings occurred or not. Electrophoresis.—Of the 14 loci exam- ined, eight showed variation (Table 3). No usable results were obtained with Acon, Fum, Gapd, Hbdh, Hk, or Mdh. The T. cal- ifornicum cultures differed from those of the other species at three loci, Acp-2, G6pd and Pgm, although each of these loci was variable among cultures of 7. californicum as well. Nei’s genetic distance was calcu- lated for the cultures analyzed in this study (Table 4), and the distances plotted in a phenogram using UPGMA clustering (Fig. 2). The distances and phenogram are in- tended as numerical and visual representa- tions of the results, and should not be in- Table 3. Allelic designation of 8 loci for the cultures examined.! Species Culture Acp-2 aGpd-1 Est-1 G6pd Idh Me Pgi Pgm T. californicum CAAD D A Cc Ec B A A B CASB Cc A B Cc B A A D CAYK (c A B D A A A B T. minutum MCVA A A null B B A A (€ T. platneri PRV1 B A B B B A A A T. exiguum EXHN E B ao A B _ — E EXSL ~- — A — — A A -— ' Relative distances traveled for electromorphs at each locus expressed as a ratio of distance between edge of cathode and homomeric band to entire gel length in alphabetical order of allelic designation: Acp-2 (0.42, 0.51, 0.58, 0.60, 0.62), aGpd-1 (0.22, 0.33), Est-1 (0.07, 0.11, 0.20), G6pd (0.09, 0.22, 0.42, 0.47), Idh (0.58, 0.64), Me (0.49), Pgi (0.51), Pgm (0.11, 0.27, 0.29, 0.33, 0.58). 38 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON T. exiguum Fig. 2. CASB CAYK CAAD MCVA PRV1 EXHN/ EXSL Phenogram (UPGMA clustering) of Nei’s genetic distances among cultures of Trichogramma ex- amined electrophoretically. See Table 1 for explanation of acronyms. terpreted phylogenetically. The phenogram does show the three cultures of 7. califor- nicum to be nearer to each other than to the cultures of the other species, but it should be noted that CASB is as near to PRV1 in the distance matrix as it is to either of the other homospecific cultures (Table 4). The results for Pgm were compared with those reported for several cultures of the 7. minutum complex (Pinto et al. 1992) using the results for MCVA and PRVI1 as stan- dards of comparison. Other than crossing unknowns with standard cultures, this locus provides the only means of separating most collections of 7. platneri and T. minutum from one another. The electromorphs for T. californicum were different from all known Pgm electromorphs in the 7. minutum com- plex and in 7. exiguum (Table 3). The data for Acp-2 and G6pd could not be compared with information from the prior study, but they are tentatively assumed to be diagnos- tic loci for T. californicum. DISCUSSION The crossing and electrophoretic results provide no basis for explaining the mor- phological intermediacy previously found between 7. californicum and T. platneri. Every one of the almost 300 reproductive pairings between the two species was neg- ative. Also, the two species have distinct allozymic profiles, as well as different ITS2 sequences (van Kan et al. 1996). Specula- tion on causes other than gene flow for this intermediacy is premature and the basis of reproductive incompatibility between the species remains unknown. The failure to detect reproductive or mo- lecular intermediacy between T. californi- cum and T. platneri in this study, of course, could be explained by limited sampling. Al- though we failed to find hybridization in the numerous heterospecific pairings, the indi- viduals of 7. californicum and T. platneri crossed represent few isofemale lines (three and two, respectively). More extensive sampling would be useful. Utilization of isofemale lines unfortunately is necessary in Trichogramma studies to insure that all replicates are homospecific (Pinto et al. 1992). The preferable approach of utilizing unrelated individuals for replications is pre- cluded by the presence of heterospecifics at most collection sites (potentially in the same host egg) coupled with problems of identification. In Trichogramma, females can not be identified unless associated with males, and slide-mounted material is re- quired for male identification. Genetic var- iation could be better estimated by utilizing a larger number of isofemale lines, but this alternative is not straightforward in uncom- mon species such as 7. californicum. The limitations of our sampling proce- dure notwithstanding, it should be noted that the species of Trichogramma studied reproductively thus far indicate that the magnitude of morphological difference sep- arating T. californicum and the T. minutum complex, as minor as it is, does correlate well with reproductive incompatibility (Pin- VOLUME 104, NUMBER 1 to 1999). If the few cases of intermediacy are due to hybridization we predict that they result from relatively uncommon events. We also should mention that although few T. californicum lineages have been studied allozymically thus far, the Pgm locus has been examined in over 100 lineages of the T. minutum complex (Pinto et al., in prep). In all cases, the alleles at this locus in both species of the complex are distinct from those reported here for 7. californicum. Both crossing and electrophoretic results indicate a high degree of intraspecific var- iation in 7. californicum as compared to that found in certain other species of 77i- chogramma (Pinto et al. 1992, 1993). The greatest Nei’s distance found among the three cultures of 7. californicum (0.693) is far greater than distances reported in all species of Trichogramma analyzed to date (Pinto et al. 1992, 1993), including that found in 7. minutum (0.486). In fact, the least distance between cultures of 7. cali- fornicum (0.470) is greater than the greatest distance between cultures of all other spe- cies previously examined except 7. minu- tum. These allozymic differences are not predicted by the bidirectional reproductive compatibility between the Adin and San Bernardino cultures of 7. californicum, or the partial compatibility between the Adin and Yakima cultures. They also are not pre- dicted by known morphological or ITS2 se- quence similarity. ITS2 sequences are use- ful in separating all morphologically dis- tinctive species examined thus far (Stou- thamer et al. 1999), but are nearly identical in the three 7. californicum cultures (Stou- thamer, pers. com.). The degree of repro- ductive disjunction within 7. californicum, however, is not completely without prece- dence in Trichogramma. Pinto et al. (1991) found similar levels of incompatibility among cultures of 7. deion. They also re- ported one-way incompatibility and re- duced two-way compatibility in cultures currently assigned to 7. minutum. Considerable morphological variation within 7. californicum already has been 39 noted and the species was divided into two forms, A and B, on this basis (Pinto 1999). The two are broadly sympatric in California but Form B is known only from museum specimens. Within Form A, populations from Baja California and western Texas also have been identified as morphological outliers (Pinto 1999). Crossing and molec- ular studies are needed to determine their relationship to the cultures investigated here. Clearly, Trichogramma californicum remains a highly variable and poorly un- derstood entity. It may constitute a unit of variation similar to or greater than the 7. minutum complex where the two compo- nent species also are morphologically sim- ilar but reproductively incompatible. As in T. californicum, these reproductive units (7. minutum and T. platneri) are electrophoret- ically distinct (Pinto et al. 1992) but do not differ in ITS2 sequence (Stouthamer et al. 2000). However, species status for 7. min- utum and T. platneri also has been sup- ported by clear-cut reproductive incompat- ibility and distinct geographic distributions. The geography of reproductive incompati- bility and allozymic variation in 7. califor- nicum is unknown and any proposal to sub- divide the species without more extensive sampling is premature. ACKNOWLEDGMENTS This study was supported by grants 96- 35312-3890 from the USDA (NRICGP) and DEB 9973150 from NSE (PEED): LITERATURE CITED Falcon, L. A. and J. Huber. 1991. Biological control of the codling moth, pp. 355-369. Jn van der Geest, L. P. S., and H. H. Evenhuis, eds. Tortricid Pests. Their Biology, Natural Enemies and Con- trol. Elsevier: Amsterdam. Kazmer, D. J. 1991. Isoelectric focusing procedures for the analysis of allozymic variation in minute ar- thropods. Annals of the Entomological Society of America 84: 332-339. Mills, N. J. and K. P. Carl. 1991. Parasitoids and pred- ators, pp. 235—252. In van der Geest, L. P. S., and H. H. Evenhuis, eds. Tortricid Pests. Their Biol- ogy, Natural Enemies and Control. Elsevier: Am- sterdam. 40 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Nagaraja, Hi Jagarkatti. 1973. A key to some New World species of Trichogramma (Hymenop- [ricl immatidae), with descriptions of our new species. Proceedings of the Entomolog- ical Society of Washington 75(3): 288-297. Nei. M. 1972. Genetic distance between populations. American Naturalist 106: 283-292. Pinto, J. D. 1999 [**1998”]. Systematics of the North American species of Trichogramma Westwood (Hymenoptera: Trichogrammatidae). Memoirs of the Entomological Society of Washington No. 22, 287 pp. Pinto, J. D., D. J. Kazmer, G. R. Platner, and C. A. Sassaman. 1992. Taxonomy of the Trichogramma minutum complex (Hymenoptera: Trichogramma- tidae): Allozymic variation and its relationship to reproductive and geographic data. Annals of the Entomological Society of America 85: 413-422. Pinto, J. D., G. R. Platner, and C. A. Sassaman. 1993. Electrophoretic study of two closely related spe- cies of North American Trichogramma: T. pre- tiosum and T. deion (Hymenoptera: Trichogram- matidae). Annals of the Entomological Society of America 86: 702-709. Pinto, J. D., R. Stouthamer, and G. R. Platner. 1997. A new cryptic species of Trichogramma (Hyme- noptera: Trichogrammatidae) from the Mojave Desert of California as determined by morpholog- ical, reproductive and molecular data. Proceedings of the Entomological Society of Washington 99: 238-247. Pinto, J. D., R. Stouthamer, G. R. Platner, and E. R. Oatman. 1991. Variation in reproductive compat- ibility in Trichogramma and its taxonomic signif- icance (Hymenoptera: Trichogrammatidae). An- nals of the Entomological Society of America 84: 37-46. Smith, S. M. 1996. Biological control with Tricho- gramma: Advances, successes, and potential of their use. Annual Review of Entomology 41: 375— 406. Sorati, M., M. Newman, and A. A. Hoffmann. 1996. Inbreeding and incompatibility in Trichogramma nr. brassicae: Evidence and implications for qual- ity control. Entomologia Experimentalis et Appli- cata 78: 283-290. Stouthamer, R., J. Hu, E J. P. M. van Kan, G. R. Plat- ner, and J. D. Pinto. 1999. The utility of internally transcribed spacer 2 DNA sequences of the nucle- ar ribosomal gene for distinguishing sibling spe- cies of Trichogramma. BioControl 43: 421—440. Stouthamer, R., Y. Gai, A. B. Koopmanschap, G. R. Platner, and J. D. Pinto. 2000. ITS-2 sequences do not differ for the closely related species Tricho- gramma minutum and T. planteri. Entomologia Experimentalis et Applicata 95: 105-111. Swofford, D. L. and R. B. Selander. 1989. BIOSYS-1. A computer program for the analysis of allelic variation in population genetics and biochemical systematics. Release 1.7. Urbana, IL (Software). van Kan, E J.P M., 1. M. M.S. Silva, M. Schilthuizen, J. D. Pinto, and R. Stouthamer. 1996. Use of DNA-based methods for the identification of mi- nute wasps of the genus Trichogramma. Proceed- ings of the section Experimental and Applied En- tomology of the Netherlands Entomological So- ciety (N.E.V.) 7: 233-237. PROC. ENTOMOL. SOC. WASH. 104(1), 2002, pp. 41-50 A NEW SPECIES OF MAEVIUS STAL FROM AUSTRALIA AND SOME NOTES ON THE FAMILY HYOCEPHALIDAE (HEMIPTERA: HETEROPTERA) HARRY BRAILOVSKY Departamento de Zoologia, Instituto de Biologia, U.N.A.M., Apdo Postal No. 70-153, México, 04510 D.E, México (e-mail: coreidae @ servidor.unam.mx) Abstract.—Maevius luridus, n. sp., is described from Australia and compared with Maevius indecorus Stal, the only previously known species of the genus. Habitus view illustrations and drawings of the male genitalia are provided to distinguish the three spe- cies now included in the family Hyocephalidae. New records for Hyocephalus aprugnus Bergroth and Maevius indecorus are presented. A key to the genera and species is given. Key Words: The endemic Australian family Hyoce- phalidae has both Coreidae and Lygaeidae features and is perhaps the most primitive of the families of the Coreoidea (Stys 1964; Schaefer 1964, 1965, 1981). Previously two genera and two species were known. The first hyocephalid, Maevius indecorus, was described by Stal (1874) from Moreton Bay in Queensland, Australia, and was included in the Division Lethaearia of the Family Lygaeidae. Scudder (1962) removed the ge- nus Maevius from the Lygaeidae and trans- ferred it to the Coreidae because of its ex- ternal similarity to Lygaeopharus Stal (Cor- eidae: Colpurini). Bergroth (1906) de- scribed the second taxon, Hyocephalus aprugnus, from a single macropterous fe- male from South Australia (Yorketown) and included this genus in a new subfamily, Hy- ocephalinae, within the Coreidae. Later Bergroth (1912) described a brachypterous female from New South Wales, Australia, and reduced the rank of this group to a mere division (Hyocephalaria) of the Coreidae. Reuter (1912) raised Hyocephalinae to fam- ily rank and this action has been followed by all subsequent authors. Stys (1964) in Hemiptera, Hyocephalidae, Maevius, Hyocephalus, taxonomy the only previous comprehensive paper on the Hyocephalidae retained them as a fam- ily, giving a detailed description, and dis- tinguishing them from Coreidae, Largidae, Lygaeidae, Stenocephalidae and Pyrrhocor- idae. In that seminal contribution only Hy- ocephalus aprugnus was available for study. Kumar (1965) analyzed the position of the hyocephalids, and concluded that the type of ovipositor in this family is a mod- ification in response to its particular mode of life. A year later Kumar (1966) studied the biology, the immature stages and the relative growth of some Australian bugs. The species studied by Kumar in both pa- pers under the name Hyocephalus sp. nov. was actually Maevius indecorus. Recently Schuh and Slater (1995) sum- marized the general morphology of the Hy- ocephalidae. In this paper I describe a second species of Maevius and give new distributional rec- ords for the other two species in the family. The following abbreviations indicate in- stitutions where specimens are deposited or which generously lent material: Australian 42 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON National In Collection, Canberra (ANIC he Natural History Museum, London (BMNH); Bernice P. Bishop Mu- seum, Honolulu, Hawaii (BPBM); Colec- cidn Entomoldégica del Instituto de Biolo- gia, Universidad Nacional AutOnoma de México (UNAM); James A. Slater Collec- tion (JAS); Australian Museum, Sydney (AMS); Queensland Museum, Brisbane (QMBA); South Australian Museum, Ade- laide (SAMA); University of Queensland Insect Collection, Brisbane (UQIC); West- ern Australian Museum, Perth (WAM). All measurements are given in millime- ters. Maevius luridus Brailovsky, new species (Pigs. 1,4, 35 11) Description.—Measurements: Male first, female second: Head length: 1.70, 1.89; width across eyes: 1.27, 1.30; interocular space: 0.80, 0.81; interocellar space: 0.59, 0.62; preocular distance: 1.30, 1.36; length antennal segments: I, 1.42, 1.39; H, 1.45, L552 Il, 0:77, 0.78: IV. 0:89) 0:92. Prono- tum: Total length: 1.42, 1.48; width across frontal angles: 1.30, 1.37; width across hu- meral angles: 1.98, 2.01. Scutellar length: 1.08, 1.20; width, 1.33, 1.36. Total body length: 9:10; 10:20: Male: Dorsal coloration: Head brown on ochre yellow background, with space between eye and ocelli and a median lon- gitudinal band running from apex of tylus to vertex light yellow; antennal segments I to Ill black, and IV yellow with basal third black; pronotum, scutellum, clavus and co- rium ochre yellow with light brown mottled patches; connexival segments III to VI light brown with anterior and posterior third yel- low, and VII light brown with posterior third yellow; dorsal abdominal segments I to VI yellow with middle third of each seg- ment black, and VII ochre yellow with light brown mottled patches. Ventral coloration: Head including rostral segments I to IV yel- low, with basal third of rostral segment II and median gular region brown: thorax light brown on ochre yellow background, with acetabulae, and metathoracic peritreme yellow; legs ochre yellow; abdominal sterna yellow with light brown mottled patches; pleural margin of abdominal sterna III to VI light brown with anterior and posterior third yellow, and VII light brown with posterior third yellow. Sculpture: Head, pronotum, scutellum, corium, clavus, thorax, and ab- dominal sterna dull, with small tubercles, bearing short, minute, curved-like hairs; an- tennal segments densely covered with ad- pressed golden setae, with some semierect longer setae, denser on segments II to IV; femora with few scattered setae; tibiae with some minute, adpressed, hardly recogniz- able hairs, and with many longer, straight setae; both ventral edges of tibiae bear one row of spine-like, thicker setae; tarsi dense- ly covered with long setae, especially on ventral surfaces. Structure: Head horizon- tal, porrect, very long and narrow; post- ocular section thicker than anteocular one; frons and vertex convex; eyes small, remote from pronotal margin; ocelli very small, sit- uated near posterolateral eye margin; tylus elevated, narrow, laterally compressed, an- teriorly considerably surpassing jugae; ju- gae short, and anteriorly pointed; antenni- ferous tubercle distinctly infericorn in po- sition, remote from eyes, occupying nearly whole area between gula and tylus, large with divergent and convex lateral sides, simple anterolateral corners, and very large apical membraneous sockets; bucculae large and long, anteriorly rounded, gradu- ally narrowing posteriorly, disappearing at level anterior eye margin; rostrum reaching anterior border of mesosternum. Pronotum: Trapezoidal; collar evident; frontal angles distinctly produced anteriorly; lateral mar- gins weakly sinuate, emarginate; humeral angles slightly exposed; posterior margin weakly straight; disc with a pair of indis- tinct large, oval callar impressions. Legs: Fore femur ventrally armed with a double row of subapical spines; middle and hind femora unarmed; tibiae sulcate. Scutellum: Triangular; disc almost flat; anterolateral corners with a pair of irregular depressions; VOLUME 104, NUMBER 1 43 Figs. 1-9. 1-3, Male genital capsule in caudal view. 1, Maevius luridus. 2, Maevius indecorus. 3, Hyoce- phalus aprugnus. 4—9, Parameres in lateral view. 4, 5, Maevius luridus. 6, 7, Maevius indecorus. 8, 9, Hyoce- phalus aprugnus. 44 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON nelytra: Staphylinoid; orium indistinguishably fused ious pad, and hemelytra meet- h other along midline for their entire length; posterior end truncated and covering only first three abdominal terga; hemelytral membrane absent. Abdomen: Connexival segments higher than terga; connexival margin complete; posterior angle of each connexival segment complete, not extend- ing as a short spine. Genitalia: Genital cap- sule with posteroventral margin complete; surface convex, laterally slightly raised al- most for their whole longitudinal length (Fig. 1). Paramere as in Figs. 4—5. Female: Coloration: Similar to male. Gen- ital plates brown with upper margin yellow. Structure: Abdominal sternite VII fissured for their entire length; paratergite VIII short, squarish, with visible spiracle; paratergite [IX larger, square; If valvifers narrow, parallel and interlocking at their entire length. Variation (in both sexes).—1, Antennal segment I chestnut orange. 2, Connexival segments IIf to VI with upper margin brown, lower margin chestnut orange, and anterior and posterior third yellow to ochre yellow. 3, Trochanters light chestnut orange. Type material.—Holotype: ¢, Australia: South Australia, Eyre Penn., near Caralue Bluff, under Spinifex sp. (Gramineae), 8 October 1964, E J. Mitchel (SAMA). Par- atypes: 1 92, same data as holotype (SAMA); 2 5, 1 2, Australia: Arden Vale, on Triodia sp. (Gramineae), 19 June 1976, Civan Dyk (SAMA, UNAM): 1-6, 1 °2, Australia: Victoria: Hattah Lakes Nat. Park, 4 April 1969, G. B. Monteith, (QMBA). Diagnosis.—This is a medium-sized spe- cies similar to M. indecorus Stal, the only previously known species of the genus. Maevius luridus is easily distinguished by the pale ochre yellow coloration in contrast with the dark reddish brown coloration of M. indecorus on which only the following areas are pale yellow or chestnut orange: antennal segment IV, anterior and posterior third of abdominal terga HI to VII, trochan- ters, basal third of tibiae, tarsi, and anterior apex subacute clavus and « into a ing ea and posterior third of pleural margins of ab- dominal sterna III to VII. The length of an- tennal segments I to IV are shorter in M. luridus and the shape of the parameres also differs (Figs. 4—7). Distribution.—Known only from the Southeastern part of South Australia and the Northwestern of Victoria. Etymology.—Named for its light color- ation; from the latin /uridus, pale yellow. Maevius indecorus Stal (Figss2,46,972110) Maevius indecorus Stal 1874: 165. The morphology and life history of this species were studied by Kumar (1965, 1966) under the name Hyocephalus sp. nov. He mentioned that it probably feeds on the seeds of Acacia (Leguminosae) or Eucalyp- tus (Myrtaceae), or both. Distribution.—Stal (1874) described this species from Moreton Bay in Queensland, Australia. Kumar (1966) recorded it from Moggill, near Brisbane. The new records list- ed below show it occurs from Rockhampton south to Newcastle in Eastern Australia and in the Southeastern Western Australia. Material examined.—Australia: Queens- land: | d, 1 2, S. E. Queensland: Brisbane, Rochedale, January 1980, V. Davies and R. Raven (UQIC, QMBA); 2 3, Lamington Nat. Park, 17-24 May 1965, G. Monteith (UQIC); 1 3d, Mt. Chalmers, 24 October 1990, R. Raven (QMBA); 1 6,4 &, Bunya Mts., 27-31 May 1962, E. A. Bernays, 17— 18 September 1966, G. Monteith (UQIC); 1 36, 2 2, Dunwich Stradbroke Is., 9 May 1964, 15-16 March 1975, 21 April 1965, G. Monteith (UQIC); | 2, Big Sand Hill at Moreton I., 1 October 1955, E. N. Marks (from litter under Banksia) (UQIC): 1 @, Dunwich, 11 April 1965, J. E. Dunwoody (UQIC); | 3, Fletcher, without date, E. Sut- ton (UQIC); 1 6d, thunderbird Park (Tam- borine Mts.), 23—29 October 1993, S. A. Slipinski and J. E Lawrence (ANIC); 2 males, | female, Brisbane, adults taken at Moggill on banks of River Brisbane and VOLUME 104, NUMBER 1 male. , 10. Dorsal view of Maevius indecorus Fig. 46 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs 11! Dorsal view of Maevius luridus, male. VOLUME 104, NUMBER 1 reared in culture by R. Kumar (BPBM); 20 3, 35 @, Brisbane, 20 May 1960, H. A. Rose, 19 August 1960, M. Smith, 15 Oc- tober 1963, December 1963, R. Kumar, 1— 9 January 1964, R. Kumar, 17 December 1964, R. Kumar (BPBM, UQIC, UNAM); So, 4 2) Mogeill, 2 Aprilmi962. TE: Woodward, 7-16 September 1963, R. Ku- mar, 4 October 1963, R. Kumar (UQIC); 1 2, Blackdown Tableland, via Dingo, 1—6 February 1981, G. B. Monteith (QMBA); 4 @, Highvale, March 1970, B. Cantrell (UQIC); | &, Brisbane, Mt. Coot-tha, 10 January 1972, J. A. Slater (JAS). Australia: New South Wales: 1 ¢, Greta, 1951, J. Sed- lacek (BPBM); 2 6, 2 2, Unumgar State Forest, via Kyogle, 12 April 1966, G. Mon- teith (UQIC); 1 2, Bonie N., Coonabara- bran, 6 September 1970, B. Cantrell (UQIC); Western Australia: | 2, Goldfields Surv, Iuake Cronin, 5 June 9S! W: FE Humphreys (WAM); 3 °, Wildlife Res., 21 Miike IN. of Perth, 6 December 19715 ).-A- Slater (adults taken under Hypocalymna an- gustifolium Endl. (Myrtaceae) (JAS, UNAM); 1 9s Mt. Cooke;"43) mi S. of Perth, 23 March 1968, E H. Uther Baker (SAMA); 1 @, 2 mi., W. of Fraser Range, 3 June 1979, EK W. Aslin (adult taken under road side rocks on edge of old road) (WAM); 2 oc, Stirling Range, 4 March 1994, Monteith and Janetzki (QMBA). Hyocephalus aprugnus Bergroth (Rigs; 3.1630 912 713) Hyocephalus aprugnus Bergroth 1906: 648. Bergroth’s (1906) original holotype was considered lost for many years and when Stys (1964) discovered 11 specimens clearly collected with the holotype in the Hungarian Natural History Museum he designated one as a neotype. Later the original holotype was discovered in the University Zoology Mu- seum Helsinki but Grant and Stys (1970) ap- plied for retention of the neotype. Stys (1964) characterized the genus and the species and commented on the mor- phology and relationship of the group. The AT macropterous (Fig. 13) and _ staphylinoid (Fig. 12) condition, has been adequately re- described by Bergroth (1912) and Stys (1964). Distribution.—This species was de- scribed from Yorketown (near Adelaide) in South Australia (Bergroth 1906), and later recorded from an unspecified locality in New South Wales (Bergroth 1912). The new records listed below show it occurs in the Southwestern Western Australia and in the Southeastern corner of Western Austra- lia. No material is available to confirm its occurrence in New South Wales. Material examined.—Australia: South Australia: 1 6, 1 2, Gum Lagoon 36°17'S— 140°02’E, 25 March 1992, J. A. Forrest (SAMA); 1 92, 9 km., N. of Long Point, 35°37'S—139°39'E, 5-9 September 1991 (SAMA); 1 6, Loftia Park, 21 September 1989, D. Hirst (SAMA); Western Australia: 1 2, Torbay in Let, 10 mi., W. of Albany, 28 December 1971, J. A. Slater (JAS); 1 &, Mt. Cooke, 43 mi., S of Perth, 23 March 1968, E H. Uther Baker (WAM); 1 a, Limestone Head, via Albany, 3 March 1970, G. W. Kendrick (UNAM); 1 6, 1 8, Garden Island, 32°12’S—115°40’E, 18 June 1969, B. Humphreys (adult taken under gal- vanized iron at the base of Pittosporum (Pittosporaceae) and 12 November 1975, S. M. Slack Smith (WAM); 1 6d, Vicinity of Devils Lair Cave, near Karridale, 20 March 1973, A. Baynes (UNAM); 1 3, 1 2, White Gum Flat, Stirling Range Nat. Park, 34°24'S=117°55'E., 1 April 1993, M.S. Harvey and J. M. Waldock (WAM); 2 6, 3 2, Thomas R., Cape Arid, 23 May 1977, R. McMillan (UNAM, WAM). PLANT ASSOCIATIONS Hyocephalids live on the ground in a va- riety of habitats from semi-desert to tall coastal eucalypt forest. They shelter under stones, logs and other debris. Maevius inde- corus has been taken under the prickly hum- mocks of Spinifex (Triodia spp.) grasses, a specialized Australian desert habitat occupied 48 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Lf 7d r f i #' Fg tt ee ts i} Fig. 12. Dorsal view of Hyocephalus aprugnus, male staphylinoid. by many invertebrates (Monteith, pers. comm. ). The natural food of hyocephalids has not been recorded but is assumed to be seeds. Kumar (1966) successfully reared Maevius indecorus through several generations in the laboratory on sunflower seeds. He spec- ulated that they may feed on seeds of Eu- calyptus (Myrtaceae) and Acacia (Mimo- saceae) in nature. This speculation was re- peated as fact (Schaefer and Mitchell 1983, Schuh and Slater 1995) but has not been 49 VOLUME 104, NUMBER 1 opterous. genus, female, macr é Dorsal view of Hyocephalus apru Fig. 13. 50 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON confirmed. Labe! data on specimens report- ed in this paper record Maevius indecorus in association with Hypocalymna angusti- folius Endl. (Myrtaceae) and Banksia (Pro- teaceae), M. /uridus in association with Spi- nifex and Triodia (Gramineae) and Hyoce- phalus aprugnus in association with Pittos- porum (Pittosporaceae). No direct feeding can be assumed from any of these. KEY TO THE KNOWN SPECIES OF HYOCEPHALIDAE 1. Antennal segment HI longer than IV; fore fe- mur unarmed or with some subapical tubercles; antennal segment I longer than 1.95 mm, par- ameres like Figs. 8—9 Sieg Vacs venta tas Hyocephalus aprugnus Bergroth — Antennal segment III shorter than IV; fore fe- mur armed with double row of apical spines; antennal segment I shorter than 1.80 mm; par- prenes Ihe lees, 4s po coop epan ease 2, . Body dark reddish brown; antennal segment I slender and longer than 1.68 mm; parameres likesBisss (6-72 he. 5 se Maevius indecorus Stal — Body ochre yellow or pale brown on a ochre yellow background; antennal segment I robust and shorter than 1.60 mm; parameres like Figs. Boies ctiey eee WE Maevius luridus, n. sp. i) ACKNOWLEDGMENTS Thanks to the following individuals and institutions for loans and other assistance: T. A. Weir (ANIC), G. Cassis (AMS), Janet Margerison-Knight (BMNH), Gordon Nish- ida (BPBM), James A. Slater (JAS), Gor- don Gross and J. Forrest (SAMA), G. B. Monteith (QMBA), Margaret Schneider and Greg Daniels (UQIC), and T. Houston and B. Hanich (WAM). E. Barrera (UNAM), J. Contreras and A. Luna (UNAM) prepared the dorsal view illustrations and genital drawings. The Sistema Nacional de Inves- tigadores, México (SNI) provided financial assistance. Special thanks to G. B. Monteith for the comments on the manuscript. LITERATURE CITED Bergroth, E. 1906. Aphylinae und Hyocephalinae, zwei neue Hemipteren-Subfamilien. Zoologischer Anzeiger, Leipzig 29: 644-649. . 1912. New or little known Hemiptera, chiefly from Australia, in the American Museum of Nat- ural History. Bulletin of the American Museum of Natural History 31: 343-348. Grant, J. A. and P. Stys. 1970. Hyocephalus aprugnus Bergroth, 1906 (Insecta, Hemiptera-Heteroptera, Hyocephalidae): request for retention of the neo- type specimen in preference to the rediscovered holotype. Bulletin of Zoological Nomenclature 27 (2): 113-114. Kumar, R. 1965. Aspects of the morphology of Cor- eoidea and their value in its higher classification. Proceedings of the Royal Society of Queensland 76 (3): 27-91. . 1966. Studies on the biology, immature stag- es, and relative growth of some Australian bugs of the superfamily Coreoidea (Hemiptera: Heter- optera). Australian Journal of Zoology 14: 895— il, Reuter, O. M. 1912. Bemerkungen uber mein neues Heteropterensystem. Ofversigt af Finska Veten- skaps-Societetens Forhandlingar 54A (6): 1—62. Schaefer, C. W. 1964. The morphology and higher classification of the Coreoidea (Hemiptera-Heter- optera): Parts I and II. Annals of the Entomolog- ical Society of America 57: 670—684. . 1965. The morphology and higher classifica- tion of the Coreoidea (Hemiptera-Heteroptera). Part II. The families Rhopalidae, Alydidae, and Coreidae. Miscellaneous Publications of the En- tomological Society of America 5: 1—76. . 1981. The morphology and relationships of the Stenocephalidae and Hyocephalidae (Hemip- tera: Heteroptera: Coreoidea). Annals of the En- tomological Society of America 74 (1): 83-95. Schaefer, C. W. and P. L. Mitchell. 1983. Food Plants of the Coreoidea (Hemiptera: Heteroptera). An- nals of the Entomological Society of America 76 (4): 591-615. Schuh, R. T. and J. A. Slater. 1995. True bugs of the world (Hemiptera: Heteroptera). Classification and Natural History. Cornell University Press, 336 PP- Scudder, G. G. E. 1962. The world Rhyparochrominae (Hemiptera: Lygaeidae). I. New synonymy and Generic changes. The Canadian Entomologist 94 (7): 764-773. Stal, C. 1874. Enumeratio Hemipterorum IV. Konglica Svenska Vetenskaps-Akademiens Handlingar 12 (1): 1-186. Stys, P. 1964. The morphology and relationship of the family Hyocephalidae (Heteroptera). Acta Zoolo- gica Hungarica 10: 229-262. PROC. ENTOMOL. SOC. WASH. 104(1), 2002, pp. 51-55 PERLESTA ETNIERI (PLECOPTERA: PERLIDAE), A NEW SPECIES OF STONEFLY FROM TENNESSEE B. C. KONDRATIEFF AND R. E KIRCHNER (BCK) Department of Bioagricultural Sciences and Pest Management, Colorado State University, Fort Collins, CO 80523, U.S.A. (e-mail: bkondrat @ceres.agsci.colostate.edu); (RFK) Water Quality Section, U.S. Army Corps of Engineers, Huntington District, P.O. Box 9, Apple Grove, WV 25502, U.S.A.! ABSTRACT.—A new species of stonefly, Perlesta etnieri is described from Maury and Williamson counties, Tennessee. It is a member of the nelsoni species group but can be distinguished from included species by the absence of two well-defined patches of sensilla basiconica on tergum 10 and by the dark body coloration. Additionally, new Tennessee state records are reported: P. adena Stark (Sumner, Trousdale, and Williamson counties) and P. lagoi Stark (Bledsoe and Cumberland counties). Key Words: The stonefly genus Perlesta now rivals Neoperla and Acroneuria in numbers of species in North America, a remarkable fact considering that during most of the 20th century only a single widespread species was recognized (Stark 1989). Currently, seventeen species are recognized (Stark 1989, Poulton and Stewart 1991, Stark and Rhodes 1997, Kirchner and Kondratieff 1997, DeWalt et al. 1998, Kondratieff and Baumann 1999). Recently, DeWalt et al. (2001) reviewed the Perlesta of Illinois, documenting eight species for the state. A new species of Perlesta was discov- ered among material from Tennessee sub- mitted for identification by Dr. David A. Et- nier, University of Tennessee. The authors collected additional adult material from the Harpeth River. This species is described be- low. The descriptive terminology follows Stark (1989). ' The views of the author do not purport to reflect the position of the Department of the Army or the Department of Defense. Plecoptera, Perlidae, Perlesta, new species Perlesta etnieri Kondratieff and Kirchner, new species (Figs. 1-6) Male.—Forewing length 8—9 mm. Gen- eral body color black in life, brown in al- cohol. Head yellow with black to brown ocellar patch and dark spot anterior to patch (Fig. 1). Wings black to brown except for pale costal margin. Femora yellow, distally and dorsally brown, tibiae brown with api- ces yellow. Abdominal terga black to brown, sterna yellow brown but darker brown apically. Cercus yellow basally, each segment posteriorly brown, distal segments brown. Tergum 10 mesal sclerite shiny dark brown, not divided, sensilla basiconica dis- tinct but small, not elevated into patches (Fig. 2). Paraproct in lateral view, long, slender, slightly curved posteriad at apex, small mesoapical tooth (Fig. 3) not visible in caudal aspect. Penis tube + sac long, caecum absent (Fig. 4). Female.—Forewing length 10-12 mm. Color pattern similar to male but paler. Sub- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON a) No Figs. 1-5. Perlesta etnieri. 1, Adult head and pronotum. 2, Male terminalia, dorsal. 3, Paraproct, lateral. 4, Penis. 5, Female subgenital plate, ventral. VOLUME 104, NUMBER | oa sy) ie] i] + 1BBy.m Fig. 6. micrograph of egg. Perlesta etnieri. Scanning electron photo- genital plate lobes short, truncate and sep- arated by small V-shaped notch (Fig. 5). Egg.—Collar sessile. Chorion surface covered with shallow irregular pits. Wide smooth eclosion line, with micropyles lo- cated at upper margin of line (Fig. 6). Nymph.—Undescribed. Material examined.—Holotype d, Ten- nessee, Williamson Co., Harpeth River at Millview, Arno Road (N35°52'41.0” W86°47'27.1"), 11 May 2000, R. FE Kirch- ner and B. C. Kondratieff. Paratypes, same data as Holotype, 18 6,4 2; same location but 19 May 1998, R. B. Evans, 46 6, 25 ee Maury =Co-, Carters) Creek,” at "Carters Creek, Carters Creek Road (N35°43'02.7" W86°59'44.7"), 19 May 1998, R. B. Evans, 82 O40 2. The holotype is deposited in the National Museum of Natural History, Smithsonian Institution, Washington, DC. Paratypes will be deposited at the following museums and individual collections: Bill P. Stark, Clinton, Mississippi (BPS), C. P. Gillette Museum of ‘Nn ee) Arthropod Diversity, Colorado State Uni- versity (CSU), Illinois Natural History Sur- vey (INHS), and Ralph E Kirchner (RFK), Huntington, West Virginia. Etymology.—We honor Dr. David A. Et- nier, University of Tennessee, a well-known North American ichthyologist and trichop- terist. Diagnosis.—Perlesta etnieri belongs to a species group including P. frisoni Banks, P. nelsoni Stark, and P. teaysia Kirchner and Kondratieff, males of which are character- ized by a long and slender penis tube + sac that lacks a caecum (Fig. 4). The following key will distinguish these species. Perlesta etnieri 1s very similar and closely related to P. nelsoni, but the male can be distin- guished by the darker pigmentation of the body and short distinct sensilla basiconica on tergum 10 (Fig. 2). The male of P. nel- soni is pale yellow and the sensilla basi- conica are reduced (see Stark 1989, fig. 65). The long and slender paraprocts of P. ef- nieri with the apex curved posteriad and a mesoapical tooth (Fig. 3) are suggestive of P. browni Stark, a species known only from Arkansas and Oklahoma (Stark 1989, Poul- ton and Stewart 1991), and P. cinctipes (Banks) known from Illinois, lowa, Kansas, Missouri, Nebraska and Oklahoma. The distinctive aedeagus of P. etnieri (Fig. 4) distinguishes it from the above species. The egg of P. etnieri is similar to P. cinc- tipes (see Stark 1989, fig. 17), but in P. et- nieri the chorion is covered with shallow irregular pits, but the collar is sessile. The egg of P. nelsoni has a smooth chorion and a short slender collar (Stark 1989). Addi- tionally, the subgenital plate of P. etnieri has a small V-shaped notch (Fig. 5), where- as the subgenital plate of P. nelsoni has a deep U-shaped notch (see Stark 1989, fig. 68). Other adult stoneflies collected at the type locality were Allocapnia granulata (Claassen), A. rickeri Frison, A. tennessa Ross and Ricker, /soperla sp., Acroneuria frisoni Stark and Brown, Perlinella ephyre 54 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (Newman) : adena Stark, and a sin- gle female of another species of Perlesta. KEY TO SPECIES OF THE PERLESTA NELSONI SPECIES GROUP MALES 1. Penis without caecum (Fig. 4) — Penis with a caecum (Stark 1989, fig. 1) .... ee. ¢ iephgsa x 2p toh eee oem pens e = not keyed here (see Stark 1989, DeWalt et al. 2001) _ T10 sensilla basiconica numerous, arranged in a pair of slightly elevated, circular patches (Stark 1989, figs. 51-52; Kirchner and Kon- i) falmainleint IUSIM/= 10s A) og eeeleio aia prone c's eta eee Bec 3 — T10 sensilla basonica sparse, not in two patch- CS LOD) ca aes Meee: Fear cara een ca ania cts 4 3. Paraprocts with spine on anteroapical margin (Stark 9 S98 ficts5 2) ares es eee: P. frisoni — Paraprocts without spine on anteroapical mar- gin (Kirchner and Kondratieff 1997, figs. 3 and C2 aia 8) ie cre ren ai tec CRN ee le gested 4. Wing membrane, veins and body pale yellow Sch oR ane cots eden cre Meme ei aaron P. nelsoni — Wing membrane, veins and body dark brown Loyal D) Yel a2ymphs (CSU, RFK, BPS); same location as Philip- pi Church, 25 May 1999, B. P. Stark, 11 ¢, 3 ¢, 2 nymphs (BPS); Williamson Co., Harpeth River at Millview, Arno Road (N35°52'41.0" W86°47'27.1"), 11 Wiay 2000, R. E Kirchner and B. C. Kondratieff, 2& (CSU): ACKNOWLEDGMENTS We thank Bill P. Stark of Mississippi Col- lege for confirming the specific status of P. etnieri and providing verification of P. ad- ena and P. lagoi. LITERATURE CITED DeWalt, R. E., B. P. Stark, and M. A. Harris. 1998. Perlesta golconda (Plecoptera: Perlidae), a new stonefly species from Illinois. Entomological News 109: 315-317. DeWalt, R. E., D. W. Webb, and T. N. Kompare. 2001. The Perlesta placida (Hagen) complex (Plecop- tera: Perlidae) in Illinois, new state records, dis- tributions, and an identification key. Proceedings of the Entomological Society of Washington 103: 207-216. Kirchner, R. E and B. C. Kondratieff. 1997. A new species of nearctic Perlesta (Plecoptera: Perlidae) from Virginia. Proceedings of the Entomological Society of Washington 99: 290-293. Kondratieff, B. C. and R. W. Baumann. 1999. Studies on stoneflies of North Dakota with the description of a new Perlesta species (Plecoptera: Perlidae). Proceedings of the Entomological Society of Washington 101: 325-331. Nelson, C. H. 1996. Placement of Helopicus rickeri Stark in Hydroperla Frison (Plecoptera: Perlodi- dae) with the description of the adult female, nymph, and egg and a cladistic anaysis of Hydro- perla. Proceedings of the Entomological Society of Washington 98: 237-244. VOLUME 104, NUMBER 1 Poulton, B. C. and K. W. Stewart. 1991. The stoneflies of the Ozark and Ouachita Mountains (Plecop- tera). Memoirs of the American Entomological Society 38, 116 pp. Stark, B. P. 1989. Perlesta placida (Hagen), an eastern nearctic species complex (Plecoptera: Perlidae). Entomologica Scandinavica 20: 263-286. Nn Nn Stark, B. P. and H. A. Rhodes. 1997. Perlesta xube, a new stonefly species from Nebraska (Plecoptera: Perlidae). Entomological News 108: 92—96. Wiersema, N. A. and L. S. Long. 2000. Plauditus grandis (Ephemeroptera: Baetidae), a new small minnow mayfly from Tennessee. Entomological News 111: 45—48. PROC. ENTOMOL. SOC. WASH. 104(1), 2002, pp. 56-62 be SCRIPTIONS OF TWO NEW SPECIES OF FULVIUS STAL (HE PTE ROPTERA: MIRIDAE: CYLAPINAE) FROM BRAZIL, WITH BIOLOGICAL AND BIOGEOGRAPHIC NOTES ON THE GENUS PAULO SERGIO FIUZA FERREIRA AND THOMAS J. HENRY (PSFF) Departmento de Biologia Animal, Universidade Federal de Vigosa, 36571-000 Vicosa, MG, Brazil (e-mail: pfiuza@mail.ufv.br); (TJH) Systematic Entomology Labo- ratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, % National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0168, U.S.A. (e-mail: thenry @sel.barc.usda.gov) Abstract.—Two new Brazilian species, Fulvius paranaensis from the state of Parana and F. vicosensis from the state of Minas Gerais, are described. Male genitalia and pho- tographs of both species are provided to assist in identification and notes on general biology and biogeography are given for the New World species of Fulvius. Key Words: biology, biogeography The cylapine genus Fulvius Stal (1862) includes about 65 species worldwide (Car- valho and Costa 1994, Schuh 1995). Schuh (1976) discussed the phylogenetic position of Fulvius within Cylapinae, and Carvalho and Ferreira (1994) provided a key to the New World cylapine genera, including Ful- vius. Carvalho and Costa (1994) revised the New World Fulvius, described 22 new spe- cies, and included an identification key to the 42 known species. Most authors have recognized the placement of Fulvius in the cylapine tribe Fulviini (e.g., Henry and Wheeler 1988, Carvalho and Ferreira 1994), but Schmitz and Stys (1973) argued to give the tribe subfamily status, an action rejected as premature by Schuh (1976). Schuh (1995) in his world catalog treated the tribe as a junior synonym of Cylapini, but more recently Gorczyca (2000), based on a cladistic analysis of the subfamily, rec- ognized four tribes, including Fulviini. A complete list of references to the species of Fulvius can be found in Henry and Wheeler Heteroptera, Miridae, Fulvius, paranaensis, vicosensis, new species, Brazil, (1988), Carvalho and Costa (1994), Schuh (1995), and Gorczyca (2000). In this paper, we describe the two new Brazilian species, F. paranaensis from Pa- rana and F. vicosensis from Minas Gerais, to provide names for an ongoing faunal study. Provided are illustrations of male genitalia and photographs of both species to assist in identification and notes on bi- ology and biogeography of the New World species of Fulvius. Fulvius Stal 1862 Fulvius Stal 1862: 322. Type species: Ful- vius anthocoroides Stal. Monotypic. Diagnosis.—Fulvius is recognized by the small to medium size, elongate body, length 1.4 to 5.5 mm; dark castaneous to brown general color, most species with whitish ar- eas at middle of clavus and base of cuneus; long, produced head; eyes extending ven- trally to gula, gula long; horizontal or near- ly horizontal frons; long rostrum, extending to metacoxae or beyond; trapeziform pron- VOLUME 104, NUMBER 1 otum, with rounded lateral margins and a sinuate posterior margin; large confluent calli, occupying anterior two-thirds; prom- inent collar; broadly exposed mesoscutum; macropterous subparallel hemelytra, with well-developed membrane and 2 closed cells; greatly reduced ostiolar opening, au- ricle, and evaporative area; long, slender legs; long metatibiae, much longer than pro- or mesotibiae; 2-segmented tarsi; and slender claws, usually with a_ subapical tooth. BIOLOGICAL NOTES While very little specific information is available on the biology, evidence suggests that members of the genus are largely pre- daceous and, possibly, mycetophagous. Large numbers of Fulvius often are attract- ed to lights (Carvalho 1956, Maldonado 1969, Schuh 1976, Paula and Ferreira 1998), including F. vicosensis. Fulvius im- becilis (Say) has been observed feeding on dipterous and coleopterous larvae and other soft-bodied arthropods found in damp areas or on fungi under loose bark of poplar (Po- pulus sp.) firewood (Kelton 1985). Fulvius quadristillatus (Stal) has been associated with fungi (Auricularia) growing on decay- ing trees at the headwaters of the Rio Ne- ero, Amazon, Brazil (Carvalho 1954), and has been taken in large numbers from litter and kitchen middens where fungi are abun- dant (Schuh 1976), but also where there is an abundance of larvae that might serve as prey. One Old World species has been reared through multiple generations on but- terfly eggs and larvae (Gorcezyca 2000). Fulvius paranaensis was found feeding on liquified material of dead vertebrate car- casses, a nutrient source perhaps not too dissimilar from that acquired from various arthropod prey. BIOGEOGRAPHIC OVERVIEW Members of the genus Ful/vius are most abundant in tropical regions of the world, with approximately 92 percent of species distributed between the Tropics of Cancer 57 and Capricorn. The majority of the species occur in the Neotropics. Prior to this study, Mexico had the largest number of species recorded (16 species), followed by Brazil (15 species) and Panama (13 species). Oth- er countries range from | to 7 species. Fif- ty-eight percent of species are known from only one country, whereas 42 percent have been reported from two or more countries. The center of diversity for the New World species appears to be Mexico and possibly adjacent Central America, radiating north- ward into the United States and Canada and southward into Central and South America. Fulvius anthocoroides Stal has the largest range, occurring in 22 countries, followed by F. bisbistillatus (Stal) (17), F. breddini Reuter (12), F. guadristillatus, and F. var- iegatus Poppius (10). Insufficient collecting probably accounts for the limited distribu- tion now known for many species. Fulvius anthocoroides is recorded from North America (United States [Florida only]), Central America (Costa Rica, El Salvador, Guatemala, Honduras, Mexico, Nicaragua, and Panama), and South America (Bolivia, Brazil, Colombia, Ecuador [including Gal- apagos Islands], French Guiana, Guyana, Paraguay, Peru, Suriname, and Venezuela), extending into the Caribbean islands. The species F. bisbistillatus, F. breddini, and F. quadristillatus are also widely distributed, radiating from Mexico southward into South America. Fulvius quadristillatus also occurs in the Caribbean islands, and F. breddini is reported from the Galapagos is- lands. These last three mentioned species are sympatric with F. anthocoroides in al- most all its distribution, except for North America. Fulvius variegatus, the species having the fourth largest geographic distri- bution, ranges in the Pacific region from Hawaii south throughout the Bismarck Ar- chipelago (New Britain, New Guinea, New Ireland, and Samoa) and Micronesian Is- land groups (Kusiae, Palau, Ponape, Truck, Yap). With the addition of the two new spe- cies presented in this paper, Brazil becomes the most speciose country, having 17 spe- 58 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1—2. cies of Fulvius, and Minas Gerais, the most speciose Brazilian state, having seven spe- cies: Fulvius paranaensis Ferreira and Henry, new species (Figs. 1—6) Diagnosis.—Fulvius paranaensis keys roughly to F. gampoensis Carvalho and Costa, but differs from it and other species of the genus in having the metacoxa brown only on basal % and pale apically (the me- socoxa is uniformly brown), antennal seg- ment II narrowly pale on apical ’;, and by the male genitalic characters, particularly the shape of elongate secondary gonopore (Fig. 4), the apically hooked phallotheca (Fig. 5), and the broad base of the left par- amere (Fig. 6). Description (Figs. 1—2).—Male (mea- surements include x, followed by ranges in parentheses; n = 4): Length 4.31 mm (3.94—4.69 mm); width 1.25 mm (1.19— 1.31 mm). Head: length 0.48 mm (0.52— 0.45 mm); width 0.60 mm (0.60—0.62 mm); distance between eyes 0.29 mm (0.26-0.31 2 Photographs of Fulvius paranaensis, 3. 1, Dorsal aspect. 2, Lateral aspect. mm). Antenna: Segment I, length 0.48 mm (0.40—0.50 mm); II, 1.24 mm (1.16—1.33 mm): II, 0.50 mm (0.48—0.52 mm); IV, 0.71 mm (0.71 mm). Pronotum: Length 0.50 mm (0.48—0.50 mm); width 1.07 (1.10-1.05 mm). Length from cuneal frac- ture to apex of membrane 0.50 mm (0.48— 0.50 mm); cuneal width 0.31 mm (0.26— 0.33 mm). General coloration dark brown with white and red areas. Head dark brown with apex of clypeus and outer margin of buc- cula red. Eye dark brown. Rostrum brown to fuscous, becoming darker apically, ex- tending to middle of abdomen. Antenna fuscous, segment I reddish brown, segment II dark brown with apical %4 white. Prono- tum dark brown, collar with two fuscous spots; calli occupying %4 of disc; posterior margin of pronotum concave with posterior angles rounded. Mesoscutum brown with lateral margins red. Scutellum brown. Hem- elytron brown; irregular yellow band on base of corium extending to embolium and apex of clavus; base of cuneus white, apex dark brown; apical area of embolium near VOLUME 104, NUMBER | Figs. 3-6. paramere. cuneus, red; membrane fuscous, iridescent, with a small pale area near inner margin of cuneus. Ventral region dark brown. Pro- and mesolegs pale brown except coxae dark brown; metaleg with apical % of coxa and trochanter whitish; setae short, erect or semierect. Head, xyphus, pronotum, pro- pleura, mesoscutum, and scutellum_ sha- greened, almost glabrous, except for very short setae on apical area of head. Antennal segments I and II with short erect or semi- erect setae ranging from shorter than, to as long as, diameter of segments; setae on seg- ments III and IV longer than diameters of their respective segments; segment IT slight- ly incrassate toward apex; relative lengths of antennal segments in ascending order I — ih =tV <1: Male genitalia? Vesica (Fig. 3) without lobe or fields of spines and spicules. Secondary gonopore long (Fig. 4), with a lobe on each side. Phallotheca re- duced (Fig. 5), membranous, apex acumi- nate, posterior region enlarged. Right par- amere very small and simple. Left paramere (Fig. 6) distally acute, with median lobe glabrous. Female (n = 3).— Length 4.69 mm (4.69 mm); width 1.54 mm (1.50—1.56 mm). Head: Length 0.55 mm (0.50—0.60 mm); width 0.66 mm (0.64—0.67 mm); distance Male genitalia of Fulvius paranaensis. 3, Vesica. 4, Secondary gonopore. 5, Phallotheca. 6, Left between eyes 0.32 mm (0.31—0.33 mm). Antenna: Segment I length, 0.49 mm (0.48— 0.50 mm); II, 1.06 mm (1.02—1.10 mm); III, 0.48 mm (0.48 mm); IV, 0.69 mm (0.62— 0.74 mm). Pronotum: Length 0.60 mm (0.60 mm); width 1.27 mm (1.24-1.31 mm). Length from cuneal fracture to apex of membrane 0.49 mm (0.48—0.50 mm); cu- neal width 0.32 mm (0.29—0.33 mm). Similar to male in color, structure, and vestiture. Type specimens.—Holotype 6, Curitiba, PR, 30/1/94, M.O. Moura (Museu Nacional, Rio de Janeiro). Paratypes: Same locality as for holotype, with following dates: 3 d: 12/ X1/1993, 30/1/1994, 20/1/1994; 3 @: 19/1/ 1994, 30/1/1994 and 27/1/1994 (all in Mu- seu Nacional and Universidade Federal de Vicosa, MG." Brazil, except’ oe 12) an National Museum of Natural History, Washington, DC). Etymology.—Named for the state of Pa- rana, Brazil, in which this species occurs. Distribution.—Curitiba, Parana, Brazil. Discussion.—According to Mr. Mauricio Moura (personal communication), the spec- imens were collected in a secondary forest at the Botanical Garden of Curitiba, feeding on the exudates of a dead animal in an ad- vanced state of decomposition. 60 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON / Figs. 7-8. Fulvius vicosensis Ferreira and Henry, new species (Figs. 7-12) Diagnosis.—Fulvius vicosensis keys to F. bisbistillatus (Stal) in Carvalho and Cos- ta (1994), but can be distinguished by the proportionately shorter second antennal segment having the apical 7 white and the broader basal width of the pronotum. It also can be distinguished from all other species of the genus by the male genitalia, partic- ularly the more elongate trough of the sec- ondary gonopore (Fig. 10), the absence of an apical notch on the phallotheca (Fig. 11), and the more bulbous base on the left par- amere (Fig. 12). Description.—Male (measurements in- clude x, followed by ranges in parentheses; n = 10): Length 3.53 mm (3.25—4.06 mm); width!.17 mm (0.94—1.38 mm). Head: Length 0.43 mm (0.33—0.52 mm); width 0.55 mm (0.52—0.61mm); distance between eyes 0.19 mm (0.17—0.21 mm). Antenna: Segment I length 0.43 mm (0.33—0.48 mm); If, 0.95 mm (0.88—1.07 mm); TH, 0.49 mm (0.47-0.59 mm); IV, 0.62 mm (0.57—0.69 Photographs of Fulvius vicosensis, 2.7, Dorsal aspect. 8, Lateral aspect. mm). Pronotum: Length 0.39 mm (0.33— 0.47 mm); width 1.27 mm (1.24—1.31 mm). Length from cuneal fracture to apex of membrane 0.36 mm (0.33—0.48 mm); cu- neal width, 0.23 mm (0.21—0.29 mm). General color brown with white areas. Head dark brown with apex of clypeus pale; rostrum pale brown, becoming fuscous api- cally; antennal segment I brown with apex paler; segment IT with apical % white, other segments dark brown. Eye dark brown. Pronotum and collar dark brown. Mesono- tum dark brown with lateral margins paler. Scutellum dark brown. Hemelytron dark brown with an irregular stripe extending through apex of clavus, basal area on co- rium and embolium yellow; base of cuneus white. Membrane darkened, slightly irides- cent, area near inner margin of cuneus pale. Legs pale brown with coxae and trochanters pale; setae short, erect or semierect. Under- side of body brown. Head, pronotum, pro- pleuron, mesoscutum, and scutellum gla- brous [or rubbed], except for short setae on apical area of head. Rostrum extending be- yond metacoxae. Antennal segment I and II VOLUME 104, NUMBER 1 61 Figs. 9-12. Left paramere. with erect or semierect setae ranging from shorter than to as long as diameter of seg- ments; setae on segments III and IV longer than diameters of their respective segments; antennal segment I longer than width be- tween eyes; antennal segment II slightly in- crassate toward apex, III and IV thinner; relative lengths of antennal segments in as- cending order I < III < IV < II. Posterior margin of pronotum concave with posterior angles rounded; collar well defined; calli occupying anterior % of pronotum. Hemel- ytron with short reclining setae. Ventral re- gion of thorax glabrous (or rubbed); abdo- men with short to relatively long simple se- tae. Male genitalia: Vesica (Fig. 9) with two fields of short spines near the secondary gonopore. Secondary gonopore (Fig. 10) short, apex acute. Phallotheca (Fig. 11) small, membranous, apical half acuminate with apex enlarged and rounded. Right par- amere small, simple. Left paramere (Fig. 12) falciform, median lobe glabrous. Female (Figs. 7, 8) (measurements in- clude x, followed by ranges in parentheses; n = 10).—Length 3.67 mm (3.81—3.84 mm); width 1.13 mm (1.00—1.19). Head: Length 0.45 mm (0.38—0.50 mm); width 0.57 mm (0.52—0.60 mm); distance between eyes 0.20 mm (0.19—-0.21 mm). Antenna: Segment I length 0.42 mm (0.36—0.48 mm); Male genitalia of Fulvius vicosensis. 9, Vesica. 10, Secondary gonopore. 11, Phallotheca. 12, II, 0.77 mm (0.76—0.98 mm); III, 0.53 mm (O.50—0.55 mm); IV, 0.71 mm (0.71 mm). Pronotum: Length 0.43. mm _ (0.40—0.45 mm); width 1.01 mm (0.93-1.05 mm). Length from cuneal fracture to apex of membrane 0.39 mm (0.36—0.45 mm); cu- neal width, 0.23 mm (0.21—0.26 mm). Similar to male in color, structure, and vestiture. Type specimens.—Holotype 6, Vigosa, Minas Gerais, Brasil, Armadilha Luminosa, Coérrego do Paraiso, 17/03/1988, P. S. E Fiuza (Museu Nacional, Rio de Janeiro) Paratypes: Same locality as for holotype, with following dates: 16 d: 23/II/1988, 09/ XI/1982, 02/11/1986, 19/1/1983, 08/1/1988, 14/1/1993, 18/11/1983, 11/1/1983, 10/XII/ 1986, 01/V1I/1983, 17/V1/82, 08/1/1988, 02/ L/19835 -22/XIV/1982); OS/MUTSS 7ale 8 : 20/1/1982, 25/11/1982, 31/11/1987, 11/11I/ 1987, 13/1/1987, 18/1/1988, 08/1/1993, 08/ 1/1988, 23/0/1988, 25/11/1986, 02/11/1983, 1O/IV/1987, 23/V1/1982, 20/1V/1983, 12/1I/ 1979, 24/11/1983 (all paratypes in Museu Nacional and Universidade Federal de Vi- cosa, MG, Brazil, except for 3 2 in Na- tional Museum of Natural History, Wash- ington, DC). Etymology.—Named for the city of Vi- cosa, Brazil, in which this species occurs. 62 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Distribution.—Vicosa, Minas Gerais, Brazil. Discussion.—The paratypes compare well with the holotype, varying in general color from paler to slightly darker. The specimens were collected in black light traps in a secondary forest at Vigosa known as ‘“‘Mata do Coérrego do Paraiso (20°46’— 20°48’S; 45°50’—45°52’W),”’ characterized by Atlantic Forest vegetation, at an altitude of 600—700 meters, an annual average tem- perature of 18° C to 19° C, and an annual average precipitation of about 131.5 cm. ACKNOWLEDGMENTS We express our appreciation to Antonio Fernando Gomes (Universidade do Vicosa, Brazil) for assistance with certain drawings, Mauricio O. Moura (Universidade Federal do Parana, Brazil) for sending specimens of F. paranaensis and additional biological in- formation, and FAPEMIG for financial sup- port to PSFE We also are grateful to Mich- ele Touchet (Systematic Entomology Lab- oratory [SEL], Agricultural Research Ser- vice, USDA, % National Museum of Natural History, Washington, DC) for the photographs of the adults, and R. C. Froes- chner (National Museum of Natural Histo- ry, Smithsonian Institution, Washington, DC), J. W. Brown (SEL), and D. R. Smith (SEL) for reviewing the manuscript. LITERATURE CITED Carvalho, J. C. M. 1954. Neotropical Miridae, LX XIV: Two new genera of Cylapinae from Brazil (He- miptera). Proceedings of the Iowa Academy of Science 61: 504-510. 1956. Insects of Micronesia. Heteroptera: Miridae. B. P. Bishop Museum, Honolulu 7(1): 1—100. Carvalho, J. C. M. and L. A. A. Costa. 1994. The Genus Fulvius from the Americas (Hemiptera: Miridae). Anales Instituto Biologia, Universidad Nacional Aut6noma México, Ser. Zool. 65(1): 63— 135. Carvalho, J. C. M. and P. S. EF Ferreira. 1994. Mirideos Neotropicais, CCCLXXXVIII: Chave para os gé- neros de Cylapinae Kirkaldy, 1903 (Hemiptera). Revista Ceres 41(235): 327-334. Gorezyca, J. 2000. A systematic study on Cylapinae with a revision of the Afrotropical Region (Het- eroptera, Miridae). Wydawnictwo Uniwersytetu Saskiego, Katowice. 174 pp. Henry, T. J. and A. G. Wheeler, Jr. 1988. Family Mir- idae Hahn, 1833 (= Capsidae Burmeister, 1835). The plant bugs, pp. 251-507. Jn Henry, T. J. and R. C. Froeschner, eds. Catalog of the Heteroptera, or true bugs, of Canada and the Continental Unit- ed States. E.J. Brill Publ, Leiden and New York. 958 pp. Kelton, L. A. 1985. Species of the genus Fulvius Stal found in Canada (Heteroptera: Miridae: Cylapi- nae). Canadian Entomologist 117: 1071—1073. Maldonado Capriles, J. 1969. The Miridae of Puerto Rico (Insecta, Hemiptera). University of Puerto Rico Agricultural Experiment Station Technical Paper 45: 1-133. Paula, A. S. and P. S. F Ferreira. 1998. Fauna de Het- eroptera de la ““Mata do Cérrego do Paraiso,” Vi- cosa, Minas Gerais, Brasil. I. Riqueza Y diversi- dad especificas. Anales Instituto Biologia, Univ- ersidad Nacional Autonoma México, Ser. Zool. 69 G)e39>si" Schmitz, G. and P. Stys. 1973. Howefulvius elytratus gen. n., sp. n. (Heteroptera, Miridae, Fulviinae) from Lord Howe Island in the Tasman Sea. Acta Entomologica Bohemoslovaca 70: 400—407. Schuh, R. T. 1976. Pretarsal structure in the Miridae (Hemiptera) with a cladistic analysis of relation- ships within the family. American Museum Noy- itates 2601: 1—39. . 1995. Plant bugs of the world (Insecta: Het- eroptera: Miridae): Systematic catalog, distribu- tions, host list and bibliography. New York En- tomological Society, New York. 1329 pp. Stal, C. 1862. Hemiptera Mexicana enumeravit spe- cies-que novas descripsit. Stettin Entomologische Zeitung 23(7—9): 289-325. PROC. ENTOMOL. SOC. WASH. 104(1), 2002, pp. 63-72 A REVIEW OF THE NEW RIODINID BUTTERFLY GENUS PANAROPSIS (LEPIDOPTERA: RIODINIDAE: SYMMACHIINI) JASON P. W. HALL Department of Systematic Biology—Entomology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0127, U.S.A. Abstract.—A new riodinid genus Panaropsis, tribe Symmachiini, is described and il- lustrated from Central and South America. Four species are recognized: elegans Schaus and semiota Bates transferred from Pterographium Stichel (n. comb.), and thyatira Hew- itson and inaria Westwood transferred from Esthemopsis C. and R. Felder (n. comb.). Key Words: The purpose of this paper is to describe and characterize a new riodinid genus in the tribe Symmachiini to provide a generic name for elegans Schaus 1920, semiota Bates 1868, inaria Westwood [1851] and thyatira Hewitson [1853], currently treated in Pterographium Stichel 1910 and Esthem- opsis C. and R. Felder 1865, for a forth- coming morphological survey of male an- droconia in the Riodinidae (Hall and Har- vey, in press). The tribe Symmachiini has been adequately diagnosed elsewhere (Har- vey 1987, Hall and Willmott 1996) and this information is not repeated here. METHODS Dissections were made using standard techniques, after abdomens were soaked in hot 10% potassium hydroxide solution for approximately five minutes, and subse- quently stored in glycerol. The terminology for male and female genital and abdominal structures follows Klots (1956) and Eliot (1973), while nomenclature for wing ve- nation follows Comstock and Needham (1918). The taxonomic status of names 1s based on the catalog of Callaghan and La- mas (in press). All the collections listed by Hall (1999) Esthemopsis, Neotropical, Pterographium, Panaropsis, Symmachiini were examined to determine the ranges of Panaropsis species. The following collec- tion acronyms are used throughout the text: AME: Allyn Museum of Entomology, Flor- ida Museum of Natural History, Sarasota, FL, U.S.A.; BMNH: The Natural History Museum, London, England; MNRJ: Museu Nacional, Rio de Janeiro, Brazil; MUNB: Museo de la Universidad Nacional, Bogota, Colombia; USNM: National Museum of Natural History, Smithsonian Institution, Washington, DC, U.S.A.; ZMHU: Zoolo- gisches Museum fiir Naturkunde, Humboldt Universitat, Berlin, Germany. SYSTEMATICS Panaropsis Hall, new genus (Figs. 1-8, 9, 10-14, 15-17, 18) Type species.—Panara elegans Schaus 1920: Etymology.—The name alludes to the superficial similarity between members of the riodinine genus Panara and the type species elegans, which was formerly placed in Panara. Description.—Male: Forewing length 17-20 mm. Wing shape: both wings typi- cally somewhat elongate; forewing costa very slightly arched at base, distal margin 64 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-8. Adults (dorsal surface on left, ventral surface on right). 1, Pterographium sicora, male, Brazil, Espirito Santo, Linhares [AME]. 2, Panaropsis semiota, male, French Guiana, Cayenne, Galion [USNM]. 3, P. elegans, male, Panama, Panama, nr. El Llano [USNM]. 4, P. elegans, female, Panama, Col6n, Colén [USNM]. 5, P. thyatira, male, Brazil, Mato Grosso, Diamantino [USNM]. 6, P. thyatira, female, Brazil, Mato Grosso, Colegio Buriti [USNM]. 7, P. inaria, male, Brazil, Pernambuco, Sao Lourengo [USNM]. 8, P. inaria, female, Brazil, Pernambuco, Sao Lourencgo [USNM]. VOLUME 104, NUMBER 1 Fig. 9. foreleg. E, Female foreleg. EK Male hindleg. slightly convex; hindwing slightly pointed at apex and tornus. Venation (Fig. 9A): four forewing radial veins. Dorsal surface: ground color of both wings black; forewing with a diagonal yellow or red postdiscal band and hindwing black or with iridescent blue patch (semiota and elegans) or both wings with a large orange patch (thyatira and inaria); patch of erectile androconial setae at base of hindwing cell Cu2, dense in semiota and elegans, diffuse in thyatira and inaria; fringe on both wings black. Ventral surface: same as dorsal surface but paler and with no iridescence. Head: labial palpus entirely black (semiota and elegans) or with white scaling ventrally (thyatira and inaria), second and third segments short (Fig. 9B); eye brown and bare, black or white scaling at margins; frons black with white lateral scaling dorsally in thyatira and inaria; antennal length approximately 70% of forewing length, segments black without white scaling at base; club long and black, tips often orange brown. Body: thorax and abdomen broad and robust; dorsal and ven- tral surface of thorax and abdomen black in 65 Morphology of Panaropsis elegans. A, Male venation. B, Male palpus. C, Female palpus. D, Male semiota and elegans, tegula black with var- iably prominent red anterior spot in ele- gans; dorsal surface of thorax black with orange tegula and orange lateral scaling, ventral surface black with patches of white scaling in thyatira and inaria, dorsal sur- face of abdomen black with remainder or- ange in thyatira and inaria, also black scal- ing ventrally in inaria; a narrowly medially divided band of concealed androconial scales on dorsal half of anterior margin of abdominal tergites four and five; tarsus of foreleg trimerous, coxa of medium length for family (Fig. 9D); legs black, midleg and hindleg with a tibial spur, a scattered group of spines along distal inner margin of tibia and two dense rows of spines along distal inner margin of tarsal segments (Fig. 9F). Genitalia (Figs. 10—14): overall large; un- cus rectangular and produced into small medial point dorsally, falces and tegumen of average size and shape for family; vin- culum evenly narrow with medium-sized rounded saccus ventrally; aedeagus short and very broad, everted vesica contains a straight row of approximately ten large an- 66 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON teriorly directed spines ventrally that be- come gradually shorter posteriorly, a large patch of large anteriorly directed spines dorsally that is slightly ventrally curved an- teriorly and a sparse lateral patch of very small spines; pedicel short, narrow and strap-like; valvae curl ventrally and slightly overlap, upper portion rectangular, lower portion produced into one or two small rounded projections, setae on valvae typi- cally only on upper rectangular portion and small raised ridge at middle. Female: Differs externally from male in following ways: forewing length 19—25 mm. Both wings more rounded (and elon- gate in thyatira and inaria); ground color of both wings paler; semiota and elegans not significantly sexually dimorphic, fe- males with no dorsal blue iridescence, broader forewing orange bands and white fringe elements on hindwing and in fore- wing apex; thyatira and inaria strongly sex- ually dimorphic, females with yellow or or- ange pattern elements divided into subapi- cal and basal areas on forewing, white sub- marginal hindwing spots present, prominent white hindwing fringe elements present in thyatira, some lateral black scaling present on each abdominal segment. Head: second segment of labial palpus slightly broader (Fig. 9C). Body: foreleg with spines at inner distal tip of tibia and tarsal segments one to four, large ovoid patch of sensilla on swol- len last tarsal segment (Fig. 9E). Genitalia (Figs. 15-17): corpus bursae elongate and slightly medially constricted, no signa pre- sent; ductus bursae short, broad and some- what sclerotized with tiny spines lining in- ner surface, becoming suddenly broader and less sclerotized anterior to broad open- ing of ostium bursae which is positioned in desclerotized invagination between eighth and seventh abdominal sternites; eighth sternite sclerotized and slightly concave medially. Diagnosis.—Having examined the inter- nal male abdominal morphology of approx- imately 95% of species in the tribe Sym- machiini (males are not known for several of the remainder) for a paper surveying male androconia in the Riodinidae (Hall and Harvey, in press), it became clear that four species which were currently treated in Pterographium (semiota and elegans) and ~Esthemopsis (thyatira and inaria) were more closely related to each other than to any other symmachiine species. These four species, which I place in the new genus Panaropsis, present a rather heterogeneous external appearance, but their morphology is very homogeneous. The only external similarities are their relatively large size for riodinds, somewhat elongate wing shapes and robust thoraces and abdomens, whereas the only interspecific genital variation lies in differences in the shapes of the valvae and dorsal cornutal band in males. There are two unique male genital synapomor- phies for Panaropsis. One is the arrange- ment of the aedeagal cornuti, with a straight row of large spines ventrally, a scattered patch of very small spines laterally and an anteriorly recurved dense band of large spines dorsally. The other is the shape of the valvae, which are curled ventrally to partially overlap and posses a lower portion produced into one or two small “‘finger- like” projections. Systematic position.—Although two Panaropsis species, semiota and elegans, externally closely resemble the sole mem- ber of Pterographium, sicora, the male gen- italia differ substantially in almost every re- spect (see Figs. 10-12), suggesting that the two genera may not be closely related. Giv- en the prevalence of aposematic and puta- tively mimetic coloration in the tribe, such wing pattern convergence would not be sur- prising. It is presently not certain what the closest relatives of Panaropsis are. Only a few Symmachia Hiibner [1819] species, all Stichelia Zikan 1949, one Mesene Double- day 1847, two Xenandra C. and R. Felder 1865, and one Esthemopsis possess medi- ally divided concealed abdominal andro- conia on segments four and five as in Pan- aropsis, but none of these taxa closely re- semble Panaropsis species in external ap- VOLUME 104, NUMBER 1 pearance or morphology. I tentatively place Panaropsis near Pterographium and _ Pir- ascca Hall and Willmott 1996, based on the presence in males of dense erectile andro- conial setae at the anal margin of the dorsal hindwing (also found in several unrelated Symmachia species) and similarities in the ultrastructure of the concealed male abdom- inal androconial scales (the ultrastructure of these scales has been examined for repre- sentatives from each symmachiine genus using scanning electron microscopy), which are elongate, narrow and apically curved in all three genera (see Fig. 18; also illustra- tion in Hall and Willmott (1996) for Pir- ascca). Those of Pterographium and Pir- ascca are particularly similar to each other. Proposed classification —Below is pre- sented a synonymic checklist for Panarop- KYAY Panaropsis Hall, n. gen. elegans (Schaus, 1920), n. comb. = chocoensis (Callaghan, 1999), n. syn. inaria (Westwood, [1851]), n. comb. = barca (Hewitson, [1853]) semiota (Bates, 1868), n. comb. = similatum (Zikan, 1949) thyatira (Hewitson, [1853]), n. comb. = isabellae (Sharpe, 1890) = perfluxa (Stichel, 1924) phlegontis (Stichel, 1910) = sarta (Stichel, 1924) Biology.—Panaropsis species are un- common to rare inhabitants of wet lowland forests below 1000 m, and little is known of their biology. Males are typically en- countered perching in hilltop forest light- gaps, often very high above the ground, and their flight is rapid. Both sexes of P. ele- gans have been recorded nectaring on Ter- minalia and Croton flowers (DeVries 1997). All species are brightly and apose- matically colored and resemble a number of distasteful moths in the families Arctiidae and Notodontidae. Distribution.—Panaropsis is a pan-Neo- 67 tropical genus whose species occur from Mexico to western Ecuador, throughout the Guianas and Amazon basin, and into south- eastern Brazil. Panaropsis semiota (Bates 1868), n. comb. (Figs. 2159) Limnas semiota Bates 1868: 380—381. Type locality: Alter do Chao, Rio Tapajos, E. Brazil. Syntype male BMNH [examined]. Pterographium similatum Zikan 1949: 536— 538, figs. 1-3. Type locality: Sao Gabriel, Rio Negro, N. Brazil. Holotype male and allotype female MNRJ [type illustrations examined ]. Identification and taxonomy.—Typical FW length: male 17 mm; female 18 mm (Zikan 1949). The taxon semiota was long treated in the riodinine genus Melanis Htib- ner [1819] (Stichel 1910, 1930—31; Bridges 1994), but was transferred, along with the newly synonymized similatum, to the monotypic Pterographium by Hall and Willmott (1996), on the basis of external similarities with the type species of that ge- nus, sicora. Panaropsis semiota differs ex- ternally from the southeastern Brazilian Pterographium sicora only by lacking prominent blue dorsal iridescence and or- ange scaling at the tip of the abdomen, and by possessing a more robust thorax and ab- domen. However, despite the remarkable external similarities, the male genitalia are completely distinct (see Figs. 10 and 11). Panaropsis semiota differs from its sister species P. elegans by its smaller size, more elongate wing shape and more distally po- sitioned postdiscal forewing band, by lack- ing an iridescent blue patch on the dorsal hindwing, and in the male genitalia by hav- ing a slightly shallower indented notch be- tween the upper and lower portions of the valvae, a lower valve portion with one in- stead of two small projections and slightly smaller cornutal spines. Biology.—This very rare species is only known from a handful of specimens in col- 68 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 10—14. thyatira. 14, P. inaria. Male genitalia. 10, Pterographium lections. Brévignon and Gallard (1998) re- port finding males in French Guiana perch- ing on forested hilltops between 1330 and 1430 h on top of leaves 3 to 4 m above the ground; their flight is rapid. Distribution.—Brazil (central and lower Amazon) and French Guiana. Panaropsis elegans (Schaus 1920), n. comb. (Figs. 3—4, 9, 12, 15) Panara elegans Schaus 1920: 108. Type lo- cality: Cayuga, Guatemala. Syntype males (7) and females (5) USNM [ex- amined]. sicora. 11, Panaropsis semiota. 12, P. elegans. 13, P. Pterographium elegans chocoensis Calla- ghan 1999: 1053, 1055, figs. 25—28. Type locality: Aguas Claras, Rio Anchicaya, W. Colombia. Holotype male MUNB [type illustrations examined]. n. syn. Identification and taxonomy.—Typical FW length: male 18 mm; female 21 mm. The taxon elegans was described (Schaus 1920) and subsequently treated (Stichel 1910, 1930-31; Bridges 1994) in the genus Panara Doubleday 1847, in the tribe Riod- inini. Harvey (1987) discovered that males of elegans possessed concealed abdominal androconia, placing it in the tribe Symma- VOLUME 104, NUMBER I 1S Figs. 15-17. chiini, but he did not transfer it to any other genus. Callaghan (1997) transferred elegans from Panara to the symmachiine genus Pterographium on the basis of it sharing ‘erectile scent hairs in cell Cu2-2A of the dorsal hindwing’’ with the type species of that genus. However, certain members of Symmachia and Pirascca also possess such hair tufts, and the fact that some members of Panaropsis possess such clearly defined hair tufts while others do not (thyatira and inaria) indicates this structure is homopla- sious even at the species-group level, and thus not a reliable generic character. Both sexes of P. elegans possess com- mon riodinid wing patterns. Within the Symmachiini, males somewhat resemble 69 Female genitalia in dorsal view. 15, Panaropsis elegans. 16, P. thyatira. 17, P. inaria. Esthemopsis pherephatte (Godart [1824]) and Pterographium sicora (Hewitson 1875), but their large size, rounded wing shape and prominent iridescent dorsal hindwing blue patch are distinctive. The fe- male of P. elegans resembles those of many species in genera such as Hypophylla Bois- duval 1836, Panara, Pirascca, Setabis Westwood [1851] and Symmachia, but its large size, entirely brown ventral surface and largely white hindwing fringe are di- agnostic. Panaropsis elegans varies slightly geo- graphically. Males from Mexico to Costa Rica typically have dorsal hindwing blue restricted to an ovoid patch in the distal third of the wing and a forewing orange 70 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON band that extends to the tornus, whereas in males from Costa Rica eastwards the blue tends to extend more proximally to the dis- cal cell and the forewing orange band does not reach the tornus. The width of the fore- wing band and its color, which varies from orange to red, and the prominence of red scaling at the anterior tip of the tegula is variable throughout the range of the spe- cies. Callaghan (1999) described the sub- species chocoensis from a small series of males and females from a single locality in western Colombia on the basis of the male forewing band being red instead of orange. However, since red-banded males occur rarely throughout the range of P. elegans (e.g., see the illustrations in d’Abrera (1994) and DeVries (1997) of specimens from Nicaragua and Costa Rica respective- ly), I synonymize chocoensis with elegans. Biology.—Panaropsis elegans is an un- common inhabitant of lowland rainforest from sea-level to 900 m. DeVries (1997) reports finding males in Costa Rica as sol- itary individuals or in small groups perch- ing along forest edges, trails, streams and in shaded forest light gaps; individuals perched between 2 and 5 m above the ground under leaves from 0800 to 1230 h and their flight was fast and erratic. Calla- ghan (1999) reports finding males in Co- lombia perching on hilltops from 1000 to 1200 h under leaves with wings outspread 2 to 3 m above the ground. Both sexes visit flowers of Terminalia and Croton (DeVries 1997). Distribution.—Mexico to western Ecua- dor. Panaropsis thyatira (Hewitson [1853]), n. comb (Figs. 5—6, 13, 16) Limnas thyatira Hewitson [1853]: pl. 59, fig. 6. Type locality: ““Amazon’’. Syntype female BMNH [examined]. Lymnas isabellae Sharpe 1890: 571, pl. 46, fig. 3. Type locality: Rio Araguaia, S. Brazil. Syntype male BMNH [examined]. Lymnas thyatira phlegontis Stichel 1910: Fig. 18. cealed male abdominal androconial scales. Scale bar = 20 pm. Scanning electron micrograph of con- 205. Type locality: Rio Songo, Bolivia. Holotype female ZMHU [examined]. Limnas inaria thyatira f. perfluxa Stichel 1924: 2. Type locality: Santarém, E. Bra- zil. Holotype male ZMHU [examined]. Limnas inaria_ thyatira f. sarta Stichel 1924: 2. Type locality: Santarém, E. Bra- zil. Holotype female ZMHU [examined]. Identification and taxonomy.—Typical FW length: male 17 mm, female 22 mm. The taxon thyatira was described (Hewit- son 1852—54) and long treated as a species in Limnas Boisduval 1836 (now a synonym of Melanis), and transferred to Esthemopsis as a subspecies of inaria by Stichel (1924). It has subsequently remained in Esthem- opsis and been treated variably as a sub- species of inaria (Bridges 1994) or as a full species (d’Abrera 1994; Callaghan and La- mas, in press). I follow Callaghan and La- mas (in press) in regarding fthyatira as a species distinct from inaria because of the substantial differences in the wing pattern and shape of the females and the fact that each species occupies a distinct biogeo- graphic zone (i.e., the Amazon basin and southeastern Brazil). The male of P. thyatira differs from that of P. inaria by having orange on the hind- wing that extends to the anal margin and predominantly orange instead of black scal- ing ventrally on the abdomen. The female has a broader wing shape, with orange in- VOLUME 104, NUMBER | stead of yellow patterning that forms rays on the forewing. The male and female gen- italia of both species do not differ. The male genitalia of thyatira and inaria differ from those of semiota and elegans only by hav- ing a dorsal cornutal patch that contains narrower spines and is restricted to a nar- row band on the left-hand side without the additional contiguous posterior patch on the right-hand side (see Figs. 12 and 13). Both sexes exhibit some wing pattern variation, which has led to the description of several unwarranted subspecific and in- frasubspecific names. In both sexes, the ex- tent of distal orange on both wings is var- iable, and in males a black interneural line in cell Cu2, as in the female, and submar- ginal white spots on the ventral hindwing may be present or absent. Biology.—Brévignon and Gallard (1998) report finding males of this uncommon spe- cies in French Guiana perching on hilltops between 1530 and 1645 h on top of leaves 5 m above the ground. In Ecuador, males were similarly encountered singly or in small groups perching on a forested hilltop 10 to 15 m above the ground on top of sun- lit leaves with their wings outspread (Hall and Willmott, unpubl. data). Notably, the female resembles a number of nymphalid heliconiine species, such as Dryas julia (Fa- bricius 1775), Eueides lineata Salvin 1868 and Eueides aliphera (Godart 1819). Distribution.—Colombia to Bolivia, Bra- zil (Amazon and southern Brazil) and the Guianas. Panaropsis inaria (Westwood [1851]), n. comb. (Figs. 7-8, 14, 17) Limnas inaria Westwood [1851]: 460, pl. (3, iee 7. Lype locality: “Brazil” .. Syn- type male BMNH [examined]. Limnas barca Hewitson [1853]: pl. 60, fig. 12. Type locality: Rio de Janeiro, S.E. Brazil. Syntype female BMNH [exam- ined]. Identification and taxonomy.—Typical FW length: male 17 mm, female 21 mm. val The taxon inaria was also described (West- wood 1850—52) and long treated as a spe- cies in Limnas and transferred to Esthem- opsis by Stichel (1910). Panaropsis inaria is distinguished from its sister species, P. thyatira, in the previous species account. Note that the substantial sexual dimorphism in both these species led to the description of each sex as separate species. Panaropsis inaria and P. thyatira externally most closely resemble Machaya obstinata Hall and Willmott 1995, from the Andes of east- ern Ecuador, but that species has reduced orange on both wings, an entirely black ventral hindwing and a black thorax and ab- domen. Biology.—Nothing is known about the biology of this uncommon species. Distribution.—Southeastern Brazil (Rio de Janeiro to Pernambuco). ACKNOWLEDGMENTS I thank Lee and Jackie Miller (AME), Phillip Ackery (BMNH), Wolfram Mey (ZMHU), and Donald Harvey and Robert Robbins (USNM) for access to riodinid col- lections and the loan of specimens, and Paul Opler for comments on the manu- script. I gratefully acknowledge The Na- tional Geographic Society (Research and Exploration Grant #5751-96) and The Smithsonian Institution (two Postdoctoral Fellowships) for financial support of field and museum research. LITERATURE CITED Bates, H. W. 1868. A catalogue of Erycinidae, a family of diurnal Lepidoptera. Journal of the Linnean So- ciety of London (Zoology) 9: 373—459. Brévignon, C. and J.-Y. Gallard. 1998. Inventaire des Riodinidae de Guyane Frangaise 1V—Riodinidae: Symmachiini, Charitini, Helicopini. Description de nouveaux taxa (Lepidoptera). Lambillionea 98(2): 304-320. Bridges, C. A. 1994. Catalogue of the Family-Group, Genus-Group and Species-Group Names of the Riodinidae and Lycaenidae (Lepidoptera) of the World. C. Bridges, Urbana, IL. 1113 pp. Callaghan, C. J. 1997. A review of the genus Panara Doubleday, 1847 (Riodinidae) in southeast Brazil, 7 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON with a d iption of two new subspecies. Journal of R on the Lepidoptera 34: 21-38. _— 199 Jew taxa of Neotropical Riodinidae Lepidoptera). Revista Brasileira de Zoologia 1): 1045-1064. Callaghan, C. J. and G. Lamas. In press. Riodinidae. In Lamas, G., ed. A Checklist of the Neotropical Butterflies and Skippers (Lepidoptera: Papiliono- idea and Hesperioidea). Atlas of Neotropical Lep- idoptera (ed. by J. B. Heppner). Scientific Pub- lishers, Gainesville, FL. Comstock, J. H. and J. G. Needham. 1918. The wings of insects. American Naturalist 32(376): 231—257. d’Abrera, B. 1994. Butterflies of the Neotropical Re- gion, Part VI. Riodinidae. Hill House, Victoria, Australia, pp. 880—1096. DeVries, P. J. 1997. The Butterflies of Costa Rica and their Natural History. Volume II: Riodinidae. Princeton University Press, Princeton, NJ. 288 pp. Eliot, J. N. 1973. The higher classification of the Ly- caenidae (Lepidoptera): A tentative arrangement. Bulletin of the British Museum of Natural History (Entomology) 28(6): 373-506. Hall, J. P. W. 1999. A Revision of the Genus Theope: Its Systematics and Biology (Lepidoptera: Riodi- nidae). Scientific Publishers, Gainesville, FL. 127 Pp. Hall, J. P. W. and D. J. Harvey. In press. A survey of androconial organs in the Riodinidae (Lepidop- tera). Zoological Journal of the Linnean Society: in press. Hall, J. BP W. and K. R. Willmott. 1996. Systematics of the riodinid tribe Symmachiini, with the de- scription of a new genus and five new species from Ecuador, Venezuela and Brazil (Lepidoptera: Riodinidae). Lambillionea 96: 637—660. Harvey, D. J. 1987. The Higher Classification of the Riodinidae (Lepidoptera). University of Texas Austin. Ph.D. Dissertation. 216 pp. Hewitson, W. C. 1852-1854. Illustrations of New Spe- cies of Exotic Butterflies, Selected Chiefly from the Collections of W. Wilson Saunders and Wil- liam C. Hewitson. 2(2—10). J. Van Voorst, Lon- don. Klots, A. B. 1956. Lepidoptera, pp. 97-110. /n Tuxen, S. L., ed. Taxonomists’ Glossary of Genitalia in Insects. Munksgaard, Copenhagen. Schaus, W. 1920. New species of Lepidoptera in the United States National Museum. Proceedings of the United States National Museum 57(2307): 107-152. Sharpe, E. M. B. 1890. Collection of Lepidoptera made by Mr. Edmund Reynolds on the rivers To- cantins and Araguaya and in the province of Goy- az, Brazil. Proceedings of the Zoological Society of London 1890(3): 552-577. Stichel, H. FE E. J. 1910-11. Family Riodinidae. All- gemeines. Subfamily Riodininae, pp. 1—452. In Wytsman, P., ed. Genera Insectorum 112A-B. P. Wytsman, Brussels. . 1924. Zur Systematik der Riodinidae (Lep., Rhop.). Neue Beitrage Systematischen Insekten- kunde® 3(1): 1—3: . 1930-31. Riodinidae., pp. 1-795. Jn Strand, E., ed. Lepidopterorum Catalogus 38—41. W. Junk, Berlin. Westwood, J. O. 1850-52. Jn Doubleday, E., ed. The Genera of Diurnal Lepidoptera: Comprising their Generic Characters, a Notice of their Habits and Transformations, and a Catalogue of the Species of each Genus. Longman, Brown, Green and Longmans, London. 534 pp. Zikan, J. EK 1949. Observagoes sobre os componentes dos géneros Phaenochitonia Stichel e Pterogra- phium Stichel, coma descrigao de uma nova es- pécie e criando um novo género (Riodinidae, Lep- idoptera). Revista de Entomologia 20(1/3): 535— 539). PROC. ENTOMOL. SOC. WASH. 104(1), 2002, pp. 73-78 NOTES ON THE RESPONSES OF HOST-SEEKING NYMPHS AND ADULTS OF THE TICKS IXODES SCAPULARIS AND AMBLYOMMA AMERICANUM (ACARI: [IXODIDAE) TO CANINE, AVIAN, AND DEER-PRODUCED SUBSTANCES J. E CARROLL U.S. Department of Agriculture, Agricultural Research Service, Livestock and Poultry Sciences Institute, Parasite Biology, Epidemiology and Systematics Laboratory, Beltsville, MD 20705-2350, U.S.A. (e-mail: jcarroll @anri.barc.usda.gov) Abstract.—In laboratory and behavioral bioassays, host-seeking nymphs and adults of the blacklegged tick, Ixodes scapularis Say, were exposed to substances rubbed from the coats of dogs, from chicken feathers and from the interdigital glands of white-tailed deer, Odocoileus virginianus (Zimmermann). Nymphs of the lone star tick, Amblyomma amer- icanum (L.) were exposed to substances from chicken feathers. Nymphs of /. scapularis became akinetic on surfaces rubbed with substances from chicken feathers, but A. amer- icanum nymphs did not. Overall, nymphs of /. scapularis appeared to avoid substances rubbed from the coats of dogs. Although hair samples were obtained from 14 dogs main- tained under the same conditions, substances from the hair of one dog elicited arrestant responses among nymphs and substances from the hair of another dog appeared to com- pletely repel 7. scapularis nymphs. Adult /. scapularis of both sexes showed high levels of arrestment when exposed to substances from the same canine hair samples. No arrest- ment was observed when /. scapularis nymphs were exposed to interdigital gland sub- stances of white-tailed deer does. Key Words: The host-finding behavior of ixodid (hard) ticks can be divided into two general types, ambushing and hunting (Waladde and Rice 1982). Typically, an ambushing tick waits on or near a vantage point, such as a twig or blade of grass, until a suitable host brushes against the tick’s outstretched legs and it catches hold. Ticks that are con- sidered hunters also spend much time wait- ing for a host, but when they detect the presence of a suitable host from chemical, visual, or other physical stimuli they leave their vantage point or shelter and move to- ward the source of the stimulus. Many spe- cies of ticks employ some of both types of behavior. The host-seeking behavior of the kairomones, blacklegged ticks, lone star tick blacklegged tick, /xodes scapularis Say, tends to be more of the sedentary ambush- ing sort, whereas lone star ticks, Ambly- omma americanum (L.), are strongly at- tracted to carbon dioxide that emanates from hosts during respiration (Wilson et al. 1972) and may be best described as hunters. Adult ticks of various species, such as I. scapularis, I. neitzi Clifford, Walker and Keirans, /. persulcatus Schulze, A. ameri- canum, and Dermacentor variabilis (Say) appear to have some ability to select opti- mum vantage points for ambushing hosts. These adult ticks may locate surfaces (e.g., grass stems) coated with substances rubbed from the pelage or external glands of hosts, 74 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON residues of host urine, and or coated wiil then remain on or near these surfaces (Re- chav et al. 1978; Carroll et al. 1995, 1996; Dobrotvorsky et al. 2000). Because many host animals repeatedly use the same trails, feeding and bedding areas, ticks that situate themselves in high host-use locations im- prove their odds of acquiring a suitable host. Adults of the relatively sedentary /. scapularis have been reported to disperse by their own locomotion as far as 10 m over a period of 3—4 wk (Carroll and Schmidt- mann 1996), and may thereby encounter host-produced kairomonal residues. In contrast to adult ticks, little is known about whether host-produced chemicals in- fluence where nymphal ticks wait for hosts. The immature stages of /. scapularis feed on a wide range of vertebrate species, in- cluding birds and lizards (Spielman et al. 1985, Durden and Keirans 1996, Keirans et al. 1996). Although adult A. americanum, like adult /. scapularis, are strongly asso- ciated with white-tailed deer, O. virginianus (Zimmerman), their immatures will also parasitize a variety of hosts, and have been reared successfully on domestic chickens (Koch 1983). Little has been reported about nymphal tick attraction to or arrestment on substances from the coats or glands of hosts or to host urine. Arrestment occurs when a tick engaged in ambulatory activity be- comes akinetic (Sonenshine et al. 1986) as a behavioral response to certain stimuli (e.g., host-produced kairomone). In labora- tory bioassays, nymphs of J. scapularis did not show an arrestant response to urine ex- creted by white-footed mice, Peromyscus leucopus (Rafinesque), a principal host of immature blacklegged ticks (Carroll 1999a). However, nymphs of a similar spe- cies, the taiga tick, /. persulcatus, were at- tracted to kairomonal constituents in canine hair (Naumov and Vitlin 1995). Nymphs of A. americanum are readily attracted to sources of carbon dioxide (Wilson et al. 1972), but their responses to other host-pro- duced chemicals are poorly understood. The purpose of this research was to evalu- ate some host-produced substances for in- ducing arrestant activity in nymphs of J. scapularis and A. americanum, and thereby gain insight as to whether the nymphs of these two medically important species might choose host ambush sites by means of chemical cues. MATERIALS AND METHODS Ticks.—Nymphs of /. scapularis and A. americanum were collected by flagging during May and June in Prince George’s County and Queen Anne’s County, Mary- land. Ticks were maintained at 22°C, 97-— 99% RH and a photoperiod of 16:8 (L:D) h. Adult 7. scapularis were collected during the fall in Prince George’s County and maintained at 7°C, 97-99% RH and a pho- toperiod of 11:13. (L:D) h. Individual ticks used in bioassays were chosen based on their physical integrity (no missing appendages) and motility when re- moved from storage vials. Substances.—To obtain samples of sub- stances associated with avian hosts, feathers were removed from 2 male 3-wk old white leghorn chickens. Feathers were removed from the back and wings of the chickens, placed in sealed plastic bags and stored at —15°C. Canine coat substances were ob- tained from hair clipped from the flanks of 14 mature dogs (4 male, 10 female) kept under uniform conditions and fed identical diets by Auburn University School of Vet- erinary Medicine, Auburn, Alabama. The hair samples were placed directly in sealed plastic bags, which were kept at —15°C. The interdigital glands of white-tailed deer are located between the toes of the deer, and their secretions are deposited on the sub- strate and leaf litter. Interdigital gland sam- ples were obtained from hunter-killed deer at a hunter check-in station in Anne Arun- del County, Maryland. The hind legs of deer were removed just above the tarsal joint, sealed in plastic bags and frozen at = @) Bioassays.—A | pl micropipette was in- serted vertically in each end of a rectan- VOLUME 104, NUMBER 1 Table 1. US Numbers of /. scapularis and A. americanum nymphs on micropipettes rubbed with chicken feather substances 24 h after nymphs were released on clay islands with treated and untreated micropipettes*. Treatment Control x? {z I. scapularis 19 6.76 <0.05 A. americanum 13 0.16 >0.05 “ Thirty nymphs of each species of tick were tested against mixed feather samples from chickens. gular clay block (2.5 cm long, | cm wide, 1 cm high) centered in a plastic petri dish (3.5 cm diameter, | cm high) containing water. The petri dish with the micropipettes was placed in a larger petri dish (9 cm di- ameter, | cm high) containing water, which was in turn placed in a transparent Plexiglas glove box (65 by 85 by 45 cm) containing water 0.5 cm deep. The water confined ticks to the clay islands and provided a high rel- ative humidity (~95%) in the glove box. Capillary tubes (1 mm diameter, 10 cm high) were substituted for the micropipettes when adult ticks were tested. Feathers or canine hair samples were rubbed between the thumb and forefingers of a vinyl-gloved hand, after which the api- cal half of one micropipette was rubbed be- tween the gloved fingers. The micropipette at the opposite end of the clay island was rubbed similarly between the thumb and the forefinger of a clean glove. When adult J/. scapularis were tested, only the apical | cm of the capillary tube was rubbed. About 5 min after interdigital gland sam- ples were removed from the freezer, they were warmed with a hair dryer for ~1 min. Glandular substances were transferred by rubbing the thumb and forefinger of a vi- nyl-gloved hand between the toes of the leg sample and then on the basal half of a mi- cropipette. The basal rather than the apical portion of the micropipettes was treated in the interdigital gland bioassays, because in nature interdigital gland substances are de- posited on the substrate. A nymph was released singly on the mid- point of a clay island. The location of the nymph was recorded at 1, 18 and 24 h after release. Ticks that were found to have fallen or crawled into the water during the 2 h after release or overnight were replaced on midpoints of the clay islands. Clay island and petri dish units were washed in soapy water and rinsed with clean water after ev- ery test. Thirty J. scapularis and 30 A. americanum nymphs were tested against feather substances. Ten J. scapularis nymphs were tested against hair substances from each of 14 dogs. An additional 10 to 20 I. scapularis nymphs were tested against hair substances from 2 dogs that produced extreme results in the initial tests. To con- firm kairomonal activity in the canine sam- ples (Carroll 1999b, Dobrotvorsky et al. 2000) and to compare the behaviors with nymphal responses, 10 female and 10 male I. scapularis were tested against substances from 10 of the same hair samples tested against nymphs. Some samples were tested against adults first and others against nymphs first. Thirty /. scapularis nymphs were tested against interdigital gland sub- stances from each of 2 white-tailed deer does. Data were analyzed by chi-square 2 x 2 contingency tables where responses of each tick were considered independent. RESULTS After their release on the clay islands, nymphs of both species tested against sub- stances from feathers, interdigital glands and canine hair wandered up and down the micropipettes. At | h after their release, nymphs were often still active and showed no tendency to be on either the treated on untreated micropipettes. By the next morn- ing (18 h after release), nymphs were large- ly akinetic. Numbers of /. scapularis that 76 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON n Treatment nymphs 140 44 male 100 70° female 100 86 ers of /. scapularis on micropipettes (or capillary tubes for adults) rubbed with dog hair frer the ticks were released on clay islands with the treated and untreated micropipettes. Control x? P 78 OS <0.05° 15 B56 <0.05 7 6722 <0.05 “ Differed from expected equal distribution between treatment and control, indicating possible tendency to avoid treatment (canine hair substances). > Some ticks remained on clay block or fell or crawled into water moat, so treatment plus control numbers may not equal n. became akinetic on the apical portions of micropipettes treated with substances from feathers were significantly greater (X? = 5.4, P < 0.05) than those on the untreated micropipettes (Table 1). No such arrestant response was observed with nymphs of A. americanum, with 13 nymphs on treated micropipettes and 11 on untreated ones. Lo- cations of nymphs at 24 h after release dif- fered little from those at 18 h after release. Canine hair substances were not ob- served to elicit arrestment among nymphs of I. scapularis when data for the samples from all 14 dogs were pooled (Tables 2—3). Instead there was a significant avoidance (X? = 8.2, P < 0.05) of the treated micro- pipettes. Samples from 2 dogs produced markedly opposite results. When /. scapu- laris nymphs were tested against hair sub- stances from the hair of dog 9656, 19 of 30 nymphs were on treated micropipettes, whereas 4 of the ticks were on the untreated (control) micropipettes. In contrast, none of the 20 /. scapularis nymphs tested against hair substances from dog 9334 were on the treated micropipettes 24 h after their release on the clay island. High levels (75-90%) of both male and female /. scapularis showed an arrestant response to substances from the same canine hair samples tested against the conspecific nymphs (Table 3). Adults re- sponded to substances from the hair of dogs of both sexes. Nymphs of /. scapularis did not exhibit an arrestment response to interdigital gland substances from white-tailed deer does. Thirty nymphs were on treated micropi- pettes and 23 on untreated micropipettes CX2 — O77. P= 0.05): DISCUSSION The responses of the /. scapularis nymphs to the chicken feathers indicate that at least certain host species can elicit ar- restment among individuals of this life stage. A variety of bird species are reported to serve as hosts for 1. scapularis nymphs (Anderson and Magnarelli, 1984, Anderson et al. 1985, Battaly et al. 1987, Durden and Keirans 1996, Keirans et al. 1996). Host- seeking nymphs of /. scapularis occur pri- marily in the leaf litter and forest floor where some species of resident birds regu- larly forage. Repeated foraging or territorial activity by birds at a location would cause oils and other substances on feathers to rub off on fallen leaves and forest floor debris. Thus, it may reward a host-seeking nymph to remain near feather-associated chemical residues on leaf litter. It may be that arrest- Table 3. Percentage of /. scapularis showing an arrestant response to canine hair samples. Sex of Dog Nymphs Females Males female 31% (n = 10) 90% (n = 7) 80% (n = 8) male 33% (n = 4) 77% (n = 3) 75% (n = 2) “n = number of hair samples from different dogs that were tested against 10 ticks each. VOLUME 104, NUMBER 1 ment in laboratory bioassays highlights a basic response of ticks to host odors that signal ticks to go no further, as if they were actually on a host. If that were the basic unit of behavior, it would account for improved host ambush site selection. Lone star tick immatures will use avian hosts in nature and have been reared on domesticated chickens in the laboratory (Koch 1983). These findings reveal basic differences in how host-seeking nymphal and adult /. sca- pularis and A. americanum respond to host- produced substances. Unlike /. scapularis, lone star ticks are equipped with eyes (all host-seeking stages) and a rapid gait suited to a more hunting type of host finding (Wal- adde and Rice 1982). It is, therefore, not surprising to observe different responses by both species to the same chemical stimuli. The results of the bioassays with feathers show that arrestment induced by _host- produced kairomones does occur in /. sca- pularis nymphs. However, the lack of the response by the nymphs to _ interdigital gland substances differs from adult conspe- cifics, which were strongly arrested by in- terdigital gland substances from both sexes of deer (Carroll 2001). Interdigital gland substances are deposited directly by deer on the stratum where host-seeking /. scapular- is nymphs occur. Most adult /. scapularis feed on white-tailed deer, but larvae and nymphs will also feed on deer if given the opportunity. Further investigations, focused on rodent and lizard hosts, are needed to as- certain whether chemically mediated behav- ior plays an important role in host-finding by I. scapularis and A. americanum nymphs. Responses of /. scapularis nymphs to ca- nine hair samples showed considerable var- iation. Against most canine hair samples /. scapularis nymphs responded indifferently or avoided them, with little indication of arrestment. However, there was a significant arrestant response to one canine sample and apparently complete avoidance of another sample. In contrast, adult /. scapularis of both sexes showed high levels of arrestment to the same canine hair samples in bioas- Ti says. Any of several explanations may ac- count for this discrepancy. Nymphs of /. scapularis may have less need to relocate themselves to areas frequented by particular species of vertebrates than adult ticks and may lack a sophisticated mechanism to ex- ploit host-produced chemical cues. A broad array of hosts from small to large mammals (Main et al. 1982, Durden and Keirans 1996, Keirans et al. 1996), birds and even lizards suffice as hosts for [. scapularis nymphs. Nymphs of J. scapularis may be most responsive to host species not tested. An unfed nymph starts seeking a host near where it dropped from its larval host, prob- ably within the home range of that host and in an area frequented by that host species. An adult /. scapularis starts seeking a host near where it dropped from its nymphal host, which may be a species unsuitable for adult feeding (adults feed on medium to large mammals). Thus, adult ticks appear to benefit more from relocating themselves to where large hosts are. This might be reflect- ed in the difference in mobility between these life stages. Adult 7. scapularis can ac- tively disperse =10 m in a few weeks, whereas nymphs of the same species have been reported to move 5 m (Carroll and Schmidtmann 1996). In order to exploit chemical cues left on vegetation and the substrate by hosts (Carroll et al. 1996), a tick must have the locomotory capability to find them, and the effort in terms of energy and risk must be commensurate with the tick’s chances of acquiring a host. Depend- ing on the degree to which a tick searches for host-produced residues in its microen- vironment, such host finding behavior could also be considered a form of hunting, albeit different from a tick rushing upwind in a carbon dioxide gradient emanating from a resting host (Waladde and Rice 1982). Fur- thermore, we cannot assume that a tick wanders continuously until it finds a resi- due. Ticks may well wait at random vantage points for a time, and, if no hosts pass by, resume wandering. Hence, host-seeking Fe PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ticks are not found just along trails and deer beds. Perhaps in differently designed bioassays more arrestment may have been observed. In any case more bioassays of substances. associated with rodent (e.g., white-footed mice) and lizard (e.g., shinks) hosts are warranted, and the threshold activity levels of host-produced kairomones need to be quantified. LITERATURE CITED Anderson, J. EF and L. A. Magnarelli. 1984. Avian and mammalian hosts for spirochete-infested ticks and insects in a Lyme disease focus in Connecticut. Yale Journal of Biology and Medicine 59: 627— 641. Anderson, J. FE, R. C. Johnson, L. A. Magnarelli, and E W. Hyde. 1985. Involvement of birds in the ep- idemiology of Lyme disease Borrelia burgdorferi. Infection and Immunity 51: 394—396. Battaly, G. R., D. Fish, and R. C. Dowler. 1987. The seasonal occurrence of Ixodes dammini Acarina: Ixodidae) on birds in a Lyme disease endemic area of southwestern New York State. Journal of the New York Entomological Society 95: 461—468. Carroll, J. E 1999a. Notes on responses of blacklegged ticks (Acari: Ixodidae) to host urine. Journal of Medical Entomology 36: 212-215. . 1999b. Responses of three species of adult tick (Acari: Ixodidae) to chemicals in the coats of principal and minor hosts. Journal of Medical En- tomology 36: 238—242. . 2001. Interdigital gland substances of white- tailed deer and the responses of host-seeking ticks (Acari: Ixodidae). Journal of Medical Entomology 38: 114-117. Carroll, J. FE, J. A. Klun, and E. T. Schmidtmann. 1995. Evidence for kairomonal influence on selection of host-ambushing sites by adult /. scapularis (Acari: Ixodidae). Journal of Medical Entomology 32: 119-125. Carroll, J. E, G. D. Mills, Jr., and E. T. Schmidtmann. 1996. Field and laboratory responses of adult /. scapularis (Acari: Ixodidae) to kairomones pro- duced by white-tailed deer. Journal of Medical En- tomology 33: 640-644. Carroll, J. EK and E. T. Schmidtmann. 1996. Dispersal of blacklegged tick (Acari: Ixodidae) nymphs and adults at the wood-pasture interface. Journal of Medical Entomology 33: 554—558. Dobrotvorsky, A. K., A. V. Tkachev, R. L. Naumov, and J. EK Carroll. 2000. Study of host odor deter- minants of questing behavior in /xodes ticks. Pro- ceedings of the 3rd International Conference “Ticks and Tick-Borne Pathogens: into the 21st Century,” Hotel Academia, High Tatra Moun- tains, Slovakia, 30 August-September, 1999, |: 235-240. Durden, L. A. and J. E. Keirans. 1996. Nymphs of the genus Ixodes (Acari: Ixodidae) of the United States: taxonomy, identification, key, distribution, hosts and medical/veterinary importance. Mono- graph, Thomas Say Publications, 95 pp. Keirans, J. E., H. J. Hutcheson, L. A. Durden, and J. S. H. Klompen. 1996. Ixodes (Ixodes) scapularis (Acari: Ixodidae): redescription of all active stag- es, distribution, hosts, geographical variation, and medical and veterinary importance. Journal of Medical Entomology 33: 297-318. Koch, H. G. 1983. Lone star ticks: oviposition, egg hatch, and molting under naturally fluctuating am- bient temperatures and humidities in the field. Southwestern Entomologist 8: 1—5. Main, A. J., A. B. Carey, M. G. Carey, and R. H. Goodwin. 1982. Immature Ixodes dammini (Acari: Ixodidae) on small mammals in Connecticut. Jour- nal of Medical Entomology 19: 655—664. Naumoy, R. L. and L. M. Vitlin. 1995. Response of taiga ticks to an attractant. Communication !: Ex- perimental study of the contact of nymphs with attractive acaricidal granules. Meditsinsaja Para- zitologija. i Parasitarnye. Bolezni 3: 49—5S0. Rechay, Y., R. A. I. Norval, J. Tannock, and J. Col- borne. 1978. Attraction of the tick /xodes neitzi to twigs marked by the klipspringer antelope. Nature (London) 275: 310-311. Sonenshine, D. E., D. Taylor, and K. A. Carson. 1986. Chemically mediated behavior in Acari: Adapta- tions for finding hosts and mates. Journal of Chemical Ecology 12: 1091-1108. Spielman, A., M. L. Wilson, J. EF Levine, and J. Pies- man. 1985. Ecology of /xodes dammini-borne hu- man babesiosis and Lyme disease. Annual Review of Entomology 30: 439—460. Waladde, S. M. and M. J. Rice. 1982. The sensory basis of tick feeding behavior, pp. 71—118. Jn Ob- enchain, EK D. and G. Galun, eds. Physiology of ticks. Pergamon, New York. Wilson, J. G., D. R. Kinzer, J. R. Sauer, and J. A. Hair. 1972. Chemo-attraction in the lone star tick (Ac- arina: Ixodidae). 1. Response of different devel- opmental stages to carbon dioxide. Journal of Medical Entomology 9: 245—252. PROC. ENTOMOL. SOC. WASH. 104(1), 2002, pp. 79-90 TWO NEW SPECIES OF OPIINAE (HYMENOPTERA: BRACONIDAE) ATTACKING FRUIT-INFESTING TEPHRITIDAE (DIPTERA) IN WESTERN KENYA SUSAN W. KIMANI-NJOGU AND ROBERT A. WHARTON (SWK-N) International Centre for Insect Physiology and Ecology, PO. Box 30772 Nairobi, Kenya; (RAW) Department of Entomology, Texas A&M University, College Station, TX 77843, U.S.A. (e-mail: rawbaw2 @tamu.edu) Abstract.—Two new species of Opiinae are described, Rhynchosteres mandibularis and Fopius okekai. Both were reared from puparia of fruit-infesting ceratitidine Tephri- tidae collected in western Kenya. The placement of these two species relative to the genera Fopius Wharton and Rhynchosteres Fischer is discussed, with emphasis on the transitional form of the clypeus. Arguments are presented for inclusion of these species in Rhyn- chosteres and the Fopius desideratus (Bridwell) species-group, respectively. This repre- sents the first host record for the genus Rhynchosteres. Evidence is presented that suggests similarity in function has lead to convergence in shape of the ovipositor tip in certain Opiinae and Doryctinae. Key Words: Fruit-infesting Tephritidae support a di- verse array of parasitoids (Clausen et al. 1965, Hoffmeister 1992, Hoffmeister and Vidal 1994, Sivinski et al. 1998, Wharton et al. 2000). Among the best-studied of these are members of the opiine Braconi- dae. Opiines oviposit in the host egg or lar- val stage and emerge from the puparium. About 100 species of opiines have been re- corded as parasitoids of fruit-infesting Te- phritidae, largely in association with bio- logical control programs (Silvestri 1914; Clausen 1978; Fischer 1971; Wharton 1997, 1999; Ovruski et al. 2000). The opiine parasitoids of tephritids are particularly diverse in the Afrotropical Re- gion, with several genera and_ species- groups represented (Fischer 1987, Wharton 1997). The apparent high diversity may in part reflect a sampling bias since extensive rearings of both wild and commercial fruits, focused specifically on parasitoids, have parasitoids, fruit, Fopius, Rhynchosteres, Ceratitis, Araceae, Flacourtiaceae been conducted on several occasions (Sil- vestri 1914, Bianchi and Krauss 1937, van Zwaluwenburg 1937, Clausen et al. 1965, Steck et al. 1986). Nevertheless, recent rearings of wild fruits in Kenya, as well as the examination of incompletely identified material from older collections, suggest that our knowledge of the diversity of parasit- oids in this host system is far from com- plete. To increase our understanding of both the alpha and beta diversity of Afrotropical te- phritid parasitoids, we describe two species whose clypeal morphology is somewhat in- termediate between members of the genus Fopius Wharton and those of the genus Rhynchosteres Fischer. We also discuss the potential impact of the discovery of these species on existing classifications proposed or used by Fischer (1987, 1999), van Ach- terberg and Maeto (1990), Wharton (1997), and van Achterberg (1999). 80 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MATERIALS AND METHODS Primary types and other material used for comparison in the present study came from the following institutions and individuals: The Hungarian Natural History Museum, Budapest, Hungary (J. Papp); Texas A&M University Insect Collection, College Sta- tion, Texas, USA (TAMU); Bernice P. Bish- op Museum, Honolulu, Hawaii, USA (G. Nishida, K. Arakawa); International Centre of Insect Physiology and Ecology, Nairobi, Kenya (ICIPE); Koninklijk Museum voor Midden Afrika, Tervuren, Belgium (E. De Coninck); and National Museum of Natural History, Smithsonian Institution, Washing- ton, D.C., USA (D. Smith). Additionally, samples of wild fruits were collected throughout Kenya from 1998—2000, and processed in the laboratory to obtain emerging flies and parasitoids. Kenyan specimens used in this study were obtained primarily from Kakamega (western Kenya), the Kenyan Highlands around Nairobi, and the coastal region north and south of Mom- basa. Collected fruits were held in cages above sand until tephritid flies emerged to pupate in the soil. Puparia were hand-sorted from the soil after about 2 weeks, and held in separate cages for emergence of adult flies and wasps. Field-collected fruit sam- ples yielding the parasitoids described here were processed at ICIPE in Nairobi. Both of the species described below were reared from tephritid puparia collected in Kaka- mega. This study represents a collaborative effort, with the senior author assuming pri- mary responsibility for preparation of the species descriptions and the junior author primarily responsible for the discussion of ovipositor and clypeal morphology. Terminology for the descriptions follows Sharkey and Wharton (1997) and Wharton (1997). Additionally, details of the ovipos- itor tip are important in distinguishing spe- cies and possibly species-groups within the Opiinae. The species described here have what has been referred to as a double dorsal node (Wharton 1997). However, this is a general description that does not adequately describe the differences in detail among species and species-groups. In the type spe- cies of Diachasmimorpha Viereck, for ex- ample, the double dorsal node is delimited basally by an abruptly elevated section that tapers gradually to the apex (Wharton 1997, figs. 87, 88). Near the middle of this tapered section, there is a second elevation that is weakly excavated on its basal face. In the species described here, the double dorsal node has a different appearance since it lacks the basal excavation on the second node (Fig. 15). To differentiate the two con- ditions, we describe the form in the species treated here as a dorsal node with a more distally placed secondary tooth. Quicke et al. (1992) also described a double dorsal node in doryctine Braconidae, suggesting that the distinctively shaped apex of the dorsal valve of the ovipositor provided a synapomorph for the Doryctinae. The shape of the nodes in the two species described below differs from those of the typical do- ryctines figured by Quicke et al. (1992). RESULTS AND DISCUSSION Morphology of the ovipositor.—The dis- tinctive doryctine double dorsal node was not found in any of the other cyclostomes nor in any of the non-cyclostome braconids examined by Quicke et al. (1992). The do- ryctine double dorsal nodes tend to be ex- cavated on the apical face, whereas the opi- ine nodes are usually excavated or truncat- ed on their basal face. However, as dem- onstrated here for opiines as well as in a few of the typical doryctines figured by Quicke et al. (1992), the forms converge in their reduced states (e.g., when the exca- vated surfaces and marginal ridges are less distinct). The condition described here for Fopius is thus somewhat intermediate be- tween that found in some of the doryctines figured by Quicke et al. (1992) and those opiines previously described as having a double dorsal node by Wharton (1997). While the findings of Quicke et al. (1992) represent an important advancement in our VOLUME 104, NUMBER 1 understanding of doryctines, more detailed surveys are obviously needed to determine the extent to which the shape and number of nodes can be used unambiguously as di- agnostic features for higher taxa. The nodes found on dorsal valves of various braconids and ichneumonids most likely represent re- ductions from the elaborately sculptured valves of siricoids, but it appears as if dif- ferent structural elements may have been lost in the doryctines relative to the opiines. The end result is convergence in form, pos- sibly for the same general function. The mechanics of drilling into hard sub- strates have been recently described in su- perb detail by Vincent and King (1996), and a good overview of the morphology of the ovipositor in Hymenoptera is provided by Quicke et al. (1994). As pointed out by both Vincent and King (1996) and Quicke et al. (1999), dorsal nodes and ventral ser- rations are important attributes for species drilling through wood. Quicke et al. (1999), summarizing previous findings on the struc- ture and function of the ovipositor, ob- served that endoparasitic ichneumonoids al- most always have the dorsal node reduced and typically have less well-developed ven- tral serrations. However, the dorsal node is clearly well-developed in the species de- scribed below, as it is in several other opi- ines attacking fruit-infesting Tephritidae. The ventral serrations in opiines do tend to be less extensive than in most doryctines, but are still well developed in species of Fopius and Diachasmimorpha attacking late instar larvae embedded in fruit. Since some endoparasitoids (such as these opi- ines) must also drill to locate hosts, it is thus not surprising to find similar structures in these species. If dorsal nodes and ventral serrations are associated more with drilling through substrates than with developmental phenomenona such as endoparasitism and ectoparasitism, we should expect to find them in other endoparasitoids that must drill through solid (or at least firm) sub- strates to find hosts. Morphology of the clypeus and the tran- rom sition from Fopius to Rhynchosteres.—Var- iation in the clypeal morphology of opiine Braconidae has received considerable atten- tion in the development of classifications for these wasps. Fischer (1972), for exam- ple, used the relative size of the clypeus and the presence of apical tubercles as the prin- ciple diagnostic features for genera within his subtribe Biosterina. He also used some of the same character states for differenti- ating subgenera of Opius as well as other genera outside the tribe Opiini. Wharton (1988, 1997) provided additional details re- garding the morphology of the opiine clyp- eus and its utility in constructing classifi- cations. Rhynchosteres was described by Fischer (1965) for three Afrotropical species with elaborate modifications of the clypeus and varying degress of exposure of the under- lying labrum. Wharton (1987) initially de- scribed Fopius as a subgenus of Rhynchos- teres Fischer, noting a number of similari- ties between the two despite obvious dif- ferences, im the form of the clypeus: Wharton (1987) also noted that within Fop- ius 8. S., the clypeus was somewhat plastic, showing variation among species in protru- sion of the ventral margin. Van Achterberg and Maeto (1990) elevated Fopius to ge- neric status, stating that the strongly pro- truding clypeus of Rhynchosteres was suf- ficient to maintain the two taxa as distinct, and suggesting that any similarities were symplesiomorphs. Van Achterberg and Maeto (1990) concluded that Fopius was more closely related to Pseudorhinoplus Fi- scher and Diachasmimorpha than to Rhyn- chosteres. Wharton (1997, 1999) main- tained Fopius as a separate genus, segre- gated and briefly diagnosed several species- groups, but noted that members of the Fopius desideratus (Bridwell) species- group shared a number of putative apo- morphic features with members of the ge- nus Rhynchosteres. Quicke et al. (1997) and Fischer (1999) have further discussed the relationships and classification of this group of tephritid parasitoids but only in general 82 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-4. Rhynchosteres mandibularis, head. 1, Dorsal view. 2, Frontal view centered on face. 3, Frontal view centered on mandible and labrum. 4, Oblique view showing rounded basal lobe of mandible and medially thickened, weakly up-turned clypeus. terms since they treated only a few of the included species. In the type species of Rhynchosteres, the middle portion of the clypeus is nearly tu- bular and protrudes like an inverted gutter from the middle of the face. As noted by Fischer (1965), van Achterberg (1983), and Wharton (1997), however, the shape of the clypeus varies among the described species. This variation in the structural details of the clypeus makes it challenging to define un- ambiguously the specific character state that supports the monophyly of Rhynchosteres. All five previously described species have the clypeus bulging medially. In R. microps Fischer, R. tuberculatus van Achterberg, and R. tubiformis Fischer, the protruding part of the clypeus is thin-walled, with a tunnel-like cavity varying from ventrally- directed to anteriorly-directly in an apparent transition series. In R. clypeatus (Bridwell) and R. brunigaster Fischer, however, the clypeus is thickened ventral-medially, re- vealing a weakly excavated, polished sur- face. The two species described below ex- hibit clypeal modifications that tend to blur the distinctions between Rhynchosteres and other opiine genera, but in different ways. In the newly discovered species reared from Rawsonia fruits, the clypeus is short and the ventral margin is thickened, slightly up- turned, and weakly protruding medially (Figs. 2—4, 9). The clypeus in this species clearly resembles that of R. clypeatus, but is less obviously protruding and conse- quently less obviously a member of Rhyn- chosteres. In the second species described below, reared from Culcasia fruits, the clypeus retains its full height along the mid- line as in Fopius s. s., but the ventral mar- VOLUME 104, NUMBER | Figs. 5-8. gin is excavated on either side of the mid- line, thus exposing a portion of the labrum (Figs. 6-8). Lateral shortening may not be a prerequisite for the development of the tube-like medial modifications seen in the microps-tuberculatus-tubiformis series, but would certainly facilitate it. The discovery of the two species de- scribed here will almost certainly require a reassessment of the generic status of Fop- ius, and minimally a transfer of the desi- deratus species-group to Rhynchosteres. The primary purpose of the present paper, however, is to make the names available so that further work on their biology and re- lationships can proceed. Thus, one of the two species described below is tentatively placed in the Fopius desideratus species- group and the other in Rhynchosteres. Shared features possessed by known spe- cies of Rhynchosteres and the desideratus Fopius okekai, head. 5, Dorsal view. 6, Frontal view centered on clypeus. 7, Frontal view centered on mandible. 8, Oblique view showing partially exposed labrum and unmodified mandible. group include transversely striate frons (usually forming a crescent-shaped pattern), a double dorsal node on the ovipositor, and a decrease in density of setae on the ovi- positor sheath (Figs. 1, 5, 14, 15). Unlike members of the genus Diachasmimorpha (as defined by Wharton 1997), the species of Fopius and Rhynchosteres possess an oblique groove/ridge on the propleuron. They also have a distinct post-pectal carina and a small second submarginal cell from which the m-cu cross-vein is excluded. The latter features are homoplastic within the Opiinae, but are useful for recognition of these genera. Rhynchosteres mandibularis Kimani-Njogu and Wharton, new species (Figs. 1-4, 9-12, 14, 15, 17, 18) Types.—Holotype 2: Top label = “KENYA: Western Prov. Kakamega, 84 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 9-12. sosoma. 11, Lateral view of petiole showing deep laterope. 12, Dorsal view of petiole. 0°13.1'N, 34°54.06'E 3.viii.1999 R. Cope- land’’. Second label = “‘reared from Te- phritidae in fruits of Rawsonia lucida’. Third label = “collection # 217 ICIPE/ USAID rearing program’’. Deposited in National Museums of Kenya, Nairobi. Par- atypes: 4 2, 2 3, same data as holotype, deposited at ICIPE and Texas A&M Uni- versity. Diagnosis.—This species closely resem- bles R. clypeatus in many respects, but the mesosoma is distinctly darker in mandibu- laris. In clypeatus, the median tooth (or tu- bercle) of the clypeus is dorsally displaced relative to the remaining portion of the ven- tral margin. In mandibularis, however, the ventral-median section of the clypeus (in frontal view, Fig. 2) extends slightly ven- trad the lateral margin. This new species Rhynchosteres mandibularis. 9, Lateral view of head and mesosoma. 10, Dorsal view of me- can be separated from nearly all other opi- ines placed in either Fopius or Rhynchos- teres by the possession of a rounded, thick- ly carinate lobe at the base of the ventral margin of the mandible (Figs. 3, 4). The rounded form of this structure is completely different from the more truncate or tooth- like lobe found in Biosteres s.s. A basal lobe is also found in clypeatus, but is not quite as rounded, nor as thickly carinate. The striate frons and double dorsal node on the ovipositor are identical to features found in members of the Fopius desidera- tus species-group. Unlike mandibularis, however, previously described members of the desideratus group do not have the clyp- eus shortened and the labrum broadly ex- posed. The clypeus of mandibularis is nei- ther as protruding nor tunnel-shaped as in VOLUME 104, NUMBER | described species of Rhynchosteres, though it approaches the form of the clypeus in cly- peatus, and is similarly shortened. Description.—Female. Length: 4.0—5.0 mm. Head: 1.45—1.50 times broader than long; 1.35—1.40 times broader than mesos- cutum. Face (Figs. 2, 3) densely setose and punctate, punctures mostly discrete with lit- tle or no indication of rugosity, setae long, mostly decumbent and medially directed, those near inner margin of eye more ven- trally directed and less decumbent; mid- ridge polished, prominent, distinctly elevat- ed, gradually narrowing dorsally and ex- tending between toruli. Distance between toruli shorter than distance from torulus to eye. Frons (Fig. |) punctate laterally and between toruli, punctures confluent or near- ly so on inner side of toruli; frons medially (including at least part of ocellar field) with crescent-shaped striae. Impressed line bor- dering ocellar triangle largely obscured by transverse striae. Vertex (Fig. 1) smooth, highly polished on either side of ocelli; sparsely punctate laterally near compound eyes and posteriorly along occiput. Occip- ital carina in lateral view (Fig. 9) promi- nent, extending dorsally from base of man- dible to a point just above middle of eye. Clypeus in profile (Fig. 9) weakly bulging dorsomedially; in frontal view (Figs. 2—4) with ventral margin protruding medially; ventral margin of clypeus thickened; la- brum broadly exposed between somewhat shortened clypeus and ventrally deflected mandibles; setae on clypeus erect or nearly so, weakly directed ventrally. Eye 3.5 times longer than temple, with a few, very short, scattered setae. Temples in dorsal view (Fig. 1) slightly receding, width of head at temples less than width at eyes. Antenna with 42—43 segments; roughly 4 times lon- ger than mesosoma; first flagellomere about 0.9 times length of second. Outer surface of mandible (Figs. 3, 4) convex basally and apically, somewhat flattened medially; ven- tral margin with broad, rounded, densely setose basal lobe, the margin of the lobe 85 thickly carinate. Maxillary palps much lon- ger than height of head. Mesosoma (Figs. 9-10): 1.35 times lon- ger than high; 1.85—1.90 times longer than broad. Median lobe of mesoscutum setose and punctate with punctures widely scat- tered posteriorly, sometimes weakly rugu- lose near midpit. Lateral lobes impunctate medially, with scattered punctures and rel- atively long inwardly directed setae around margins. Notauli broad at base, narrow pos- teromedially, distinctly crenulate through- out length, meeting posteriorly in a clearly defined midpit that extends narrowly to- wards posterior margin. Midpit broadly and very shallowly depressed. Space between midpit and scutellar sulcus smooth with scattered long setae directed outwardly to- wards lateral lobes. Scutellar sulcus broader medially than laterally, the posterior margin with a distinct median excavation; with 5 widely spaced longitudinal ridges. Scutel- lum densely setose, setae directed posteri- orly; distinctly but finely punctate over most of surface, punctures elongate; trans- verse band of rugulose sculpture present posteriorly. Metanotum with distinctly ele- vated median ridge. Propodeum rugose, with a short median longitudinal carina on basal 0.25 and with crescent-shaped, trans- verse carina usually discernible among ru- gose sculpture; with long, well-spaced setae laterally, nearly bare medially; propodeum separated from metapleuron by well defined crenulate groove. Metapleuron smooth, at least on dorsal half; densely covered with ventrally-directed setae. Sternaulus broad, with deep pits, crenulate throughout; ex- tending from anterior margin of mesopleu- ron at least 0.75 the distance to mid coxa. Wings (Figs. 17-18): Fore wing stigma broad, 3.8—4.0 times longer than wide; r arising slightly distad its midpoint; 2RS weakly sinuate, nearly straight, 1.4—1.5 times longer than 3RSa; 3RSa 2.5 times longer than r; 3RSb ending nearly at wing tip; (RS+M)a sinuate; (RS+M)b present, m-cu arising distinctly basad 2RS; 1M strongly bent posteriorly; Icu-a inclivous, rele) ~ROCEEDINGS OF 1 ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 13-16. along middle of ovipositor sheath. 15, R. mandibularis, apex of ovipositor. 16, Fopius sp. near silvestrii, setal pattern along middle of ovipositor sheath. postfurcal relative to 1M. Hind wing m-cu very weakly curved towards wing base; ex- tending nearly to wing margin as a faint crease, very weakly pigmented anteriorly, barely visible posteriorly. Petiole “(Figss li, len times longer than apical width, apex 1.8 times wider than base; finely striate; dorsal carinae well-developed over basal 0.8, in- Metasoma: distinct toward posterior margin; dorsope absent, laterope present as a deep pit. Me- tasoma unsculptured beyond petiole. Hy- popygium strongly narrowed and pointed posteriorly but short, not greatly attenuated. Ovipositor tip (Fig. 15) with a dorsal node, and a small, more distally placed secondary tooth. Ovipositor sheath sparsely setose ba- sally (Fig. 14), with about 3 rows of short to very short, well-spaced setae, setae more dispersed apically, with only two apparent 13, Fopius okekai, dorsal view of mesosoma. 14, Rhynchosteres mandibularis, setal pattern rows; sheath longer than body, 2.5 times longer than mesosoma. Color: Head, mesosoma, ovipositor sheath, wing veins, and last three metaso- mal terga brown to very dark brown; hind tibia and tarsi dorsally brown; metapleuron, propodeum laterally and most of metano- tum orange; petiole, basal segments of gas- ter, scape, annellus, and remainder of legs yellow. Male.—About as in female except anten- na 42—45 segmented; mandibles not as strongly deflected ventrally, thus labrum only narrowly exposed; scutellar sulcus with 3 ridges; transverse carina of propo- deum not discernible; metasomal terga 4 and following black. Discussion.—The plant host is a member of the Flacourtiaceae. Three species of Te- phritidae have been reared from this host VOLUME 104, NUMBER 1 87 Figs. 17-20. Fore and hind wings. 17, 18, Rhynchosteres mandibularis. 19, 20, Fopius okekat. plant in Kenya: Ceratitis anonae Graham, C. rosa Karsch var. fasciventris (Bezzi), and an undescribed species of Trirhithrum Bez- zi. Only one of these (the Trirhithrum) was reared from the same batch of puparia that yielded the type series. These puparia were isolated from a collection of 165 fruits. The undescribed species of Trirhithrum (family Tephritidae) is the first confirmed host re- cord for the genus Rhynchosteres. Etymology.—The species name refers to the enlarged basal lobe of the mandible. Fopius okekai Kimani-Njogu and Wharton, new species (Figs. 5—8, 13, 19, 20) Types.—Holotype 2: Top label = “KENYA: Western Prov. Kakamega, along Yala River Trail, 0°12.34'N, 34°53.57’E 88 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Copeland’. Second label = from Tephritidae in fruits of Cul- casia falcifolia’’. Third label = “collection # 639 ICIPE/USAID rearing program’’. De- posited in National Museums of Kenya, Nairobi. Paratypes: 2 ¢, same data as ho- lotype, deposited at ICIPE and Texas A&M University. Diagnosis.—This species is recognized as a member of the Fopius desideratus spe- cies-group by the enlarged clypeus plus the combination of features noted in the mor- phology section above that are shared by Rhynchosteres and the desideratus species- group. The ventral margin of the clypeus is strongly protruding medially as a truncate tooth in this species (Figs. 6—8). A median projection is also present (though not as well developed) in F. ottotomoanus and F. niger, the two species with which it is most likely to be confused. In okekai, the ventral margin of the clypeus on either side of the projection is weakly concave, exposing a portion of the labrum and making the pro- jection appear more prominent. In okekai, the projection is directed ventrally whereas in ottotomoanus and niger it is directed more anteriorly and thus protrudes more from the face. The mandible is normal in okekai, lacking the rounded basal lobe on the ventral margin found in mandibularis. It is also a darker species than mandibular- is, has a more distinctly punctate scutellum, a better-developed hind wing m-cu, and a less prominent secondary tooth on the ovi- positor. Description.—Female. Length: 4.0 mm. Head: 1|.35 times broader than long; 1.45 times broader than mesoscutum. Face (Figs. 6, 7) densely setose and punctate through- out, pattern variable, space between most punctures greater than diameter of punc- tures, but some punctures nearly confluent, suggesting rugulose sculpture; setae as in mandibularis; midridge distinctly elevated, polished, narrower dorsally, extending be- tween toruli. Distance between toruli al- most equal to distance from torulus to eye. Frons (Fig. 5) with sculpture and setal pat- tern about as in mandibularis, but striae much weaker, not extending posteriorly to median ocellus, thus surface appearing more extensively smooth and polished. Ocellar triangle almost completely mar- gined by an impressed line. Vertex and oc- cipital carina as in mandibularis. Clypeus in profile weakly bulging dorsomedially; ventral margin of clypeus (Fig. 6) with a strongly protruding rectangular projection; labrum (Fig. 8) partially exposed between somewhat shortened clypeus and ventrally deflected mandibles; setae on clypeus sparse, at least twice as long as those in middle of face, erect or nearly so. Eye ap- parently bare; large, 3 times longer than temple. Temples in dorsal view receding, width of head at temples less than width at eyes. Antenna 35 segmented; roughly 3.7 times longer than mesosoma. Ist flagellom- ere about 0.8 times length of second. Man- dible (Fig. 8) normal, outer surface distinct- ly convex, ventral margin without expanded lobe basally. Maxillary palps much longer than height of head. Mesosoma (Fig. 13): 1.4 times longer than high; 1.9 times longer than broad. Me- dian lobe of mesoscutum setose as in man- dibularis, with unevely spaced punctures; lateral lobes with numerous, relatively long setae around margins, bare and impunctate medially. Notauli distinctly crenulate throughout, meeting posteriorly in a clearly defined midpit that extends slightly onto median lobe anteriorly as a_ sculptured groove, midpit with irregular sculpture. Me- soscutum otherwise as in mandibularis. Scutellar sulcus parallel-sided or nearly so. Scutellum uniformly setose; with deep, elongate punctures clearly visible beneath weakly decumbent setae; transverse band of sculpture distinctly rugose. Metanotum about as in mandibularis, with distinctly el- evated median ridge but more prominently carinate lateral fields. Propodeum densely rugose, with median carina on basal 0.5 and with irregular, transverse carina discernible among rugose sculpture; setal pattern as in mandibularis; propodeum not distinctly VOLUME 104, NUMBER 1 separated from metapleuron laterally, the demarcation represented only by the tran- sition across a shallow groove to the weakly sculptured dorsal portion of the metapleu- ron. Metapleuron and sternaulus about as in mandibularis. Wings (Figs. 19, 20): Forewing stigma broad, 3.2 times longer than wide, with r arising slightly distad its midpoint; 2RS nearly as sinuate as (RS+M)a, 1.3 times longer than 3RSa; 3RSa 2.25 times longer than r; 3RSb ending almost at wing tip; (RS+M)b present and fairly long, roughly 0.2 times length of m-cu; IM _ weakly curved posteriorly; Icu-a inclivous, post- furcal relative to 1M by 0.35-0.40 its length. Hind wing m-cu reclivous, but not recurved, distinctly pigmented slightly more than halfway to wing margin. Metasoma: Petiole 1.3 times longer than apical width; apex about 1.8 times wider than base; sculpture, dorsope, and laterope as in mandibularis. Metasomal terga un- sculptured beyond petiole. Hypopygium as in mandibularis. Ovipositor tip with dorsal node and a barely perceptible, more distally placed secondary tooth. Ovipositor sheath with setae about as in mandibularis, with setae slightly more closely spaced in lon- gest row; sheath distinctly shorter than body, about 1.6 times longer than mesoso- ma. Color: Dark brown to black; petiole brown basally, yellow-brown apically; te- gula, T2, trochantellus dorsally, and apices of fore and mid femora yellow; palps whit- ish. Male.—Length 3.8—4.5 mm. About as in female except temple larger, eye 2.5 times longer than temple; antenna 37 segmented; first flagellomere 0.85—0.90 times length of second; yellow coloration on legs and me- tasoma slightly more extensive, with petiole completely yellow. Discussion.—The plant host is a member of the Araceae. The type series came from a sample of tephritid puparia, isolated from 200 fruits, that also produced Trirhithrum 89 inscriptum (Graham) and T. nigrum (Gra- ham). Etymology.—The species is named for Wilberforce Okeka, who assisted in the col- lection of fruits and identification of plants in Kakamega forest. ACKNOWLEDGMENTS This work could not have been accom- plished without the logistic support and fa- cilities provided by ICIPE and the person- nel in the division of forestry in Kakamega Forest who kindly facilitated the sampling program there. We also express our thanks to Ian White and Marc DeMeyer for iden- tification of the flies and especially to Rob- ert Copeland for conducting and managing the sampling program. Milicent Okumu as- sisted with sample processing and Peterson Nderitu provided a sample of Cucumis dip- saceus that produced parasitoids crucial for comparison with the newly described spe- cies. Special thanks to Jim Ehrman at the Digital Microscopy Facility, Mount Allison University, Sackville, NB, Canada for do- ing all of the SEM work, including arrang- ing the plates, and to Matt Buffington and Terry Junek for help with the wing figures. This work was supported primarily by USAID grant No. PCE-G-00-98-00048-00 to Texas A&M University (in collaboration with ICIPE and the African Fruit Fly Ini- tiative) and in part by USDA/NRI (grant no. 9703184) and NSF (DEB9712543), both to Texas A&M. LITERATURE CITED Bianchi, F A. and N. H. Krauss. 1937. Fruit fly inves- tigations in East Africa. The Hawaiian Planters’ Record 41: 299-306. Clausen, C. P. 1978. Tephritidae (Trypetidae, Trupa- neidae), pp. 320—335. Jn Clausen, C. P, ed. Intro- duced parasites and predators of arthropod pests and weeds: A world review. United States De- partment of Agriculture, Agricultural Handbook 480, 545 pp. Clausen, C. P., D. W. Clancy, and Q. C. Chock. 1965. Biological control of the Oriental fruit fly (Dacus dorsalis Hendel) and other fruit flies in Hawaii. United States Department of Agriculture, Techni- cal Bulletin 1322: 1-103. 90 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Zwei neue Opiinen-Gattungen aus (1. Beitrag ueber die Opiinae des Mu- Royal de |’Afrique Centrale) (Hymenoptera, Braconidae) Revue de Zoologie et de Botanique \fricaines 71: 309-323. —. 1971. Hymenoptera Braconidae. Index of World Opiinae, pp. 1-187. Jn Delucchi, V. and G. Remaudiére, eds. Index of Entomophagous In- sects. Le Francois, Paris. . 1972. Hymenoptera: Braconidae (Opiinae 1). Das Tierreich 91: 1—620. ; 1987. Hymenoptera: Opiinae [—athiopis- che, orientalische, australische und ozeanische Re- gion. Das Tierreich 104: 1-734. . 1999. Zur Evolution und zum System der Op- ius-verwandten Gattungen der Unterfamilie Opi- inae mit einer erweiterten Aufteilung dieses Gat- tungs-Komplexes (Hymenoptera, Braconidae, Opiinae). Linzer Biologische Beitraege 31: 277— 336. Hoffmeister, T. 1992. Factors determining the structure and diversity of parasitoid complexes in tephritid fruit flies. Oecologia 89: 288-297. Hoffmeister, T. S. and S. Vidal. 1994. The diversity of fruit fly (Diptera: Tephritidae) parasitoids, pp. 47— 76. In Hawkins, B. A. and W. Sheehan, eds. Par- asitoid community ecology. Oxford Science Pub- lications, Oxford University Press, 516 pp. Ovruski, S., M. Aluja, J. Sivinski, and R. Wharton. 2000. Hymenopteran parasitoids on fruit-infesting Tephritidae (Diptera) in Latin America and the southern United States: diversity, distribution, tax- onomic status and their use in fruit fly biological control. Integrated Pest Management Reviews 5: 81-107. Quickes Da. I> b. G2 ricken)andiMeGabitton= 1992" New diagnostic ovipositor characters for dorycti- ne wasps. Journal of Natural History 26: 1035— 1046. Quicke, D. L. J., M. G. Fitton, J. R. Tunstead, S. N. Ingram, and P. V. Gaitens. 1994. Ovipositor struc- ture and relationships within the Hymenoptera, with special reference to the Ichneumonoidea. Journal of Natural History 28: 635-682. Quicke, D. L. J., A. Leralec, and L. Vilhelmsen. 1999. Ovipositor structure and function in the parasitic Hymenoptera with an exploration of new hypoth- eses. Atti dell’Accademia Nazionale Italiana di Entomologia 47: 197-239. Quicke, D. L. J., K. van Achterberg, and H. C. J. God- fray. 1997. Comparative morphology of the ven- om gland and reservoir in opiine and alysiine braconid wasps (Insecta, Hymenoptera, Braconi- dae). Zoologica Scripta 26: 23-50. Sharkey, M. J. and R. A. Wharton. 1997. Morphology and terminology, pp. 19-37. In Wharton, R. A., P. M. Marsh, and M. J. Sharkey, eds. Manual of the New World Genera of the Family Braconidae (Hymenoptera). International Society of Hymen- opterists. Special Publication 1, 439 pp. Silvestri, EF 1914. Report of an expedition to Africa in search of the natural enemies of fruit flies (Try- paneidae) with descriptions, observations and bi- ological notes. Territory of Hawaii Board of Ag- riculture and Forestry Division of Entomology Bulletin 3: 1-176. Sivinski, J., K. Vulinec, E. Menezes, and M. Aluja. 1998. The bionomics of Coptera haywardi (Og- lobin) (Hymenoptera: Diapriidae) and other pupal parasitoids of tephritid fruit flies (Diptera). Bio- logical Control 11: 193-202. Steck, G. J., E 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. Van Achterberg, C. 1983. A new species of Rhyn- chosteres Fischer from Zaire (Hymenoptera, Bra- conidae). Zoologische Mededelingen, Leiden 57: 91-95. . 1999. The Palaearctic species of the genus Diachasmimorpha Viereck (Hymenoptera: Bra- conidae: Opiinae). Zoologische Mededelingen, Leiden 73: 1-10. Van Achterberg, C. and K. Maeto. 1990. Two new and aberrant species of Braconidae (Hymenoptera) from Japan. Zoologische Mededelingen, Leiden 64: 59-70. Van Zwaluwenburg, R. H. 1937. West African Notes. The Hawaiian Planters’ Record 41: 57-83. Vincent, J. E V. and M. J. King. 1996. The mechanism of drilling by wood wasp ovipositors. Biomimet- ics 3: 187-201. Wharton, R. A. 1987. Changes in nomenclature and classification of some opine Braconidae (Hyme- noptera). Proceedings of the Entomological Soci- ety of Washington 89: 61-73. . 1988. Classification of the braconid subfamily Opiinae (Hymenoptera). The Canadian Entomol- ogist 121: 333-360. . 1997. Generic relationships of opiine Bracon- idae (Hymenoptera) parasitic on fruit-infesting Te- phritidae (Diptera). Contributions of the American Entomological Institute 30(3): 1—S3. . 1999. A review of the Old World genus Fop- ius Wharton (Hymenoptera: Braconidae: Opiinae), with description of two new species reared from fruit-infesting Tephritidae (Diptera). Journal of Hymenoptera Research 8: 48—64. Wharton, R. A., M. K. Trostle, R. H. Messing, R. S. Copeland, S. W. Kimani-Njogu, S. Lux, W. A. Oy- erholt, S. Mohamed, and J. Sivinski. 2000. Para- sitoids of medfly, Ceratitis capitata, and related tephritids in Kenyan coffee: a predominantly ko- inobiont assemblage. Bulletin of Entomological Research 90: 517-526. PROC. ENTOMOL. SOC. WASH. 104(1), 2002, pp. 91-96 A NEW SPECIES AND NEW RECORDS OF MICROCERELLA MACQUART (DIPTERA: SARCOPHAGIDAE) FROM ARGENTINEAN PATAGONIA JUAN C. MARILUIS ANLIS “Dr. C. Malbran,’’ Vectores. Av. Vélez Sarsfield 563, (1281) Buenos Aires, Argentina (e-mail: vectores@ malbra.sld.ar) Abstract.—Microcerella antonioi, n. sp. (Sarcophagidae), from Santa Cruz, Argentina, is described and illustrated. The following four species are recorded for the first time from Argentinean Patagonia: Microcerella aulacophyto Pape, M. chilena (Dodge), M. penai (Lopes), and M. rusca (Hall). Resumen.—Se describe e ilustra una nueva especie de Sarcophagidae, Microcerella antoniot, de Santa Cruz, Argentina. Asimismo se registran por primera vez para la Pa- tagonia Argentina a M. aulacophyto Pape, M. chilena (Dodge), M. penai (Lopes) y M. rusca (Hall). Key words: Fifteen species of Microcerella Macquart 1851 are presently known in Patagonia, (Pape 1996). During 1997 and 1998, sev- eral collecting trips to Patagonia were car- ried out by the author, and a large number of specimens of Microcerella were cap- tured. A detailed study of this material re- vealed an undescribed species, which is herein described and illustrated. Also, sev- eral new localities in the Argentinean Pa- tagonia are provided for M. aulacophyto Pape, M. chilena (Dodge), M. penai (Lopes), and M. rusca (Hall). For general Sarcophagidae terminology see Lopes (1978, 1982). Illustrations were made by using a camera lucida attached to a stereoscopic microscope. The holotype and paratypes of the new species are pinned. The holotype and some paratypes are deposited in the collection of the Museo de La Plata, Buenos Aires, Ar- gentina. Other paratypes are deposited in the collections of the Museo Argentino de Ciencias Naturales “Bernardino Rivada- Diptera, Sarcophagidae, Microcerella, new species, Argentinean Patagonia via,” Buenos Aires, Argentina, and Fun- dacion Miguel Lillo, Tucuman, Argentina. The specimens of the other four species are in Museo de La Plata, Buenos Aires, Ar- gentina. Microcerella antonioi Mariluis, new species (Figs. 1-6) Diagnosis.—Male with a pair of preac- rostichal bristles; first pair of ocellar bristles situated at center of ocellar triangle; lower part of sternopleura with spot of silvery pollinosity; scutellum without discal scutel- lar bristles; sternite IV without hump; ster- nite V without evident humps at medial part of both arms; inner border of cerci not un- dulated; paralobi slender; and apical and lateral plate with a single lobe. Holotype male.—Length: 7-11 mm. Head: Black with silvery pollinosity, al- most quadrate. Epistoma_ protuberant. Cheeks, clypeus, parafrontalia, parafacialia, cheek grooves, occipital fringe, and post- 92 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-6. Microcerella antonioi, male. 1, Cerci rear view. 2, Right paralobi lateral view. 3, Sternite IV lateral view. 4, Sternite V lateral view. 5, Right paramere + gonopod lateral view. 6, Apex of aedeagus lateral view. Scale = 0.5 mm. VOLUME 104, NUMBER 1 gena black with silvery pollinosity. Fron- talia velvety black, upper part with silvery pollinosity and black hairs. Ocellar triangle black, with two pairs of the strong ocellar bristles. Parafrontalia and parafacialia with black hairs. Upper parafacialia with small hairs and low parafacialia with long hairs twice size of upper ones. Parafacialia 0.44 of distance between vibrissa. Front at nar- rowest part about 0.29 of head width. With 5 to 7 frontal bristles, to antennal base 4 to 5, rest pass antennal base, first pair faces backwards, all rest converge. Inner vertical bristles twice size of outer vertical bristles. Facialia black, with black hairs, lower part shining and upper part with silvery polli- nosity. Antenna black, segments I and II shining black, HI with brown pollinosity, arista bare. Thorax: Black with silvery pol- linosity. In lateral view, notopleuron and mesopleuron with spot of golden pollinos- ity. Pteropleuron with spot of silvery pol- linosity. Sternopleuron with two stigma of silvery pollinosity, one in foremost upper part around first sternopleural bristles and other at lower part. Thorax in dorsal view a bit inclined with head down and abdomen up, black with stripes of silvery pollinosity. Prescutum and postcutum with two stripes in area of dorsocentral bristles. Following marks of silvery pollinosity: all humeri; on inferior part of prescutum and one major on inferior central part of postscutum, these marks between intraalar and supraalar bris- tles. Postalar callus with apical spot of sil- very pollinosity. Scutellum black, with three marks of silvery pollinosity, two lat- eral and one distal between posterior mar- ginal lateral bristles. Acrostichals 1:0; dor- socentrals 2:3; intraalar 1:2; supraalar 1:3 Scutellum with two pairs of marginal lateral bristles, one anterior and other posterior, without discal bristles. Wings, epaulet, base of stem vein and point of union of R3 with RS black. Subepaulet and rest of veins or- ange. Base of R5 with black hairs. Legs black, hind femur with row of bristles on anterior and anterodorsal sides. Middle and posterior tibiae with 2, 3 or 4 bristles on 93 anterodorsal and posterodorsal sides. Ab- domen: Black. In posterior view, tergite I+II without pollinosity, II to V with two marks of silvery pollinosity dorsally, with one mark of golden pollinosity laterally and other silvery pollinosity ventrally. Sternites I to V black with black hairs. Sternite IV (Fig. 3) without hump. Sternite V (Fig. 4) without evident humps at medial part of both arms. First genital segment shining black without silvery pollinosity and with six marginal bristles. Second genital seg- ment orange reddish and with black hairs. Cercus (Fig. 1) slender, curved forward, or- ange reddish on proximal part, black api- cally, with black hairs. Paralobi (Fig. 2) slender. Apical and lateral plates of disti- phallus well developed, with a single lobe (Fig. 6); styli curved, with a crest on con- vex border (Fig. 6). Female.—Unknown Distribution.—Argentina, Santa Cruz. Type material—Holotype ¢ and 24 6 paratypes from Argentina, Santa Cruz: Pto. Santa Cruz, II-1998, J. C. Mariluis; 1 par- atype ¢ same locality but XI-1997, J. C. Mariluis; 2 paratype d same locality but I- 1998, J. C. Mariluis; 3 paratype d from Ar- gentina, Santa Cruz: Rio Gallegos, I-1998, J. C. Mariluis. Etymology.—Named after my father Mr. Antonio Leoncio Mariluis. Discussion.—Microcerella antonioi 1s similar to M. rusca (Hall 1937). However, the male of M. rusca differs from the male of Microcerella antonioi by the first pair of ocellar bristles situated behind or at the same level of the two anterior ocelli, ab- sence of preacrostichals bristles, lower part of sternopleuron with spot of golden polli- nosity, scutellum with a pair of discal scu- tellar bristles, sternite [TV (Fig. 13) with a conspicuous hump behind middle, sternite V (Fig. 14) with evident humps on both arms, cerci (Fig. 11) with inner border un- dulated, paralobi (Fig. 12) heavy, and apical and lateral plate of distiphallus with two lobes. No material in reference to this new species has been found in both of the most 94 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON important Natural History Museums in Ar- gventina (Museo de La Plata and Museo Ar- gentino de Ciencias Naturales ‘‘Bernardino Rivadavia ’’ ). Microcerella aulacophyto Pape (Figs. 7-8) Aulacophyto auromaculata Townsend L919; 1ss'(d, 2) Perm: "Oroya; junior secondary homonym of Euparaphyto au- romaculata Townsend 1919); Hall 1937: 350,360! (key: 6, 93 "Chile" Santiago; “East side’? Andes presumably near Po- trerillos, Mendoza, Argentina); Lopes 1968: 57 (3 paratype, figs. 25—27; Boliv- ia: La Paz, Chile: Iquique and Manina); Lopes 1969: 7 (in catalog; Peru, Chile); Wopes 1978: 757; 758 (oe 2s hese a9: Argentina: Jujuy, Bolivia: La Paz, Chile: Antofagasta and I[quique, Peru: Camacani and Puno); Lopes 1982: 362 (key); Lopes and Tibana 1982: 136 (Chile: Arica); T1- bana and Lopes 1987: 372 (key). Microcerella aulacophyto Pape 1990: 49 (new name for Aulacophyto auromacu- lata Townsend 1919); Pape 1996: 253 (in catalog; Argentina: no further data, Bo- livia, Chile: Biobio and Tarapaca, Peru). Distribution.—Argentina (Jujuy and San- ta Cruz), Bolivia (La Paz), Chile (Antofa- gasta, Arica, Biobio, Iquique, Manina, San- tiago and Tarapaca), Peru (Camacani, Oroya and Puno). New records.—Argentina, Santa Cruz, Pto. Santa Cruz, XII-1997, J. C. Mariluis, 9 36,2 2; same data I-1998, J. C. Mariluis, 5 3, 1 2; same locality but II-1998, J. C. Mariluis, 45 ¢, 2 2; Rio Gallegos, I-1998, J. C. Mariluis, 4 d. Discussion.—The studied specimens agree with the description of A. auroma- culata by Lopes (1968, 1978). Microcerella chilena (Dodge) (Fig. 10) Aulacophyto chilena Dodge 1967: 679 (6, ?; Chile: Antofagasta, Longuén and San- tiago); Lopes 1978: 757, 758 (d para- type, 2, figs. 1O—23, Argentina: Mendo- za, Chile: Aconcagua, Alichahue, Anto- fagasta, Coquimbo and Curico, Peru: Ca- macani, Cuzco and Puno); Lopes 1982: 362 (key); Tibana and Lopes 1987: 372 (key, Chile). Microcerella chilena: Pape 1990: 49 (com- bination), 1996: 254 (in catalog; Chile: Antofagasta and Santiago, Peru). Distribution.—Argentina (Mendoza and Rio Negro), Chile (Aconcagua, Alichahue, Antofagasta, Coquimbo, Curico, Longuén and Santiago), Peru. New records.—Argentina, Rio Negro, Ruta 305, among S. A. Oeste and Pomona, 15-I-1977, J. C. Mariluis, 1 36; Choele- Choel, 18-I-1977, J. C. Mariluis, 1 &. Discussion.—The studied specimens agree with the description of A. chilena by Lopes, (1978): Microcerella penai (Lopes) (Fig. 9) Aulacophyto penai Lopes 1978: 757, 762 (3, &, figs. 28-39, Argentina: Jujuy, Bo- livia: La Paz); Tibana and Lopes 1987: 372 (key; Chile: no further data); Verves 1989: 534 (list). Microcerella penai: Pape 1990: 49 (com- bination), 1996: 257 (in catalog; Argen- tina: Jujuy). Distribution.—Argentina (Jujuy and San- ta Cruz), Bolivia (La Paz), Chile. New records.—Argentina, Santa Cruz, Pto. Santa Cruz, I-1998, J. C. Mariluis, 2 3. Discussion.—The studied specimens agree with the original description. Microcerella rusca (Hall) (Figs. 11-14) Aulacophyto rusca: Hall 1937: 350, 361 (od; Argentina: Bariloche); Lopes 1969: 7 (in catalog; Argentina); Lopes 1978: 757, 758 (key); Tibana and Lopes 1987: 371, 372 (3, 2; figs. 6-12, Chile: Antofagas- ta, Cautin, Coquimbo, Curico, Malleco and Santiago). VOLUME 104, NUMBER | 95 Ss — — ——_ i mae ee ig op ie a ah ee — SA fie oat Figs. 7-14. Microcerella aulacophyto, male. 7, Cerci rear view. 8, Right cerci lateral view. 9, M. penai, male, cerci rear view. 10, M. chilena, male, cerci jiew. Figs. 11-14. M. rusca, male. 11, Cerci rear view. 12, Right paralobi lateral view. 13, Sternite IV lateral view. 14, Sternite V lateral view. Scale = 0.5 mm. 96 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON isca: Pape 1990: 49 (com- 1996: 257 (in catalog; Argen- tina: Rio Negro, Chile: Antofagasta, Cau- tin, Coquimbo, La Araucania, Maule and Micro @} Jinat } bu Santiago). Distribution.—Argentina (Rfo Negro, Santa Cruz), Chile (Antofagasta, Cautin, Coquimbo, Curico, Malleco, Maule and Santiago). New records.—Argentina, Santa Cruz, Pto. Santa Cruz, XII-1997, J. C. Mariluis, 2. G9 .2 same. locality ‘but. 1-19938.-Jz.C. Mariluis, 5 6, 15 2; same locality but II- 1998, J. C. Mariluis, 57 3, 81 2; Rio Ga- llegos, XII-1997, J. C. Mariluis, 3 6; same locality but I-1998, J. C. Mariluis, 7 3d, 8 2; same locality but H-1998, J. C. Mariluis, 16-6622 Discussion.—The studied specimens concur with the description of A. rusca by Tibana and Lopes (1987). ACKNOWLEDGMENTS I thank Dr. Salvador V. Peris, who kindly provided the checklist of Microcerella and for his suggestion to study Neotropical Sar- cophagidae; Drs. Sixto Coscaron, Gustavo R. Spinelli, and Juan A. Schnack for their critical reading of this paper; Drs. Thomas Pape and Rita Tibana for sending me 1m- portant literature about Sarcophagidae; Consejo Nacional de Investigaciones Cien- tificas y Técnicas, Argentina, for the finan- cial support of this research and anonymous reviewers for improving the quality of the manuscript. LITERATURE CITED Dodge, H. R. 1967. Some new American Sarcophag- idae (Diptera). Pacific Insects 9: 679-686. Hall, D. G. 1937. Sarcophaginae, pp. 347—375, 1 pl. In Diptera of Patagonia and south Chile. Part 7, 384 pp., 1 pl. British Museum (Natural History) London. Lopes, H. S. 1968. Notes on paratypes of some Sar- cophagid flies described by C. H. T. Townsend (Diptera). Revista Brasileira de Biologia 28: 51— 60. . 1969. Family Sarcophagidae. Jn Papavero, N., ed. A catalogue of the Diptera of the Americas south of the United States 103: 1.88. Sao Paulo; Departamento de Zoologia, Secretaria da Agricul- tura. . 1978. On Aulacophyto Townsend and Aula- cophytoides n. gen. (Diptera, Sarcophagidae). Re- vista Brasileira de Biologia 38(4): 757—766. . 1982. The genera of Microcerellini (Diptera, Sarcophagidae). Revista Brasileira de Biologia 42(2): 359-369. Lopes, H. S. and R. Tibana. 1982. Sarcophagid flies of Tarapaca, north of Chile (Diptera). Revista Brasileira de Biologia 42: 135-145. Macquart, J. 1851. Dipteres exotiques nouveaux ou peu connus. (Suites du 4 e. supplément). Mémo- ires de la Société des Sciences, de 1 Agriculture, et des Arts de Lille (1850): 134—282. Pape, T. 1990. Revisionary notes on American Sarco- phaginae (Diptera: Sarcophagidae). Tijdschrift voor Entomologie 133: 43—74. . 1996. Catalogue of the Sarcophagidae of the world (Insecta: Diptera). Memoirs on Entomolo- gy, International 8: 1—558. Tibana, R. and H. S. Lopes. 1987. On Aulacophyto and Euparaphyto (Diptera, Sarcophagidae. Microcer- ellini), with descriptions of four new species. Me- morias do Instituto Oswaldo Cruz 82(3): 371-378. Verves, Yu. G. 1989. Prof. Hugo de Souza Lopes and the modern system of Sarcophagidae (Diptera). Memé6rias do Instituto Oswaldo Cruz 84 (Suppl. 4): 529-545. Townsend, C. H. T. 1919. New muscoid genera, spe- cies and synonymy (Diptera). Insecutor Inscitiae Menstruus 6: 157-182. PROC. ENTOMOL. SOC. WASH. 104(1), 2002, pp. 97-105 NEW SPECIES OF THE PLANT BUG GENERA KELTONIA KNIGHT AND PSEUDATOMOSCELIS REUTER (HETEROPTERA: MIRIDAE: PHYLINAE) THOMAS J. HENRY Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart- ment of Agriculture, % National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0168, U.S.A. (e-mail: thenry @sel.barc.usda.gov) Abstract.—The new phyline plant bugs, Keltonia wheeleri from Georgia and South Carolina in the southeastern United States, and Pseudatomoscelis nubila from Province Pedernales in the Dominican Republic, are described. A dorsal habitus illustration and selected scanning electron micrographs of K. wheeleri, male genitalia of both species, and modified identification keys, including photographs of the keyed species [K. rubrofemor- ata (Knight); K. wheeleri, n.sp.; P. flora (Van Duzee); P. insularis Henry; P. nubila, n.sp.; and P. seriata (Reuter)], are provided to help distinguish the new species. Key Words: Heteroptera, Miridae, Phylinae, Keltonia wheeleri, Pseudatomoscelis nubila, new species, Nearctic, Neotropical The mirid genus Keltonia Knight con- tains 12 species ranging from Massachu- setts to Colorado in the United States, south to Colombia, and into the West Indies (Ba- hamas, Jamaica), and the genus Pseudato- moscelis Reuter contains three species, in- cluding the cotton fleahopper [P. seriata (Reuter)], ranging from southern Canada, south to Venezuela, and into the West In- dies (Puerto Rico, St. Thomas) (Henry 1991). Henry (1991) revised these two phy- line genera, described six new species of Keltonia and one new species of Pseuda- tomoscelis, and hypothesized that they rep- resented sister genera, based on the shared stout C-shaped vesica bearing a large sick- le-shaped spicule, clumps or tufts of acu- minate sericeus or scalelike setae on the dorsum, one or two patches of dark bristle- like setae along the inner margin of the cu- neus, and the large dark area on the mem- brane near the apex of the cuneus. Syna- pomorphies defining the species of Keltonia are the dorsal spots limited to the hemely- tra; the conspurcate hemelytral membrane; sericeus or scalelike setae in distinct clumps and, often, in rows along the midline of the head and pronotum; the pale tibiae, with dark spots at bases of spines; and the stout- ly formed, weakly twisted, C-shaped vesi- ca, with a slender spicule and cuplike apical process. Synapomorphies defining the spe- cies of Pseudatomoscelis are the three or more spots on antennal segment II; the brown to black hemelytral membrane, with a black mark just posterior to the apex of the cuneus; the sericeus or scalelike setae in patches or clumps, but not in rows along midline (as in species of Keltonia); the stoutly formed, C-shaped vesica (lacking an apical cuplike process); and the phallotheca with a subapical spine (except in P. nubila MSPs): In this paper, I describe K. wheeleri to provide a name for a forthcoming paper on its seasonal history, host, and distribution (Wheeler, in preparation), and P. nubila to provide a name for my forthcoming list of 98 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON the Miridae of the Dominican Republic. An adult dorsal habitus illustration and scan- ning electron micrographs of selected struc- tures of K. wheeleri; figures of the male genitalia for both species; and modified keys, including photographs of species keyed in Henry (1991) [K. rubrofemorata (Knight); K. wheeleri, n. sp.; P. flora (Van Duzee); P. insularis Henry; P. nubila, n. sp.; and P. seriata (Reuter)], are provided to help distinguish the new species. Acronyms for type depositories as fol- lows: AMNH (American Museum of Nat- ural History, New York, NY); CNC (Ca- nadian National Collection, Ottawa, Ontar- io); TAMU (Texas A & M University, Col- lege Station, TX); and USNM (National Museum of Natural History, Smithsonian Institution, Washington, DC). Keltonia wheeleri Henry, new species (Figs. 1-8, 10—13) Diagnosis.—Keltonia wheeleri (Figs. 1, 8) is distinguished from all other species of the genus in having a shiny yellow dorsum (including the head, antenna, and pro- and mesofemora), a large centrally positioned fuscous spot on the hemelytra, an apically orange or red-tinged metafemur, and a fus- cous or black membrane sprinkled with white spots. Description—Male (n = 10): Length 2.45—2.85 mm, width 1.02 mm. Head (Figs. 2, 3): Width 0.56—0.59 mm, vertex 0.34— 0.38 mm. Rostrum: Length 0.74—0.77 mm, extending to mesocoxae. Antenna: Segment I, length 0.18—0.19 mm; II, 0.58—0.66 mm; II, 0.43-0.50 mm; IV, 0.37—-0.40 mm. Pronotum: Length 0.34—0.35 mm, _ basal width 0.78—0.88 mm. Female (n = 10): Length 2.50—2.72 mm, width 1.08—1.10 mm. Head: Width 0.56— 0.59 mm, vertex 0.35—0.37 mm. Rostrum: Length 0.77—0.83 mm, extending to meso- coxae. Antenna: Segment I, length 0.16— 0.19 mm; IJ, 0.64—0.67 mm; III, 0.43—-0.51 mm; IV, 0.42 mm. Pronotum: Length 0.32— 0.34 mm, basal width 0.82—0.83 mm. Overall coloration shiny yellow to green- ish yellow. Head, antenna, and pronotum uniformly yellowish to greenish yellow. Hemelytron yellow to greenish yellow, api- cal half of clavus and inner area of corium from middle of clavus to base of cuneus forming a large, dark patch or cloud. Ven- tral surface uniformly greenish yellow. Legs mostly yellowish green; metafemur tinged with orange or red on apical % and anterior half finely brown spotted; pale tib- ial spines with vague pale brown spots at bases; tarsi yellowish; claws brown. Vesti- ture of two types: simple, erect and semi- erect, pale brown setae on head, pronotum, and on greenish-yellow areas of hemely- tron, simple fuscous to black setae on dark central area; and silvery sericeus or slender scalelike setae scattered on head, pronotum, and dark areas of hemelytron, a few clusters of sericeus setae along inner margins of eyes (Fig. 3), but without a median row of scalelike setae on head, pronotum, and scu- tellum typical of most other species in the genus. Male genitalia: Genital capsule (Figs. 6, 7); vesica (Fig. 10); left paramere (Fig. 11); right paramere (Fig. 12); phallotheca (Fig. 1)) Etymology.—This species is named in honor of Dr. A. G. Wheeler, Jr., long-time friend, colleague, and discoverer of this at- tractive new species. Host.—Polygonella americana (Fischer- Meyer) Small [Polygonaceae]. Distribution.—Known only from the southeastern United States (Georgia and South Carolina). Type material.—Holotype ¢: South Car- olina, Aiken Co., Henderson Heritage Pres[erve]., N. of Aiken, 30 May 1998, A. G. Wheeler, Jr., taken on Polygonella amer- icana (USNM). Paratypes: GEORGIA: 2 2, Burke Co., Yuchi Wildlife Mgt. Area, 4 mi. NNE of Girard, 1 July 1998, AGW, on P. americana (USNM); 4 6, 2 2, Colum- bia Co., Rt. 383, E of Lewiston, 5 July 1998, AGW, on P. americana (USNM); 1 3,5 2, Glascock Co., Rt. 80, NE of Gib- son, 5 July 1998, AGW, on P. americana VOLUME 104, NUMBER 1 Fig. 1. Dorsal habitus illustration of Keltonia wheeleri °. 99 100 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 2-7. (122). 3, Clusters of scalelike setae bordering compound eye (805%). 4, Metathoracic scent gland auricle and evaporatory area (326). 5, Claw (1,170). 6, Lateral aspect of genital capsule, with vesica extended (114%). 7, Caudal aspect of genital capsule, with vesica extended (118%). (USNM); 5 6, 3 2, Jefferson Co., Rt. 1, 3.5 mi NE of Wrens, | July 1998, AGW, on P. americana (USNM); 13 6, 17 2, Mc- Duffie Co., Rt. 221, 5 mi. S of Harlem, 11 Oct. 1998, AGW, on P. americana (AMNH, CNC, USNM, and TAMU); 2 6, 5 2, McDuffie Co., CR [county road]-86, nr jct rts 78 & 278, W of Dearing, 5 July 1998, AGW, on P. americana (USNM); 5 3,2 2, Richmond Co., Rt. 1, Blythe, 1 July 1998, A. G. Wheeler, Jr. [AGW], taken on Scanning electron micrographs of Keltonia wheeleri, 3. 2, Head and pronotum, lateral aspect P. americana (USNM); 4 6, 6 @, Rich- mond Co., Wheeler Rd., Augusta, | July 1998, AGW, on P. americana (USNM); 4 3, 5 %, Richmond Co., Augusta, 18 Oct. 1998, AGW, on P. americana (USNM). SOUTH CAROLINA: 7 6, 32 ¢%, same data as for holotype (AMNH, CNC, USNM); 1 3, 6 2, same data as for holo- type, except taken on 5 June 1998 (USNM); 3 9, Aiken Cos. Rt. 1 and*CR=1512 Weer Monetta, 28 June 1998, A. G. Wheeler, Jr. VOLUME 104, NUMBER 1 101 Bigs: 8, 9: coll., on P. americana (USNM); 1 3,7 &, PMikciee@oe ke 25. 177 mi. SE of Beech Island, 5 July 1998, AGW, on P. americana (USNM); 4 6, Aiken Co., Rt. 20w, exit 33, NW of New Holland, 28 June 1998, AGW, on P. americana (USNM); 2 2, Aiken Co., CR-1121, nr. Rt. 39, 4 mi. W of Wagener, 28 June 1998, AGW, on P. americana COSNM): 2. o-5' 2s Aiken Co., Rt. 19), 1 mi. S of Eureka, 18 Oct. 1998, AGW, on P. americana (USNM); 2 3,5 2, Barnwell Co, Rie39) 02 mr N of Rt. 278, SW of Williston, 18 Oct. 1998, AGW, on P. amer- jeana (USNM);1°G; 2 2; Edgefield Co, Bettis Acad. Rd., 4.5 mi. SSW of Trenton, 5 June 1998, AGW, on P. americana (USNM); 1 6, 1 2, Edgefield Co., S-429 at junc Rt 25, 5.5 mi. N. of Augusta, 5 July 1998, AGW, on P. americana (USNM); 1 2, Edgefield Co., Rt. 25, 6.5 mi. N of Ai- ken Co. line, S of Trenton, 28 Nov. 1998, AGW, on P. americana (USNM); 4 6,8 @, Edgefield Co., Mt. Zion Rd., NE of North Augusta, 5 June 1998, AGW, on P. ameri- cana (USNM); 5 6, 6 @, Lexington Co., CR-445, E of Rt. 178, SE of Batesburg, 28 June 1998, AGW, on P. americana (USNM); 2 2, Lexington Co., Rt. 178, 0.3 mi E of Rt. 20, N of Steedman, 28 June Dorsal photographs of Keltonia spp. 8, K. wheeleri, 3.9, K. rubrofemorata, 3. 1998, AGW, on P. americana (USNM); 8 3,7 2, Orange Co., CR-1609, E of Rt. 39, 2 mi. SE of Salley, 28 June 1998, AGW, on P. americana (USNM, TAMU). Discussion.—Keltonia wheeleri (Figs. 1, 8) is one of the more distinct species of the genus in having a bright, shiny yellow, dor- sal coloration, with a large, dark, centrally located mark through the middle of the co- rium and across the apex of the clavus. It is most closely related to K. rubrofemorata (Fig. 9), the type of the genus, based on the overall coloration, male genital structures, and the unusual shared host genus (Poly- gonella Michx. [Polygonaceae]). Keltonia wheeleri has the head, all antennal seg- ments, and the pro- and mesofemora yel- lowish, with only the apex of the metafe- mur tinged with pale red or orange, has a shorter antennal segment II (0.58—0.66 mm in 6; 0.64—0.67 mm in @), has a larger he- melytral spot (Figs. 1, 8), and is found only on Polygonella americana, whereas K. rub- rofemorata has the head, first antennal seg- ment (and often segments II-IV), and all femora dark red, has a proportionately lon- ger antennal segment II (0.68—0.70 mm in 3, 0.68—0.76 mm in 2), has a smaller he- 102 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 10-13. Phallotheca. melytral spot (Fig. 9), and is found only on P. myriophylla (Small) Horton. Couplet 2 of my key to the species of the genus (Henry 1991) may be modified as follows to accommodate K. wheeleri: 2a. Dorsum bright, shiny yellow; hemelytra lack- ing small brown spots or speckles surround- ing brown cloud or solid dark area at middle; metafemur dark red or tinged with orange or regkapicallivgem sey aie sees es i ee eee 2b — Dorsum dull, sometimes phosphorescent yel- low; hemelytra with numerous small spots or speckles surrounding brown cloud or dark area at middle; femora pale yellow to green- ish yellow, never tinged with orange or red 2b. Head, antennal segment I, and all femora dark red (Fig. 9) K. rubrofemorata Knight — Head, antennal segment I, and pro- and meso- femora pale yellow (Figs. 1, 8); metafemur tinged with orange or red only on apical % K. wheeleri, n. sp. Pseudatomoscelis nubila Henry, new species (Figs. 16, 18-21) Diagnosis.—Pseudatomoscelis nubila (Fig. 16) is distinguished from all other spe- cies of the genus by the uniformly brown- spotted, yellow to greenish-yellow dorsum, with an irregular, transverse, brown band Male genitalia of Keltonia wheeleri. 10, Vesica. 11, Left paramere. 12, Right paramere. 13, across the middle of the corium, and the mostly fuscous to black hemelytral mem- brane, having a pale or white spot near the apex of the cuneus and a narrower spot just beyond first spot. Description.—Male (n = 10): Length 2.48—2.72 mm, width 1.13—1.16 mm. Head: Width 0.64—0.66 mm, vertex 0.29—0.30 mm. Rostrum: Length 1.06—1.12 mm, ex- tending to middle of abdomen. Antenna: Segment I, length 0.18—0.20 mm; II, 0.67— 0.77 mm; III, 0.40—0.43 mm; IV, 0.33—0.35 mm. Pronotum: Length 0.50—0.53> mm, basal width 0.96—1.16 mm. Female (n = 10): Length 2.56—2.80 mm, width 1.20—-1.24 mm. Head: Width 0.60— 0.62 mm, vertex 0.32—0.34 mm. Rostrum: Length 1.02—1.14 mm, extending to base of ovipositor. Antenna: Segment I, length 0.20—0.21 mm; II, 0.65—0.77 mm; II, 0.36— 0.38 mm; IV, 0.29-0.30 mm. Pronotum: Length 0.48—0.49 mm, basal width 0.98— 1.02 mm. Overall coloration golden yellow, evenly brown spotted on pronotum and hemelytra. Head golden yellow, paler along median line, with scattered brown or fuscous spots, mostly along inner margins of eyes and across base of vertex. Antenna yellow; seg- VOLUME 104, NUMBER 1 103 Figs. 14-17. Dorsal photographs of Pseudatomoscelis spp. 14, P. flora, 2. 15, P. insularis, 3. 16, P. nubila, OF 7a FP. seriata, 6. ment I with a partial to complete subapical and subbasal black band; segment II yel- low, with four or five distinct black spots; segment III yellow to yellowish brown, nar- rowly fuscous at base; segment IV brown, narrowly black at base. Pronotum evenly brown spotted, less so on anterior 3 across calli; mesoscutum and scutellum evenly brown spotted. Hemelytron evenly brown spotted, with spots coalescing to form an 104 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 18-21. 21, Phallotheca. irregular band across middle of corium; membrane fuscous or black, with two pale spots, one near apex of cuneus and a second narrower one just beyond first spot, some- times slightly paler between areoles; veins yellowish, becoming fuscous on basal half or more of large areole. Ventral surface pale yellow, with a few scattered spots on ab- domen; pleural area of pronotum often with an orange tinge or dash just above proace- tabula. Legs yellow; coxae with a few scat- tered brown spots; femora evenly brown spotted; tibiae with large fuscous spots at bases of black spines; tarsi yellow; claws brown. Vestiture of two types: simple, erect, black setae arising from dorsal spots, black setae on head considerably longer than those on pronotum and hemelytra; and scattered clusters containing three to nine silvery, sericeus or scalelike setae on pron- otum, scutellum, hemelytra, and along inner margin of eyes and across vertex of head. Male genitalia: Vesica (Fig. 18); left paramere (Fig. 19); right paramere (Fig. 20); phallotheca (Fig. 21). Male genitalia of Pseudatomoscelis nubila. 18, Vesica. 19, Left paramere. 20, Right paramere. Etymology.—The specific name is from the Latin “‘nubila,”” meaning cloud, to draw attention to the mottled brownish band or cloud across the middle of the corium. Host.—This species was collected on a shrublike species of Croton (Euphorbi- aceae) growing in an arid habitat, contain- ing native cacti, along the coast near Cabo Rojo, Dominican Republic. Distribution—Known only from south- western Dominican Republic (Pedernales). Type material—Holotype ¢, Dominican Republic, Prov. Pedernales, 5 km N Cabo Rojo, nr sea’ level) 1757 59 Nal7 39 Oly. 12 April 2000, T. J. Henry & R. E. Wood- ruff, taken on Croton sp. (USNM). Para- types: 17 5d, 11 @, same data as for holo- type (1 6, 12, AMNH; 1 6, 1 2 CNG; 15 3, 9°22 USNM): Discussion.—This new species is most similar to P. insularis Henry (Fig. 15) and P. seriata (Reuter) (Fig. 17) in the overall brown-spotted dorsum. From P. seriata, it can be distinguished by the transverse brown band on the corium and the fuscous VOLUME 104, NUMBER | membrane and the stouter, more curved ve- sica (Fig. 18), having more pronounced spi- culi. From P. insularis, it can be separated by the larger size, transverse corial band, and shape of the vesica. All of the above brown-spotted species are distinct from the uniformly yellowish-orange P. flora (Van Duzee) (Fig. 14), which lacks dorsal spots. KEY TO SPECIES OF PSEUDATOMOSCELIS 1. Dorsum immaculate, without spots (Fig. 14); body yellow to yellowish orange; Arizona and Mexico flora (Van Duzee) — Dorsum always with dark spots; body pale yel- low to dark green, sometimes tinged with yel- lowishtoranpemefrts: .ttat AbAs Seles. J 8 2 i) . Inner margin of cuneus without or with only vague spots bearing 2 or 3 bristlelike setae (Fig. 15); vesica of male short and stout; Puerto Rico and St. Thomas, West Indies ........ insularis Henry — Inner margin of cuneus with | or 2 distinct dark spots bearing numerous dark bristlelike setae; vesica of male more slender ........ 3 3. Dorsum uniformly brown spotted (Fig. 17); he- melytral membrane mostly hyaline, often tinged with brown basally and with only a black dash just beyond apex of cuneus:; wide- spread from Saskatchewan, Canada, to Vene- zuela, and the West Indies .... . seriata (Reuter) — Dorsum uniformly brown spotted, but also usu- ally with an irregular brown band across mid- dle of corium (Fig. 16); hemelytral membrane mostly fuscous or black, with two pale areas, one just beyond apex of cuneus and a second 105 more narrow one slightly beyond the first; Do- Mimican Republic ua eee eee nubila, n. sp. ACKNOWLEDGMENTS I am grateful to A. G. Wheeler, Jr. (De- partment of Entomology, Clemson Univer- sity, Clemson, SC) for providing me with the specimens of the interesting new species of Keltonia and R. E. Woodruff (Gaines- ville, Florida) for inviting me to join him on a productive, month-long collecting ex- pedition to the Dominican Republic that yielded numerous interesting and several new species of Heteroptera, including Pseu- datomoscelis nubila. 1 also thank Kathleen Schmidt (Hillsdale, New York) for the adult habitus illustration of K. wheeleri and Michele Touchet (Systematic Entomology Laboratory [SEL], ARS, USDA, % USNM) for the dorsal photographs used in this paper. M. G. Pogue (SEL), D. R. Smith (SEL), and A. G. Wheeler, Jr. kindly re- viewed the manuscript and offered useful comments. LITERATURE CITED Henry, T. J. 1991. Revision of Keltonia and the cotton fleahopper genus Pseudatomoscelis, with the de- scription of a new genus and an analysis of their relationships (Heteroptera: Miridae: Phylinae). Journal of the New York Entomological Society 99: 351-404. PROC. ENTOMOL. SOC. WASH. 104(1), 2002, pp. 106-110 NEW SPECIES OF NOTHOTRICHIA FLINT (TRICHOPTERA: HYDROPTILIDAE) FROM BRAZIL AND COSTA RICA RALPH W. HOLZENTHAL AND STEVEN C. HARRIS (RWH) Department of Entomology, University of Minnesota, St. Paul, MN 55108, U.S.A. (e-mail: holze001 @tc.umn.edu); (SCH) Department of Biology, Clarion Univer- sity, Clarion, PA 16214, U.S.A. (e-mail: harris @clarion.edu) Abstract.—The genus Nothotrichia was thought to be amphitropical in distribution. Two species, N. cautinensis Flint and N. illiesi Flint, are known from Chile and a third, N. shasta Harris and Armitage was described from California in the United States. The addition of two new species, N. tupi, from Brazil, and N. munozi, from Costa Rica, indicates that the genus occurs sporadically across the Neotropics. Key Words: distribution Flint (1967) erected the genus Nothotri- chia for the Chilean species N. illiesi. A second species from Chile, N. cautinensis Flint, was added in 1983 (Flint 1983). The genus was thought to be endemic to the Chilean subregion until a third species, N. shasta Harris and Armitage, was discovered in North America from northern California. It was suggested that the genus displayed an amphitropical distribution (Harris and Armitage 1997). The discovery of two ad- ditional new species from Brazil and Costa Rica, collected as part of ongoing invento- ries of the caddisfly faunas of those coun- tries, expands the distribution of the genus substantially. Nothotrichia are infrequently collected and specimens within collections are few, thus the genus may occur in inter- vening areas. Specimens are deposited in the collec- tions of the Museu de Zoologia da Univ- ersidade de Sao Paulo, Brazil (MZUSP), the University of Minnesota Insect Collec- tion, St. Paul, Minnesota (UMSP), and the National Museum of Natural History, Smithsonian Institution, Washington, D.C. (NMNH). Trichoptera, Brazil, Costa Rica, new species, Nothotrichia, microcaddisfly, Nothotrichia munozi Holzenthal and Harris, new species (Fig. 1) This new species shares with its conge- ners features of the head and thorax typical for the genus (Harris and Armitage 1997). Although quite different in many details of the male genitalia, this species is more closely aligned with N. cautinensis Flint and N. shasta Harris and Armitage than with WN. illiesi Flint. With N. cautinensis, N. munozi shares the feature of long narrow processes from the dorsum of segment IX; and with N. shasta, the elongate process from the base of the inferior appendage. The new species is immediately distin- guished by the longer inferior appendage, which also has a large dorsal spine at mid- length and a very long inner process from the base. Male.—Length 2.3 mm. Antenna simple with 25 segments. Preserved in alcohol with no evident pattern to the wings. Ab- dominal segment VII with short ventrome- sal process. Segment VIII annular, dorsum slightly wider than venter. Segment IX VOLUME 104, NUMBER 1 107 Fig. 1. Nothotrichia munozi, 3 genitalia. A, Lateral. B, Ventral. C, Dorsal. D, Phallus, lateral. E, Phallus, dorsal. 108 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON rae , 2D Fig. 2. Nothotrichia tupi, 3 genitalia. A, Lateral. B, Ventral. C, Dorsal. D, Phallus, lateral. E, Phallus, dorsal. VOLUME 104, NUMBER | largely enclosed within VIII, in lateral view narrowing dorsally, with elongate thin pro- cesses extending posteriorly, posteroven- trally with setose lobe; in dorsal and ventral views, rounded anteriorly; ventrally with posterior setose lobe divided mesally. Seg- ment X elongate and rectanguloid in lateral view; dorsally a narrow membranous lobe, incised distally. Inferior appendages thin and elongate in lateral view, heavy sclero- tized spinose process at midlength extend- ing dorsad; in ventral view narrow over length, thin processes originating mesally at base and extending beyond midlength, apex tipped with pair of setae. Phallus thin and tubular, apex divided into two processes, one sickle-shaped, the other tubular and bearing the ejaculatory duct. Female.—Unknown Type material—Holotype, 6. COSTA RICA: Guanacaste: Area de Conservacion Guanacaste, Parque Nacional Guanacaste, Estaci6n Maritza, Rro Tempisquito, 10.958°N, 85.497°W, el. 550 m, February 1994, Fdo. Munoz-Q (UMSP). Etymology.—Named for Fernando Mu- noz, who collected the specimen and con- tributed much to our knowledge of the Cos- ta Rican caddisfly fauna. Nothotrichia tupi Holzenthal and Harris, new species (Fig. 2) In overall appearance of the genitalic structures, particularly the inferior append- ages, this new species is very similar to N. munozi. Both have elongate inferior ap- pendages, with a dorsal process arising near midlength, which is spinose in N. munozi and rounded in the new species, and both have an elongate process originating at the base of the appendages. The new species lacks the narrow dorsal processes from seg- ment IX, seen in both N. munozi and N. cautinensis, and as well segment X tapers distally in the new species, similar to that seen in N. cautinensis and N. shasta, as op- posed to the truncate distal shape seen in N. munozi. 109 Male.—Length 2.4—2.7 mm. Antenna simple with 25 segments. Brown in color with diffuse silver hairs on forewing and patches of silver hair on thorax and head. Abdominal segment VII annular, lacking ventromesal process. Segment VIII annular, posterior margin emarginate ventrally. Seg- ment IX largely enclosed within segments VII and VIII, in lateral view narrowing dor- sally and tapering anteriorly; in dorsal view triangular, and seemingly fused with X pos- teriorly, series of setae posterolaterally. Seg- ment X short in lateral view, tapering dis- tally to knob-like apex, anteriorly with elon- gate sclerite within segment IX; dorsum di- vided posteriorly with sclerous lateral margins, anteriorly with sclerotized bands laterally at juncture with segment IX. Infe- rior appendages thin and elongate in lateral view, tipped apically with stout spine, rounded dorsal process near midlength, elon- gate process originating at base and extend- ing dorsad; in ventral view thin over length and curving inward, basal process appearing to be short, midlength process squarish. Phallus tubular with ribbon-like sheath orig- inating near midlength, apex bearing pair of short lateral processes, one of which is as- sociated with the ejaculatory duct. Female.—Unknown. Type material—Holotype, ¢. BRAZIL: Minas Gerais: Parque Estadual Itacolomi, Rio Belchior, 20°25.041'S, 43°25.633’W, el. 725 m, 2.11.1998, Holzenthal & Paprocki (MZUSP). Paratypes: Same data as holo- type, | 6 (UMSP), 1 d (NMNH). Etymology.—Named for the Tupi-Gua- rani, the indigenous people of southeastern Brazil. ACKNOWLEDGMENTS We thank David Houghton and Fernando Munoz, University of Minnesota, for re- viewing the manuscript. Appreciation is also extended to Brian J. Armitage, Ohio Biological Survey, and Oliver S. Flint, Jr., Smithsonian Institution, for their comments on the manuscript. This research was sup- ported by NSF grants DEB-9400632 to 110 RWH for research in Costa Rica, and DEB- 9971885 to RWH and R.J. Blahnik for re- search in Brazil. This support is gratefully acknowledged. LITERATURE CITED Flint, O. S., Jr. 1967. Studies of Neotropical caddis flies, Il: Trichoptera collected by Prof. Dr. J. Ilies in the Chilean subregion. Beitrage zur Neotro- pischen Fauna 5: 45—63. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON . 1983. Studies of Neotropical caddisflies, XXXIII: New species from austral South America (Trichoptera). Smithsonian Contributions to Zo- ology 377: 1-100. Harris, S. C. and B. J. Armitage. 1997. New member of the Chilean genus Nothotrichia from North America (Trichoptera: Hydroptilidae), pp. 123— 128. In Holzenthal, R. W. and O. S. Flint, Jr., eds. Proceedings of the 8th International Symposium on Trichoptera. Ohio Biological Survey, Colum- bus, Ohio. PROC. ENTOMOL. SOC. WASH. 104(1), 2002, pp. 111-118 A NEW GENUS AND A NEW SPECIES OF DALADERINI (HEMIPTERA: HETEROPTERA: COREIDAE) FROM MADAGASCAR HARRY BRAILOVSKY Departamento de Zoologia, Instituto de Biologia, U.N.A.M., Apdo Postal No. 70-153, México, 04510 D.E, México (e-mail: coreidae @ servidor.unam.mx) Abstract.—Kerzhnercryptes perinetus, n. gen., n. sp., from Madagascar are described in the tribe Daladerini (Coreidae). A habitus view illustration and drawings of the anten- nae, pronotum, legs, abdomen, and male and female genitalia are provided. A key to the known genera of Daladerini from Madagascar is included. Key Words: agascar The Coreidae fauna of Madagascar is rich and diverse but has been studied little by modern workers. The tribe Daladerini at present is composed of 8 genera and 28 nominal species distributed throughout the Old World tropics, with the majority of spe- cies and the greatest diversity in southern Africa. The Madagascar Daladerini have never been revised. The first species recorded was by Stal (1873), who described a new genus and one new species, Odontorhopala cal- losa. Distant (1879) studied the Madagascar Hemiptera and added one new genus, Par- abrachytes, with two new species, P. co- loratus and P. obscurus, and later Distant (1893) described Odontorhopala bergrothi and discussed the generic position of Odon- torhopala and Parabrachytes. Bergroth (1906) described a new genus and one new species, Rhombolaparus tardigradus, and later Bergroth (1912) added the new spe- cies, Odontorhopala geminata. Garcia Var- ela (1913) described Parabrachytes longi- cornis and gave new distribution records for Odontorhopala callosa and Parabrach- ytes coloratus. Here I add a new genus and a new spe- Insecta, Heteroptera, Coreidae, Daladerini, new genus, new species, Mad- cies in order to make the name available for discussion of the zoogeography of the tribe. The following abbreviations are used for the institutions cited in this paper: MNHN (Muséum National d’Histoire Naturelle, Paris, France), UNAM (Instituto de Biolo- gia, Universidad Nacional Aut6noma de México), ZISP (Zoological Institute, St. Pe- tersburg, Russia). All measurements are given in millime- ters. CHECKLIST OF THE GENERA AND SPECIES OF MADAGASCAR DALADERINI Kerzhnercryptes Brailovsky, new genus. K. perinetus Brailovsky, new species. Odontorhopala Stal 1873: 55—56. O. bergrothi Distant 1893: 54. O. callosa Stal 1873: 56. O. geminata Bergroth 1912: 82-83. Parabrachytes Distant 1879: 213-214. P. coloratus Distant 1879: 214. P. longicornis Garcia Varela 20-21. P. obscurus Distant 1879: 214-215. Rhombolaparus Bergroth 1906: 17-18. R. tardigradus Bergroth 1906: 18. 1913: 2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Kerzhnercryptes Brailovsky, new genus Diagnosis.—This genus appears to be closely related to Parabrachytes Distant (1879) particularly due to the relatively nar- row, non-expanded abdomen (Fig. 9), and the humeral angles of the pronotum not conspicuously expanded as a wing-like lobes (Figs. 7-8). Kerzhnercryptes has an- tennal segment II almost cylindrical without distal spines (Fig. 1), the rostrum elongate, slender and reaching abdominal sternite II, each femur slender and armed ventrally with two small subapical spines (Fig. 16), the humeral angles of the pronotum sub- acute (Fig. 7), and the tibiae longer, and more slender. Parabrachytes is distin- guished by having antennal segment II dis- tally clavate and armed with strong spines (Fig. 3), the rostrum short, robust and not extending beyond the middle third of the mesosternum, the femora robust and ven- trally armed with two rows of stout spines (Fig. 5), the humeral angles rounded, and obtuse (Fig. 8), and the tibiae short, and robust. Rhombolaparus Bergroth (1906) is clear- ly distinguished by having the abdomen conspicuosly dilated, rhomboidal, and the humeral angles of pronotum laterally ex- panded. Generic description.—Body medium sized to large, moderately elongate. Head: Across eyes wider than long, subquadrate, dorsally flat; tylus conspicuously deflexed, unarmed, apically truncated, and shorter than juga; juga produced forward on a strong conical tubercle, extending anteriorly to tylus and slightly raised in lateral view; vertex with deep longitudinal groove along midline; antenniferous tubercle large, prom- inently produced, wide, separated by dis- tance equal to their own width; sides of head in front of eyes almost straight; anten- nal segment I slightly robust, thickest and much longer than head; segments IIT and III cylindrical, slender; segment IV fusiform; antennal segment I longest, II longer than Ill, IV the shortest (Fig. 1); ocelli scarcely tuberculate; preocellar pit deep, diagonally excavated; eyes protruding, hemispherical; postocular tubercle protuberant; buccula rounded, raised, short, not extending be- yond antenniferous tubercle, with short spine anterior projection; rostrum reaching anterior third of abdominal sternite III; mandibular plate unarmed. Thorax: Pronotum wider than long, tra- peziform, moderately declivent; collar not distinctly marked; anterolateral margins obliquely straight, coarsely nodulose; fron- tal angles obtuse; humeral angles produced laterad, directed upward and each margin finely nodulose (Fig. 7); posterolateral mar- gin sinuate, with upper half nodulose and inner half smooth; posterior margin smooth, concave; callar region indistinct, transverse- ly flat, separated along midline by an ob- scure longitudinal groove; posterior margin with transverse ridge, distinctly raised; prosternum with deep excavation; meso- sternum with a broad, medio-longitudinal groove; metasternum entire; anterior lobe of metathoracic peritreme elevated, reniform, posterior lobe sharp, small. Legs: Femora slightly incrassate, surface smooth, armed with two small anteapical spines; tibiae cylindrical, sulcate, unarmed. Scutellum: ‘Triangular, flat, relation length-width, with two conditions indepen- dent of the sex, longer than wide, or wider than long, and transversely striated; apex short, acute. Hemelytra: Macropterous, reaching apex of last abdominal segment; costal margin with a shallow groove; apical margin obliquely straight, with short apical angle almost reaching middle third of hemelytral membrane. Abdomen: Connexival segments higher than margin of hemelytron at rest; upper margin smooth; posterior angles simple, not spinose; posterior third of connexival seg- ments III, [V and VII straight, and posterior third of segments V and VI depressed, con- cave or diagonally truncated; abdominal spiracle closer to anterior margin than to posterior margin. VOLUME 104, NUMBER 1 113 Figs. 1-5. 1-3, Antenna. 1, Kerzhnercryptes perinetus. 2, Odontorhopala callosa. 3, Parabrachytes color- atus. 4, 5, Hind leg. 4, O. callosa. 5, P. coloratus. 114 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 6-10. 6-8, Pronotum. 6, Odontorhopala callosa. 7, Kerzhnercryptes perinetus. 8, Parabrachytes co- loratus. 9, 10, Abdomen. 9, K. perinetus. 10, O. callosa. VOLUME 104, NUMBER 1 Integument: Body surface rather dull, al- most glabrous. Dorsal head, pronotum, scu- tellum, clavus, corium, propleura, meso- pleura, metapleura, abdomen, and exposed parts of genital segments of both sexes punctate. Ventral head, calli, connexival segments, prosternum, mesosternum and metasternum impunctate; antenniferous tu- bercles granulate. Male genitalia: Genital capsule: Poste- rior margin simple, convex (Fig. 13). Par- ameres: Shaft robust; anterior lobe convex, posterior lobe short and slender (Figs. 11— Wa Female genitalia: Abdominal sternite VII with plica and fissura; plica triangular, reaching anterior third of sternite VII; gon- ocoxae I enlarged dorso-ventrally, in caudal view closed, in lateral view convex, with upper margin rounded; paratergite VIII tri- angular, with spiracle visible but hard to see; paratergite IX squarish, larger than par- atergite VIII (Fig. 15). Spermatheca: Distal bulb spherical; sclerotized duct leading from bulb moderately coiled; chamber elon- gate with acute lateral spines, distal duct straight and widely membranous duct (Fig. 14). Etymology.—Named for I. M. Kerzhner distinguished Russian hemipterist. Type species.—Kerzhnercryptes perine- tus, New species. Kerzhnercryptes perinetus Brailovsky, new species (Bigs. 100729 ala), Description.—Measurements: Male: Head length 2.00; width across eyes 2.23; interocular space 1.30; interocellar space 0.62; preocular distance 1.32; length anten- nalasesments: 1, 4-71; I, 4.96; IL, 3.96; TV, 2.91. Pronotum: Total length 3.78; width across frontal angles 1.73; width across hu- meral angles 6.69. Scutellar length 2.31; width 2.29. Total length 21.10. Female: Head length 2.12; width across eyes 2.41; interocular space 1.50; interocellar space 0.65; preocular distance 1.44; length anten- nal segments: I, 5.08; I, 5.56; I, 4.46; IV, ts) 3.10. Pronotum: Total length 4.34; width across frontal angles 1.98; width across hu- meral angles 7.87. Scutellar length 2.88; width 2.91. Total length 24.80. Male coloration: Dorsal coloration: Head, pronotum, clavus and corium chest- nut orange; scutellum yellow with apex chestnut orange and a black square at basal middle third; corium with black spots scat- tered along costal border; connexival seg- ments black with anterior third or anterior half orange yellow; dorsal abdominal seg- ments shining orange yellow; hemelytral membrane pale orange brown; antennal segment I yellow, Hl and III yellow with apical third black, and IV creamy yellow. Ventral coloration: Pale orange yellow with following areas black: apex of rostral seg- ment IV, anterior and posterior lobe of metathoracic peritreme, 1 to 3 discoidal spots on coxae, dense discoidal spots on femora and tibiae and few scattered spots on abdominal sterna III to VII. Female coloration: Similar to male. Dorsal abdominal segments VIII and IX and genital plates pale orange yellow. Types.—Holotype: ¢, Madagascar, An- alamasotra, Pr. Perinet, December 1930 (without collector) (ZISP). Paratypes: 3 6, 4 2, same data as holotype (UNAM, ZISP). 2 36,2 ¢, Madagascar, Perinet, December 1932 (without collector) (ZISP). 6 6, 6 @, Madagascar, Dist. Mananara, N. Mont An- tampona, July 1965, Vadon, Peyrieras (MNHN, UNAM). Etymology.—Named for the type locali- ty; a noun in apposition. Distribution.—Known only from Mada- gascar. KEY TO THE KNOWN GENERA OF DALADERINI FROM MADAGASCAR 1. Antennal segment II clavate, with distal third armed with stout spines (Figs. 2—3); hind femur robust, ventrally armed with two rows of stout andilarseuspines: (Rissa— 5) sien ene 2, — Antennal segment II almost cylindrical without distal spines (Fig. 1); hind femur elongate, slender, slightly thickened toward apex, and 116 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Riese i>) Kerzhnercryptes perinetus. 11 Spermatheca. 15, Female genital plates. . 12, Parameres. 13, Male genital capsule in caudal view. 14, 117 VOLUME 104, NUMBER 1 oF ut Sys ete chutou inetus. ptes perin hnercry VEZ Dorsal view of Ke Fig. 16. 118 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON two minute or indistinct spines ifrably with eral angles obtuse, rounded, not expand- ed; abdominal segments not expanded laterally (Fig. 8) Parabrachytes Distant Humeral angles strongly produced laterally and anteriorly into rounded wing-like lobes (Fig. 6); abdomen strongly and abruptly dilated (Fig. LO) eee See tore Sos chore eae Odontorhopala Stal 3. Abdomen strongly expanded in middle, rhom- boidal in outline ..... Rhombolaparus Bergroth — Abdomen not expanded, relatively narrow, par- allel-sided (Fig. 9) Kerzhnercryptes Brailovsky, new genus ACKNOWLEDGMENTS Sincere thanks is given to Dra. Domi- nique Pluot (MNHN) and Dr. I. M. Ker- zhner (ZISP) for the loan of specimens. Special thanks to Albino Luna and Ernesto Barrera for the preparation of the illustra- tions. LITERATURE CITED Bergroth, E. 1906. Neue Hemiptera aus Madagaskar. Wiener Entomologische Zeitung 25: 17-19. . 1912. Notes on Coreidae and Neididae. An- nales de la Société Entomologique de Belgique 56: 76-93. Distant, W. L. 1879. Contributions to a knowledge of the Hemipterous Fauna of Madagascar. Transac- tions of the Entomological Society of London 1879(2): 209-218. 1893. Notes on Two Genera of Coreidae found in Madagascar. Annals and Magazine of Natural History 6(11): 53—54. Garcia Varela, A. 1913. Contribuci6n al estudio de los Hemipteros de Africa. Notas sobre coreidos del Museo de Madrid. Trabajos del Museo Nacional de Ciencias Naturales. Serie Zoologica 12: 1—33. Stal, C. 1873. Enumeratio hemipterorum. 3. Kongliga Svenska Vetenskaps Akademiens Handlingar 9(1): 1-163. PROC. ENTOMOL. SOC. WASH. 104(1), 2002, pp. 119-125 A NEW SPECIES OF GLYPHIDOCERA WALSINGHAM (LEPIDOPTERA: GELECHIOIDEA: GLYPHIDOCERIDAE) FROM COSTA RICA DAVID ADAMSKI Department of Entomology, National Museum of Natural History, Smithsonian Insti- tution, Washington, DC 20560-0127 (e-mail: dadamski @sel.barc.usda.gov) Abstract.—Glyphidocera guaroa, n. sp., is described from Costa Rica. A photograph of the imago and illustrations of wing venation, male abdominal sex scales, and male and female genitalia are provided. Key Words: Limon Glyphidocera Walsingham (1892) are small to medium-sized moths that vary in color from pale yellowish brown to dark brown with few, if any, diagnostic mark- ings. Limited to the New World, their great- est diversity is in the tropics where the numbers of species of Glyphidocera are about ten times greater than that of the tem- perate region. Host associations are un- known for all but G. juniperella, which feeds on Juniperus horizontalis Moench (Cupressaceae) (Adamski and Brown 1987). Glyphidocera was first recognized by Walsingham (1892) who described G. au- dax from Saint Vincent Island in the West Indies. Originally placed the Gelechiidae (Walsingham 1892), Glyphidocera later was transferred (Hodges 1978), with sev- eral other genera, to the Symmocinae (Blas- tobasidae). Hodges later (1998) transferred Symmocinae to Autostichidae, except for Glyphidocera, which he elevated to family rank based on two apomorphies: 1) fore- wing with CuA, and CuA, stalked and downcurved from posterodistal angle of cell, and 2) forewing with Rs terminating on the outer margin. Becker (1999), follow- Lepidoptera, Gelechioidea, Costa Rica, Puntarenas, Heredia, Guanacaste, ing Hodges (1998), recognized two gele- chiid genera, Ptilostonychia Walsingham and Stibarenches Meyrick, as junior syno- nyms of Glyphidocera. Glyphidoceridae usually can be recog- nized by a combination of the following characters: male 4th flagellomere with tuft of setiform scales on dorsal surface extend- ing over concavity with sex scales on the lateral surface of 5th and 6th flagellomeres; male abdomen with squamiform or piliform sex scales on intersegmental membrane be- tween terga 2—3 and/or terga 3—4; valva narrowed basally, abruptly widened apical- ly, apex protracted, base of costa with dig- itate process; sacculus twisted apically; gnathos projecting dorsally from beneath tuba analis, juxta basally narrow and fused to vinculum, distolaterally expanded and forming a ventral support for the aedeagus; aedeagus with medium to large cornutus or cornuti; ductus ejaculatoris spiralled; fe- male ductus bursae large, sclerotized, with funnel-shaped antrum; ductus seminalis spi- ralled from middle or posterior end of cor- pus bursae and with internal sclerotized support; and corpus bursae spinulate, es- pecially at anterior end, and with a sclero- 120 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ¢ Fig. 1. Holotype of Glyphidocera guaroa. tized plate opposite base of ductus semin- alis. Kornerup and Wanschner (1978) is used as a color standard for the description of the adult vestiture. Genitalia were dissected as described by Clarke (1941), except that mercurochrome and chlorazol black were used as stains. Measurements of wings and genitalia were made using a calibrated oc- ular micrometer. The following species is described as part of several major studies planned by the author, including the Glyphidocera of Costa Rica. Glyphidocera guaroa Adamski, new species (Figs. 1-6) Diagnosis.—Glyphidocera guaroa can be distinguished from other species of Gly- phidocera by the following combination of characters: squamiform sex scales on the intersegmental membrane between abdom- inal terga 2—3 of the male, an elongate un- cus, a bulbous gnathos; a widely bifurcate apex of valva, an aedeagus with a large cor- nutus; female antrum with long longitudinal ridges, and ductus seminalis spiralled, ex- tremely wide basally. Glyphidocera guaroa is most similar to G. alexandrae Adamski and Brown (2001) but differs from the latter species by having a bifurcate costal apex of the valva. Description, adult——Head: Vertex and frontoclypeus pale brownish orange inter- mixed with brown marginal scales tipped with pale brownish orange; outer surface of labial palpus brown intermixed with few pale brownish-orange scales, inner surface pale brownish orange intermixed with few brown scales; scape, pedicel and flagellom- eres 1—6 brownish orange, flagellomeres brown distally; 4th flagellomere with a tuft of setiform scales on dorsal surface in male, extending over concavity on 5th and 6th flagellomeres; proboscis pale brownish or- ange. Thorax: Mesonotum and _ tegula pale VOLUME 104, NUMBER 1 brownish orange intermixed with few brown scales; legs pale brownish orange in- termixed with brown scales; some speci- mens with areas at midsegment and apical areas of all segments and tarsomeres pale brown; forewing (Figs. 1—2) length 5.9—7.5 mm (n = 121), brownish orange intermixed with pale brownish-orange scales tipped with brown and few brown scales; costa and outer margin brown intermixed with few pale-brown scales; discal cell with two spots, one near middle and one near distal end; one brown spot or streak from base of CuP to near midcell spot (spots may be faint in rubbed specimens); fringe scales brown; venation (Fig. 2) with R;, Ry, and R; approximate to distoanterior part of cell; M, and M, approximate to distoposterior part of cell; M, absent; CuA, and CuA, branched distad of cubitus beyond cell, ex- tending in a slight curve to margin; CuP slightly evident basally; undersurface pale brown except for pale yellowish-brown cos- ta; hindwing pale gray, with two acanthae in female; venation (Fig. 2) with Rs and M, stalked about % length beyond end of cell; M, closer to M, than to M,; M, branched with CuA, beyond cell, about %4 distance as Rs and M,; CuA, arising from cell about % length of cubitus; 1A and 2A separate ba- sally, forming an elliptical support. Abdomen (Fig. 3): Male with squami- form sex scales on intersegmental mem- brane between abdominal terga 2—3. Male genitalia (Figs. 4—5): Uncus nar- row and elongate with few subapical setae; gnathos enlarged distally and forming a bulbous projection with subapical setae on lateral surface; genital capsule slightly ta- pered from base; valva with apicoventral area beyond sacculus membranous and se- tose, apex widely bifurcate, base of costa with digitate projection with several apical setae; sacculus curved apically; vinculum narrow, bifurcating at base to nearly 3 times width at % length; juxta fused ventrally with vinculum, widening distolaterally forming a ventral support for aedeagus; ae- deagus widest near middle, slightly tapering 121 toward apex; ductus ejaculatoris with sev- eral spirales; vesica with many microspi- nules and a large cornutus. Female genitalia (Fig. 6): Papillae ana- les conical, with short and long setae inter- mixed; apophyses anteriores bifurcating into two thin posterior arms, one fusing with eighth tergum, and one fusing with eighth sternum; eighth sternum spinulate and setose; antrum funnel-shaped, spinula- te, and with posterior margin entire, ante- rior *% with longitudinal ridges above two internal flanges; inception of spiralled duc- tus seminalis near middle of corpus bursae; larger spirales at base, each gradually nar- rowing toward apical end; ductus seminalis with a spiralled internal support; internal support enlarged basally; accessory lobe opposite base of ductus seminalis, juxta- posed to large, elongate plate; corpus bur- sae sparsely spinulate, larger spinules near base of ductus seminalis. Holotype.— 03472067 = 05469037"; 0625780"; =0347168" 03469237; Id 1 Same except, Mar 1995, 1G: Varela”: 1409702”; “1409698”; “INBio Genitalia 122 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 2-5. Glyphidocera guaroa. 2, Wing venation, female; scale line = 1.0 mm. 3, Male abdominal sex scales; scale line = 1.0 mm. 4—5, Male genitalia; scale line = 0.5 mm. VOLUME 104, NUMBER I Fig. 6. Female genitalia of Glyphidocera guaroa. Scale line = 1.0 mm. 123 Slide, Sex’ 2, No. 4273 1 "6 "samc except, “R. Zuniga, Mar 1991”; “0648407”; 1 6, same except, “May 1991”; “1319436”; 1 6, Same wexcept, ~ Jul) 1992. "Gi Warelae: 0950374’; ““INBio Genitalia Slide, Sex, °, No. 426’’; 1 6, same except, “R. Zuniga, Feb 199%; —0347090.> 5) 6, same ex- cephiMeyAso: OOS” Gi rVarelae : rT LEOSOO13" = =aIG D602 3-7 os G0USe: + 1636012" EF W6ES6082 "> ING. samerex- cepty 1992” 2 09406147 td | Same exe cept, May 199, SREeZunieas - SIBI9A72 7 1S Oe Seliplt 19925 Gavan ela ar FOS I5605 =e) “G isame, excep MODS: SEN OO8 326 2a salle vexcepe, 1992). Ge Zuniga, 0935656) <7) 1% same except, ““G. Varela & R. Zuniga, 990" = 950223069" 23 GY same "except, ““Nov 1990’; ““0227948"’; 6 ¢, same ex- Cept, ) Dic 9502238703 02290104=5 ~O229072°% ~ 0229266. “02287 70% **0583981’’; ““INBio Genitalia Slide, Sex, 6, No. 466’; 8 6, “‘Est[acion] Magsasay, P[arque] N[acional] Braulio Carrillo, 200 m, Prov[incia] Here[dia], COSTA RICA, R. Aguilar, Nov 1990, L-N-264600, 36 LOOM Ce: 0226705 FOES 0226077) 34 0206735" ae O220738 826 HO22672 I: | 0226729745202 266093 2 AS31542 6 same except, Ra Aguilas Abr loor <0 2750995027 5070" 925G,, Sametexcept, “M. Zumbado, Oct 1990”; “0180818”; “0180807”; 2 3, same except, “A. Fernan- dex? 0179675" 7 O17 9638; > I iG-same except, Ri Aguilar Dic 19907; 0701559"; 1 3, same except, “May 1991, M. A. Zum- bado’’, “1300398”; 1 6, “‘Est[acion] Pitilla, 700 m 9 km §S Sta[tion] Cecilia, Prov[incia] Guan|[acaste], COSTA RICA, P. Rios & C. Moraga, Oct 1990, L-N-330200, 380200”; “0197960”; 1 3, same except, “C. Moraga, Jul 1991”; ‘0300184’; 1 3, same except, “Ene 1995, #43547.) 2931715" 21 oAsame except, ““P[arque] N[acional] Guanacaste, 700 my "1923 un 1998, -PiP Rios’: 1835123”; “INBio Genitalia Slide by D. Adamski” “Sex? 4G.) Nov) 479i 4.286; ‘‘Est[acion] Sirena, 0-100 m, P[arque] N[acional] Corcovado, Prov[incia] Punt- 124 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON [arenas], COSTA RICA, G. Fonseca, Se[p]t 1991, L-S-270500, 5083007; “03575717; “0357617”: | 0357448”: 3085750745 20c. same, except, 219905. s.Oli20797 “01791877;; “1. Sn same Mexcepr poole: 0643420’; 2 d, same except, “Nov 1990”’; “0183423”: “O1832517 ssl, sametexcept: “1093, #24907; “1625857 ;2.6.,same x= cepts “G:Fonseca; (Diew1992" =" 0783307": “0783813";. 1) d, “samenexcepiz 19907: 0297295": 1%, Sametexcept,. “Je Sabon, 19917? 505266357; (Ge sametexcept, 1nG: Fonseca, Feb 1992’’; “0621647”; 3 3, same except, ja Mar), 1992°eiipe -0780383:7¢ “0788594”; “0780480”; 2 3, same except, “Abr 19917 3-704 75866; 20475887 2489S “same except, May yli9Oilk-: 0587949. 0588 1147350587822” Vin © same except ~1992735"0914822>"32"6%, same except, Jun 1991”, “0646450”; ‘*0646447”; 1 d, same except. #19927)? S0708208** 4° 671 jcSector Cerro Cocori, F[inJca de E. Rojas, 150 m, Prov[incia] Limon, Costa Rica, E. Rojas, 31 Ene—21 Feb 1992, L-N-286000, 567500”; 07853187 107 750964 0775089 4.7 IN- Bio Genitalia Slide by D. Adamski, Sex, d, No. 471” [yellow label]; “0785137”; “‘IN- Bio Genitalia Slide by D. Adamski, Sex, 6, No.47/07 226, samenrexcepiye 19937: “0402859”; 2 6, same except, “Feb 1993’; “0998918”; “0998790”; “INBio Genitalia Slide by D. Adamski, Sex 6, No. 425”; 5 6, Same except... Mar 19927: 0363635: 7; B0S03719" 2” 650363754. 29 408635934: =03036297: ling, samemexcepty lOOLy; “0181437”; 6 3, same except, “Abr 1992”; “OTSS94472" 07855 ING O7 S55 18r8 “0786302”; “07855017; “INBio Genitalia Slide by D. Adamski, Sex 6, No. 482”; 0786366"; “‘INBio Wing Slide by D. Adamski, Sex 6, No. 429”; 1 3, same ex- cept, saMay, 1992”: “03735607: 2° Gssame except, “26 Jun a 16 Jul 1992”; “0703169”; 0745136”; 1 6, same except, ‘1993’’; “1699131”; “INBio Genitalia Slide by D. Adamski, Sex ¢, No. 472”; 1 3, same ex- cept, |. “Sefipiti 1991"; “05989382 hick Same, except “selp}t 1993°" 411413703": lemare:.| wtSamMem except, “Oct, 1992722 0819686"; 6 <6, Same except, “Nav POON. “0501430%: **0501449”’; 0460128’; “‘04601517; “‘0460169”’; ‘0460144’; 2 3, same except, “9 a 30 Noy 1992”:) 709314 189" 093 14654 “INBio Wing Slide by D. Adamski, Sex 6; No. 4913-2 df same except, Wie L991; “03612692. 0361205 Sexier same except, ““30 Km N de Cariari, A.C. Tortuguero, Ene-19947";.” 185657 Is @ine hundred and five paratypes deposited in INBio and I5 paratypes deposited in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA. Distribution.—Glyphidocera guaroa is widespread in Costa Rica, ranging from the coastal Pacific to the coastal Caribbean, and from the western province of Guanacaste east to the southeastern province of Puntar- enas near Panama. It is likely that this spe- cies occurs elsewhere in Central America. Etymology.—Glyphidocera guaroad is named for a Costa Rican liquor, giiaro, that is made from sugar cane. ACKNOWLEDGMENTS I thank Eugenie Phillips Rodriguez of The National Biodiversity Inventory Divi- sion, Instituto Nacional de Biodiversidad, Santo Domingo, Heredia, Costa Rica for the loan of specimens; Kuniko Arakawa for Figs. 2—6; and Carl Hansen of the Office of Imaging, Printing and Photographic Servic- es, Smithsonian Institution, Washington, DC, USA, for the photograph of the holo- type of Glyphidocera guaroa. LITERATURE CITED Adamski, D. and R. L. Brown. 1987. A New Nearctic Glyphidocera with descriptions of all stages (Lep- idoptera: Blastobasidae: Symmocinae). Proceed- ings of the Entomological Society of Washington 89(2): 329-343. . 2001. The Glyphidocera of Cerro de la Neb- lina and adjacent areas in Amazonas (Lepidoptera: Gelechioidea: Glyphidoceridae). Proceedings of the Entomological Society of Washington 103: 968-998. Becker, Vitor O. 1999. Family reassignments and syn- onymy of some taxa of Neotropical Microlepidop- VOLUME 104, NUMBER 1 tera. Revista Brasileira de Zoologica 16(Suppl. 2): 141-170. Clarke, J. EF G. 1941. The preparation of slides of the genitalia of Lepidoptera. Bulletin of the Brooklyn Entomological Society 36: 149-161. Hodges, R. W. 1978. Gelechioidea: Cosmopterigidae. In Dominick, R. B. et al. The Moths of America north of Mexico, fasc. 6.1 + X + 166 pp., text figs. 1-53, pl. 1-6. . 1998. Gelechioidea, pp. 131—158. /n Kristen- 1725) sen, N. P. ed. Lepidoptera: Moths and Butterflies. Handbook of Zoology/Handbuch der Zoologie IV, Part 35. Berlin & New York. Kornerup, A. and J. H. Wanschner. 1978. Methuen Handbook of Colour. 2nd ed. Methuen and Co., Ltd., London, 243 pp. Walsingham, Lord (Thomas de Grey). 1892. On the Micro-Lepidoptera of the West Indies. Proceed- ings of the Zoological Society of London 1892: 492-549. PROC. ENTOMOL. SOC. WASH. 104(1), 2002, pp. 126-137 PHE VEOPERLA CLYMENE (NEWMAN) COMPLEX (PLECOPTERA: PERLIDAE) IN ILLINOIS, NEW STATE RECORDS, DISTRIBUTIONS, AND AN IDENTIFICATION KEY R. EDWARD DEWALT, DONALD W. WEBB, AND Amy M. SOLI (RED, DWW) Illinois Natural History Survey, Center for Biodiversity, 607 E. Peabody Drive, Champaign, IL 61820, U.S.A. (e-mail: edewalt@inhs.uiuc.edu); (AMS) University of Illinois, Natural Resources and Environmental Sciences, Champaign, IL 61820, U.S.A. Abstract.—Stoneflies of the Neoperla clymene (Newman) species complex in Illinois were reexamined from museum and newly collected adult material. Seven species have historically been present in the state. The previously known species, N. clymene, is joined by six new state records including N. catharae Stark and Baumann, N. harpi Ernst and Stewart, N. mainensis Banks, N. occipitalis (Pictet), N. robisoni Poulton and Stewart, and N. stewarti Stark and Baumann. Four species (N. mainensis, N. occipitalis, N. robisoni, and N. stewarti) are considered extirpated since all records predate 1930. Two species (N. catharae and N. clymene) have had extensive range reductions, while N. harpi has main- tained its historical distribution in the state. Large rivers at the periphery of Illinois and small streams in the Shawnee Hills of southern [linois still maintain populations of Neo- perla. Key Words: Neoperla in the Nearctic Region were viewed until recently as a single, wide- ranging and variable species, Neoperla cly- mene (Newman) (Stark 1990). Stark and Baumann (1978), Stark and Lentz (1988), and Ernst et al. (1986) split this complex into two species groups, clymene and choc- taw, based on adult genitalic and egg struc- tures, with 12 recognized species. Subse- quent researchers (Ernst et al. 1986, Stark 1995, Smith and Stark 1998) have de- scribed three new species to bring the cur- rent North American total to 15. Neoperla reside in relatively clean streams throughout the eastern half of North America (Stewart and Stark 1988). Vaught and Stewart (1974) found that WN. clymene exhibited a univoltine-slow (no egg diapause) life cycle, while Ernst and Stewart (1985) recorded a late emergence Plecoptera, Neoperla, Illinois, identification key, extirpation for Neoperla spp. compared to other cohab- itating perlids and a preference for trees as adult habitat. These life history attributes leave them susceptible to changes in local stream health including increases in sum- mer water temperature, reduction in dis- solved oxygen, increased imbeddedness of mineral substrates, and removal of natural riparian vegetation. The Illinois Natural History Survey (INHS) established natural history collec- tions late in the nineteenth century as a re- pository to document the fauna and flora of Illinois, the Midwest, and across the conti- nent. The age of these collections makes them irreplaceable as documentation of the natural communities found before major degradation of the native landform took place. This is fortunate since extinctions, local and regional extirpations, and shrink- VOLUME 104, NUMBER 1 127 Table 1. Spatial accuracy codes used for Neoperla localities in [linois (Figs. 1-4). Spatial Accuracy Code Location Description from Labels 1 Known exact location 2 Small municipality (<50,000 population) or park, point usually placed at cen- ter of town or at waterfront if stream name included on label. 3 Land survey data (principal meridian, township, range, section) or large munic- ipality (>50,000), taken as center of municipality or section. 4 County, plotted as center of county. age of historical range are common re- sponses of stonefly species across the con- tinent. Frison (1935) investigated the Illinois stonefly fauna for over two decades early in the twentieth century. Many sites were repeatedly visited and their stonefly assem- blage was well documented. Among these collections are sufficient numbers of Neo- perla, then identified exclusively as N. cly- mene, to accurately document the original species assemblage of this environmentally sensitive genus. Reevaluation of this mate- rial has recently been completed and the re- sults presented herein. The objective of this study is to document the species of Neo- perla present in Illinois. We hope, through providing a fully illustrated key to adults and eggs of regional species, that others lo- cate additional populations of this environ- mentally sensitive genus. This study is part of a larger effort to update Frison’s (1935) The Stoneflies, or Plecoptera, of Illinois over 60 years after its publication. METHODS Illinois Neoperla in the INHS insect col- lection were reevaluated using current lit- erature. The task of tracking information about each specimen was simplified by a recently completed Plecoptera database that links species names, label data, and location information with a catalogue number. This database is now available in simplified form on the Internet (URL: www.inhs.uiuc.edu/ cbd/EPT/index.html). Poulton and Stewart (1991) reared and associated nymphs of six of seven Interior Highlands species. However, there are still eight species whose nymphs are either un- described or inadequately known. Until fur- ther associations have been completed, the adults are necessary to definitively identify species in the complex. The tergal process, paraproct, and internal aedeagus are useful for identification of males. Relative length and degree of sclerotization of the sper- mathecum and the ornamentation of the egg chorion are important female characteris- tics. Adult internal structures were viewed by clipping the abdomen from the body at about the fourth segment, then placing it in boiling 10% KOH until clear. Eggs were extracted before boiling and viewed using a compound microscope. In addition to the historical material, new specimens were ob- tained by rearing of nymphs, ultra-violet light trapping, and by sweeping streamside vegetation from many locations across the state. Latitude and longitude coordinates for each unique location were recorded from a digital DeLorme® Street Atlas USA. Many records had vague localities, the exact lo- cations of which were impossible to deter- mine. So as not to lose any location data, locations were coded one through four for accuracy (Table 1). This accuracy was de- noted on range maps as different sized sym- bols, usually with closed points and open circles of increasing size. Collection loca- tions were stored in electronic format using FileMakerPro® software. Mapping of these locations was produced using ArcView® software. 128 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON i litionally, location information has been provided in the text of works such as these. We have opted to rely on the Plecop- tera database web site for dissemination of this detailed and highly repetitive informa- tion. Those who do not have access to the Internet may request the information from the senior author in other formats. RESULTS Adult Neoperla accounted for 92 of the current 4,880 Illinois and 18,885 total Ple- coptera records at the INHS (Table 2). A total of 411 adult specimens was reevalu- ated that represented 32 unique historical and contemporary locations (Fig. 1). Seven of the 15 species in the genus are now known to occur in Illinois (Table 2). Six belong to the clymene species group, while one belongs to the choctaw group. Six of these species are new state records and some represent sizable range extensions. The greatest concentration of both species and specimens were from the state’s south- ern tip where stream gradients were high and overall stream health best. Despite the large range in latitude, 37.00° to 42.16°N, the majority of records indicated a mid- June through late July emergence for all species encountered. [Illinois distributions, brief notes on general ecological require- ments, and additional information on taxo- nomic characters are provided for each spe- cies. Neoperla catharae Stark and Baumann (Biss. S255 al2) This species provided 7.6% of Illinois Neoperla records and 2.7% of all specimens (Table 2). This is the only member of the N. choctaw Stark and Baumann species group to be taken in Illinois. Its presence is a new state record. Historically, specimens were taken as far north as Champaign County, in the headwaters of the Embarras and Vermilion rivers (Fig. 3). The exact collection location has been coded with an accuracy code of two, which straddled streams in both drainages. All populations in the central Illinois Grand Prairie natural division (Schwegman 1973) appear to have been extirpated since no specimens have been taken since 1889. Extant populations survive only in high quality streams in the Lusk Creek drainage of southern Illinois. The Shawnee National Forest buffers this area from agriculture and other development; therefore, the species is afforded some measure of protection against water quality degradation. Other populations may eventually be found fur- ther west in similar uplands of the Ozark Hills natural division along the Mississippi River. Its current range includes Alabama, Arkansas, Kentucky, Missouri, Ohio, Oklahoma, Pennsylvania, Tennessee, and Virginia (www.mce.edu/~stark/stonefly. html). No detailed life history information is available for this species (Stewart and Stark 1988). Ernst and Stewart (1985) found that it emerged later than its conge- ners, August through September, with males and females having negatively skewed and extended emergence patterns, respectively. Neoperla clymene (Newman) (Figs. 2.1.0; 1:3) Harris and Webb (1995) listed this spe- cies from Illinois. Neoperla clymene pro- vided 35.9% of the records and 14.4% of the specimens and was the most widely dis- tributed Neoperla in Illinois (Table 2, Fig. 2). It was most abundant in larger rivers such as the Wabash, Ohio, and Mississippi rivers, Where it was recently common in light trap samples from a reach south of Rock Island. Several specimens were recently taken from the Vermilion River near the Indiana border. These represent the only contemporary spec- imens from small-to-medium sized _ rivers throughout middle Illinois (DeWalt et al. 199): Historically, the species was abundant in the Embarras, Kaskaskia, and Vermilion rivers (Fig. 3) of the Grand Prairie and Southern Till Plain of central Ilinois. Stark (1990) reported it from throughout the 129 VOLUME 104, NUMBER 1 Illino is. 1, All Neoperla locations in Figs. 1-4. Neoperla locations in N. harpi, N. robison rtt. ccipitalis, N. stewa i, 4, N. mainer 130 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 2. Neo INHS insect verla species found, number of adult records in Plecoptera database, number of individuals in ollection, and the number of unique sites across Illinois from which Neoperla species were taken. Unique Species Records Individuals Sites Neoperla catharae Stark and Baumann 1978 if 11 4 Neoperla clymene (Newman) 1839 33 59 20 Neoperla harpi Ernst and Stewart 1986 18 235 Wl Neoperla mainensis Banks 1948 2 2 ! Neoperla occipitalis (Pictet) 1841 l l | Neoperla robisoni Poulton and Stewart 1986 3 18 3 Neoperla stewarti Stark and Baumann 1978 28 85 4 Total 92 411 3y southeastern United States and indicated that some older eastern and midwestern lit- erature records should be viewed with doubt until specimens have been reexam- ined. He implied that they might have been confused with a newly named species, N. osage Stark and Lentz. Vaught and Stewart (1974) recorded an univoltine-slow life cy- cle for N. clymene in the Brazos River of west central Texas. Ernst and Stewart (1985) reported emergence characteristics for this species, but Stark and Lentz (1988) showed their studied population to be an undescribed species, N. falayah Stark and Lentz. Neoperla harpi Ernst and Stewart (Figs. 3, 7, 14) This species is restricted to the southern- most tip of Illinois, in the Shawnee and Ozark Hills natural divisions (Fig. 3). Its presence represents a new state record. De- spite representing only 19.6% of the rec- ords, it provided a disproportionate share, 57.2%, of the specimens examined. A ro- bust population exists in the Hutchins Creek drainage near Wolf Lake, Illinois. Part of this drainage is protected within the La Rue—Pine Hills Ecological Area, a Na- tional Natural Landmark. Historical records also indicate its presence in the Shawnee Hills, farther to the east. Poulton and Stewart (1991) considered this species an Ozark endemic. Ross (1944) spoke of the Ozark and Shawnee Hills nat- ural divisions as extensions of the Missouri Ozarks, and therefore, it is not surprising to find this species in southern I[]linois. Stark (1990) listed Arkansas, Missouri, and Oklahoma as locations for this species. Poulton and Stewart (1991) collected adults from late May through late August, but no additional life history data are available. Neoperla mainensis Banks (Figs. 4, 8, 15) The species is represented by two records consisting of two females (Table 2) from Frison’s (1935) famous Oakwood, Illinois location on the Salt Fork of the Vermilion River (Fig. 4). Neoperla mainensis is the only known member of the clymene group that has a notched, triangular subgenital plate, thus, we feel confident in our deter- mination. Its presence in Illinois represents a new state record and a significant range extension from the known localities of Ohio, Maine, and Ontario (Stark 1990). No specimens have been collected since 1927; therefore, we feel that it has been extirpated from Illinois. No life history data are avail- able for this species (Stewart and Stark 1988). Neoperla occipitalis (Pictet) (Figs. 4, 9, 16) This species was taken as a single female (Table 2) from the Oakwood location (Fig. 4). Its presence represents a new state rec- ord. It has not been collected since 1925; therefore, it appears to have been extirpated from [linois. It is known from much of the VOLUME 104, NUMBER 1 131 ventral Fig. 5-8. Male structures of Neoperla spp. A = abdominal terga 7-10; B = aedeagus, lateral. 5, N. catharae. 6, N. clymene. 7, N. harpi. 8, N. mainensis. Abbreviations: b = bulb, t = tube, s = sac. Figures 5A and 7A modified from Poulton and Stewart (1991); Figures 6A and 8A modified from Stark and Baumann (1978). 132 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 11 A Fig. 9-11. Male structures of Neoperla spp. A = abdominal terga 7-10; B = aedeagus, lateral. 9, N. occipitalis. 10, N. robisoni. 11, N. stewarti. Figure 10A modified from Poulton and Stewart (1991); Figures 9A and 11A modified from Stark and Baumann (1978). VOLUME 104, NUMBER 1 Great Lakes region, where it is apparently common, and from as far south as northern Alabama and Mississippi (Stewart and Stark 1988). No life history information is available for this species (Stewart and Stark 1988). Neoperla robisoni Poulton and Stewart (Figs. 3; TOF) This species represented 3.3% of records and 4.4% of Illinois Neoperla specimens (Table 2). Its presence constitutes a new state record, but since it has not been col- lected since 1939, it is probably extirpated from the state. Historically, this species re- sided in the Vermilion and Wabash river ba- sins of east central [Illinois (Fig. 3). Its pub- lished distribution includes Arkansas, Mis- sissippi, Missouri, Oklahoma, and Tennes- see (Stark 1990). Stewart et al. (1974) listed N. clymene from New Mexico, which was subsequently identified by Stark (1990) as N. robisoni. Stark (1990) has raised doubt as to the validity of the label data associated with any Neoperla specimens found in New Mexico. Poulton and Stewart (1991) and Ernst et al. (1986) found that N. robisoni primarily inhabited Gulf Coastal Plains streams, emerging in late May and through June. No detailed life history data are avail- able for this species (Stewart and Stark 1988). Neoperla stewarti Stark and Baumann Gigs: 4.11 ahs) This species represented 30.4% of rec- ords and 20.7% of the individuals (Table 2). It is also a new state record. It has not been collected in Illinois since 1930, and is feared extirpated from the state. Historical- ly, it was locally abundant in the Vermilion River drainage of eastern Illinois (Fig. 4). Previous records are for Alabama, Ken- tucky, Maine, Minnesota, Mississippi, Ohio, Pennsylvania, Tennessee, Virginia, and Wisconsin (Stark 1990). Difficulty in identification may arise with some females due to the presence of a par- asitoid wasp, the pupa of which usually de- 133 velops in the spermathecum. Distortion of this organ gives the mistaken appearance of a choctaw complex species by darkening and expanding the very tip of the sperma- thecum. Parasitism rates ranging from 10 to 50% were observed. The pupa was large enough to completely block the sperma- thecum and questions the efficacy of mating for afflicted specimens. No life history data are available for this species (Stewart and Stark 1988). DISCUSSION The number of stonefly species known from Illinois has changed considerably since Harris and Webb (1995) published an updated nomenclature and listed 63 species for the state. DeWalt and Stark described Perlesta golconda (in DeWalt et al. 1998) and DeWalt et al. (2001) added six state record Perlesta species for a new total of 70. The addition of the new Neoperla spe- cies brings the total to 76 species. The perlid stonefly genus Perlesta was found to contain at least eight species in Illinois (DeWalt et al. 2001). These species appear to be relatively unaffected by the large-scale anthropogenic disturbance of the Illinois landscape. In fact, one species has moved into Illinois over the past three decades. A rather different scenario has un- folded for Neoperla species. It appears that four of the seven species now known from the state, have been extirpated (VN. mainen- sis, N. occipitalis, N. robisoni, and N. ste- warti). Additionally, two have experienced apparent range reductions (N. clymene and N. catharae), while only one (N. harpi) ap- pears to have maintained its historical range. This differential response to changes in the Illinois landscape is intriguing. Neoper- la typically reside in relatively clean streams in I]linois and in the Interior High- lands (Poulton and Stewart 1991). Hilsen- hoff (1987) and Lentz (1993) have provided pollution tolerance values that rank Neo- perla as highly sensitive to environmental degradation. Neoperla clymene exhibits an 134 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 7 : | ieee Fig. 12-15. Female and egg structures of Neoperla spp. A = abdominal sternite 8 and subgenital plate; B = spermathecum; C = egg: D = detail of egg chorion. 12, N. catharae. 13, N. clymene. 14, N. harpi. 15, N. mainensis. Abbreviations: sc = sclerite. sp = spermathecum, str = striae, su = sulci, v = vagina. Figures 12A, 12B, 14A, and 14B modified from Poulton and Stewart (1991): 13A-D from Stark (1990); 12C and 14C from Ernst et al. (1986); and 15A and ISC from Stark and Baumann (1978). VOLUME 104, NUMBER 1 univoltine-slow life cycle, eggs hatch soon after being laid, and nymphs grow through- out the majority of the year (Vaught and Stewart 1974). In that study, no egg dia- pause was present, unlike what has been shown for Perlesta decipiens (Walsh) (Snellen and Stewart 1979). Diapause pre- sumably permits this species to carry the generation through poor environmental conditions such as high summer water tem- peratures, low dissolved oxygen, or inter- mittent flow. The presumed lack of dia- pause in the rest of Neoperla species, and any other stonefly with a similar life his- tory, would make them highly susceptible to changes in local land use including stream channelization and removal of ripar- ian trees. This very scenario has taken place in streams across the rich fertile prairies of Illinois, where most of the species’ loss and range reductions have taken place for Neo- perla. It is doubtful that the extirpated spe- cies will again live in Illinois streams with- in the near future. The *islands’ of suitable habitat from which recolonization could oc- cur, even if the conditions of streams in the state were to improve dramatically, are so distant that the species may not bridge the gap. The kind of stream habitat fragmen- tation that exists in Illinois, and elsewhere in the Midwest, puts the healthy islands at risk of high extinction rates and limits rates of emigration (Zwick 1992). Thankfully, some protection is afforded to those re- maining populations of Neoperla in south- ern Illinois, since much of the land is in the Shawnee National Forest. To facilitate identification of Neoperla in Illinois and neighboring states, a key to adults and eggs of the seven known species is provided. This key is modified from Poulton and Stewart (1991) and Stark and Lentz (1988). KEY TO ADULTS AND EGGS OF ILLINOIS NEOPERLA 1. Male aedeagal tube with external spines (Fig. 5B): anterior margin of female sternum 8 en- i) ie . Subgenital tire (Fig. 12A); female spermathecum short, not coiled (Fig. 12B); egg chorion smooth (Fig. 12C) N. catharae Aedeagal tube without external spines (Fig. 6B): female sternum 8 with rectangular, tri- angular, or sub-circular indentation along an- terior margin (Fig. 13A); female spermathe- cum long, atleast partially coiled (Fig. 13B): egg striate or punctate (Fig. 13C, 14C):; Neo- pera clymene STtoup’ 2. seo ee 2 Tergum 10 with anteriorly directed genital hooks (Fig. 6A), males Tergum 10 without genital hooks, females.. 8 Aedeagal tube abruptly bent ventrad near modlensthy (Eis ssi es eee enn 4 Aedeagal tube straight or slightly sinuate without abrupt bend at midlength (Fig. 6B) SP SE: i Sane ne nated OR oer ni 5) Aedeagal tube apex abruptly constricted (Fig. DIB) etre lobe sat Ee Susas tees ene betes ue N. harpi Aedeagal tube apex as wide as or wider than subapical area (Fis, 10B)) S32 5.) - N. robisoni Aedeagal tube short and plump, length about 3X width at bulb (Fig. 11B) N. stewarti Aedeagal tube slender, length about 4—6x width at bulb (Fig. 6B) Aedeagal tube apex with prominent knob (Fig. 8B) Aedeagal tube apex without knob (Fig. 6B) N. mainensis Dorsal midlength of tube with prominent patch of spicules, sac armed with very small spines (Fig. 9B) Dorsal midlength of tube without prominent patch of spicules, sac with large spines (Fig. 6B) N. clymene Posteromesal margin of sternum 8 recessed, N. occipitalis not projecting beyond posterolateral margins (Fig. 13A) Posteromesal margin of sternum 8 projecting Sheet eedeen eon Veen seein N. clymene beyond posterolateral margins (Fig. l14A) .. 9 Spermathecal stalk lining forming an irregu- lar, jagged inner margin (Fig. 14B); egg non- striate, but coarsely punctate (Fig. 14C) sans. Gy sists, aeeuTe Suet cee oa er nie EO ele N. harpi Spermathecal stalk lining not forming jagged margin (Figs. 17B, 18B); egg striate (Figs. 1SC-18C) plate triangular and notched (Bigs A) Subgenital plate a small truncate tab or oth- apically awoke & N. mainensis erwise slightly convex surface, not notched at apexe (Bigs: HGASSIS AN ae sneer 11 Setal lining of spermathecum restricted to a small apical patch (Fig. 18B) N. stewarti Setal lining not restricted to apical patch, usu- ally extends >% length of spermathecum (Fics¥Vil6Byel7B) indict Ae ee 12 136 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON chy ' ‘ Fig. 16-18. Female and egg structures of Neoperla spp. A = abdominal sternite 8 and subgenital plate; B = spermathecum; C = egg; D = detail of egg chorion. 16, N. occipitalis. 17, N. robisoni. 18, N. stewarti. Figures 17A and 17B modified from Poulton and Stewart (1991); 17C and 17D from Ernst et al. (1986): and 16A—D and 18A-C from Stark and Baumann (1978). 12. Spermathecal stalk lining tapered to a round- ACKNOWLEDGMENTS ed apex and without darkened evn eune ali) . We thank B. P. Stark, Mississippi Col- mdges (Big. VOB)N-.. Aine ene N. occipitalis fe wee dardized illwet — Spermathecal stalk lining truncate apically lege, for providing standar ora we ae and with 2—3 folds forming dark longitudinal tions of Neoperla males and for confirming maeesn(Rig ali) ay-tns och hed. ie N. robisoni the identities of some specimens. C. E. VOLUME 104, NUMBER 1 Warwick, of the INHS publication office, provided technical assistance with scanning of other Neoperla illustrations. M. Water- straat provided original illustrations of an N. mainensis spermathecum and of egg chorionic detail for N. stewarti. This work was partially funded by the INHS Critical Trends Assessment Program, by a State- wide Biosurvey contract from the Hlinois Department of Transportation, and by a Na- tional Science Foundation collection com- puterization grant numbered DBI-9876756. LITERATURE CITED DeWalt, R. E., B. P. Stark, and M. A. Harris. 1998. Perlesta golconda (Plecoptera: Perlidae), a new stonefly species from Illinois. Entomological News 109: 315-317. DeWalt, R. E., D. W. Webb, and M. A. Harris. 1999. Summer Ephemeroptera, Plecoptera, and Trichop- tera (EPT) Species Richness and Community Structure in the lower IIlinois River basin of Illi- nois. The Great Lakes Entomologist 32: 115-132. DeWalt, R. E., D. W. Webb, and T. N. Kompare. 2001. The Perlesta placida (Hagen) complex (Plecop- tera: Perlidae) in Hlinois, new state records, dis- tributions, and an identification key. Proceedings of the Entomological Society of Washington 103: 207-216. Ernst, M. R. and K. W. Stewart. 1985. Emergence pat- terns and an assessment of collection methods for adult stoneflies (Plecoptera) in an Ozark foothills stream. Canadian Journal of Zoology 63: 2962-— 2968. 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. Annals of the Entomo- logical Society of America 79: 645—661. Frison, T. H. 1935. The stoneflies, or Plecoptera, of Illinois. Bulletin of the Illinois Natural History Survey 20: 1—471. Harris, M. A. and D. W. Webb. 1995. The stoneflies (Plecoptera) of Illinois revisited. Journal of the Kansas Entomological Society 67: 340-346. Hilsenhoff, W. L. 1987. An improved biotic index of organic stream pollution. Great Lakes Entomolo- gist 20: 31—39. Lenat, D. R. 1993. A biotic index for the southeastern United States: deviation and list of tolerance val- ues, with criteria for assigning water-quality rat- 137 ings. Journal North American Benthological So- ciety 12: 279-290. Poulton, B. C. and K. W. Stewart. 1991. The stoneflies of the Ozark and Ouachita Mountains (Plecop- tera). Memoirs of the American Entomological Society 38. 116 pp. Ross, H. H. 1944. The caddis flies, or Trichoptera, of Illinois. Hlinois Natural History Survey Bulletin 23(1): 1-236. Schwegman, J. E. 1973. Comprehensive plan for the Illinois Nature Preserves System. Part 2. The Nat- ural Divisions Of Illinois. Illinois Nature Pre- serves Commission, Springfield, [linois. 32 pp. Smith, A. D., and B. P. Stark. 1998. Neoperla coosa (Plecoptera: Perlidae), a new stonefly species from Alabama. Entomological News 109: 153-158. Snellen, R. K. and K. W. Stewart. 1979. The life cycle and drumming behavior of Zealeuctra claasseni (Frison) and Zealeuctra hitei Ricker and Ross (Plecoptera: Leuctridae) in Texas, USA. Aquatic Insects 1: 65—89. Stark, B. P. 1990. Neoperla clymene revisited: system- atics of the Nearctic species complexes (Plecop- tera: Perlidae), pp. 299-310. In Campbell, I. C., ed., Mayflies and Stoneflies: Life Histories and Biology. Proceedings of the Fifth International Ephemeroptera Conference and the 9th Interna- tional Plecoptera Conference. Kluwer Academic Publishers, Dordrecht, The Netherlands. 366 pp. . 1995. A new species of Neoperla (Insecta: Plecoptera: Perlidae) for Mississippi. Proceedings of the Biological Society of Washington 108: 45— 49. Stark, B. P. and D. L. Lentz. 1988. New species of Nearctic Neoperla (Plecoptera: Perlidae). Annals of the Entomological Society of America 81: 371— 376. Stark, B. P. and R. W. Baumann. 1978. New species of Nearctic Neoperla (Plecoptera: Perlidae), with notes on the genus. Great Basin Naturalist 38: 97— 114. Stewart, K. W., Baumann, R. W., and B. P. Stark. 1974. The distribution and past dispersal of southwest- ern United States Plecoptera. Transactions of the American Entomological Society 99: 507-546. Stewart, K. W. and B. P. Stark. 1988. Nymphs of North American Stonefly Genera (Plecoptera). The Thomas Say Foundation, Entomological Society of America 12: 1—460. Vaught, G. L. and K. W. Stewart. 1974. The life his- tory and ecology of the stonefly Neoperla clymene (Newman) (Plecoptera: Perlidae). Annals of the Entomological Society of America 67: 167—178. Zwick, P. 1992. Stream habitat fragmentation—a theat to biodiversity. Biodiversity and Conservation 1: 80-97. PROC. ENTOMOL. SOC. WASH. 104(1), 2002, pp. 138-141 NEW DISTRIBUTIONAL AND HOST-PLANT RECORDS FOR THE HETEROPTERANS AUFEIUS IMPRESSICOLLIS STAL (RHOPALIDAE) AND PRIONOSOMA PODOPIOIDES UHLER (PENTATOMIDAE) IN THE SOUTHEASTERN UNITED STATES, WITH NOTES ON THEIR WESTERN UNITED STATES HOSTS A. G. WHEELER, JR. Department of Entomology, Clemson University, Clemson, SC 29634-0365, U.S.A. (e-mail: awhlr@clemson.edu) Abstract.—The rhopalid Aufeius impressicollis Stal and pentatomid Prionosoma po- dopioides Uhler, native to western North America, were not known east of the Mississippi River until the early twentieth century and were first reported from the Southeast (South Carolina) in 1988. New state records for A. impressicollis are Georgia and North Carolina, and North Carolina is a new state record for P. podopioides; additional South Carolina records are given for both species. Most collections of A. impressicollis in the Southeast were from Amaranthus palmeri S.Wats.; host plants in California were other species of Amaranthaceae plus Chenopodiaceae. Southeastern collections of P. podopioides were from crowns of bunchgrasses growing near its main hosts, Plantago aristata Michx. and P. wrightiana Dene. Adults in Oklahoma and Texas were taken on P. patagonica Jacq. and P. rhodosperma Dcne.; Oklahoma and Texas also are new state records. Key Words: The rhopalid Aufeius impressicollis Stal and pentatomid Prionosoma podopioides Uhler are litthe-known, mainly western North American heteropterans. Eastern U.S. records for both species are scant. Ohio, the first eastern record for A. impressicollis (Os- born and Drake 1915), remained the eastern- most record until it was reported from Mary- land (Wheeler 1984). The first southeastern record was South Carolina (Wheeler 1988). Blatchley’s (1926) manual of eastern U.S. Heteroptera did not include P. podopioides; at the time, its easternmost records—Iowa and Missouri (Stoner 1916, 1920)—were west of the area treated by Blatchley (east of 90th meridian). Previously published re- cords of P. podopioides east of the Missis- sippi River now include Michigan (McPher- son 1970), Illinois (McPherson and Cuda 1975), and South Carolina (Wheeler 1988). insect distribution, Rhopalidae, Pentatomidae, host-plant relationships Here I provide additional southeastern re- cords of A. impressicollis and P. podopioi- des and, based on observations in eastern and western states, notes on their habitats and host plants. Adults of both species were collected at all sites; sites at which nymphs of A. impressicollis were observed with adults are indicated by asterisks. Voucher specimens have been deposited in the Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. Aufeius impressicollis Stal 1870 New southeastern U.S. records—GEOR- GIA: *Jefferson Co., Rt. 1, 2.8’ km-NE of Wrens (33°13.5'N, 82°22.0'W), 15 Oct. 2000; McDuffie Co., Rt. 78, 2.7 km W of Harlem (32°25.2’N, 82°20:5°W)s 15) Oct.72Z008; *Richmond Co., Rt. 1, 3.8 km NNE of Bly- the (33°19.4'’N, 82°11.1'W), 15 Oct. 2000. VOLUME 104, NUMBER 1 NORTH CAROLINA: Montgomery Co., Rt. P20 or kimres: Voie Emerys) (35: 12sMN, 79°43.6'W), 16 May 1999. SOUTH CARO- LINA: *Aiken Co., Rt. 25, 0.1 km S of Edge- field Co. line, 9 km N of North Augusta (33°35.2'N, 81°56.5'W), 15 Oct. 2000; Cal- houn Co., Rt. 6, 16 km NW of St. Matthews (33°41.8'N, 80°56.4'W), 21 Oct. 2000; Dar- lington Co., Rt. 34, 4.7 km W of Darlington (34°18.0'N, 79°55.4’W), 4 Nov. 2000; Dillon Co., Rt. 917, 3.6 km W of Latta (34°20.3’N, 79°28.2'W), 4 Nov. 2000; Edgefield Co., CR 407 nr jct. CR 37, 8.5 km SW of Eureka (33°39.5'N, 81°51.6’W), 28 Nov. 1998; Rt. 95,/13.9 ikniSSW ot Trenton’''(G3"42.7'N, 81°51.3'W), 15 Oct. 2000; Greenville Co., SR 50, 0.3 km N of Rt. 418, 4.5 km NW of Fork Shoals (34°38.4'N, 82°21.1'W), 29 Oct. 2000; Kershaw Co., Rt. 34, 5.5 km E of Camden (34°14.3’N, 80°32.9'W), 4 Nov. 2000; Lee Co., jct. Rt. 34 & CR 39, 0.5 km W of Una (34°16.1'N, 80°08.2’W), 4 Nov. 2000; Rt. 15, 2 km SW of Bishopville (34°12.4'N, 80°15.9'W), 4 Nov. 2000; Lex- ington Co., Rt. 302, 2 km NNE of Pelion (33°46.8'N, 81°14.3"W), 21 Oct. 2000; Mar- ion, Co,. Kt. 76, Marion (64°10:7'N, 79°22.6'W), 2 Nov. 2000; “Orangeburg Co., Rt. 178, 2.3 km NNE of North (33°37.0’N, 81°04.7'W), 21 Oct. 2000; Sumter Co., Rt. 261, 4.2 km S of Wedgefield (33°51.4'N, 80°30.9'W), 26 May 1999; Wedgefield (3753:5(N, 80°31.17 W),.21, Oct., 2000. Habitats and host plants.—Southeastern collections were from disturbed sites such as roadsides and cotton and soybean fields, mainly in the fall-line sandhills. The collec- tion from Greenville Co., S.C., was in the piedmont. An adult was beaten from the crown of weeping lovegrass (Eragrostis curvula [Schrad.] Nees; Poaceae) in North Carolina, and adults were on fruits of Mexican tea or wormseed (Chenopodium ambrosioides L.; Chenopodiaceae) southwest of Eureka, Edgefield Co., S.C. All other collections were from Palmer’s amaranth (Amaranthus palmeri S. Wats.; Amaranthaceae), the host 139 plant on which this rhopalid was found ini- tially in the Southeast (Wheeler 1988). In the western United States, an adult was beaten from the crown of weeping lovegrass in late April 2000 at each of three sites: Cart- er (34°11.2'N, 97°00.6'W) and Woodward (36°15:5'N, 99°09.7'W). counties in Oklahoma (new state record) and Gaines Co., Tex. (32°37.2'’N, 102°38.4’W). In late May 2001, an adult was found in the crown of weeping lovegrass in Terry Co., Tex. (33°13.7'N, 102°09.1'W). In California in mid-August 1998, nymphs and adults were on tumble pigweed (A. albus L.) in Fresno County (36°51.7’N, 120°27.8'W), and on prostrate pigweed (A. blitoides S. Wats.) in San Benito County (36°56.8'N, 121°23.4’W). Host plants—that is, species on which nymphs were found—in a sparsely vegetated gravel lot at the Port of Stockton, San Joaquin County (37°57.1'N, 121°19.3'W), in mid-Au- gust 1998 were A. albus, A. blitoides, C. am- brosioides, and Russian thistle (Salsola tra- gus L.; Chenopodiaceae). Nymphs and adults appeared to be more numerous on and under the amaranths A. a/bus and A. blitoides than they were on the chenopods C. ambrosioides or S. tragus. Prionosoma podopioides Uhler 1863 New southeastern U.S. records.— NORTH CAROLINA: Moore Co., Aber- deen (35°07.9'N, 79°26.0’W), 25 Aug. 1996; Richmond Co., Rt. 1, 2.4 km NNE of Mar- ston (35°00.3'N, 79°34.0’W), 25 Aug. 1996, 12 July 1998. SOUTH CAROLINA: Ches- terfield Co., Rt. 1, Middendorf (34°31.3'N, 80°09.5'W), 11 July & 6 Sept. 1998; Edge- field Co., CR 407 nr jet. CR 37, 8.5 km SW of Eureka (33°39.5'N, 81°51.6’W), 27 June & 28 Nov. 1998, 12 May 2001; Fairfield Co., Rt. 34, 0.6 km W of Kershaw Co. line, 12.6 km SSE of Ridgeway (34°16.4'N, 80°49.9'W), 20 May 2000. Habitats and host plants.—AII southeast- ern collections were from the fall-line sand- hills. Plantago aristata Michx., a known host plant (Wheeler 1988), was present at all sites in North Carolina and South Car- 140 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON olina, but the pentatomid was collected from bunchgrasses within one or two m of bracted plantain, except in May 2001 when two third instars were found on a new host, Wright’s plantain (P. wrightiana Dene.), in Edgefield Co., S.C. In North Carolina, four adults were beaten from crowns of weeping lovegrass in Richmond County, and an adult was beaten from weeping lovegrass in Moore County. In South Carolina, crowns of weeping lovegrass yielded three adults in Chesterfield County and an adult (28 Nov.) in Edgefield County. An adult also was beaten from the crown of broomsedge (An- dropogon virginicus L.; Poaceae) in Fair- field County. In Edgefield County, an adult also was swept from the inflorescences of southern jointweed (Polygonella americana [Fisch. & Mey.] Small; Polygonaceae). In the western United States, I collected an adult of P. podopioides in Oklahoma (new state record) from the crown of Andro- pogon sp. in Dewey County (35°58.0'N, 98°58.6'W) on 26 April 2000 and three adults from woolly plantain (Plantago pa- tagonica Jacq.) in Woodward County (36°19.2'N, 99°20.1'W) on 25 June 1999; in Texas (new state record) an adult was found on redseed plantain (P. rhodosperma Dene.) in Scurry Co. (32°53.4’N, 101°03.9'W), on 25 May 2001. If nymphal development oc- curs on redseed plantain and woolly plan- tain, these plants are new host records for Prionosoma podopioides. DISCUSSION Although they belong to different heter- opteran families, A. impressicollis and P. po- dopioides are similar in having been de- scribed more than 130 years ago and in being seldom collected and poorly known through- out their current ranges. Both the rhopalid and the pentatomid also belong to monotypic genera, are considered western species that are recent additions to the fauna of the south- eastern United States, develop frequently on western plants that are naturalized in the East, and are narrowly host specific. Amaranthus palmeri, the only plant spe- cies on which nymphs of Aufeius impressi- collis have been collected in the Southeast (Wheeler 1988), is indigenous to the South- west or southern Great Plains (e.g., Muensch- er 1980, Great Plains Flora Association 1986). Correll and Johnston (1970) remarked that A. palmeri is rarely adventive in the east- ern United States, but since 1974, this plant has become an increasingly important weed in crops such as cotton, peanut, and soybean in the southeastern states (Webster and Coble 1997). Because nymphs of A. impressicollis were found on several chenopods in Califor- nia, the host range of this rhopalid appears also to include the Chenopodiaceae, a family closely related to the Amaranthaceae (e.g., Heywood 1993, Takhtajan 1997). I assumed that one of the rhopalid’s chenopodiaceous_ hosts in California was Salsola tragus and did not submit the plant for identification. In California, however, Russian thistle consists of genetically di- vergent populations, one of which is un- named. The more widespread species is true S. tragus (Ryan and Ayres 2000). An important host of P. podopioides in the Southeast is bracted or buckhorn plantain (Plantago aristata; Plantaginaceae) (Wheel- er 1988), a plant of disturbed habitats, par- ticularly dry roadsides and sandy soils (e.g., Harper 1944, Johnson 1981). This plant might be native to the southwestern states or South America (Harper 1944). Gleason and Cronquist (1991) listed the original U.S. range of bracted plantain as Illinois to Lou- isiana and Texas, noting that it is naturalized farther east. Plantago aristata was not re- corded from the southeastern states until the 1860s (Harper 1944, Fernald 1950). Another western heteropteran that recent- ly became established in the eastern United States, Catorhintha mendica Stal, apparently tracked the eastern spread of its principal host, wild four 0’ clock (Mirabilis nyctaginea [Michx.] MacMill.; Nyctaginaceae) (Balduf 1957, Hoebeke and Wheeler 1982). Whether the eastward spread of either A. impressi- collis or P. podopioides was as host depen- dent as that of C. mendica is unknown. VOLUME 104, NUMBER | Both the rhopalid and pentatomid also were beaten from the crowns of bunchgrass- es, especially those of Eragrostis curvula. In the southeastern states, the well-developed crowns of this African grass provide shelter and overwintering sites for numerous native heteropteran species (Wheeler 1999; AGW, unpublished data). ACKNOWLEDGMENTS I thank G. FE Hrusa (California Depart- ment of Food and Agriculture, Sacramen- to), PB D. McMillan (Clemson University), and J. EF Townsend (formerly Clemson Uni- versity) for identifying plants; L. A. Allen (San Joaquin County Department of Agri- culture, Stockton, Calif.) for facilitating field work in his county; T. J. Henry (Sys- tematic Entomology Laboratory, USDA, Washington, D.C.) for companionship dur- ing field work in California (1998) and Oklahoma (1999); D. A. Rider (North Da- kota State University, Fargo) for sharing his knowledge of pentatomid distribution; and PH. Adler (Clemson University) for re- viewing the manuscript. LITERATURE CITED Balduf, W. V. 1957. The spread of Catorhintha men- dica Stal (Coreidae, Hemiptera). Proceedings of the Entomological Society of Washington 59: 176-185. Blatchley, W. S. 1926. Heteroptera or True Bugs of Eastern North America, with Especial Reference to the Faunas of Indiana and Florida. Nature Pub- lishing Co., Indianapolis, Ind. 1116 pp. Correll, D. S. and M. C. Johnston. 1970. Manual of the Vascular Plants of Texas. Texas Research Foundation, Renner, Tex. 1881 pp. Fernald, M. L. 1950. Gray’s Manual of Botany, 8th ed. American Book Co., New York. 1632 pp. Gleason, H. A. and A. Cronquist. 1991. Manual of Vascular Plants of Northeastern United States and Adjacent Canada, 2nd ed. New York Botanical Garden, Bronx, N.Y. 910 pp. Great Plains Flora Association. Great Plains. University Press of Lawrence. 1392 pp. Harper, R. M. 1944. Notes on Plantago, with special reference to P. cordata. Castanea 9: 121—130. Heywood, V. H., ed. 1993. Flowering Plants of the World. Oxford University Press, New York. 335 pp. 1986. Flora of the Kansas, 141 Hoebeke, E. R. and A. G. Wheeler, Jr. 1982. Cator- hintha mendica, a Great Plains coreid now estab- lished on the Atlantic Coast (Hemiptera: Corei- dae). Entomological News 93: 29-31. Johnson, M. FE 1981. Phrymaceae and Plantaginaceae in Virginia. Virginia Journal of Science 32: 12-16. McPherson, J. E. 1970. A key and annotated list of the Scutelleroidea of Michigan (Hemiptera). Michi- gan Entomologist 3: 34—63. McPherson, J. E. and J. P. Cuda. 1975. The first record in Illinois of Prionosoma podopioides (Hemiptera: Pentatomidae). Journal of the Kansas Entomolog- ical Society 48: 371—373. Muenscher, W. C. 1980. Weeds, 2nd ed. Cornell Uni- versity Press, Ithaca, N.Y. 586 pp. Osborn, H. and C. J. Drake. 1915. Additions and notes on the Hemiptera-Heteroptera of Ohio. Ohio Nat- uralist 15: 501—508. Ryan, FE J. and D. R. Ayres. 2000. Molecular markers indicate two cryptic, genetically divergent popu- lations of Russian thistle (Salsola tragus) in Cal- ifornia. Canadian Journal of Botany 78: 59-67. Stal, C. 1870. Enumeratio Hemipterorum. Bidrag till en foreteckning 6fver alla hittills kanda Hemip- tera, jemte systematiska meddelanden. Part 1. Kongliga Svenska Vetenskaps-Akademiens Han- dlingar 9(1): 1-232. Stoner, D. 1916. Distributional notes on some Iowa Pentatomoidea. Proceedings of the lowa Academy of Science 23: 303-307. . 1920. The Scutelleroidea of lowa. University of Iowa Studies in Natural History 8: 1—140. Takhtajan, A. 1997. Diversity and Classification of Flowering Plants. Columbia University Press, New York. 643 pp. Uhler, P. R. 1863. Hemipterological contributions. No. 2. Proceedings of the Entomological Society of Philadelphia 2: 361—366. Webster, T. M. and H. D. Coble. 1997. Changes in the weed species composition of the southern United States: 1974 to 1995. Weed Technology 11: 308— Siliv/e Wheeler, A. G., Jr. 1984. Aufeius impressicollis (He- miptera: Rhopalidae): Easternmost U.S. record, host plant relationships, and laboratory rearing. Journal of the New York Entomological Society 92: 174-178. . 1988. First southeastern U.S. records of the western heteropterans Prionosoma podopioides (Pentatomidae) and Aufeius impressicollis (Rho- palidae), with a review of distribution and host plants. Journal of the New York Entomological Society 96: 304—309. . 1999. Oncozygia clavicornis Stal and Allo- podops mississippiensis Harris and Johnston: As- sociation of rarely collected Nearctic turtle bugs (Heteroptera: Pentatomidae: Podopinae) with an introduced African grass. Proceedings of the En- tomological Society of Washington 101: 714—721. PROC. ENTOMOL. SOC. WASH. 104(1), 2002, pp. 142-159 DESCRIPTIONS OF TEPHRITIS FOOTEI AND T. HEADRICKI, NEW SPECIES (DIPTERA: TEPHRITIDAE), WITH NOTES ON THEIR LIFE HISTORIES IN SOUTHERN CALIFORNIA RICHARD D. GOEDEN Department of Entomology, University of California, Riverside, CA 92521, U.S.A. (e-mail: richard.goeden @ucr.edu) Abstract.—Tephritis footei Goeden, n. sp., is a fruit fly (Diptera: Tephritidae) pro- ducing a single, annual generation in the flower heads of Artemisia tridentata Nuttall (Asteraceae) belonging to the subtribe Artemisiinae of the tribe Anthemideae in south- ern California. The egg, first-, second- and third-instar larvae and adults are described and figured, and selected characteristics of these stages are compared with those of other southern California Tephritis. Another new species of Tephritis belonging to the “araneosa complex’’, like 7. footei, also is described: T. headricki Goeden, n. sp. Host plants of 7. headricki belong to the genus Solidago of the subtribe Solidagininae of the tribe Astereae. The adults of these two new species are distinguished by a combination of characters involving oviscape lengths, wing patterns, wing lengths, and leg colors. The egg of T. footei is covered by a smooth, membraneous sheath of unknown function. The partial description of the first instar of 7. footei is the second such for this instar in the genus Tephritis. Noteworthy for the first instar is confirmation of the fusion of the integumental petal with the stomal sense organ of 7. footei, like that of 7. teerinki Goeden. Similarly, the partial description of the second instar also is only the second for the genus Tephritis. The third instar has the fewest number of minute acanthae and the most integumental petals circumscribing the prothorax among four other described species. The lower, lateral, integumental petal of the cephalothorax continues around the oral cavity, like that of the third instar of 7. teerinki. The life cycle is of the aggregative type and overwintering occurs as long-lived, sexually im- mature adults. Key Words: Insecta, Tephritis, Artemisia, Solidago, Asteraceae, nonfrugivorous Tephri- tidae, biology, taxonomy of adults and immature stages, flower-head feed- ing, aggregative life cycle, seed predation, parastoids To date, the life histories and immature (Coquillett), will be described in my next stages of four species of Tephritis have been described in detail from southern Cal- ifornia: 7. arizonaensis Quisenberry (Goe- den et al. 1993), 7. baccharis (Coquillett) (Goeden and Headrick 1991), T. joanae Goeden (Goeden 1993, 2001b), and T. teer- inki Goeden (Goeden 2001c). The imma- ture stages of a fifth species, 7. stigmatica paper, and its biology is relatively well known (Tauber and Toschi 1965; Goeden 1988a, 1993). This paper describes the life history, adults, and immature stages of the new species, 7. footei Goeden, along with the adults of another new species, 7. head- ricki Goeden, both segregates from the “‘ar- aneosa complex” (Foote et al. 1993). VOLUME 104, NUMBER 1 MATERIALS AND METHODS The present study was based in part on specimens of adults belonging to the araneo- sa complex of Tephritis (Foote et al. 1993) reared from 1-liter samples of mature flower heads of various species of Asteraceae col- lected from throughout California since 1980 (Goeden 1993). The life history study and description of the immature stages of 7. footei was based in large part on dissections of sam- ples of mature and immature flower heads of Artemisia tridentata Nuttall (Asteraceae) col- lected north of the Hitchcock Ranch at the junction of U.S. Forest Service roads 3N16 and 3N54 in the San Bernardino National Forest (North Section) at 2,210-m elevation, southwestern San Bernardino Co., during 1995. One-liter samples of excised, immature and mature flower heads containing eggs, lar- vae, and puparia were transported in cold- chests in an air-conditioned vehicle to the lab- oratory and stored under refrigeration for subsequent dissection, photography, descrip- tion, and measurement. Two eggs, two first-, six second-, and nine third-instar larvae dis- sected from flower heads were preserved in 70% EtOH for scanning electron microscopy (SEM). Prepuparia and puparia were placed in separate, glass shell vials stoppered with absorbant cotton and held in humidity cham- bers at room temperature for adult and par- asitoid emergence. Specimens for SEM were hydrated to distilled water in a decreasing se- ries of acidulated EtOH. They were osmicat- ed for 24 h, dehydrated through an increasing series of acidulated EtOH and two, 1-h im- mersions in hexamethyldisilazane (HMDS), mounted on stubs, sputter-coated with a gold- palladium alloy, studied and digitally photo- graphed with a Philips XL-30 scanning elec- tron microscope in the Institute of Geophys- ics and Planetary Physics, University of Cal- ifornia, Riverside. Most adults reared from isolated prepu- paria and puparia were individually caged in 850-ml, clear-plastic, screened-top cages with a cotton wick and basal water reser- voir and provisioned with a strip of paper 143 toweling impregnated with yeast hydroly- zate and sucrose. These cages were used for studies of longevity and sexual maturation in the insectary of the Department of En- tomology, University of California, River- side, at 25 + 1°C, and 14/10 (L/D) photo- period. Sixteen arenas each consisting of a clear-plastic, petri dish provisioned with a flattened, water-moistened pad of absorbant cotton spotted with honey (Headrick and Goeden 1994). Each arena contained a vir- gin male and female obtained from emer- gence cages that were used for observations of courtship and copulation behavior. Plant names used in this paper follow Hickman (1993) and Bremer (1994); tephritid names and adult terminology follow Foote et al. (1993). Format used to describe the adults follows the format and method of measure- ment of Jenkins and Turner (1989), as used and modified by Goeden (1993, 2001c). Ter- minology and telegraphic format used to de- scribe the immature stages follow Goeden (2001a, b, c), Goeden et al. (1993), Goeden and Headrick (1991), Goeden and Teerink (1999), Teerink and Goeden (1999), and our earlier works cited therein. The holotypes and allotypes together with 74 and 25 reared par- atypes of T. footei and T. headricki, respec- tively, have been deposited in the National Museum of Natural History, Smithsonian In- stitution, Washington, D.C. (USNM). The ho- lotypes, allotypes, and 22 and 35 paratypes were used for measurements to describe 7. footei and T. headricki, respectively. All re- maining paratypes and voucher specimens not designated as paratypes, voucher speci- mens of immature stages, and reared parasit- oids of 7. footei reside in my research col- lections. Means + SE are used throughout this paper. Digitized photographs used to con- struct text figures were processed with Adobe Photoshop® Version 5. RESULTS AND DISCUSSION Taxonomy The following key couplets modifying couplet 6 and replacing couplet 17 in the 144 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON key of Foote et al. (1993), and replacing couplets 10 and 17 as previously modified by Goeden (1993, 2001c), enables one to distinguish these two new species. The three-digit, figure numbers in couplets 6, 10a, 18, and 19 refer to Foote et al. (1993), not to the present publication. The strike- throughs indicate parts of the original key that I suggest deleting. 6. Preapical brown area of wing interrupted by numerous round subhyaline spots (fig. 428, c); dark prtlhera ret eel rao Heattedtch peste FIOF to PLr atta cst be woth at least | vd spet-present anterior arm of Y-shaped mark at wing apex may be broken or conspicuously narrower than posterior arm (fig. 429, d)... 7 — Preapical brown area of wing usually with at most a few round subhyaline spots, its ap- pearance never obscured by them (fig. 430, bo. darh—pathern + -eelt ep ectiattel y poste roy bosphereetHetet ate ee shaped mark in wing apex rarely broken as ADOVE™ oe iets Gita eee ees soa aees te 10 10. Anal lobe with dark markings of wing usually extendinoatonwin Sum aroin alee eee 10a — Anal lobe with dark markings of wing usually not, or only very faintly and narrowly, ex- tenGine toOnwinPemarcin ese ee sien 1] 10a. Wing with hyaline spot in basal end of cell t,,5 large, oval to quadrate, and extended from vein R,,; to vein M (often broadly based on the latter), usually contiguous with the hyaline area in cell r,,, directly anterior to it; at least some of hyaline spots along margin of anal lobe wider than half the width of anal lobe; wing pattern as in Goeden (1993) = Wing with hyaline spot in basal end of cell cell r,,; anterior to vein dm-cu, smaller, round or oval, and not extended from vein R,,;, but sometimes touching vein M (fig. Joanae Goeden 430, c); hyaline spots along margin of anal lobe usually less than half the width of this lobe 17. Wing length of females usually under 2.6 mm; of males, usually under 2.3 mm (Fig. IWAN) Ane ie tot oid CR eee 3 = Wing length of females usually exceeding 2.6 mm; of males, usually exceeding 2.3 mm signatipennis Foote footei Goeden, n. sp. 18. Wing with hyaline spot in basal end of cell T4,5 large, oval to quadrate, and extending from vein R,,; to vein M (often broadly based on the latter), usually contiguous with the hyaline area in cell r,,, directly anterior WO (Gites Ais) Bp ooo oe araneosa (Coquillett) - Wing with circular, elliptical, ovate, or bell- shaped spot in cell r,,; extending anteriorly from vein M, but usually not touching vein ante reece renee PIB at atts Rita eas c Ne) 19. Femora, especially those of hind legs, dark tomentose; frons—reddish—browntomentese: wing pattern as in fig. 438 . . ovatipennis Foote - Femora all or mostly yellow—frens—mesty whiteto-yeHow-tomentese 2. .....22.. 20 20. Oviscape two to three times as long as ter- minal abdominal tergite; hyaline spot in cell r,,5 anterior to vein dm-cu elliptical, oval, or bell-shaped and usually extending anteriorly from vein M more than two-thirds across cell 4,5; Wing pattern as in Goeden (200Ic)... MAO LES | te tee eo eee lc teerinki Goeden = Oviscape only slightly longer than the ter- minal abdominal tergite; hyaline spot in cell r,,5 anterior to vein dm-cu, smaller, round or oval, and may not touch vein M (Fig. 1B) SS Oe Pdeer Se headricki Goeden, n. sp. Tephritis footei Goeden, new species (Figs. 1A, 2—6) Adult female.—Head: In profile, 1.1 to 1.3 times as high as long, face distinctly protruding below antennae, face and frons meeting at an angle of ca. 120°, gena below eye 0.18 to 0.24 times eye height, genal bristle and most genal setulae light brown; occiput slightly swollen; frons white to light yellow pollinose, white to greyish pol- linose mid-dorsally and laterally, about 0.4 mm wide at vertex, narrowing to 0.3 mm at antennal bases, and 0.2 to 0.3 mm long; the two frontal setae dark brown to black; posterior orbital seta white, 0.4 to 0.7 times as long as anterior orbital seta; inner verti- cal seta dark brown, 0.5 to 0.7 times as long as head height, outer vertical bristle white, 0.2 to 0.3 times as long as head height; face, including antennal foveae, white; pal- pi and labellum light yellow to dark ocher- ous, sometimes tinged reddish, with four to seven, prominent, dark brown to black se- tulae apically; antenna 0.6 to 0.8 times as long as face at midline, yellow to dark ocherous, sometimes reddish, arista dark brown except base ocherous. Thorax: Scutum, scutellum, and pleural VOLUME 104, NUMBER | A (A) Right wing of Tephritis footei female; (B) Right wing of 7. headricki female. Lines = | mm. lan, I. sclerites gray pollinose over shiny black ground-color, sometimes with single, slight- ly darker, faint, narrow, medial dorsolon- gitudinal stripe and faint lateral stripe through anterior dorsal central bristle; short, white setulae invest entire scutum; comple- ment of thoracic bristles usual for the ge- nus, all dark brown or black except poste- rior notopleural bristle, white; subscutellum and mediotergite black pollinose; scutellar setulae shorter and inserted closer to each other than are scutal setulae; scutellum bare centrally, setulae present only laterally; hal- teres yellow. Legs, especially femora, dark tomentose; hind tibia dark with parallel rows of black setulae; hind femur with black setulae on posterior sixth. Wing length 2.0—2.2 mm, wing pattern as in Fig. 1A, with a prominent hyaline area imme- diately distad of pterostigma extended from costal margin posterior to and touching vein R,,;, but never extending to vein M; in- stead, usually round to oval, hyaline spot in basal end of cell r,,; arising on and nearly always touching vein M, and usually ex- tending at least halfway across cell r,,5; dark area in pterostigma extending posteri- orly to vein M and covering crossvein r-m, very rarely with hyaline markings; cross- vein r-m removed from crossvein dm-cu by about its own length; large hyaline mark- ings occupy most of cell m and extend pos- teriorly to posterior wing margin, as do large hyaline markings in cell cua,, which 145 do not cross vein CuA,, but anteriorly di- vide into hyaline spots in cell dm; anal lobe mostly hyaline. Abdomen: All tergites gray pollinose ex- cept T;, which sometimes has shiny, black spot posteromedially, but otherwise is con- colorous with mediotergite, other tergites without pattern; densely covered with col- orless setulae inserted much closer to each other than their average length, becoming longer laterally and posteriorly; two or three long, hyaline to light or dark brown setae laterally along posterior margin of last abdominal tergite; oviscape flat, dark brown or black, with setulae on basal % similar to those on abdominal tergites, apical third with extremely fine, short black setae; ovi- scape in dorsal view 1.1 to 1.9 times as long as last abdominal tergite and 1.2 to 1.7 times as wide at base as long. Male.—Similar to @ in all respects ex- cept wing length 1.7—2.1 mm; genitalia ocherous brown to black, subshining. Variation.—Examination of reared 7. footei with intact setation confirmed that the holotype, allotype, and most adults had two pairs of black frontal setae; however, one (1%) of 80 2 paratypes had five frontal se- tae, i.e., two pairs plus a third, weak, black seta inserted ventral to the other two pairs. One (1%) of 85 d paratypes had three fron- tal setae, and another d paratype had two pairs of frontal setae, one of which in the ventral pair was weak and white. 146 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON rhe right wings of the 80 ¢ paratypes varied from 1.7 to 2.6 mm in length, but 77 (96%) 2 paratypes measured = 1.8 and = 2.4 mm: whereas, the right wings of the 85 } paratypes varied from 1.7 to 2.3 mm in length. Only one (1%) of the 80 2 paratypes had a single, faint, hyaline spot in cell r, within the dark area extending posteriorly from the stigma in one wing. Likewise, one (1%) of the 85 d paratypes had one, faint, hyaline spot in cell r,,, within this dark area in one wing. The antennae of three (3.8%) of the 80 2 paratypes were tinged reddish, as were the first flagellomeres only of another 14 (17.5%) of the 2 paratypes and the scape and peduncle of another 2 paratype (1%). The labellum and palpi of four of these same 18 @ paratypes also were tinged red- dish, as was the labellum alone of two more of these 18@ paratypes, and the palpi of another one of the 18 2 paratypes. The an- tennae of four (4.9%) of the 85 d paratypes were tinged reddish, as were the first fla- gellomeres only of one (1%) of the 85 d paratypes and both the scape and peduncle of another (1%) d paratype. The labellum and palpi of three of these same six d par- atypes also were tinged reddish, as was the labellum alone of the other three of these same six d paratypes. An additional four gd of the 85 d para- types examined bore the thalli of a benign, ectoparasitic fungus, Stigmatomyces sp. (Ascomycetes: Laboulbeniaceae), a genus reported from California by Goeden and Benjamin (1985). Diagnosis.—The main morphological characters distinguishing the adults of 7. footei from all other North American spe- cies north Mexico are its short wing length and the dark band in the wing extending obliquely from the pterostigma to cover vein r-m (Fig. 1A), which usually is free of hyaline spots. The wing length of T. footei is at least | mm shorter than that of T. head- ricki Goeden n. sp. described below, and the femora of T. footei are dark, not yellow like 7. headricki. An elliptical, ovate, or bell-shaped, hyaline spot in cell r,,; extend- ing anteriorly from vein M, but not touch- ing vein R,,, (Fig. 1A), instead of a large oval to quadrate, hyaline spot in basal end of cell r,,; extending from vein R,,; to vein M (often broadly based on the latter), and usually contiguous with the hyaline area in cell r,,; directly anterior to it, further dis- tinguishes 7. footei from the smallest spec- imens of 7. araneosa (Coquillett). The host relations of 7. footei, T. head- ricki, and T. araneosa apparently also dif- fer, as discussed in the next section. Indeed, as' Foote et al. (1993) stated (ps 196) see araneosa belongs to a complex of closely related species, the precise identification of which may never be attained without exten- sive biological studies.’” Foote (1960) de- scribed three of these, 1.e., candidipennis Foote, ovatipennis Foote, signatipennis Foote, to distinguish those larger species with longer oviscapes, and Blanc (Foote and Blanc 1979) described another, /eavit- tensis Blanc, on the basis of additional characters. Goeden (1993, 2001c) described T. joanae and T. teerinki, (respectively) two more of those Tephritis species with longer oviscapes. I also describe below in addition to T. footei, another new species with a shorter oviscape belonging to the “‘araneo- sa complex,” i.e., 7. headricki. Unfortu- nately, I have not been able to study its life history and immature stages, but am able to distinguish it morphologically, as noted above, as adults reared from a separate, dif- ferent tribe of host plants in California. Types.—Holotype: 2; N(orth) of Hitch- cock Ranch at 2,207-m (7,240-feet) (eleva- tion), S(an) Bernardino Nat(ional) F(o)r(e)st, S(an) Bernardino Co(unty); T(ownship) 3N(orth), R(ange)lW(est), S(ection) 25; 27.x.1993; R. D. Goeden, coll. (hereafter RDG, coll.)/J. A. Teerink, coll. (hereafter JAT, coll.); reared from flower head of Ar- temisia tridentata. Allotype; ¢, same data as holotype (USNM). Paratypes: CALIFOR- NIA: 8 ¢ and 6@; same data as holotype (5 6 and 5 2 to USNM). 1 @; same data as VOLUME 104, NUMBER 1 holotype, except 31.vi11.1995; RDG/JAT, coll. (1 2 to USNM). 1 ¢ and 6 2; same data as holotype, except 13.1x.1995; RDG/ WARscoll.G, 2 to USNM):3 <@ andi2°2; same data as holotype, except 12.x.1995; RDG/JAT, coll. (1 6 and 1 2 to USNM). Unless otherwise indicated the following specimens also were reared from flower head of A. tridentata. 16 3 and 14 2; Onyx Summit at 8,370 ft (2,551 m), S. Bernardino Nat. Forest, S. Bernardino Co.; TIN, R3E, S12; 27.x.1993; RDG/JAT, coll. (6 6 and 6 2 to USNM). 6 o and 4 &; St. Hwy. 38 N(orth)W(est) of Onyx Peak at 2,323 m (7,620 feet); S. Bernardino Nat. Frst., S. Bernardino Cos T2N,.R2E, S353.27:%:1993; RDG/JAT, coll. (3 6 and 2 2 to USNM). 6 3d and 8 @; along road to Big Pine Flat (U. S. F(orest) S(ervice) Road 3N14) at 1,747 m (5,730 feet), S. Bernardino Nat. Forest, S. Bernardino Co.; T2N, R2W, S1; 27.x.1993; RDG/JAT, coll. (3 d and 4 @ to USNM). 2 2; along road to Big Pine Flat (USFS Road 3N16) at 1,902 m (6,240 feet), S. Bernardino Nat. Forest., S. Bernardino Co., T2N, R1IW, S690 272419932. RDG/JAT, coll...) 2 sto USNM). 17 6 and 12 9; E(ast) of Big Pine Flat Station (along) USFS Road 3N16 at 2,039 m (6,690 feet); S. Bernardino Nat. Forest., S. Bernardino Co., TSN, RIW, S29; 27.x.1933; RDG/JAT, coll. (7 6 and 4 @ to USNM). 4 6; same data as preceding entry, only all infected with Stigmatomyces; 9.viii.1995; (2 d6 to USNM). 11 6 and 12 2: E(ast) of Big Pine Flat Station at 2,149 m (7,050 feet); S. Bernardino Nat. Forest, S. Bernardino Co., T5N, RIW, S28; 27.x.1993; RDG/JAT, coll. (6 6 and 6 & to USNM). 6 6 and 5 @; Arrastre Flat at 2,225 m (7,300 feet); S. Bernardino Nat. Forest, S. Bernar- dine Gos T3N; RIE, S34; 27:x:1993;, RDG/ JAT, coll. (3 3 and 2 2 to USNM). 6 6 and 6 @; Junction (State) H(igh)w(a)y 38 and Heartbar R(oa)d (U. S. Forest) S(ervice) Road 1NO2) at 2,051 m (6,730 feet), S. Ber- nardino Nat. Forest, S. Bernardino Co.; TSN, RIW, S29; 27.x.1993; RDG/JAT, coll. (3 36 and 3 2 to USNM). 2 6 and 1 2; SE of Holcomb Valley Campground at 2,170 m 147 (7,190 feet); S. Bernardino Nat. Forest, S. Bernardino Co., T2N, RIE, S4; 27.x.1933; RDG/JAT, coll. (1 d to USNM). 1 3; Coon- creek Jumpoff sat 25192, mv@7, 190) feet)ses. Bernardino Nat. Forest, S. Bernardino Co.; RIN; R2E-6S223127/ 321993: RDGAATseoll. (1 6 to USNM). Etymology.—This tephritid is named for an early mentor, Richard H. Foote, who en- couraged my study of the nonfrugivorous Tephritidae of California, and whose many taxonomic writings provided examples of careful craftsmanship and a firm foundation and guidance for my own studies, especial- ly those on Tephritis and Trupanea spp. Tephritis headricki Goeden, new species (Fig. 1B) Adult female.—Head: In profile, 1.1 to 1.4 times as high as long, face distinctly protruding below antennae, face and frons meeting at an angle of ca. 120°, gena below eye 0.13 to 0.18 times eye height, genal bristle and most genal setulae light brown, some hyaline; occiput slightly swollen; frons white to ocherous pollinose, contrast- ing white to dark ocherous mid-dorsally and laterally, 0.5—0.6 mm wide at vertex, narrowing to about 0.4 mm at antennal ba- ses, and 0.3 to 0.4 mm long; the two frontal setae dark brown to black; posterior orbital seta white, 0.6 to 0.8 times as long as brown anterior orbital seta; inner vertical seta brown, about 0.7 times as long as head height, outer vertical bristle white, 0.2 to 0.3 times as long as head height; face, in- cluding antennal foveae, white to reddish; palpus and labellum light yellow to dark ocherous, sometimes tinged reddish, with four to seven, prominent, brown setulae apically; antenna about 0.7 as long as face at midline, yellow to dark ocherous, some- times reddish, arista brown except base ocherous. Thorax: Scutum and pleural sclerites light-gray pollinose anteriorly, scutum darkening to golden brown pollinose pos- teriorly, all over shiny, dark brown to black ground-color; short, white setulae invest en- 148 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tire scutum:; complement of thoracic bristles usual for the genus, all light to dark brown except posterior notopleural, white; scutel- lum golden brown medially, dark anteriorly along juncture with scutellum and dark api- cally, ocherous posteriolaterally, ventral half of subscutellum light-gray pollinose, dorsal half pale ocherous, mediotergite shiny black laterally and ventrally, dorso- centrally light-gray pollinose; scutellar se- tulae similar in length, but inserted closer to each other than scutal setulae; scutellum bare centrally, setulae present only laterally; halteres yellow, ocherous basally. Legs, yellow; hind tibia with parallel rows of brown setulae; hind femur with brown se- tulae on posterior fifth. Wing length 3.4 to 3.7 mm, wing pattern as in Fig. 1B, witha prominent hyaline area immediately distad of pterostigma extending from costal mar- gin posterior to and touching vein R,,;, with round to oval, hyaline spot in basal end of cell r,,; just touching or slightly sep- arated from vein M and usually only half or two-thirds as wide as cell r,,, (Fig. 1B); dark area in pterostigma extending posteri- orly to vein M and covering crossvein r-m, usually with no or one or two, faint to small, but discrete, hyaline spots in cell br; crossvein r-m removed from crossvein dm- cu by about its own length; large hyaline markings occupy most of cell m and extend posteriorly to posterior wing margin, as do large hyaline markings in cell cua,, which may cross vein CuA,, or occupy as separate hyaline spots, most of cell dm; anal lobe usually hyaline along wing margin or oc- casionally very faintly patterned with large, hyaline spots. Abdomen: All tergites light-gray polli- nose, concolorous with dorsocentral part of mediotergite, without pattern; densely cov- ered with colorless setulae inserted much closer to each other than their average length, becoming longer laterally and pos- teriorly; two or three, long, hyaline to light or dark brown setae along posterior margin of last abdominal tergite; oviscape flat, dark-brown to black, with setulae on basal *%; similar to those on abdominal tergites, apical third with extremely fine, short dark brown setae; oviscape in dorsal view 1.2 to 1.8 times as long as last abdominal tergite and 1.2 to 1.9 as wide at base as long. Male.—Similar to 2 in most respects ex- cept wing length 3.0—3.6 mm; genitalia ocherous to dark brown, subshining. Variation.—Examination of reared 7. headricki with intact setation confirmed that the holotype, allotype, and all paratypes ex- cept one 2 had two pairs of black frontal setae; this female had a third pair of short, white setae between the other two pairs of frontal setae. Eight (50%) of the 16 @ types had no hyaline spot, however faint, in cells r,, r5,3, or br within the dark area extending pos- teriorly from the stigma in both wings. Two (13%) of the 16 @ types had one faint hy- aline spot in cell r,,, within this dark area in only one wing. Eight (50%) of the 16 @ types had one or two, faint to small, but discrete, hyaline spots in the apex of cell br, basad of crossvein r-m. In comparison, eight (35%) of 22 d types had no hyaline spot, however faint, in cells r,, 1,3, or br within the dark area extending posteriorly from the stigma. Only one (5%) of the 22 d types had one, small, discrete and two, faint hyaline spots in cell r,,; within each dark area, respectively, in its wings. And, 15 (68%) of the 22 6 types had one or two, faint to prominent, hyaline spots in the dark area at the apex of cell br, basad of cross- vein r-m. The hyaline spot in the basal end of cell t,,; reaches vein M posteriorly in seven (44%) of 16 @ types, but was slightly sep- arated from vein M in the remaining nine (56%) 16 & types. Likewise, the hyaline spot in the basal end of cell r,,; reaches vein M posteriorly in 13 (59%) of 22 d types, but was slightly separated from vein M in another eight (36%) of 22 d types. The one remaining ¢ type had one wing in each category. The anal lobe of only one (6%) of the 16 2 types contained a very faint pattern with VOLUME 104, NUMBER 1 large hyaline spots that extended to and along the wing margin; whereas, three (14%) of the 22 d types showed this con- dition. The first flagellomeres of the antennae of nine (56%) of 16 2 types were tinged red- dish, as were both pedicels of still another (6%) of the 2 types. The labellum and pal- pus of three of these same 10 @ types also were tinged reddish, as was the labellum or palpi alone of two other, separate @ para- types. The antennae of four (18%) of the 22 d types were tinged reddish, as were the first flagellomeres only or pedicels only of another two each (9 and 9%) of the 22 3 types. The labellum and palpus of 10 (45%) of these 22 ¢ types also were tinged red- dish, as was the labellum alone or palpi alone of one each (5 and 5%) of these 22 d types. As some, but not all specimens of T. araneosa and T. ovatipennis (unpub- lished data) as well as T. footei and T. head- ricki had reddish colored head parts, this character is removed from couplet 19a de- scribing 7. ovatipennis in my revision above of the key to Tephritis in Foote et al. (1993). This reddish coloration apparently lies on a continuum of colorations from white to yellow to ocherous to reddish to brown to black seen in Tephritis. Diagnosis.—The main morphological characters distinguishing the adults of 7. headricki from all other North American species of Tephritis north Mexico are a combination of a round or ovate, hyaline spot in cell r,,; anteriorad of vein M, and not touching vein R,,; (Fig. 1B); the dark band from pterostigma to vein M extending obliquely to cover vein r-m (Fig. 1B); yel- low femora; and an oviscape only about twice as long as the last abdominal tergite. The femora of 7. ovatipennis, especially those of the hind legs are dark grey to black tomentose, not yellow (Foote et al. 1993). Moreover, T. ovatipennis and T. teerinki be- long to those larger congeners in the “ar- aneosa complex”’ (Foote 1960, Foote et al. 1993) with long oviscapes, not like those of T. araneosa with an oviscape “*... only 149 slightly longer or shorter than, or equal to, the terminal abdominal tergite ...”. In this last regard, of 62 7. araneosa reared from four species of Artemisia other than A. tri- dentata (see discussion of hosts below), none had oviscapes shorter than the termi- nal abdominal tergite, only one (5%) was equal, and the remainder had oviscapes from 1.1 to 1.7 times as long as the terminal abdominal tergite. Another grouping of 104 T. araneosa reared from three Chrysotham- nus spp. and one Ericameria spp. (which I could not distinguish morphologically as another new species; see discussion of hosts below) also contained none with an ovi- scape shorter than or equal to the terminal abdominal tergite. The oviscapes of these 104 females varied from 1.1 to 2.3 times as long as the terminal abdominal tergites. No obvious distinctions in measurements or perceived characters of these two groups could be correlated with their distinctive host-plant taxonomic affinities. These two groups represent what remains of 7. ara- neosa after the removal of segregates with longer oviscapes by Foote (1960), Foote and Blanc 1979), and Goeden (1993, 2001c), and two more species with shorter Oviscapes in the present paper. This “‘resid- ual” 7. araneosa probably contains at least one more undescribed species. Types.—Holotype: 2; Dead Man Creek at 2,500 m (elevation), Inyo National For- est, Mono County; T(ownship)3S(outh), R(ange)27E(ast), S(ection)5; 9.x.1986; R. D. Goeden, coll. (hereafter RDG, coll.); reared from flower head of Solidago cana- densis L. Allotype: d, same data as holo- type (USNM). Paratypes: CALIFORNIA: 8 3d and 82; same data as holotype (5 ¢ and > 2 to" USNM)4'7 6 Vandi 5 92> %Beasore Meadow off Beasore Road 25 km N(orth)E(ast) of Bass L(ake) at 1,960 m, Sierra National Forest, Madera County; T6SSsRQSES So 1osxe19ssy RDG? coll® reared from flower heads of S. canadensis (4 ¢ and 3 2 to USNM). 2 @; along Upper Deadman Creek at 2,496 m; Inyo National Forest, Mono County; T3S, R27E, S6; 150 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Egg of Tephritis footei: (A) habitus, anterior to left; (B) pedicel showing pattern and shapes of aeropyles. 18.1x.1991; RDG/JAT, coll.; reared from flower heads of S. canadensis (1 @ to USNM). 4 6; Antelope Spring; Inyo Na- tional Forest., Inyo County; T17S, R35E, S24; 15.ix.1982; RDG, coll.; reared from flower head of S. confinis Nuttall (2 d to USNM). 1 3; Kennedy Meadows, Sequoia National Forest; Inyo County; T22S, R35E, S24; 25.ix.1980; RDG, coll.; reared from flower head of Euthamia (formerly Solida- go) occidentalis Nuttall (1 ¢ to USNM). 1 3; Round Valley Reservoir at 1,372 m; Plu- mas National Forest; Plumas County; 10.ix.1986; RDG, coll.; reared from flower head of E. occidentalis (1 3d to USNM). Etymology.—Tephritis headricki 1s named for my friend, last Ph.D. student, and research collaborator, David H. Head- rick, who as my successor in California te- phritidology hopefully will one day study and publish the life history and describe the immature stages of his tephritid namesake. Immature stages.—The egg, first-, sec- ond-, and third-instar larvae of Tephritis footei are described below. Egg.: Five eggs measured in situ in field- collected, preblossom flower heads plus a to- tal of five ova dissected from two females were white, opaque, smooth, elongate-ellip- soidal, 0.52 + 0.007 (range, 0.48—0.56) mm long, 0.16 + 0.005 (range, 0.14—0.18) mm wide, smoothly rounded at tapered basal end (Fig. 2A); pedicel button-like, 0.02 mm long, circumscribed apically by different-sized, semicircular aeropyles arranged singly or in rows of two parallel to the long axis of the egg (Fig. 2B). The egg of T. footei (Fig. 2A), like those of T. joanae (Goeden 2001b) and T. teerinki (Goeden 200Ic), differs from eggs of 7. bac- charis (Goeden and Headrick 1991) and T. arizonaensis (Goeden et al. 1993) by lacking prominent polygonal reticulation of the cho- rion. Also, the egg of T. footei, like those of T. teerinki and T. joanae, apparently is cov- ered by a smooth, membraneous sheath (Figs. 2A, B; Goeden 2001b, c), which remains 1n- tact and is not partly shed and peeled back during oviposition as in the other two species. The function of this membraneous sheath re- mains unknown. It was first reported for T. arizonaensis by Goeden et al. (1993), who then belatedly recognized it in 7. baccharis, and apparently it only has been reported to date from the eggs of these five species of Tephritis. In T. arizonaensis (Goeden et al. 1993), this membraneous sheath also is prominently, longitudinally striated. Weak longitudinal striations are present at the an- terior, pedicellar end of the egg of T. footei (Figs. 2A, B), but otherwise are not seen on the rest of the egg body. First instar larva: White, cylindrical (Fig. 3A); gnathocephalon conical; dorsal sensory organ well-defined, round, flattened (Fig. 3B- 1); anterior sensory lobe (Fig. 3B-2) with ter- VOLUME 104, NUMBER 1 151 Fig. 3. 1—dorsal sensory organ, 2—anterior sensory lobe, 3—terminal sensory organ, 4 First instar of Tephritis footei: (A) habitus, anterior to left; (B) gnathocephalon, ventrofrontal view, lateral sensory organ, 5— supralateral sensory organ, 6—stomal sense organ, 7—“‘lateral” integumental petal, 8—*‘median” integumental petal, 9—mouthhook, 10—median oral lobe. minal sensory organ (Fig. 3B-3), lateral sen- sory organ (Fig. 3B-4), supralateral sensory organ (Fig. 3B-5), and pit sensory organ not seen; stomal sense organ (Fig. 3B-6) ventro- laterad of anterior sensory lobe and fused with flattened, protrudent, “lateral” integu- mental petal (Fig. 3B-7) dorsad of each mouthhook, one “median” integumental pet- al between anterior sensory lobes (Fig. 3B- 8); mouthhook (Fig. 3B-9) bidentate (not shown); median oral lobe laterally com- pressed, apically rounded (Fig. 3B-10). Re- maining characters could not be seen on the few specimens examined. Only the first instar of 7. teerinki has been described in detail (Goeden 2001c) for which this partial description for 7. footei provides at least some basis for comparison. For ex- ample, it is worth noting that an integumental petal fused with the stomal sense organ (Figs. 3B-6, -7) also distinguished the first instar of T. teerinki (Goeden 2001c) and the first in- stars of at least five species of Neaspilota (Goeden 2001la) from subsequent instars. This character was first reported for the first instar of Trupanea vicina (Wulp) (Goeden and Teerink 1999). Second instar larva: White, cylindrical, rounded anteriorly, truncated posteriorly, body segments well-defined (Fig. 4A); gnath- ocephalon not seen; anterior thoracic spiracle with five, subglobose or subquadrate papillae (Fig. 4B); lateral spiracular complexes not seen; posterior spiracular plate bears three ovoid rimae (Fig. 4C-1), ca. 0.02 mm long, and four interspiracular processes (Fig. 4C- 2), each with one or two, lanceolate branches, each with one or two apical teeth, longest branch measuring 0.01 mm; stelex sensillum ventrolaterad (Figs. 4C-3, D-1) of posterior spiracular plate, dorsolateral stelex sensillum observed, but not shown; no other sensillum seen at lateral position; intermediate sensory complexes (Fig. 4D-2) with a stelex sensil- lum (Fig. 4D-3) and a medusoid sensillum (Fig. 4D-4). The habitus of the second instar of T: foot- ei (Fig. 4A) approximates those of 7. bac- charis (Goeden and Headrick 1991), T. ari- zonaensis (Goeden et al. 1993), 7. joanae (Goeden 2001b), and 7. teerinki (Goeden 2001c). Again, the partial description for 7. footei allowed limited comparison with the second instars of 7. joanae (Goeden 2001b) and T. teerinki (Goeden 2001c), the only spe- cies described in sufficient detail to date to allow full comparison of characters. The an- terior spiracle of second instar 7. footei bears five, subglobose or subquadrate papillae (Fig. 4B), like that of 7. joanae (Goeden 2001b), not four, doliform papillae like 7. feerinki (Goeden 2001c). Only minor differences in 152 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 4. segment, 1—rimae, 2—interspiracular processes, 3 Second instar of Tephritis footei: (A) habitus, anterior to right; (B) anterior spiracle; (C) caudal ventrolateral stelex sensillum; (D) 1—ventrolateral stelex sensillum, 2—intermediate sensory complex, with 3—stelex sensillum and 4—medusoid sensillum. the number of branches on the interspiracular processes, 1.e., one to two for 7. footei (Fig. 4C-2), one to three for 7. teerinki (Goeden 2001c), versus two to four for T. joanae (Goeden 2001b), were noted. Also, the branches of the interspiracular processes of 7. footei are lanceolate (Fig. 4C-2), not foliose, like those of the second instars of 7. joanae (Goeden 2001b) and T. teerinki (Goeden 200 1c). Third instar larva: White, ellipsoidal, dis- tinctly segmented, tapered anteriorly, truncat- ed posteriorly (Fig. 5A); gnathocephalon con- ical, anteriorly flattened, and medially divided by a vertical suture (Fig. 5B-1); posteriorly directed, spinose, minute acanthae incom- pletely circumscribe prothorax anteriorly (Figs. 5B-2, C-12), instead many round to elongate-rounded, integumental petals cir- cumscribe central half of prothorax (Fig. 5B- 3); dorsal sensory organ well-defined, hemi- spherical (Fig. 5C-1); anterior sensory lobe (Fig. SC-2) bears terminal sensory organ (Fig. 5C-3), lateral sensory organ (Fig. 5C- 4), supralateral sensory organ (Fig. 5C-5), and pit sensory organ (Fig. 5C-6); two me- dial, papilliform integumental petals (Fig. 5C- 7) and two, lateral, spatulate, integumental petals (Fig. 5C-8) in two rows above each mouthhook (Fig. 5C-9), lower lateral petal continuing laterally around oral cavity, and separate from stomal sense organ (Fig. 5C- 10) ventrolaterad of anterior sensory lobe; at least three complete oral ridges (Fig. SC-11) laterad of each anterior sensory lobe; mouth- hook apparently bidentate; median oral lobe Nn eS) VOLUME 104, NUMBER | l Fig. 5. Third instar of Tephritis footei: (A) habitus, anterior to left; (B) gnathocephalon and prothorax, frontolateral view, |—vertical medial suture of gnathocephalon, 2—minute acanthae, 3—integumental petals on prothorax, 4—anterior spiracle; (C) gnathocephalon, frontolateral view, 1—dorsal sensory organ, 2—anterior sensory lobe, 3—terminal sensory organ, 4—lateral sensory organ, 5—supralateral sensory organ, 6—pit sensory organ, 7—medial integumental petal, 8—lateral integumental petal, 9—mouthhook, 10—stomal sense organ, 1 1—oral ridges, 12—minute acanthae; (D) anterior spiracle; (E) caudal segment, 1—spiracular plates, 2—dor- solateral stelex sensillum, 3—ventrolateral stelex sensillum; 4—intermediate sensory complex, with 5—stelex sensilla and 6—medusoid sensillum; (F) posterior spiracular plate, 1—rimae, 2—interspiracular processes. 154 present, but only partially seen and not pic- tured: anterior thoracic spiracle on posterior margin of prothorax bears three (not shown) or four subglobose or subquadrate papillae (Figs. 5B-4, 5D); mesothoracic, metathoracic, and abdominal lateral spiracular complexes not seen; each posterior spiracular plate (Fig. 5E-1) surrounded by a pair of dorsolateral stelex sensilla (Fig. 5E-2) and ventrolateral pair of stelex sensilla (Fig. 5E-3); each pos- terior spiracular plate bears three ovoid rimae (Fig. 5F-1), ca. 0.02 mm in length, and four, three- to four-branched, single-, bi- or trifur- cately-tipped, interspiracular processes, each ca. 0.05 mm long (Fig. 5F-2); intermediate sensory complex (Fig. 5E-4) with a stelex sensillum (Fig. SE-5) and a medusoid sensil- lum (Fig. 5E-6). The habitus of the third instar of 7. footei differs from those of four other described congeners in at least two ways. The elongate- ellipsoidal shape of the third instar of 7. foot- ei (Fig. 5A) appears intermediate to the ovoi- dal shape of the third instars of 7. joanae (Goeden 2001b) and T. teerinki (Goeden 2001c) and the cylindrical shape ascribed to third instars of 7. baccharis (Goeden and Headrick 1991) and 7. arizonaensis (Goeden et al. 1993). The prothorax (Fig. 5B-1) is cir- cumscribed by many more integumental pet- als than the much smoother prothoracic seg- ments of the four other species of Tephritis examined to date (Goeden and Headrick 1991; Goeden et al. 1993; Goeden 2001b, c). Fewer minute acanthae anteriorly circum- scribe the prothorax of the third instar of 7. footei (Fig. 5B-2) than those of 7. baccharis (Goeden and Headrick 1991), 7. arizonaensis (Goeden et al. 1993), 7. joanae (Goeden 2001b), and T. teerinki (Goeden 2001c). On the other hand the gnathocephalon, or at least the anterior sensory lobes of all five species are separated by a vertical medial suture (Goeden and Headrick 1991; Goeden et al. 1993; Goeden 2001b, c; Fig. 5B). The integumental petals in the third instars of all five congeners examined to date are arranged in a double row above each mouth- hook, but those of 7. footei (Fig. 5C-8) occur PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON in two rows of two like 7. teerinki (Goeden 2001c), with an additional, medial pair of dif- ferent shape in 7. footei (Fig. SC-7). How- ever, the integumental petals of both 7. teer- inki (Goeden 2001c) and T. footei (Figs. SC- 7, 8) are fewer in number than those of 7: baccharis (Goeden and Headrick 1991), 7: arizonaensis (Goeden et al. 1993), and T. joanae (Goeden 2001b). Both 7. footei and T. teerinki also apparently lack the additional integumental petals found in a vertical double row above these papillae, one pair to each side of the medial depression separating the anterior sensory lobes (Fig. 5C-2; Goeden and Headrick 1991; Goeden et al. 1993; Goe- den 2001b, c). The integumental petals in- crease in number between the last two instars of T. footei (Figs. 5C-7, 8), T. joanae (Goe- den 2001b), and T. teerinki (Goeden 200Ic). The mouthhooks of the third instars of T- footei (Fig. 5C-9), like those of 7. teerinki (Goeden 2001c), appear bidentate, unlike the tridentate mouthhooks of 7. baccharis (Goe- den and Headrick 1991), 7. arizonaensis (Goeden et al. 1993), and T. joanae (Goeden 2001b). However, most of the nine specimens of third instar 7. footei examined by SEM had their mouthparts hidden, which precluded examination of the oral cavity and mouth- hooks in ventral view. The anterior spiracle of the third instar of T. footei (Fig. 5D) bore three or four papillae, one or two less than the second instar, but not three in the second instar and three or four in the third instar, like 7. joanae (Goeden 2001b), or four in the second instar and five in the third instar as reported for 7. teerinki (Goeden 200Ic). Goeden (2001c) discussed the stelex sen- silla surrounding the posterior spiracular plates of the third instars that apparently dif- fer in number among the Tephritis species examined to date. However, the caudal seg- ment of the third instar of 7. footei, like those of T. joanae (Goeden 2001b) and T. teerinki (Goeden 2001c), apparently are surrounded by a dorsolateral and ventrolateral pair of ste- lex sensilla (Figs. 5SE-2, 3) as well as a ventral pair of intermediate spiracular complexes VOLUME 104, NUMBER 1 (Fig. 5E-4), the medusoid sensilla of each bears short apical papillae typical of this type of chemosensillum (Goeden 2001a, b, c; Goeden and Teerink 1999, and references therein). Puparia: All puparia prepared for SEM unfortunately turned out to be Campiglossa clathrata (Loew), which imperfectly, tempo- rally partitions and thus shares the flower heads of A. tridentata with T. footei, 1.e., symphagy (Goeden 1997). Other puparia ex- amined, identified, and measured in situ pro- vided the basis for the following abbreviated description. Dull black, ellipsoidal, and smoothly rounded at both ends. Thirty-four puparia av- eraged 1.87 + 0.02 (range, 1.63—2.13) mm in length; 0.98 + 0.013 (range, 0.85—1.14) mm in width. DISTRIBUTION AND Hosts To date, Tephritis footei is only known from California and from flower heads of Ar- temisia tridentata; however, it has been re- ported elsewhere by me (Goeden 1993), and probably by others, as 7. ovatipennis, and this should be corrected. Accordingly, 7. footei, possibly may be a nearly monophagous, widely distributed species on A. tridentata, which itself is widely distributed throughout the western United States (Hickman 1993), and belongs to the subtribe Artemisiinae of the Tribe Anthemideae (Bremer 1994). This monophagy is presumed because 7. araneo- sa, aS now constituted, has been reared from flower heads of four other California species of Artemisia, 1.e., A. californica Lessing, A. douglasiana Besser, A. dracunculus L., and A. ludoviciana Nuttall, the last-named plant formerly was reported as a host of 7. ovati- pennis by Goeden (1993), which now should be deleted; whereas, A. dracunculus is a new host record for 7. araneosa (Goeden 1993). As noted above, 7. araneosa, or what is prob- ably a separate new species near araneosa, also has been reared by me from Chryso- thamnus nauseosus (Pallas) Britton, C. parryi (A. Gray) E. Greene, C. teretifolius (Durand and Hilgard) H. M. Hall, C. viscidiflorus 155 (Hooker) Nuttall, and Ericameria (as Hap- lopappus) bloomeri (A. Gray) J. EF Mcbride, most of which were reported as hosts of 7. araneosa by Goeden (1993), except for this newly confirmed host record for C. nauseo- sus in Wasbauer (1972). Chrysothamnus and Ericameria both belong to the subtribe Soli- dagininae of the tribe Astereae (Bremer 1994). Also as noted above, 7. headricki is an oligophagous tephritid, reared solely from the flower heads of two species of Solidago, 1.e., S. canadensis and S. confinis and Eu- thamia occidentalis, all three of which belong to the subtribe Solidagininae of the tribe As- tereae (Bremer 1994). The first two plant spe- cies were reported as hosts of 7. ovatipennis by Goeden (1993), which now need to be deleted, and the latter plant represents a new host-plant genus and species record for Te- phritis. Tephritis headricki undoubtedly has additional hosts as yet undiscovered, and probably is a widespread species like its widespread and diverse host-plant genus, Sol- idago (Hickman 1993). Goeden (2001c) also removed Hulsea spp. from those hosts re- ported by Goeden (1993) as T. ovatipennis, when he described 7. teerinki. The three specimens identified and reported by Goeden (1993) for T. ovatipennis reared from Ma- chaeranthera canescens (Pursh) A. Gray, upon re-examination were re-identified and confirmed with other reared specimens as 7. michiganesis Quisenberry; this finding ex- tends its distribution across the western Unit- ed States to southern California from western Minnesota/eastern North Dakota, and repre- sents the first host-plant record for this te- phritid (Foote et al. 1993). Therefore, as pres- ently constituted, 7. ovatipennis represents an oligophagous species reared from two species of Erigeron, 1.e., E. foliosus Nuttall and E. glaucus Ker-Gawler, and Trimorpha loncho- phyllus (Hooker) G. Nesom, all of which be- long to the subtribe Asterinae of the tribe As- tereae. Of the last three species, the first named is a new host-plant record; the second and third species were reported as hosts of 7. ovatipennis by Goeden (1993), the third also as another species of Erigeron (Hickman, 156 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1993). Recent revisions by plant and insect taxonomists (Hickman 1993; Bremer 1994; Goeden 2001a, b) have helped to clarify and better interpret the host affinities of these Cal- ifornia Tephritis since Goeden (1993), but continued study is needed of these and other Tephritis belonging or not to the “araneosa complex.” BIOLOGY Egg.—In each of 11, closed, preblossom, immature flower heads of Artemisia triden- tata a single egg of T. footei was inserted pedicel-last through one or more phyllaries into an ovule (Fig. 6A). Thus, the 11 eggs each rested with their long axes at a 45° to 60° angle to the receptacles of the young flower heads and an average of one ovule/ floret was damaged per flower head by ovi- position. Thus, about 25% of an average total of 4.3 + 0.2 (range, 3-7) ovules/florets counted in 27 infested, closed preblossom flower heads were damaged by oviposition. The receptacles of preblossom heads. that contained eggs averaged 0.36 + 0.02 (range, 0.28—0.4) mm in diameter. Larva.—Upon eclosion, the single first in- stars found feeding in seven, closed, preblos- som flower heads tunneled immediately into a floret and then continued to feed parallel to the receptacle on one or more ovules (Fig. 6B). The receptacles of these seven infested flower heads averaged 0.39 + 0.02 (range, 0.28—0.46) mm in diameter and an average of 1.8 + 0.49 (range, 1-3) ovules was dam- aged in these seven flower heads. No recep- tacle was abraded or pitted by larval feeding. Thus, about 42% (range, 25-70%) of an av- erage total of 4.3 ovules/florets counted for the above-mentioned, 27 flower heads were damaged by the first instars. Second instars (Fig. 6C) continued feeding on ovules in closed, preblossom flower heads. They fed with their bodies perpendicular to the receptacles, but always well above the re- ceptacles (Fig. 6C). Receptacles of 25 flower heads containing second instars averaged 4.30 + 0.22 (range, 2.28—6.84) mm in di- ameter. These eight flower heads each con- tained a single larva that had damaged a sin- gle ovule, or again, about 25% of the average total of 4.3 ovules/florets per flower head counted within the above-mentioned, 27 flower heads. Third instars in flower heads fed with their long axes oriented perpendicular to the re- ceptacles, and with their mouthparts com- monly directed towards (Fig. 6D), or less commonly, away from the receptacles (Fig. 6E). All of the ovules/florets in each of the heads were destroyed (Figs. 6D, E). The re- ceptacles were abraded or pitted in two (64%) of nine, closed flower heads containing third instars, or the larvae were found feeding on the basal fragments of the ovules connected to the receptacles, which suggested that sap constituted at least part of the diet of third instars of T. footei, probably towards the end of the third stadium. Goeden (1988b), Head- rick and Goeden (1990), Goeden and Head- rick, (1992), Goeden et al. .’'993,199s)) Headrick et al. (1996), Goeden and Teerink (1997) first noted, described, and discussed sap feeding by florivorous species of Tephri- tidae in the genera Trupanea, Paracantha, Neaspilota, Tephritis, Urophora, Dioxyna, and Xenochaeta, respectively. Upon complet- ing feeding, the larvae oriented with their an- terior ends away from the receptacles, retract- ed their mouthparts, and formed puparia (Fig. 6F). Pupa.—The receptacles of 43 flower heads, each of which contained a single pu- parium (Fig. 6F) averaged 0.6 + 0.04 (range, 0.42—1.42) mm in diameter. The receptacles were abraded or pitted in 19 of 43 (44%) flower heads containing puparia, further con- firming that sap constituted part of the diet of the third instars. All of the contents of these flower heads were destroyed and the puparia occupied most of the flower heads (Fig. 6F). Adult.—Adults (Fig. 6G, H) apparently are long-lived and constitute the only overwin- tering stage in southern California. Under in- sectary conditions, six unmated females (Fig. 6G) lived an average of 87 + 8 (range, 71— 120) days, and five virgin males (Fig. 6H) averaged 64 + 12 (range, 18-91) days. Such VOLUME 104, NUMBER 1 57 Fig. 6. Life stages of Tephritis footei in flower heads of Artemisia tridentata: (A) egg (arrow) in closed, preblossom flower head inserted into floret; (B) first instar (arrow) feeding within floret in preblossom flower head; (C) second instar (arrow) feeding in adjacent florets in preblossom flower head: (D) third instar feeding on receptacle in closed, preblossom flower head; (E) full-size, third instar positioned for pupariation; (F) single puparium occupying interior of flower head; (G) adult female; (H) adult male; (I) mating pair with female forming droplet. Lines = 1 mm. 158 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON lengthy longevities are commensurate with the aggregative type of life history possessed by this tephritid (Headrick and Goeden 1994, 1998). The premating and mating behaviors of 7. footei were not studied in the field, but were limitedly observed in petri dish arenas of the type otherwise found to be so useful with many other nonfrugivorous, tephritid species (Headrick and Goeden 1994). Premating be- haviors occasionally observed with T. footei were tracking and side-stepping by males (Headrick and Goeden 1994). The most com- mon wing display was asynchronous supi- nation by both sexes, both spontaneous and in response to other individuals (Goeden et al. 1993; Headrick and Goeden 1994, 1999). Males did not exhibit any of the common tephritid courtship displays, including regular, abdominal pleural distension. Four pairs were observed to mate (Fig. 61) once or twice per day for a total of 10 matings that lasted an average of 208 (range, 84-316) min. Copu- latory induction behavior (CIB) (Headrick and Goeden 1994, 1999), and the copulatory positions attained by each sex, generally were as described for 7. arizonaensis (Goeden et al. 1993). Separation of a male and female was observed three times, with the male turn- ing and rapidly walking off and away from the female while pulling free his genitalia. Seasonal history.—The life cycle of T. footet on Artemisia tridentata in southern California follows an aggregative pattern (Headrick and Goeden 1994, 1998) in which the adult is the principal overwintering stage. Consequently, come late-spring/early summer (May to June), adults aggregate on preblos- som shoots and subsequently oviposit in the small, newly-formed, closed, preblossom flower heads. The larvae feed until fully grown, then pupariate in flower heads and emerge in midsummer (late-June to July). They spend the rest of the summer and fall, probably as non-reproductive adults, feeding in mountain meadows and riparian habits where they subsequently overwinter. Or, pos- sibly two overlapping, nondescrete genera- tions are produced starting earlier on lower- elevation, or later on higher-elevation, host plants. Natural enemies.—Halticoptera sp. and Mesopolobus sp. (Hymenoptera: Pteromali- dae) were reared from separate puparia of 7. footei as solitary, larval-pupal endoparasit- oids. Many additional specimens of both spe- cies were reared from caged mature flower heads of A. tridentata as probable parasitoids. ACKNOWLEDGMENTS I thank Andrew C. Sanders, Curator of the Herbarium, Department of Botany and Plant Sciences, University of California, Riverside, for identifications of plants mentioned in this paper. Krassimer Bozhiloy in the Institute of Geophysics and Planetary Physics, Univer- sity of California, Riverside, greatly facilitat- ed my scanning electron microscopy. Harry Andersen, now deceased, identified the par- asitoids. I also am grateful to Allen Norrbom and Gary Steck for their taxonomic help, to Jeff Teerink for his technical assistance, and to David Headrick and Jeff Teerink for their helpful comments on earlier drafts of this pa- per. This paper, like the name of the principal tephritid treated, is dedicated to Dick Foote, in special recognition and thanks. LITERATURE CITED Bremer, K. 1994. Asteraceae Cladistics & Classification. Timber Press, Inc. Portland, Oregon. Foote, R. H. 1960. The genus Tephritis Latreille in the Nearctic Region north of Mexico: Descriptions of four new species and notes on others. Journal of the Kansas Entomological Society 33: 71—85. Foote, R. H. and FE L. Blanc. 1979. New species of Te- phritidae (Diptera) from the western United States, Mexico, and Guatemala, with revisionary notes. Pan- Pacific Entomologist 55: 161—179. Foote, R. H., E L. Blanc, and A. L. Norrbom. 1993. Handbook of the Fruit Flies (Diptera: Tephritidae) of America North of Mexico. Cornell University Press, Ithaca, New York. Goeden, R. D. 1988a. Gall formation by the capitulum- infesting fruitfly, Tephritis stigmatica (Diptera: Te- phritidae). Proceedings of the Entomological Society of Washington 90: 37—43. . 1988b. Life history of Trupanea imperfecta (Co- quillett) on Bebbia juncea (Bentham) Greene in the Colorado Desert of southern California (Diptera: Te- phritidae). Pan-Pacific Entomologist 64: 345-351. VOLUME 104, NUMBER | . 1993. Analysis of known and new host records for Tephritis from California, and description of a new species, 7. joanae (Diptera: Tephritidae). Pro- ceedings of the Entomological Society of Washing- ton 95: 425—434. . 1997. Symphagy among florivorous fruit flies (Diptera: Tephritidae) in southern California. Chap. 3. In Dettner, K., G. Bauer, and W. Volkl, eds. Ver- tical Food Web Interactions: Evolutionary Patterns and Driving Forces. Ecological Studies 130: 39-52. Springer-Verlag, Heidelberg, Germany. . 2001a. Life history and description on immature stages of Neaspilota footei Freidberg and Mathis (Diptera: Tephritidae) on Aster occidentalis (Nuttall) Torrey and A. Gray (Asteraceae) in southern Cali- fornia. Proceedings of the Entomological Society of Washington 103: 191—206. . 2001b. Life history and description on immature stages of Tephritis joanae Goeden (Diptera: Tephri- tidae) on Ericameria pinifolia (A. Gray) H. M. Hall (Asteraceae) in southern California. Proceedings of the Entomological Society of Washington 103: 586— 600. . 2001c. Life history and description on immature stages of Tephritis teerinki Goeden (Diptera: Tephri- tidae) on Hulsea vestita A. Gray (Asteraceae) in southern California. Proceedings of the Entomolog- ical Society of Washington 103: 807-825. Goeden, R. D. and R. K. Benjamin. 1985. New records for Stigmatomyces verruculosus Thaxter (Ascomy- cetes: Laboulbeniaceae), a fungal parasite of adult Tephritidae in southern California. Proceedings of the Entomological Society of Washington 87: 392— 394. Goeden, R. D. and D. H. Headrick. 1991. Life history and descriptions of immature stages of Tephritis bac- charis (Coquillett) on Baccharis salicifolia (Ruiz & Parvon) Persoon in southern California (Diptera: Te- phritidae). Pan-Pacific Entomologist 67: 86-98. . 1992. Life history and descriptions of immature stages of Neaspilota viridescens Quisenberry (Dip- tera: Tephritidae) on native Asteraceae in southern California. Proceedings of the Entomological Society of Washington 94: 59-77. Goeden, R. D., D. H. Headrick, and J. A. Teerink. 1993. Life history and descriptions of immature stages of Tephritis arizonaensis Quisenberry (Diptera: Tephri- tidae) on Baccharis sarothroides Gray in southern California. Proceedings of the Entomological Society of Washington 95: 210-222. . 1995. Life history and description of immature stages of Urophora timberlakei Blanc and Foote (Diptera: Tephritidae) on native Asteraceae in south- 59 ern California. Proceedings of the Entomological So- ciety of Washington 97: 779-790. Goeden, R. D. and J. A. Teerink. 1997. Life history and description of immature stages of Xenochaeta albi- florum Hooker in central and southern California. Proceedings of the Entomological Society of Wash- ington 99: 597-607. . 1999. Life history and description of immature stages of Trupanea vicina (Wulp) (Diptera: Tephri- tidae) on wild and cultivated Asteraceae in southern California. Proceedings of the Entomological Society of Washington 101: 742-755. Headrick, D. H. and R. D. Goeden. 1990. Resource uti- lization by larvae of Paracantha gentilis (Diptera: Tephritidae) in capitula of Cirsium californicum and C. proteanum (Asteraceae) in southern California. Proceedings of the Entomological Society of Wash- ington 92: 512-520. . 1994. Reproductive behavior of California fruit flies and the classification and evolution of Tephri- tidae (Diptera) mating systems. Studia Dipterologica 1(2): 194-252. . 1998. The biology of nonfrugivous tephritid fruit flies. Annual Review of Entomology 43: 217-241. . 1999. Behavior of flies in the subfamily Te- phritinae, pp. 671-707. In Aluja, M. and A. L. Norr- bom, eds. Fruit Flies (Tephritidae): Phylogeny and Evolution of Behavior. CRC Press, Boca Raton, London, New York, Washington, D.C. Headrick, D. H., R. D. Goeden, and J. A. Teerink. 1996. Life history and description of immature stages of Dioxyna picciola (Bigot) (Diptera: Tephritidae) on Coreopsis spp. (Asteraceae) in southern California. Proceedings of the Entomological Society of Wash- ington 98: 332-349. Hickman, J. C. (ed.). 1993. The Jepson Manual. Univer- sity of California Press. Berkeley and Los Angeles. Jenkins, J. and W. J. Turner. 1989. Revision of the Bac- charis-infesting (Asteraceae) fruit flies of the genus Tephritis (Diptera: Tephritidae) in North America. Annals of the Entomological Society of America 82: 674-685. Tauber, M. J. and C. A. Toschi. 1965. Life history and mating behavior Tephritis stigmatica (Coquillett) (Diptera: Tephritidae). Pan-Pacific Entomologist 41: 73-79. Teerink, J. A. and R. D. Goeden. 1999. Description of the immature stages of Trupanea imperfecta (Co- quillett). Proceedings of the Entomological Society of Washington 101: 75-85. Wasbauer, M. C. 1972. An annotated host catalog of the fruit flies of America north of Mexico (Diptera: Te- phritidae). California Department of Agriculture, Oc- casional Paper 19: 134. PROC. ENTOMOL. SOC. WASH. 104(1), 2002, pp. 160-163 GEOICA SETULOSA (PASSERINI) (HEMIPTERA: APHIDIDAE): NEW DISTRIBUTION RECORDS FOR NORTH AMERICA GARY L. MILLER, MANYA B. STOETZEL, ROLANDO LOPEZ, AND DANIEL A. POTTER (GLM, MBS) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, Bldt. 005, BARC-West, Beltsville, MD 20705, U.S.A. (e-mail: gmiller@sel.barc.usda.gov; mstoetze @sel.barc.usda.gov); (RL, DAP) Department of Entomology, University of Kentucky, Lexington, KY 40546 U.S.A. (e-mail: rlopez@ pop.uky.edu; dapotter@ca.uky.edu) Abstract.—We report the first eastern United States records of the fordine aphid Geoica setulosa (Passerini). Description and illustrations of the apterous viviparous female and a key to species of Geoica in the United States are provided. Key Words: The genus Geoica (Hemiptera: Aphidi- dae) is comprised of at least 14 species worldwide with the possibility of Geoica utricularia (Passerini) representing a com- plex of species (Eastop and Hille Ris Lamb- ers 1976, Remaudiére and Remaudiére 1997). Geoica is considered an Old World genus (Blackman and Eastop 2000) and, until recently, four species of Geoica were recognized from North America: Geoica lu- cifuga (Zehntner), Geoica pellucida (Buck- ton), Geoica squamosa Hart, and Geoica utricularia (Passerini) (Smith and Parron 1978). However, Geoica pellucida (Buck- ton) and Geoica squamosa Hart are now both considered synonyms of Geoica utri- cularia (Passerini) (Remaudiere and Re- maudiere 1997). Certain members of this genus exhibit a complex, 2-year life cycle with alternation between galls on Pistacia, pistachio (Anacardiaceae), and migration to and from their secondary host where pop- ulations feed on the roots of Gramineae and occasionally Cyperaceae (Brown and Blackman 1994, Blackman and Eastop 2000). Geoica setulosa (Passerini) has been re- Aphidoidea, North America, key, turfgrass corded from northwestern and central Eu- rope, Italy, Iran, Turkey, and has recently been reported as occurring in the United States from Idaho (Blackman and Eastop 2000). Recent collections of G. setulosa from Kentucky indicate that it has a much greater distribution than its first collection from Idaho would indicate. In Iran, G. se- tulosa is holocyclic with its primary host, Pistacia khinjuk Stocks. In Europe, it is ex- clusively anholocyclic on grass roots (Blackman and Eastop 2000). Collections of G. setulosa in the United States have been made from roots of grasses and En- glish plantain. MATERIALS AND METHODS Synoptic descriptions are based on pub- lished descriptions as well as material from the Aphididae portion of the National Col- lection of Insects (USNM), Beltsville, Maryland. In Specimens Examined, apter- ous adults are abbreviated as “‘ap. ad.”’ For specimens collected at the same locality, on the same date, and from the same host plant as previously listed, the duplicate informa- tion is not repeated. Unless otherwise not- VOLUME 104, NUMBER 1 161 Figs. 1-5. ed, voucher material consists of a single slide (sl.). Specimens were measured using a Nikon Eclipse E600® compound micro- scope. Measurements are presented in mil- limeters (mm) as minimum and maximum ranges of representative specimens. Geoica setulosa (Passerin1) (Figs. 1—5) Tychea setulosa Passerini 1860: 40. Tycheoides setulosa: Theobald 1916: 52. Geoica setulosa: Theobald 1929: 197. Geoica herculana Mordvilko 1935: 215. Geoica setulosa: Eastop and Hille Ris Lambers 1976: 204. Geoica setulosa: Heie 1980: 198. Geoica setulosa: Remaudiere and Remau- diére 1997: 240. Geoica setulosa. 1, Dorsal (left) and ventral (right) aspects apterous adult female head and an- tennae. 2, Variations of body setae. 3, Ultimate rostral segment. 4, Acuminate ventral abdominal seta with polygonal reticulate pattern. 5, Anal plate region. Field characters.—Coloration of apterous viviparous females range from “‘... dirty whitish or greenish grey ...” (Heie 1980) to “*... yellowish-white to greyish-white, some creamy > (hcobaldhl929) soe antennae dusky, apices darkened; legs brown; rostrum dark at apex ...” (Theo- bald 1929). In specimens preserved in EtOH the head, prothorax, legs, antennae, and rostrum are crineous, the last segment of the rostrum is fuscous, and the base of most body setae is delineated by slight am- ber markings. Body form is globose. Recognition characters from. slide- mounted specimens.—Apterous viviparous female (Figs. 1—5): Body ovoid without wax gland plates, length 1.500—2.136 mm; width through eyes, 0.279—0.409 mm. Eye 162 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON with three ommatidia. Antenna 5-segment- ed (Fig. 1), shorter than body, slightly dark- er than rest of body, without secondary sen- soria, primary sensoria elongate and evert- ed; length of segment: I, 0.062—0.086 mm; II, 0.062—0.081 mm; III, 0.093—0.123 mm; IV, 0.062—0.068 mm; base of V, 0.056-— 0.074 mm; terminal process of V, 0.019— 0.031 mm. Setae on body and appendages variable, ranging from acuminate, multifid, capitate, to flabellate (Fig. 2), with dense random placement on dorsum. Rostrum ex- tending to metacoxae; length of ultimate segment (Fig. 3), 0.136—0.167 mm, with 8— 10 accessory setae, shorter than hind tarsal segment II. Length of metafemur, 0.223— 0.279 mm; metatibia, 0.217—0.310 mm; metatarsus II, 0.087—0.093 mm. Abdominal venter with polygonal reticulate pattern (Fig. 4). Cornicles absent. Anus positioned dorsally, anal plate rectangular with numer- ous small setae dorsally and a median row of paired setae (Fig. 5). Biology.—Geoica setulosa is anholo- cyclic in Northern and Central Europe with alatae being extremely rare or absent (Heie 1980). It feeds on the roots of grass species such as Agrostis, Alopecurus, Corynepho- rus, Festuca, and Holcus (Heie 1980) and has also been recorded from Setaria sp., Oryza sativa L., and Triticum vulgare Vill. [= Triticum aestivum L.| (Theobald 1929). It was reported in Idaho (Blackman and Eastop 2000) where it was collected on Panicum capillare L., Poa pratensis L., Hordeum jubatum L., Echinochloa crus- galli (L.) Beauvois, Phalaris arundinaceae L., Avena fatua L., Setaria lutescens (Wei- gel) EK T. Hubbard [= Pennisetum glaucum (L.) R.Br.], and “‘?crabgrass”’ (Halbert, per- sonal communication). Geoica setulosa has also been associated with ants including Lasius niger (L.) (Theobald 1929) and Las- ius flavus (FE) (Heie 1980, Blackman and Eastop 2000). Geoica setulosa collected in Kentucky always were found associated with the ant Lasius neoniger Emery on the roots of grass. Alatoid nymphs were col- lected in the population of G. setulosa and, while they could not be identified with cer- tainty to species, we can assume that winged adults of G. setulosa may occur in Kentucky. Geoica setulosa is probably also anholocyclic in the United States. Specimens examined.—U.S.A.: Ken- tucky, Lexington, University of Kentucky lawn #1, on Kentucky bluegrass [Poa pra- tensis|, R. Lopez collector, USNM: V-4- 1999 (20 ap. ad. 2 on 6 sl.); Spindletop Research Farm, on Kentucky bluegrass [Poa pratensis], R. Lopez collector, USNM: V-24-1999 (1 ap. ad. 2); Spindletop Re- search Farm, on Falcon II tall fescue [Fes- tuca arundinarea Schreber], R. L6pez col- lector, USNM: VI-19-1999 (16 ap. ad. 2 on 6 sl.); on Aurora hard fescue [Festuca lon- gifolia Thuill], R. Lopez collector, USNM: VI-19-1999 (14 ap. ad. @ on 5 sl.); on Adelphi Kentucky bluegrass [Poa pratensis L. adelphi|, R. Lopez collector, USNM: VI- 19-1999 (1 ap. ad. ¢); on Palmer II peren- nial ryegrass [Lolium perenne L.], R. Lopez collector, USNM: VI-19-1999 (4 ap. ad. 2 on 3 sl.); on Kentucky bluegrass [Poa pra- tensis|, L., R. L6pez collector, USNM: VI- 25-1999 (1 ap. ad. 2); on KY 31 tall fescue w/endophyte, R. Lopez collector, USNM: VII-02-1999 (2 ap. ad. @). KEY TO APTERAE OF NORTH AMERICAN GEOICA (Modified from Blackman and Eastop 2000) 1. Anal plate with numerous subequal setae ran- domly dispersed ..... Geoica utricularia group — Anal plate with median row of paired, long se- tae in addition to shorter dorsal setae (Fig. 5) tO . Dorsal body setae randomly placed: primary sensorium elongate; last rostral segment 0.136— 0.167 mm long . Geoica setulosa (Passerini) — Dorsal body setae forming transverse rows; primary sensorium subcircular; last rostral seg- amare OI TO =0)-2400) taavan ION Soc cose ccu soe Geoica lucifuga (Zehntner) DISCUSSION Although members of Geoica are not considered economically important (Foottit and Richards 1993), their association with VOLUME 104, NUMBER 1 grass and aphid-tending ants can be prob- lematic. This is especially true where close- ly-mowed turfgrass is managed on golf courses. Associated ants, e.g., L. neoniger, tend to build mounds around the plants that serve as hosts for Geoica. These ant mounds are not only unsightly but they smother the grass, dull mower blades, and disrupt the smoothness and uniformity of playing surfaces, including tees, fairways, and putting greens (L6pez and Potter 2000). ACKNOWLEDGMENTS We thank Susan E. Halbert, Florida De- partment of Agriculture, Gainesville, FL, for her collection records of Geoica setu- losa from Idaho. We also thank Andrew S. Jensen, Moses Lake, WA, and Eric Grissell, USDA, ARS, Systematic Entomology Lab- oratory, Washington, DC, for their reviews and comments on the manuscript. Collec- tions of Geoica in Kentucky were support- ed in part by a grant from the U.S. Golf Association to D. A. Potter. LITERATURE CITED Blackman, R. L. and V. EK Eastop. 2000. Aphids on the World’s Crops: An Identification and Infor- mation Guide, second edition. John Wiley & Sons, Ltd., Chichester, 466 pp. Brown, P. A. and R. L. Blackman. 1994. Morphometric variation in the Geoica utricularia (Homoptera: Aphididae) species group on Pistacia (Anacardi- aceae), with descriptions of new species and a key 163 to emigrant alatae. Systematic Entomology 19: 119-132. Eastop, V. EK and D. Hille Ris Lambers. 1976. Survey of the World’s Aphids. Dr. W. Junk, Publishers, The Hague, 573 pp. Foottit, R. G. and W. R. Richards. 1993. The genera of the Aphids of Canada. Homoptera: Aphidoidea and Phylloxeroidea. Part 22 of the Insects and Arachnids of Canada. Centre for Land and Bio- logical Resources Research, Publication 1885, Ot- tawa, Ontario, 765 pp. Heie, O. E. 1980. The Aphidoidea (Hemiptera) of Fen- noscandia and Denmark. I. General part. The Families Mindaridae, Hormaphididae, Thelaxidae, Anoeciidae, and Pemphigidae. Fauna Entomolo- gica Scandinavica 9, 236 pp. L6pez, R. and D. A. Potter. 2000. Ant predation on eggs and larvae of the black cutworm (Lepidop- tera: Noctuidae) and the Japanese beetle (Cole- optera: Scarabaeidae) in turfgrass. Environmental Entomology 29: 116-125. Mordvilko, A. K. 1935. Die Blattlause mit unvollstan- digem Generationszyklus und ihre Entstehung. Er- gebnisse und Fortschritte der Zoologie 8: 36-328. Passerini, G. 1860. Gli Afidi con un prospetto dei ge- neri ed alcune specie nuove Italiane. Parma, 46 pp: Remaudiere, G. and M. Remaudiere. 1997. Catalogue of the World’s Aphididae Homoptera Aphidoidea. Institut National de la Recherche Agronomique, Paris, 474 pp. Smith, C. F and C. S. Parron. 1978. An Annotated List of Aphididae (Homoptera) of North America. North Carolina Agricultural Experiment Station Technology Bulletin 255, 428 pp. Theobald, E V. 1916. Notes on Aphididae found in ants’ nests. The Entomologist 49: 49—52. Theobald, E V. 1929. The Plant Lice or Aphididae of Great Britain, Vol. III. Headley Brothers, London, 364 pp. PROC. ENTOMOL. SOC. WASH. 104(1), 2002, pp. 164-167 COMPARATIVE ECOLOGY OF THE GENUS LECONTELLA WOLCOTT AND CHAPIN (COLEOPTERA: CLERIDAE: TILLINAE), WITH NOTES ON CHEMICALLY DEFENDED SPECIES OF THE BEETLE FAMILY CLERIDAE JONATHAN R. MAWDSLEY Department of Entomology, National Museum of Natural History, Smithsonian Insti- tution, Washington, DC 20560-0187, U.S.A. (e-mail: mawdsley.jonathan@nmnh. si.edu) Abstract.—The genus Lecontella Wolcott and Chapin currently contains two species, L. brunnea (Spinola) from eastern North America and L. gnara Wolcott from southwestern North America. The results of the author’s laboratory and field studies of L. gnara Wolcott are summarized, and this species’ biology compared to that of L. brunnea. Larvae of L. brunnea are parasites in nests of solitary bees and wasps (Hymenoptera: Eumenidae, Sphecidae, and Megachilidae); adults have been collected at lights at night. The larva of L. gnara, which preys on immature stages of Cerambycidae and Buprestidae (Coleoptera), was described and illustrated by earlier workers under the name Cymatodera morosa. Adults of L. gnara are commonly collected at lights at night and have been reared from Quercus arizonica Sargent, Prosopis sp., and Juglans sp. Adult feeding, antennal groom- ing, and copulatory behaviors are described for L. gnara, and the presence of a chemical defense in adults of this species is noted for the first time. Key Words: The genus Lecontella Wolcott and Chap- in (1918) currently contains two species, L. brunnea (Spinola) and L. gnara Wolcott, both of which are large, nondescript, brown tilline clerids. Lecontella brunnea is widely distributed throughout eastern North Amer- ica, while L. gnara is found in deserts of the southwestern United States and México. Despite the small size of this genus, its tax- onomic history is quite complex, and for those interested in the details a review was provided by Ekis (1975). The taxonomy of Lecontella and its relatives is actively being studied by W. FE Barr (in litt.), and additions to this genus are expected in the near future. In contrast to the wealth of taxonomic writings on the genus Lecontella, little has been published on the ecology of its spe- cies. The present paper includes substantial Cleridae, Lecontella, chemical defense, ecology, larvae, behavior new information on the ecology of both species currently classified in Lecontella. SEPARATING SPECIES OF LECONTELLA The two species currently placed in Le- contella can be readily separated from spe- cies in most genera of Tillinae by the elon- gate, slender form of the terminal anten- nomere and the presence of rows of large elytral punctures that continue until the ely- tral apices. The sympatric genus with which Lecontella shows greatest affinity (and with which its species are most often confused in collections) is Cymatodera Gray, a large New World genus that has been the subject of considerable careful study by W. E Barr for the past fifty years. The combination of antennal form and elytral punctures will separate L. gnara and L. brunnea from sympatric species of Cymatodera. VOLUME 104, NUMBER 1 Characters for separating the two cur- rently recognized species of Lecontella were discussed by Wolcott (1927). Of the characters listed by Wolcott, I have found the relative size of the pronotal punctures (large and dense in L. gnara, small and sparser in L. brunnea) to be the most reli- able feature for separating adult specimens of these two species. Further information on the identification and relationships of Le- contella species will be provided by W. FE Barr in a forthcoming paper. ECOLOGY OF LECONTELLA BRUNNEA (SPINOLA) References.—Rau (1944): Bitner (1972); Foster and Barr (1972). Larva.—Larvae of L. brunnea are para- sites in the nests of solitary bees and wasps in the families Eumenidae, Megachilidae, and Sphecidae. Bitner (1972) found larvae of L. brunnea in nests of the following spe- cies of Hymenoptera in Indiana, USA: Eu- menidae: Monobia quadridens (L.); Me- gachilidae: Megachile sp., Osmia coerules- cens (L.), Osmia lignaria Say; Sphecidae: Isodontia auripes (Fernald), Trypargilum clavatum (Say), Trypargilum striatum (Pro- vancher). Adult habits.—Labels accompanying adult specimens of L. brunnea in NMNH indicate that adults are most frequently col- lected at lights at night. Adults also have either been reared from or found dead in the nests of bees and wasps. Rau (1944) reported finding adults of this species in the cells of nests of the mud-dauber wasp Sce- liphron caementarium (Drury) (Hymenop- tera: Sphecidae). Foster and Barr (1972) noted that R. M. Bitner successfully reared adults of L. brunnea from nests of Osmia lignaria, O. coerulescens, Megachile rotun- data (FE), Monobia quadridens, and Trypar- gilum striatum. Adult specimens in NMNH were reared from an abandoned Polistes nest (Hymenoptera: Vespidae), nest of Me- gachile rotundata, nest of Trypoxylon stri- atum, and a “‘mud dauber nest.’ The rear- ing record from an abandoned Polistes nest 165 probably indicates parasitism of a mega- chilid bee species, as megachilids common- ly reuse cells of abandoned Polistes nests. ECOLOGY OF LECONTELLA GNARA WOLCOTT References.—B6ving and Champlain (1920) (as Cymatodera morosa LeConte). Larva.—The mature larva of this species was described by Béving and Champlain (1921) under the name Cymatodera morosa LeConte. This larva was found in dead branches of Quercus arizonica Sargent in- fested by species of Cerambycidae and Bu- prestidae (Coleoptera); the adult reared from this larva (in NMNH) is unquestion- ably the species now known as L. gnara Wolcott. Rearing records in NMNH indi- cate that larvae of L. gnara are also found in Prosopis and Juglans spp. infested by buprestid and cerambycid beetles. Adult habits.—This species is one of the most frequently encountered clerids at lights at night in deserts of southwestern North America. In my experience, adults of L. gnara are most abundant at lights on overcast, humid nights preceding summer ‘“‘monsoon”’ thunderstorms. Adult speci- mens in NMNH were reared from Quercus arizonica, Prosopis sp., and Juglans sp. Live material studied.—Eleven adult L. gnara collected at ultraviolet lights at night in Vail, Arizona, between 7 and 14 Septem- ber, 1999, were kept alive for laboratory studies. Adult feeding behavior.—Adults in cap- tivity feed readily on small beetles of a wide range of families. Prey items are cap- tured and manipulated using the first two pairs of legs. The mandibles are used to re- move the legs of the prey, which are dis- carded. The mandibles then serve to nearly divide the prey’s body in two at the junction of the prothorax and the mesothorax. Con- tents of the prothorax and head are con- sumed, and the prothorax and attached head is discarded. Contents of the mesothorax, metathorax, and abdomen are consumed last. Often the elytra and metathoracic wings are removed to reach the metathorax 166 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON and abdomen. In Arizona, the bostrichid beetle Dendrobiella aspera (LeConte) (Co- leoptera: Bostrichidae) was especially pre- ferred, but beetles of the families Bupres- tidae, Cerambycidae, Dermestidae, Mordel- lidae, and Scraptiidae were also consumed. In the laboratory in New York, adults fed on small beetles of the families Buprestidae and Dermestidae, and also readily imbibed sweet syrupy liquids such as honey and ma- ple syrup. Wolcott (1921) noted that species of the related genus Cymatodera were found on mesquite imbibing sap flowing from wounds. Given the fact that adults of L. gnara readily consumed sweet syrupy liquids in captivity, it is probable that adults of this species also feed on sap flowing from wounds on mesquite and other desert shrubs. Antennal grooming behavior.—Adults clean their antennae in the same manner as species of Cymatodera (C. bicolor (Say), C. hurdi Barr, C. inornata (Say), C. sobara Barr, and C. undulata (Say)) studied by the author. Oral secretions are transferred to the ventral pads of one of the protarsi; that pro- tarsus steps on the middle of the antenna on the same side of the body, pinning it to the substrate; and the head is moved in the op- posite direction, pulling the antenna through. To clean the base of the antennae, the protarsus is rubbed along the basal half of the antenna beginning at the base of the scape. Reproductive behavior.—Three male and female pairs of L. gnara were placed in sep- arate containers. Repeated short (<1 mi- nute) copulations were observed; as in most clerids, the male mounts the female from behind and the male’s ventral surface is kept in proximity to the female’s dorsal sur- face throughout copulation. Several likely oviposition substrates were presented to fe- male L. gnara after copulation, including dead branches of Prosopis and Quercus spp. However, no oviposition was observed, and no first instar larvae were obtained, in- dicating that the conditions necessary for Oviposition in this species were not met in the laboratory. Chemical defense.—Adults of L. gnara emit a foul-smelling odor when handled. This odor was not emitted by adult beetles at rest and seems to be associated with dis- turbance. The odor is similar in smell to the defensive secretions of certain stink bugs (Hemiptera: Pentatomidae) and ladybird beetles (Coleoptera: Coccinellidae) from eastern North America, and undoubtedly could be used as a chemical defense against predation. CHEMICAL DEFENSES IN CLERIDAE The first report of a chemical defense in any species of Cleridae was provided by Marshall and Poulton (1902) in their classic study of the bionomics of South African mimetic insects. Marshall and Poulton (1902) reported that a bright orange clerid species (subsequently identified by Maw- dsley (1994b) as Aphelochroa fulva Que- denfeldt, subfamily Clerinae) emitted a strong, ““verbena-like”’ smell. Hawkeswood (1987) provided the second example of pos- sible chemical defense in Cleridae, noting that Australian species of the genus Euna- talis (again, subfamily Clerinae) emitted strongly-smelling chemicals when handled. Lecontella gnara is the first clerid in the subfamily Tillinae identified as possessing a chemical defense, suggesting that chemi- cal defenses may be more widespread in this family than previously thought. The possible existence of chemical de- fenses against predation in adult beetles of the family Cleridae is of particular interest given the large numbers of clerid species that mimic venomous or chemically defend- ed insects (Mawdsley 1994a). It is quite possible that many of these mimetic clerids are themselves chemically defended, and that the mimetic interactions in which these clerids participate are Miillerian, rather than Batesian. ACKNOWLEDGMENTS Funding for travel associated with this project was provided by a grant from the VOLUME 104, NUMBER 1 Theodore Roosevelt Memorial Fund of the American Museum of Natural History, New York. Corinne M. Carter and William and Mary Webber are to be thanked for their valuable assistance with the fieldwork por- tion of this project. Natalia J. Vandenberg of the USDA Systematic Entomology Lab- oratory graciously permitted access to col- lections of adult and larval Lecontella spec- imens in the National Museum of Natural History, Smithsonian Institution (NMNH). The author was supported by a Smithsonian Institution Postdoctoral Research Fellow- ship, sponsored by Terry L. Erwin. William F. Barr provided helpful comments and sug- gestions on earlier drafts of the manuscript. LITERATURE CITED Bitner, R. M. 1972. Predation by the larvae of Lecon- tella cancellata (LeConte) (Coleoptera: Cleridae) on seven species of aculeate Hymenoptera. Ento- mological News 83: 23-26. Boving, A. G. and A. B. Champlain. 1921. Larvae of North American beetles of the family Cleridae. Proceedings of the United States National Muse- um 57: 575-649 + pls. 42-53. Ekis, G. 1975. Taxonomic and nomenclatural status of clerid taxa described by Massimiliano Spinola 167 (1780-1857) (Coleoptera: Cleridae). Bollettino Museo Torino 1: 1—80. Foster, D. E. and W. E Barr. 1972. Notes on the dis- tribution and bionomics of some North American Cleridae. Journal of the Kansas Entomological So- ciety 45(1): 122-125. Hawkeswood, T. 1987. Beetles of Australia. Angus and Robertson, North Ryde, N. S. W. viii + 248 pp. Marshall, G. A. K. and E. B. Poulton. 1902. Five years’ observations and experiments (1896-1901) on the bionomics of South African insects, chiefly directed to the investigations of mimicry and warning colours. Transactions of the Entomolog- ical Society of London 1902(3): 287-584 + 15 pls. Mawdsley, J. R. 1994a. Mimicry in Cleridae (Cole- optera). Coleopterists Bulletin 48(2): 115-125. . 1994b. A revision of the genus Aphelochroa Quedenfeldt (Coleoptera: Cleridae: Clerinae). Journal of African Zoology 108: 121-132. Rau, P. 1944. A note on Lecontella cancellata LeConte (Coleoptera: Cleridae) in cells of the mud-daubing wasp. Entomological News 55(8): 197. Wolcott, A. B. 1921. North American predaceous bee- tles of the tribe Tillini in the United States Na- tional Museum. Proceedings of the United States National Museum 59: 269-290 + pl. 43. . 1927. Descriptions of a new genus and four new species of American Cleridae. Coleopterol- ogical Contributions 1(1): 105-110. Wolcott, A. B. and E. A. Chapin. 1918. Notes on Cler- idae. Bulletin of the Brooklyn Entomological So- ciety 13: 107-108. PROC. ENTOMOL. SOC. WASH. 104(1), 2002, pp. 168-173 (IMMATURE STAGES OF MONTINA CONFUSA (STAL) (HETEROPTERA: REDUVITDAE: HARPACTORINAE) P.M. DELLAPE, M. DEL C. COSCARON, AND B. E AMARAL FILHO (PMD, MCC) Departamento Cientifico de Entomologia, Facultad deCiencias Naturales y Museo, Paseo del Bosque, 1900 La Plata, Argentina (e-mail: coscaron @netverk.com.ar); (BFAF) Departamento Zoologia, Instituto de Biologia, Universidad del estado de Cam- pinas, CP 1170—Campinas, 13100 SP, Brazil Abstract.—The egg and five instars of the harpactorine Montina confusa (Stal) are described and illustrated. Descriptions include morphological and monphometric charac- ters of specimens reared in the laboratory. Key Words: The genus Montina Amyot and Serville is comprised of ten species (Maldonado Ca- priles 1990). Pais Bueno and Berti Filho (1984a, b) described the nymphs and adult male and female of Montina confusa Stal and gave notes on biological aspects. Frei- tas et al. (1991), Rosa et al. (1991), Consol and Amaral Filho (1992) and Freitas (1995) evaluated prey consumption by this species reared under laboratory conditions. In this paper, we redescribe the egg and five nymphal instars of Montina confusa because previous descriptions are inade- quate and the morphological terminology does not follow that used for Reduviidae. MATERIALS AND METHODS The material used in this study consists of eggs and specimens reared in the labo- ratory at the Instituto de Biologia, UNI- CAMP (Universidad do Campinas), Cam- pinas, Sao Paulo, Brazil. The material was preserved in 75% ethanol. Terminology used for morphology follows that of Miller (1971) and Swadener and Yonke (1975). The measurements are given in millimeters (a total of 10 specimens for each instar). Illustrations were made with a drawing tube Heteroptera, Reduviidae, Harpactorinae, Montina confusa, eggs, nymphs on a Wild M-stereomicroscope. Scanning electron micrographs of eggs and fifth in- stars were made from specimens mounted on stubs, sputter-coated with a gold palla- dium alloy, and studied with a JEOL T-100 SEM. DESCRIPTION Montina confusa Stal Egg. (Figs. 1—-2)—Length 2.23—2.34 (X = 2.29), diameter of operculum 0.386—0.466 (X = 0.416). Ova consist of regular masses of dark brown eggs cemented together by their margins. Egg cylindrical and elongated (Fig. 1). Chorion provided with well-devel- oped perforations near apex. Most of chori- on and entire surface of operculum covered with a mucilaginous substance. Operculum with a medial pore (Fig. 2). First instar (Fig. 3).—Total length 3.00— 3.40 (X = 3.22). Length of head 0.92—1.00 (X = 0.97), width of head 0.50—0.57 (X = 0.52). Head: Pyriform, yellowish, stained with dark brown behind eyes; setae short, longer posteriorly. Postocular region sur- rounded. Labrum and clypeus brown dis- tally; clypeus and gena pilose. Eyes intense red, prominent, rounded anteriorly, truncate VOLUME 104, NUMBER 1 169 Figs. 1-2. operculum, 150. posteriorly, not passing superior or inferior edge of the head. Width of eye 0.07—0.10 (X = 0.08), interocular space 0.33—0.38 (X = 0.36). Rostrum yellow, segment I nearly glabrous, length 1.15—1.28 (X = 1.19), ratio of segment lengths ca. 1:1.41:0.48. Antenna elongate, filiform, segments I and IV pale brown, segments I] and HI dark brown, with finer and more abundant setae apical- ly; length 4.32—4.94 (X = 4.71), ratio of segment lengths ca. 1:0.37:0.41:1.18. Thorax: Pronotum pale brown, length 0.28—0.33 (X = 0.31), width 0.50—0.52 (X = 0.51), with two anterolateral tubercles; median sulcus with scattered setae. Pro- pleuron pale brown; mesopleuron pale brown, dark brown ventrally; metapleuron dark brown, tinged with pale brown. Pros- ternum, mesosternum, and metasternun white, tinged with yellow. Wings pads ab- sent. Legs yellowish, with irregular dark stains and scattered short setae. Abdomen: Length 1.33—1.60 (X = 1.42), width 0.60—0.72 (X = 0.66); fusiform, pale brown; pilose. Second instar (Fig. 4).—Total length 4.00—-4.94 (X = 4.47). Length of head 1.38-1.55 (X = 1.49), width of head 0.67— OFS Ci 0:71): “Head: Pyritorm, pale brown, similar to instar V; without spines or small stains in front of eye; ocelli out- lined in red. Eyes similar to instar V. Width of eye 0.12-0.13 (X = 0.12), interocular Micrographs of Montina confusa eggs. 1, General aspect in lateral view, 35. 2, Chorion and space 0.45—0.48 (X = 0.46). Rostrum pale brown. Rostral length 1.63—1.78 (X = 1.68), ratio of segment lengths ca. 1:1.24: 0.47. Antenna elongate, filiform, dark brown, segments III and IV yellowish; seg- ment I with two whitish bands; all segments setose; lengths 6.62—7.09 (X = 6.86), ratio of segment lengths ca. 1:0.35:0.51:1.01. Thorax: Pronotum dark brown, length 0.42—0.53 (X = 0.49), width 0.67—-0.73 (X = 0.70), with two anterolateral tubercles. Median sulci present. Pleural area dark brown, tinged with pale brown, setose. Stridulatory sulci whitish. Prosternum, me- sosternum, and metasternum pale brown. Area between wing pads pale brown. Wings pads 0.57-0.67 long (X = 0.60), dark brown. Legs yellowish to pale brown with irregular dark stains, tarsi darker brown; all segments setose; trochanter I densely pi- lose, black ventrally. Abdomen: Length 1.15—1.33 (X = 1.26), width 0.70—1.25 (X = 1.06). fusiform, pale brown, with setae; scent glands on abdom- inal segments 3—5 pale brown; ventrally with black spots. Third instar (Fig. 5).—Total length 5.70— 6.33 (X = 6.00). Length of head 1.58—2.05 (X = 1.87), width of head 0.92—1.02 (X = 0.95). Head: Pyriform, pale brown, similar to instar V without spines; ocelli outlined in red. Eyes similar to instar V. Width of eye 0.15—0.22 (X = 0.18), interocular space 170 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 0.52—0.63 (X = 0.59). Rostrum similar to instar V, paler, especially first segment. Rostral length 2.19—2.42 (X = 2.32), ratio of segment lengths ca. 1:1.24:0.42. Antenna similar to instar V, with whitish bands on segment II and III. Antennal length 8.84— 9.71 (X = 9.36), ratio of segment lengths ca. 1:0.36:0:63:0.87. Thorax: Pronotum similar to instar V. Length of pronotum 0.67—0.77 (X = 0.73), width of pronotum 0.93—1.05 (X = 0.97). Pleural areas pilose. Stridulatory sulci whit- ish. Propleuron and mesopleuron pale brown tinged with orange except for dark margins on propleuron, posterior and infe- rior fringe dark on mesopleuron, metapleu- ron dark brown, pale brown medially. Pros- ternum, mesosternum, and metasternum whitish, tinged with pale brown. Area be- tween wing pads pale brown, tinged with orange, whitish anterior and _ posteriorly. Wings pads 0.80—0.93 long (X = 0.86), dark brown, tinged with pale brown. Legs whitish, except coxae and femora pale brown distally. Coxae, trochanters, femora, and tibiae stained with dark brown. Tro- chanter I, femora I basally, and tibia I dis- tally with dense black setae ventrally. Abdomen: Length 3.00—4.00 (X = 3.48), abdominal width 0.95—1.42 (X = 1.21). Abdomen fusiform, pale brown, tinged with orange and whitish; setose; scent glands well developed on abdominal segments 3— 5. Segment 1 whitish; segment 2 pale brown tinged with white; segment 3 dark brown tinged with white; segment 6 dark brown, with a black spot expanded laterally. Yellowish ventrally, tinged with pale brown; segments 4-6 red and white stained, with small black spots medially; segment 8 brown stained. Fourth instar (Fig. 6).—Total length 8.10-9.12 (X = 8.37). Length of head 2.63—2.93 (X = 2.79), width of head 1.17— 30 CS= 1:21). Head: Similar tounstar VW; with spines; ocelli less distinctly outlined in red, in some specimens superior border of gena dark. Eyes as instar V. Width of eye 0.20—0.27 (X = 0.23), interocular space 0.68—0.83 (X = 0.75). Rostrum similar to instar V, segment I paler and stains on edg- es of III absent. Rostral length 2.83—3.47 (X = 3.12), ratio of segment lengths ca. 1:1.14: 0.36. Antenna as instar V; length 11.89— 13.08 (X = 12.35), ratio of segment lengths eae 120°38:0:81-0.82. Thorax: Pronotum as for instar V. Length of pronotum 0.97—1.17 (X = 1.06), width of pronotum 1.30—1.55 (X = 1.41). Pleural areas pilose. Stridulatory sulci whit- ish. Propleuron and mesopleuron pale brown, tinged with orange, edges of pro- pleuron dark brown, posterior and inferior fringe of mesopleuron and metapleuron dark brown, except medially pale brown. Prosternum, mesosternum and metasternum whitish, tinged with pale brown. Posterior lobe of pronotum pale brown, tinged with orange, posterior process whitish, with two brown spots laterally. Wing pads 1.38—1.75 long (X = 1.58). Legs whitish or. pale brown, except coxae, femora distally, and tibiae basally pale brown, tinged with red. Dark stains more distinct in instar V; all segments setose. Trochanter I, femur I, and tibia I densely pilose, black ventrally. Abdomen: Length 4.18—4.68 (X = 4.36), width 1.67—2.43 (X = 2.06). Abdomen fu- siform, pale brown, tinged with orange; se- tose; segments | and 2 with whitish stains medially; scent glands well developed. Seg- ment 6 with a black spot expanded laterally in same position as scent glands of anterior segments. Two whitish stains lateral to scent glands. Segments 4—6 with a red fringe distally and whitish stains centrally, in some specimens. Connexivum whitish distally and white continuing to the medial part of the abdomen. Segment 7 with a rect- angular dark stain; segment 8 with a dark stain expanded laterally. Pale brown with lateral white lateral spots bordered in red. Ventrally pale brown (some specimens with black points), medially pale. Apical seg- ments darker externally (Fig. 7). Fifth instar (Fig. 8).—Total length 12.41-15.19 (X = 13.98). Length of head 3.47—4.07 (X = 3.70), width of head 1.50— VOLUME 104, NUMBER 1 171 Figs. 3-10. Montina confusa. 3, First instar. 4, Second instar. 5, Third instar. 6, Fourth instar. 7, Fourth instar, ventral aspect of last segments of abdomen. 8—10, Fifth instar. 9, Head, lateral view. 10, ventral aspect of last segments of abdomen. (72 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 11-14. Montina confusa, micrographs of antennal segment 2 trichobothria. 11, Antennal segment 2, 75%. 12, Medial trichobothrium, 350. 13, Distal trichobothria, 750. 14, Medial trichobothrium, 750. 1.73 (X = 1.58). Head: Pyriform, pale brown, darker brown behind eyes, paler on neck, ocelli outlined in red; laterally (Fig. 9) with a stripe from antenna to eye, a small stain in front and an other stain posteriorly brown; labrum and clypeus dark brown, buccula distally dark brown. Eyes promi- nent, rounded anteriorly and truncated pos- teriorly, intense red, not passing superior or inferior edge of head. Width of eye 0.28— 0.65 (X = 0.47), interocular space 0.90— 1.10 (X = 0.99). Rostrum pale brown, stained dark brown (in some specimens joint of I and II darker); II and HI paler than I and with more setae. Stain basally on I, II, and along edges on III. Rostral length 3.67—4.64 (X = 4.12), ratio of segment lengths ca. 1:1.12:0.40. Antenna elongate, filiform, dark brown to almost black, paler on Ill and IV; two whitish bands on seg- ment I, one on segments II and III; setae finer, more abundant distally. Segment I with a trichobothrium as seen in Figs. 11— 14. Antennal length 15.08-17.19 (X = 16.04), ratio of segment lengths ca. 1:0.38: 019220) 7A Thorax: Pronotum orange (tinged with red in some specimens), with two antero- lateral tubercles and anterior margin dark brown; with setae. Median sulci present, with sparse, short setae. Length of prono- tum 1.47—2.00 (X = 1.71), width of pron- otum 1.85—2.53 (X = 2.17). Propleuron and mesopleuron pale brown, tinged with or- ange, except dark brown margins, with nar- row, pale borders. Stigma whitish interiorly. VOLUME 104, NUMBER 1 Metapleuron dark brown, except basally and medially pale brown. Pleura pilose. Prosternum dark brown except stridulatory sulci whitish. Mesosternum and metaster- num whitish, tinged with pale brown, a cen- tral fringe of black setae on mesosternum in some specimens. Scutellum pale brown, tinged with orange, posterior process whit- ish. Wings pads 3.64—4.37 long (X = 3.93), pale yellow, dark brown basally and along edge. Legs pale brown, tinged with pale yellow; coxae, femora distally, and tibiae basally pale brown, tinged with red; tarsi brown. Trochanters, femora, and tibiae tinged with dark brown. Trochanter I, femur I, and tibia I densely setose, black ventrally. Abdomen: Length 6.52—7.91 (X = 7.28), abdominal width 3.17—3.92 (KX = 3.53). Abdomen fusiform, pale brown, tinged with orange; setose; scent gland well developed on abdominal segments 3—5, consisting of black papilas, except base of first papila pale brown. Segment 6 with a black spot expanded laterally in same position as scent glands of anterior segments. Two whitish stains lateral to scent glands. Segments 3— 6 with a red fringe distally, surrounded by whitish stains centrally. Connexivum dis- tally and continuing to the medial part of the abdomen whitish; the remainder brown, tinged with orange. Pale brown ventrally, tinged with red surrounded by whitish spots laterally, last segments darker externally (Fig. 10). ACKNOWLEDGMENTS We express our gratitude to Dr. T. J. Hen- ry (Systematic Entomology Laboratory, USDA, Washington, DC) for reading the manuscript and to UNICAMP technicians for rearing the reduviids. This work was supported by the Consejo Nacional de In- 7/3} vestigaciones Cientificas y Técnicas (CON- ICET) and the Universidad Nacional de La Plata, Argentina. LITERATURE CITED Consoli, E L. and B. E Amaral Filho. 1992. Ciclo biol6gico de Montina confusa (Stal, 1859) (He- miptera: Reduviidae) alimentado com diferentes presas. Revista Brasileira de Entomologia 36(3): 697-702. Freitas, S. 1995. Capacidade de predacao, sobreviven- cia e ciclo biol6gico do predador Montina confusa Stal (Heteroptera: Reduviidae) alimentado com la- gartas da broca da cana-deacucar, Diatraea sac- charalis Fabr. (Lepidoptera: Pyralidae). Anais da Sociedade Entomologica do Brasil 24(2): 195— 199% Freitas, S., S. A. De Bortoli, and Z. A. Ribeiro. 1991. Aspectos biologicos de Montina confusa (Hem.- Reduviidae) alimentada com larvas de Diatraea saccharalis (Lep.-Pyralidae). XIIl Congresso Brasileiro de Entomologia. Resumos., Vol. II, p. 489. Maldonado Capriles, J. 1990. Systematic catalogue of the Reduviidae of the World. Caribbean Journal of Sciences (special edition). Mayaguez, P. R., 694 PP: Miller, N. C. E. 1971. The Biology of the Heteroptera. Second (Revised) Edition. E. W. Classey, Hamp- ton, England, 206 pp. Pais Bueno, V. H and E. Berti Filho. 1984a. Montina confusa (Stal, 1859) (Hemiptera: Reduviidae, Ze- linae): II. Aspectos morfol6gicos de ninfas e ad- ultos. Revista Brasileira de Entomologia. 28(3): 355-364. . 1984b. Montina confusa (Stal, 1859) (Hemip- tera: Reduviidae, Zelinae): I. Aspectos bioldgicos. Revista Brasileira de Entomologia 28(3): 345— 353. Rosa, M. P, A. G. Damasceno, and V. H. P. Bueno. 1991. Avaliacao da actividade predatoria do Re- duvidae predador Montina confusa (Hemiptera: Reduviidae, Zelinae) sobre lagartas de Galleria mellonella (Lepidoptera: Pyralidae). XIII Con- gresso Brasileiro de Entomologia. Resumos. Vol. I, p. 268. Swadener, S. O. and T. R. Yonke. 1975. Inmature stag- es and biology of Pselliopus cinctus and Psellio- pus barberi (Hemiptera: Reduviidae). Journal of the Kansas Entomological Society 48: 477—492. PROC. ENTOMOL. SOC. WASH. 104(1), 2002, pp. 174-194 \ REVIEW OF THE SIRICID WOODWASPS AND THEIR IBALITD PARASITOIDS (HYMENOPTERA: SIRICIDAE, IBALITDAE) IN THE EASTERN UNITED STATES, WITH EMPHASIS ON THE MID-ATLANTIC REGION DAVID R. SMITH AND NATHAN M. SCHIFF (DRS) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. De- partment of Agriculture, % National Museum of Natural History, Smithsonian Instituion, Washington, D.C. 20560-0168, U.S.A. (e-mail: dsmith@sel.barc.usda.gov); (NMS) U.S. Forest Service, U.S. Department of Agriculture, Southern Research Station, Center for Bottomland Hardwood Research, PO. Box 227, Stoneville, MS 38776, U.S.A. (e-mail: nschiff @ fs.fed.us) Abstract.—Keys are presented for the five genera and 15 species of adult Siricidae and one genus and two species of their parasitoids of the family Ibaliidae that occur in or may be adventive in eastern United States. Siricid larvae are wood borers in conifers and broadleafed trees. Notes on their biology, fungal symbionts, distributions and host asso- ciations are given. Data from collections in the mid-Atlantic states include seasonal oc- currence of both Siricidae and Ibaliidae. Key Words: Worldwide, there are 85—100 species of Siricidae in two subfamilies and 11 genera (Smith 1978, 1993). The subfamily Tremi- cinae is associated with angiosperms and Siricinae with gymnosperms. The family is widespread in the deciduous and coniferous forests of the northern hemisphere, extend- ing south to Cuba, northern Central Amer- ica, New Guinea, Philippines, Viet Nam, northern India, and northern Africa; one ge- nus with two species is Afrotropical. No Siricidae are native to Australia and South America. All species for which larvae are known bore into weakened or dying trees (Middle- kauff 1960, Smith 1979). Most species in their native range are considered to be of minor importance except for decreasing the value of lumber; however, introduced, ex- otic species can be very damaging. Because larvae of siricids feed and develop in wood, Sirex, Urocerus, Tremex, Xerix, Eriotremex, Ibalia, fungal symbionts with several years required for their life cy- cle, they are commonly transported in lum- ber by commerce. Thus, non-native species may emerge in buildings constructed of lumber that originated in other parts of the country or from other nations. For example, Sirex noctilio (E) is a European species that became a major pest of Pinus radiata, an American species, when P. radiata was planted in New Zealand and Australia (Gil- bert and Miller 1952, Rawlings 1955, Gaut 1970). A major control effort in the 1960’s and 1970’s used parasitic nematodes to con- trol S. noctilio (Bedding and Akhurst 1974). Sirex noctilio is now considered to be the most important threat to new P. radiata plantations in South Africa, Brazil, and Ar- gentina (Stival et al. 1993; Iede et al. 1998; Tribe 1995, 1997). In the early 1970’s, an Asian species, Eriotremex formosanus (Matsumura), was accidentally introduced VOLUME 104, NUMBER I into the southeastern United States where it has spread rapidly (Smith 1975, 1996). It attacks hardwoods, including oaks, but it is not considered to be a major pest at this time. More recently, a Palearctic species, Urocerus sah (Mocsary), was reported in New Hampshire (Smith 1987). The Ibaliidae (Cynipoidea) are part of the parasitoid complex of Siricidae. We include them here because we recorded their pres- ence in collections from the mid-Atlantic states and are able to present their season- ality in relation to that of their hosts. Keys for the identification of the eastern United States species of Siricidae have been non-existent since Bradley’s (1913) revi- sion, although Smith (1987) published a key to North American Urocerus, and Stan- ge (1996) gave a key to the six species of Siricidae in Florida. Johnson (1928) gave some notes on New England species and illustrated them, but he did not give a key to species. For the Ibaliidae, Liu and Nor- dlander (1992, 1994) published a revision of world and North American Ibaliidae. Here, we give keys for identification of the eastern U.S. Siricidae and Ibaliidae, sum- marize their distribution and hosts, and pre- sent seasonal activity of some species with emphasis on collections from the mid-At- lantic states. General biology Like other wood-boring insects, siricids do not make the enzymes that digest cel- lulose, the major energy source for wood feeders (Kukor and Martin 1983). To utilize cellulose, wood-boring insects live in sym- biotic relationships with other organisms that produce cellulases (Buchner 1965, Francke-Grosmann 1939). Siricids use ba- sidiomycetous wood-decay fungi to break down cellulose (Buchner 1928; Cartwright 1929, 1938). The relationship between sir- icids and these fungi is true symbiosis, as organisms derive benefit. The siricids gain the ability to utilize a large energy source, cellulose, and the fungus benefits because it is not only carried to a specific host tree, 175 but it is also injected underneath the bark, past the tree’s first line of defense. Female siricids, except in the genus Xeris, carry oidia (hyphal fragments) of the fungus in specialized, abdominal glands called my- cangia that have ducts leading to the repro- ductive tract (Buchner 1928). When the wasp lays her eggs, oidia are also deposited. The fungus grows rapidly and secretes di- gestive enzymes onto the substrate. When the larvae hatch, they commence feeding on the fungus and digested wood. There is some disagreement about whether or not the insects actually consume the wood or feed on the fungi alone (Gilbertson 1984). In either case, however, wood is the ulti- mate source of energy for the larvae. Ex- actly how adult females acquire the fungus after the larvae pupate is also uncertain (Gilbertson 1984), but female larvae have specialized hypopleural organs (Parkin 1941, 1942; Stillwell 1965) that may play a role in scraping fungus from the tunnel walls which is then somehow incorporated into the mycangia. Larvae take from one to three or more years to reach pupation and emerge as adults (Middlekauff 1960, Stillwell 1967, Smith 1993). Males are commonly reared but seldom seen in the field. They are oc- casionally seen in small groups at the tops of trees (Middlekauff 1960; Schiff, personal observation). In the western United States, many species are attracted to forest fires, where females lay eggs into charred trees. Forest fires are much less common in the eastern United States, and there are no data on eastern siricid species being attracted to fires in this region. Species of the genus Xeris do not appear to carry a wood decay fungus in mycangial glands; instead, they limit oviposition to substrates that have al- ready been inoculated with a wood decay fungus by another species of siricid (Francke-Grosmann 1939, Fukuda and Hijii L997): FUNGI The identity of the fungal symbionts used by siricids has been complicated by both 176 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON the difficulty in inducing the fungi to pro- luce fruiting bodies in vitro and in some ases by misidentifications of the siricids involved (Gilbertson 1984, Gaut 1970). The fungal symbionts of only two tremicine species have been identified: Tremex col- umba (L.) from North America and Tremex longicollis Konow from Japan, both of which harbor Cerrena (Daedalea) unicolor Bull. ex Fries (Stillwell 1964, Tabata and Abe 1995). The siricines use fungi in the genus Amylostereum, and there are three species worldwide: A. chailettii (Pers.: Fr.) and A. laevigatum (Fr.) that occur in the United States, and A. areolatum (Fr.) that is not known from the United States (Farr et al. 1995). All United States siricines whose symbionts have been determined (Sirex cy- aneus, S. nigricornis, S. edwardsit, S. lon- gicauda, S. juvencus californicus, Urocerus albiconris, U. gigas flavicornis, and U. cal- ifornicus) use A. chailettii (see Gilbertson 1984 review). American isolates of fungus from Sirex juvencus have not been deter- mined, but European examples use A. ar- eolatum (Gaut 1970). If each siricid is faith- ful to a specific symbiont (Gaut 1970), then Sirex Juvencus in America must use A. ar- eolatum, and the fungus has simply not been found in the field as yet, or, the Amer- ican Sirex juvencus is not the same as the European species. Amylostereum areolatum should also have been introduced with Sirex noctilio, its wasp symbiont, in places where that species was accidentally introduced. MALAISE TRAPPING AND VIRGINIA SPECIES This study is supplemented by material collected in Virginia and adjacent states over the past 18 years and specimens in the collections of the National Museum of Nat- ural History, Smithsonian Institution, Washington, D.C. (USNM), and Virginia Polytechnic Institute and State University, Blacksburg, Virginia (VPI). Collections were made using Malaise traps which were set up and in continual use in different localities from approxi- mately March through the first of Novem- ber. Siricidae and Ibaliidae are not com- monly collected this way, but enough have accumulated over the years of collecting to help verify what occurs in this region and indicate the seasonal activity of some spe- cies. Totals collected are 163 specimens of 5 species of Siricidae and 149 specimens of 2 species of Ibaliidae. The Siricidae are rep- resented mainly by 133 specimens of Tre- mex columba (L.), associated with broad- leaved trees, and 126 specimens of its par- asitoid, [balia anceps Say. The other Siri- cidae, 30 specimens of the genera Urocerus and Sirex, are associated with conifers, and 23 specimens of their parasitoid /balia leu- cospoides (Hockworth). All Siricidae col- lected in Malaise traps were females. Seven species of Siricidae and two spe- cies of Ibaliidae occur in Virginia. We have been able to verify several exotic species, e.g., Sirex longicauda Middlekauff, but there are undoubtedly others. Another non- native species 1s Eriotremex formosanus (Matsumura), an introduced species in southeastern U.S. which has been found in extreme southeastern Virginia. MATERIALS AND METHODS The keys are valid for eastern United States, east of the Rockies, and eastern Can- ada. Species not native to this region but are possible adventives in imported lumber are included. There are no confirmed col- lections of Sirex noctilio from eastern U.S., but it is included it in the key because the species has a history of introductions in other parts of the world and specimens are occasionally intercepted at ports-of-entry in the U.S. Recorded distributions and host plants are from the literature and examined spec- imens. The following are given for each species: distribution (state/province from which recorded); Virginia records; collec- tion records (from Malaise trapping in Maryland, Virginia, and West Virginia); hosts (as recorded); and remarks. Records in the collections sections are cited only by county. Specific data are as VOLUME 104, NUMBER 1 177 Fig. 1. Sirex nigricornis (arrow points to cornus). follows: MARYLAND: Allegheny Co., Green Ridge State Park.; Prince George’s Co., Beltsville Agricultural Research Cen- ter. VIRGINIA: Clarke Co., University of Virginia Experimental Farm and State Ar- boretum of Virginia, 2 mi S Boyce; Essex Co., 1 mi SE Dunnsville; Fairfax Co., near Annandale; Louisa Co., 4 mi S Cuckoo; Loudoun Co., nr. jct. of Sycolin Road and Goose Creek. WEST VIRGINIA: Hardy Co., 3 mi NE Mathias, 38°55’N, 78°49'W; Tucker Co., Fernow Experimental Forest, south of Parsons. Literature references to original descrip- tions and synonymies are found in Smith (1978, 1979). Only significant or subse- quent literature is presented here. The use of pits on the lancet for species separation were first used by Viitasaari (1984) and Viitasaari and Midtgaard (1989). The size, form, and location of the lateral ridges and pits may be significant for species identification. They are used here mainly to separate Sirex noctilio from the native North American species of Sirex. Males are more difficult to separate than females. Use caution in the keys to males since color variation may be more extensive than we have observed. RESULTS Siricidae KEY TO GENERA OF NORTH AMERICAN SIRICIDAE 1. Hindtibia with two apical spurs; antenna with 18 or more segments, usually more than 22 i) — Hindtibia with one apical spur; antenna with more than 22 segments in Xeris, less than 22 segments in 7remex and Eriotremex ....... Head usually with large white spot behind each eye; female cornus long and slender, constrict- ed at base (Fig. 2); male hindtarsus slender, first segment 4 or more times longer than broad, commonly 5—6 time longer, second and third segments 2 times or more longer than broadh(Eve216) area hee ee Urocerus — Head black; female cornus short, triangular, not constricted at base (Figs. 1, 3, 4); male hind- tarsus stout, laterally flattened, first segment 4 times or less longer than broad, second and third segments triangular, slightly longer or about as long as broad (Fig. 5) i) Go apt Se eet. Sirex 178 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Antenna with 15-21 segments; head without ge- nal carina behind eyes; female ovipositor shorter than forewing; hindwing with anal cell ...... 4 Antenna with 23 or more segments; head with a genal carina behind each eye (Figs. 7, 8); female ovipositor longer than forewing; hind- Naver AwatloCul cioell Gell’ 4 ces oe noose cse Xeris 4, Antenna with 14-15 segments; body without lone coldenwhaiiss restrain a eee Tremex — Antenna with 20-21 segments; body with long coldenwhairs yen an ae eee LOUFEIMe: Comments.—Use of the white spot be- hind each eye to separate Urocerus and Sir- ex is commonly used in keys but is not in- fallible. Although most specimens can be separated by this, a series of Urocerus cres- soni from Virginia varies from an entirely black head to some with a distinct white spot. Examination of the shape of the fe- male cornus and the male hind tarsus should be checked for determination. The more slender, more rounded hindtarsus (Fig. 6) of Urocerus vs. the stout, laterally flattened hindtarsus (Fig. 5) of Sirex males is a good character for the eastern species. SUBFAMILY SIRICINAE Genus Sirex Linnaeus The most recent key is to the species of California by Cameron (1967). Two spe- cies, S. edwardsii and S. nigricornis, are widespread in eastern U.S. Sirex cyaneus is northern but extends south in the Appala- chians to North Carolina, S. juvencus is northern and transcontinental, S. behrensii has only been recorded from Virginia and emerging from imported wood in Florida, and two species, S. areolatus and S. longi- cauda, are adventive in the eastern states emerging from wood. KEY TO SPECIES [is = Remlal emer .asvacc0st aos tk aha ots ED ce 2 Nialemrpeyal Mees ro aos, i ee 9 2. Ovipositor longer than forewing ......... 3 — Ovipositor shorter than forewing ........ 4 3. Tibiae and tarsi red ... longicauda Middlekauff — Legs entirely black or blue black areolatus (Cresson) 4. Abdomen mostly red, but may be infuscated to black at base, laterally or ventrally Figs. 2-4. Female cornus. 2, Urocerus cressoni. 3, Sirex cyaneus. 4, Sirex longicauda. — Abdomen entirely black or blue black ..... 6 5. Legs black, only tarsi reddish brown; wings darkly, uniformly infuscated (Fig. 1)...... A SD ie OA ie Re Sate §: nigricornis (Fabricius) — Legs with tibia and tarsi reddish brown; wings hyaline, forewing with fuscous band below stigma and fuscous apical margin .. . ft NEA RS Ga One, On: aoe ee ee behrensii (Cresson) Gn) egsiblacktiiar 2s: See edwardsii Brullé Legs largely orange 7. Apical tarsal segment black; sawsheath (val- vula 3) shorter than oblong plate (valvula 2); length of ovipositor as long as distance from base of wing to base of radial cell; mesopleu- ron densely punctured, in the middle with in- terspaces mostly smaller than punctures (Fig. 9); pits of lancet large and close together (Fig. 1): santenna black). ss.) noctilio (Fabricius) — Tarsi yellow to orange; sawsheath (valvula 3) as long as or longer than oblong plate (valvula 2); length of ovipositor as long as distance VOLUME 104, NUMBER | Figs. 5-6. . Femora and rest of hindleg black from base of wing to beyond base of radial cell; punctures at middle of mesopleuron far- ther apart, with shining flat interspaces mostly broader than punctures (Fig. 10); pits of lan- cet small and far apart (Fig. 12); antenna black or brown at base Sawsheath (valvula 3) longer than oblong plate (valvula 2); length of ovipositor as long as distance from base of wing to apex of ra- dial cell; antenna black cyaneus (Fabricius) Sawsheath (valvula 3) equal to oblong plate (valvula 2); length of ovipositor as long as distance from base of wing to middle of radial cell; basal segments of antenna often reddish brown jJuvencus (Linnaeus) Abdomen red, with only basal 1 or 2 seg- mMentsablacksoyr .f hon Soaks eet ee 10 Abdomen black at both base and apex, black at apex may be only on dorsum or venter of apicaliSeSMentt == assoc 6 235 a eee oes 12 . Femora, tibiae and tarsi red; antenna pale at base; wings hyaline...... behrensii (Cresson) Hindleg black or black with femur red; mid- leg black; antenna black; wings yellowish 11 . Hindleg black, apical 2 tarsal segments yel- lowish nigricornis (Fabricius) Hindleg with femur red, tibia and tarsus black cyaneus (Fabricius) . Legs entirely black (abdominal segments 2—7 orange; wings hyaline, stigma of forewing black Wes) oh eh areolatus (Cresson) Legs partly reddish brown to orange .... . 13} Hindtarsus. 5, Sirex edwardsii. 6, Urocerus cressoni. — Femora orange; apical 2—3 hindtarsal seg- ments orange 14. Abdominal segments 5 and 6 and sometimes part of 7 orange; wings yellow, stigma of forewing yellow edwardsii Brullé — Abdominal segments 3—7 and sometimes 8 orange; wings clear, hyaline, stigma of fore- wing black longicauda Middlekauft 15. Basal antennal segments orange; apical tarsal segments yellow to orange; punctures of me- sopleuron separated by flat, shining interspac- es, usually broader than punctures (Fig. 10) Juvencus (Linnaeus) — Antenna black; apical two tarsal segments blackish; punctures of mesopleuron close to- gether, interspaces usually shorter than punc- tures (Fig. 9) noctilio (Fabricius) Sirex areolatus (Cresson) Distribution.—Alabama, Arizona, Ar- kansas, British Columbia, California, Col- orado, Florida, New Mexico, Oregon, Vir- ginia, Washington (Middlekauff 1960, Smith 1979). The Florida specimens emerged from fir wood imported from western United States (Stange 1996). In Al- abama, specimens were “‘found boring into sheet rock inside home,” and in Arkansas, ‘“‘ex sheet rock.”’ It was found in Kauai, Ha- wali ‘“‘ex cedar wall.” PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Xeris spectrum. 7, Head lateral view. 8, Head, dorsal view (arrows point to carina). VOLUME 104, NUMBER | Virginia records.—Cape Henry, IX- 13023, J.N. Knull, bald cypress (1 female, 2 males); Cape Henry, 9-9-24, W.S. Fisher (1 male). Hosts.—Cupressus macrocarpa, Junipe- rus occidentalis, J. scopulorum, Lioboced- rus decurrens, Pinus contorta, P. jeffreyi, P. lambertiana, P. radiata, Psuedotsuga menziesil, Sequoia sempervirens, Taxodium distichum, Thuja sp. Commonly attacks redwood, cypress, and cedars; found less frequently on pines (Middlekauff 1960, Smith 1979). Remarks.—The only eastern records known for this species are from Virginia and Florida. The Virginia collections appear authentic and are probably not from im- ported wood. Inasmuch as S. areolatus commonly attacks redwood, cypress, and cedars and is found less frequently on pines (Middlekauff 1960), this species could be established on baldcypress in the East. De Leon (1952) recorded it from Sequoia sem- pervirens in California, and Westcott (1971) from Juniperus occidentalis in Oregon. Sirex behrensti (Cresson) Distribution.—California, Idaho, Neva- da, Oregon, Washington. Hosts.—Cupressus macrocarpa, Pinus Jeffreyi, P. lambertiana, P. ponderosa, P. radiata, Japanese pine (Middlekauff 1960, Smith 1979). Remarks.—This species has been found in Ohio emerging from imported lumber: “Cleveland O., ex plaster wall.” Sirex cyaneus Fabricius (Fig.3) Distribution.—Alabama, Alaska, Alber- ta, British Columbia, California, Colorado, Georgia, Idaho, Illinois, Kansas, Maine, Michigan, Minnesota, Montana, Nebraska, New Brunswick, New Hampshire, New Jer- sey, Newfoundland (Labrador), Nevada, New Mexico, New York, North Carolina, Nova Scotia, Oregon, Quebec, Utah, Wash- ington, West Virginia, Wyoming (Middle- kauff 1960, Smith 1979). 181 Hosts.—Abies balsamea, A. concolor, A. fraseri, A. grandis, A. lasiocarpa, A. mag- nifica, Larix laricina; Picea engelmanni, P. glauca, P. sitchensis, Picea sp., Pinus con- torta, P. ponderosa, Pseudotsuga menziesii, Tsuga heterophylla (Middlekauff 1960, Kirk 1975, Smith 1979). Remarks.—This species is known as the “blue horntail.”” No specimens have been collected in Virginia, but S. cyaneus prob- ably occurs in higher elevations in the Ap- palachians between New York and North Carolina (Amman 1969) where its hosts, Abies and Picea, occur. Johnson (1930) recorded hundreds of emergences from balsam fir, Abies balsa- mea, in Randolph, N.H., from July 29 to August 29. He discussed variation in size and color. Forty-one specimens emerged from a section of the tree 11” long and 434” in diameter. Also, several specimens of /ba- lia emerged. Amman (1969) reared S. cy- aneus from Fraser fir in North Carolina. Blackman and Stage (1918), reporting it as S. abbotii, reared it from Larix laricina in New York. The life history was reported on by Chrystal (1928, 1930) and Chrystal and Meyers (1928). It has often been confused with, or sometimes considered a subspecies of, S. juvencus. Sirex edwardsii Brullé (Biss. ko) Distribution.—Alabama, Arkansas, Dis- trict of Columbia, Georgia, Louisiana, Maryland, Massachusetts, Mississippi, North Carolina, Oklahoma, Quebec, Sas- katchewan, South Carolina, Texas, West Virginia, Wisconsin, Virginia. The Louisi- ana record is from Chapin and Oliver (1986). Virginia records.—Lynhaven Inl., City of Virginia Beach, X-27-43; Virginia Beach, Pinus, Hopk. 69202; W. Minor Hills, Falls Church, pine, Hopk. 11339al1; Montgomery Co., Oct. 2, 1964; Blacksburg, X-11-47. Collection records.—VIRGINIA: Essex Co., X-24—XI-17-95 (1). 182 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON “a ee 4 i) 4 “4 Figs. 9-10. Mesopleuron. 9, Sirex noctilio. 10, Sirex juvencus. Arrows point to area where density of punctures differ. Hosts.—Picea abies; Pinus echinata, P. elliottii, P. palustris, (Py rigida, RP. strobus. P. virginiana (Kirk 1974, Smith 1979). Remarks.—This species was recorded as Sirex abbotii by Kirk (1974). In Essex Co., where one specimen was collected, only two species of pines were present (Pinus virginiana and P. taeda). Sirex Juvencus (Linnaeus) (Figs. 10, 12) Distribution.—Eurasia; British Colum- bia, New Brunswick, Newfoundland (insu- lar and Labrador), New Jersey, Nova Sco- tia, Yukon Territory. Hosts.—Abies balsamea, A. lasiocarpa, Larix sp., Picea sp., Pinus contorta. Remarks.—Benson (1962) first recog- nized that S$. juvencus occurs in North America, recording it from ‘“‘Labrador and Newfoundland (? introduced).’’ Subse- quently, specimens have been identified as S. juvencus from across North America, and it is believed to be widespread. Its distinc- tion from §. cyaneus and S. californicus (Ashmead) is not clear, but we have sepa- rated S. juvencus and S. cyaneus in the key using traditional characters pending a thor- ough study of this complex. Both S. juvencus and S. cyaneus can be separated from S. noctilio, the three species most likely to be confused, by the much smaller pits on the lancet (Fig. 12; S. noc- tilio, Fig. 11). Sirex juvencus and S. cy- aneus also differ by the size of the pits, with those of S. cyaneus being much small- er (breadth of pit less than a third of di- ameter of ovipositor), but this distinction is not as evident (see Viitasaari and Midtgard 1989). Sirex longicauda Middlekauff (Fig. 4) Distribution.—California, Colorado, Ida- ho, Kansas, Montana, Nebraska (“‘emerg. wood in new home;”’ probably west coast wood), Nevada, New Mexico, Ohio (?), Oregon, Utah, Virginia, West Virginia (Monongalia Co., ex 2 X 4’s in new house, lumber cut in Pacific Northwest). Adven- tive in eastern states emerging from lumber. The records from Kansas, Ohio, and Vir- ginia are probably from imported lumber. Virginia records.—Campbell Co., Lynch- burg, 6-31-64 (1). This specimen is at VPI&SU. Probably emerged from building materials imported from western United States, though this is not given on the la- bels. Hosts.—Abies concolor, A. magnifica; Pinus ponderosa, Pseudotsuga menziesii (Kirk: 1975, Smuth979): VOLUME 104, NUMBER 1 183 Figs. 11-12. Remarks.—So far as is known, this spe- cies is not established outside its native range in western North America. Sirex nigricornis Fabricius (Bigs. ol, 116) Distribution.—Alabama, Arkansas, Dis- trict of Columbia, Florida, Georgia, Florida, Louisiana, Maryland, Massachusetts, Mis- sissippi, North Carolina, Ohio, Pennsylva- nia, Quebec, Saskatchewan, South Caroli- na, Texas, Virginia, West Virginia, Wiscon- sin (Kirk 1974, Smith 1979). Virginia records.—Falls Church, Oct. 14, 1970, Oct. 28, 1916; Virginia Beach, Nov. 11, 1907; Norfolk, 1909; Lurray, Page Co., X-5-13; Lynhaven Inl., City of Virginia Beach, X-27-43; Montgomery Co., X-6-61. Collection records.—VIRGINIA: Essex Co., IX-29—X-27-92 (1). Hosts.—Picea sp., Pinus clausa, P. echinata, P. palustris, P. rigida, P. strobus, P. taeda, P. virginiana (Kirk 1974, Smith nO79). Remarks.—Webster (1895) received specimens, called the ‘“‘black horned horn- tail,’ reported as being numerous on peach and apple trees in Tennessee. He was prob- ably referring to S. nigricornis. Pits on lancet. 11, Sirex noctilio. 12, Sirex juvencus. Sirex noctilio (Fabricius) (Figs. 9, 11) Distribution.—Eurasia; introduced and established in Australia, New Zealand, Bra- zil, Uruguay, Argentina, and South Africa. Hosts.—Abies spp., Larix spp., Picea spp., Pinus radiata, Pinus spp., Pseudot- Suga Menziesil. Remarks.—There is a questionable re- cord for this species in North America by Benson (1962) from ‘‘Manitoba (? intro- duced).’” No other information is available and we have not seen confirmed records ex- cept the Wyoming record by Smith (1979) from a specimen in the USNM which is la- beled “Old Faithful, Yellowstone, Wyo., VITI/26.25, Satterthwait collector.”” We will not consider this species established in North America until there are more con- firmed records. It has been intercepted oc- casionally in wood and dunnage at ports- of-entry in the United States. This species is a major pest of pine plantations where it has been accidentally introduced (see ref- erences in Introduction). We have included it in the preceding key so it can be distin- guished from native North American spe- cies, and it will be helpful to identifiers at ports-of-entry. 184 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Genus Urocerus Geoffroy Smith (1987) gave a key to North Amer- ican species. Two species, Urocerus cres- soni and U. taxodii, are widespread in the East, though the latter is seldom collected; one species, U. albicornis, is northern and extends south in the Appalachians to North Carolina; one species, U. gigas flavicornis, is transcontinental in northern U.S. and Canada; and U. sah, an introduced species, has been recorded only from New Hamp- shire. KEY TO SPECIES — Male Wings black; antennal flagellum partly white with NO some basal and/or apical segments black ..... 3 — Wings yellow, only apical margins may be slightly blackish; antenna yellow, scape and pedicelimaysbeablackaser: set sean ane 5 3. Abdomen mostly red, sometimes basal and/or apical segments blackish Abdomen black, only cornus may be orange .. 4 Be Geet cressoni Norton 4. Cornus orange, contrasting with black abdo- men; fore- and midlegs usually all black .... taxodii (Ashmead) — Cornus black; basal half of tibiae and basal half of tarsi of each leg whitish yellow albicornis (Fabricius) 5. Yellow on head continuous across top, at most separated by a narrow black line at center of postocellar area; pronotum and upper half of mesepisternum yellowish orange (legs mostly yellow with hindfemur and apical two-thirds of lnidaChilloylan IRC). 5 o5c550ne acne sah (Mocsary) — Yellow on head separated into a spot on each side by a black band usually as broad as dis- tance between eyes; thorax black gigas flavicornis (Fabricius) 6. Head mostly yellow to yellow orange; wings yellowish (legs usually with hindfemur and apical two-thirds of hindtibia black; apical ab- dominal segment may be black) . . sah (Mocsary) — Head largely black, with a broad black band separating yellow spots on each side of head; Winessblackish or hyaline sel enienene ree q 7. Abdomen reddish, may be black at base, but basically unicolorous; wings black ........ 8 — Abdominal segments | and 2 or | to 3, and 7 to apex black, segments 2 or 3 to 7 red to or- AINSS VIS lowell Sig oe ooo bn bale Ba boo: 9 St lessiblacks Wetas secant. e cressoni (Norton) — Fore- and midtibiae and tarsi dark orange; bas- al third of hindtibia and basal half of hindbas- itarsus white taxodii (Ashmead) 9. Hindbasitarsus 4.0—5.5 < longer than broad gigas flavicornis (Fabricius) — Hindbasitarsus 6.5—8.0 X longer than broad albicornis (Fabricius) Urocerus albicornis (Fabricius) Distribution.—British Columbia, Califor- nia, Georgia, Idaho, Louisiana, Michigan, Minnesota, Missouri, Montana, New Bruns- wick, Newfoundland, New Mexico, New York, North Carolina, Nova Scotia, Ontar- i0, Oregon, Pennsylvania, Rhode Island, Virginia, Washington, West Virginia. Virginia records.—Montgomery Co., VII-8-74; Arlington, VI-10-1952; Wash- ington Co., Bristol, 7-24-72. Hosts.—Abies amabilis, A. balsamea, A. fraseri, A. lasiocarpa; Larix laricina, L. oc- cidentalis; Picea engelmanni, P. mariana, P. sitchensis; Pinus spp.; Pseudotsuga men- ziesil; Thuja plicata; Tsuga heterophylla (Smith 1978). Remarks.—Some of the Virginia records may represent emergence from imported lumber or firewood in buildings; the species is probably native to only the higher ele- vations. For the life cycle, see Belyea (1952). Amman (1969) reared this species from Fraser fir, Abies fraseri, in North Car- olina. Blackman and Stage (1918) reared it from Larix laricina in New York and gave notes on its life history; it was found in dying or recently felled conifers. Urocerus cressoni Norton (Figs. 2, 6, 16) Distribution.—Florida, Georgia, lowa, Minnesota, Nebraska, New Brunswick, North Carolina, Nova Scotia, Ohio, Ontar- i0, Quebec, Virginia, Wisconsin. Virginia records.—Falls Church, IX-2- 17; Montgomery Co., VIII-9-1974. Collection records.—VIRGINIA: Essex Co., VH-27—VHUI-9-91 (1), VITI-27-IX-16- 91 (1); VU-3—16-96 (1); VH-17—VII-2-96 (2); VII-18—VIHI-1-97 (4); VIII-2—18-97 (2); IX-9—X-6-97 (2); VIH-4—21-98 (3); VITI-22-IX-11-98 (3); IX-12—X-5-98 (1); VOLUME 104, NUMBER 1 VII-17—VIII-2-99 (3); VIII-3—20-99 (3). Fairfax Co., [IX-16—22-90 (1). Hosts.—Abies balsamea, A. fraseri; Pi- cea sp.; Pinus rigida, P. taeda (Kirk 1974, Smith 1979). Remarks.—Most all collections in Essex Co., VA, were from traps in or adjacent to mixed coniferous hardwood forests. The predominant conifers were Pinus taeda and P. virginiana. All collections were from July to early October (Fig. 16). Sixteen of the 26 specimens from this collection had no white behind the eyes or a very small and faint white spot slightly lighter than the black on the rest of the head; thus, they could be confused with Sirex if the female cornus and male hind tarsus are not checked carefully. Bradley (1913) recognized several vari- eties based on the amount of red on the ab- domen and wing darkness: (1) abdomen en- tirely red and wings dark fuliginous; (2) basal six dorsal segments of abdomen dark brown; and (3) abdomen red with a black band on the third, fourth, and fifth dorsal segments. All specimens collected would belong to variety (2), except three had the basal 5 segments black, one had the basal 7 segments black, and one had only the bas- al 3—4 segments black. In a number of spec- imens, there is a central red mark on the second segment. Amman (1969) reared this species from Fraser fir, Abies fraseri, in North Carolina. Urocerus gigas flavicornis (Fabricius) Distribution.—Alaska, Alberta, Arizona, California, Colorado, Idaho, Montana, New Mexico, Northwest Territories, Ontario, Oregon, Washington, Wyoming, Yukon Territory. Hosts.—Abies concolor, A. lasiocarpa, Larix occidentalis, Picea engelmanni, P. sitchensis, Pinus contorta, Pseudotsuga menziesii (Middlekauff 1960, Kirk 1975). Remarks.—The North American form is considered a subspecies of the Eurasian Urocerus gigas (Linnaeus). It is transcon- 185 tinental in Canada, but we have not seen specimens from eastern United States. Urocerus sah (Mocsary) Distribution.—North Africa, Asia Minor and the Near East, southern Russia; New Hampshire. Hosts.—Probably Abies spp., Picea spp.., Pinus spp. Remarks.—This species was first record- ed in North America by Smith (1987) from several specimens taken in New Hamp- shire. No other specimens have been dis- covered. It is close to the European Uro- cerus augur (Klug) and sometimes has been treated as a subspecies of it. Urocerus taxodii (Ashmead) (Fig. 16) Distribution.—District of Columbia, Florida, Mississippi, Missouri, Virginia. Virginia records.—City of Virginia Beach, Cape Henry, VI-8-24. Collection records.—VIRGINIA: Essex Co., V-14—24-91 (1). Host.—Taxodium 1979). Remarks.—It was unusual to find a spec- imen in Essex Co., VA. No Taxodium were anywhere near the trap in which it was col- lected. distichum (Smith Genus Xeris Costa Xeris spectrum (Linnaeus) (Pigss 728) Distribution.—Holarctic; Alaska, Arizo- na, British Columbia, California, Colorado, Connecticut, Maine, Michigan, Montana, Nevada, New Brunswick, New Hampshire, Nova Scotia, Oregon, Utah, Washington. Hosts.—Abies concolor, A. lasiocarpa, Larix occidentalis, Picea engelmanni, P. pungens, Pinus contorta, P. ponderosa, Pseudotsuga menziesii. Pinus contorta 1s the favorite host in California (Middlekauff 1960, Kirk 1975). Remarks.—This species 1s transcontinen- tal in Canada and northern United States. 186 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON The most southern records in the East are Connecticut and Michigan. Adults of this species mate at the highest points on moun- tain tops. Many specimens were hand col- lected at the top of Mount Rigaud, Quebec (H. Goulet, personal communication). Subfamily Tremicinae Genus Tremex Jurine Tremex columba (Linnaeus) (Fig: 15) Distribution.—Alabama, Arizona, Ar- kansas, California (one record from River- side Co.), Colorado, Connecticut, Georgia, Florida, Illinois, Indiana, Iowa, Kansas, Maryland, Massachusetts, Michigan, Min- nesota, Mississippi, Missouri, Montana, Nebraska, New Brunswick, New Jersey, New Mexico, New York, Nova Scotia, Ohio, Ontario, Pennsylvania, Quebec, Ten- nessee, Texas, Utah, Virginia, West Virgin- ia, Wisconsin, Wyoming (Middlekauff 1960, Smith 1979). Virginia records.—Widespread_ through- out the state. We have seen specimens from the following: Appomattox Co., Arlington Co., Bedford Co., City of Norfolk, Clarke Go. Essex Co;, Eamiax Co} Fredericks Co; Giles Co., Hanover Co., Louisa Co., Lou- doun Co., Montgomery Co., Nottaway Co., Roanoke Co., Smythe Co., Warren Co., Washington Co. Collection records.—MARYLAND: Prince George's Co. (4).” VIRGINIA: Clarke Co. (12): Essex (Co: (84): Bairtax. Co. (7); Loudoun Co. (11); Louisa Co. (19). WEST VIRGINIA: Tucker Co. (3). See Fig. 15 for dates of collection. Hosts.—Acer spp., Ulmus spp., Quercus spp., Carya spp., Fagus grandifolia, Fagus spp. (Smith 1979); Celtis laevigata, Car- pinus sp. Also reared or collected from the following: apple, box elder, hackberry, pear, and sycamore (Middlekauff 1960). Proba- bly in other angiospermous trees. Usually in dead or dying trees. Remarks.—This species is known as the “pigeon tremex.” Adults oviposit in dead or weakened deciduous trees or those dying as a result of disease or other cause. Feed- ing of larvae in the wood aids in the dis- integration of trees. Stillwell (1967) noted a minimum two year life cycle in New Brunswick where the primary host was weakened or injured Fagus grandifolia, and occasionally Acer saccharum. Adult emer- gence and oviposition was from August to October, and 2—7 eggs were laid in each oviposition hole. Eggs hatched in 2—4 weeks or hatched the next May or June. Larvae tunnel in wood infected with the fungus Daedalea unicolor, a fungus asso- ciated with each stage of the female devel- opment. Stillwell (1965) also studied the hypopleural organs in larvae and the asso- ciated fungus. Eggs in absence of the fun- gus hatch but larvae do not develop past the first instar. Oviposition tunnels are at a right angle to the bark surface and 2-15 mm deep in the sapwood; eggs are laid at inter- vals along the tunnel. Several hundered eggs are deposited at intervals in the gal- lery. Packed sawdust and frass are in the galleries. Galleries may be 1—2 m long. Pu- pation is in the sapwood or up to 30 cm deep in the heartwood and may be oriented in any direction. The females may tunnel up to | m before emergence. Laurent (1931) found numerous larvae in dead or nearly dead maple during May in Pennsylvania. From a cross section of trunk, 20” long and 10” in diameter, 96 males and 10 females emerged June 16—26; 7 males and 22 females from June 27 to July 6; 5 females and | female from July 7-16; and about the same number through August. There were no specimens after Au- gust. In all, 162 Tremex emerged. Fattig (1949) observed Megarhyssa (Ich- neumonidae) parasitoids in Georgia. From a dead tree, 86 specimens of Megarhyssa emerged during July. From July 29 to Au- gust 22, he collected 28 specimens of 7. columba. He reported his observations on Oviposition of the parasitoid. Harrington (1887) gave some biological notes for 7. columba on maple in Ontario. VOLUME 104, NUMBER 1 Fig. 13. Eriotremex formosanus. This species is extremely variable in col- oration of the body and wings. Because of this, Bradley (1913) recognized several col- or races: (1) Entire body fulvous, legs be- yond femora yellow, and wings dark red- dish brown; (2) abdomen marked with black and yellow with ground color black and wings brown or yellowish; and (3) ab- domen marked with black and yellow with ground color yellow and wings yellow. From collections of 133 specimens, 21% corresponded with variety (1) with the body entirely dark yellowish and the wings dark- ly and almost uniformly black, and 79% corresponded with variety (2) having the abdomen mostly black with yellow bands which vary in size. All had blackish wings, but some were slightly paler than others. This second form (2) is the predominate form in the northeastern states, while the pale form (1) is most common in the south- eastern states. Genus Eriotremex Maa Eriotremex formosanus (Matsumura) (Fig. 13) Distribution.—Alabama, Georgia, Flori- da, Louisiana, Mississippi, North Carolina, South Carolina, Texas, Virginia (Smith 1996). Virginia records.—City of Virginia Beach, Seashore State Park, maintenance area, 30-X-1989. Hosts.—Quercus nigra; Q. phellos; Car- ya sp. Also recorded from dead oak trees, laurel oak, in trunk of Quercus alba, fire- wood of Liquidambar styraciflua, dead hickory, and firewood of water oak. Adult collection records, mostly from label data, 188 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 14. Ibalia anceps, \ateral. are on Quercus laurifolia, Liquidamber styraciflua, Pinus palustris, Pinus taeda, ‘on bark of longleaf pine” (Pinus palus- tris), ““ovipositor imbedded in slash pine” (Pinus elliottii), “‘sitting on Pinus taeda.” Remarks.—See Smith (1996) for distri- bution and spread of this species in the United States. Most adult collection records are from April to June and September to November. Very few have been collected in July, August, December, and February. Eriotremex formosanus may be confused with Tremex columba. In E. formosanus, the antennae are 20—21 segments (14—15 in T. columba), there are dense long golden hairs on the head and body (short and not as bright or grayish in 7. columba), and the color pattern is consistent, with the prono- tum and band on the abdomen yellow and the wings hyaline with their apices infus- cated and with a dark spot below the stigma (Fig. 13) (variable in 7. columba, but not with this color pattern). Ibaliidae Genus /balia Latreille Liu and Nordlander (1992) gave a key to the severn North American species and Liu and Nordlander (1994) keyed the 13 world species. The genus is Holarctic, with two species in the eastern United States. KEY TO SPECIES 1. Abdomen red; head and thorax black pe Men abo aes Tilak iain oY 2 leucospoides (Hockworth) — Orange yellow with black stripes or markings (Fig. 14) anceps Say Ibalia anceps Say (Figs. 14, 15) Distribution.—Arkansas, Colorado, Flor- ida, Georgia, Illinois, Maryland, Massachu- setts, Michigan, Mississippi, Missouri, New Brunswick, New Jersey, New York, Nova Scotia, Ontario, Pennsylvania, Tennessee, Texas, Virginia, West Virginia, Wisconsin. Virginia records.—The only records are from the collections below. VOLUME 104, NUMBER 1 189 36 he —C———— — a | o £ (3) 18 4 —_—— - ee ® a a 2 15 — _— ® a = 3 12 ae - — — 9 = a= = L ——— = — = 3 i 2 = = tl 4 ty Y% Y% yy Y %e YG Fig. 15. and West Virginia. Collection records—MARYLAND: AI- legheny Co. (9); Prince George’s Co. (28). VIRGINIA: Clarke Co. (9); Essex Co. (45); Fairfax Co. (4); Loudoun Co. (7). WEST VIRGINIA: Hardy Co: (7); Tucker Co: (17). All collections are from the end of April to the first of July with the peak flight time from mid-May to mid-June. The two records in September are from Allegheny Co. See Fig. 15 for flight times. Host.—A parasitoid of Tremex columba in deciduous trees (Liu and Nordlander 1994). Remarks.—Adult flight activity is limited to spring, whereas Tremex columba flies from July through October (Fig. 15). Since T. columba takes two or more years to com- plete its life cycle, the /balia possibly par- asitizes the young larvae from eggs laid by the Tremex from the previous year. Liu and Nordlander (1994) recognized variation in wing and body coloration of this species. The three predominate wing color forms are: (1) wings maculate with _ Milbalia anceps 11 Tremex columba Seasonal occurrence of /balia anceps and Tremex columba from collections in Maryland, Virginia, dark and almost clear areas; (2) same, but more or less dark; and (3) evenly dark. The first 1s the most common in the northeastern states, and all specimens collected appear to be of this form (Fig. 14). Ibalia leucospoides (Hochworth) (Fig. 16) Distribution.—Europe; Alberta, Arizona, California, Colorado, Florida, Georgia, Ida- ho, Maine, Maryland, Michigan, Minneso- ta, Mississippi, New Brunswick, North Car- olina, Oregon, Pennsylvania, South Caroli- na, Utah, Virginia, Washington, West Vir- ginia. Virginia records.—City of Virginia Beach, Cape Henry, IX-18-23. Collection records.—VIRGINIA: Essex Co. [V-9-—21-94 (1), IV-12—26-95 (2), X-24—XI-17-95 (1), IX-28—X-23-95 (1), VIII-19-IX-8-97 (1), [IX-9—X-6-97 (3), X- 7-31-97 (1), VIII-4—21-98 (1), VUI-22-Ix- 11-98 (2), IX-12—X-5-98 (3), X-6—XI-7-98 (1); Fairfax Co. VH-12—16-82 (1), VH-6-85 190 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 9 |. leucospoides | NU. taxodii | DU. cressoni | | £3IS. nigricornis | 6 + -— —— - _ EIS. edwardsii |- _ 5 i ® 25 So ZO ro) 3 | at eee ma Ee ie ie ili IL ! Duh ~S s\ @ ss ~ N < < ye wv sy > S&S < SS < ye 3 o a og = Fig. 16. Seasonal occurrence of /balia leucospoides and Sirex edwardsii, S. nigricornis, Urocerus cressoni, and U. taxodii from collections in Maryland, Virginia, and West Virginia. (1), VII-8—14-85 (1), [X-6—12-98 (1); Loui- sa Co. VI-29-85 (1), VI-8-19-89 (1), VI- 22—VII-3-89 (1). Hosts.—A parasitoid of woodwasps of the genera Sirex, Urocerus, and Xeris living in various conifers (Liu and Nordlander 1994). Remarks.—Chrystal (1930) studied this species in Europe. Eggs are inserted in the egg of the host before hatching or on the first larval instar. They hatch after 2-3 months to a year later. The third stage leaves the interior of the host larva and Starts to tunnel toward the surface of the log. Pupation is 5—6 weeks. The complete life cycle is not less than three years. Adults were found from August to October ovi- positing in spruce and larch infested with Sirex cyaneus. Kirk (1974) collected this species from trees infested with Sirex edwardsii (as ab- botii) and S. nigricornis in Alabama, Geor- gia, South Carolina, and Florida. Infested trees included Pinus virginiana, P. taeda, P. clausa, P. elliottii, and P. palustris. In the mid-Atlantic states, adults were collected from April to October, with most from August to October (Fig. 16). There was no clear, narrow emergence time as for Ibalia anceps, and correlations with the emergence times of the host siricids could not be determined. Host LIst FOR SIRICIDAE OF EASTERN UNITED STATES Abies amabilis Douglas ex J. Forges (Pa- cific silver fir)—Urocerus albicornis. Abies balsamea (L.) Mill. (balsam fir).— Sirex cyaneus, S. juvencus, Urocerus al- bicornis, U. cressonti. Abies concolor (Gord. & Glend.) Lindl. ex EH. Hildebr. (white fir).—Szrex cyaneus, S. longicauda, Urocerus gigas flavicor- nis, Xeris spectrum. Abies fraseri (Pursh) Poir. (Fraser fir).— VOLUME 104, NUMBER | Sirex cyaneus, Urocerus albicornis, Uro- cerus cressoni. Abies grandis (Douglas ex. D. Don) Lindl. (grand fir).—Sirex cyaneus. Abies lasiocarpa (Hook.) Nutt. (subalpine fir).—Sirex cyaneus, S. juvencus, Uroce- rus albicornis, Urocerus gigas flavicor- nis, Xeris spectrum. Abies magnifica A. Murray (California red fir).—Sirex cyaneus, S. longicauda. Abies sp. (fir).—Sirex noctilio, Urocerus sah. Acer negundo L. (boxelder).—Tremex col- umba. Acer spp. (maple).—Tremex columba. Carpinus sp. (hornbeam).—Tremex colum- ba. Carya spp. (hickory).—Tremex columba, Eriotremex formosanus. Celtis laevigata Willd. (lowland hackber- ry).—Tremex columba. Cupressus macrocarpa Hartw. ex Gordon (Monterey cypress).—Sirex areolatus, S. behrensii. Fagus grandifolia Ehrh. beech).—Tremex columba. Fagus spp. (beech).—Tremex columba. Juniperus occidentalis Hook. (western ju- niper).—Sirex areolatus. Juniperus scopulorum Sarg. (Rocky Moun- tain juniper).—Sirex areolatus. Larix laricina (Du Roi) K. Koch (tama- rack).—Sirex cyaneus, Urocerus albicor- nis. Larix occidentalis Nutt. (western larch).— Urocerus albicornis, U. gigas flavicornis, Xeris spectrum. Larix sp. (larch).—Sirex juvencus, S. noc- tilio. Libocedrus decurrens Torr. (incense-ce- dar).—Sirex areolatus. Liquidamber styraciflua L. (sweetgum).— Eriotremex formosanus. Picea abies (L.) Karst (Norway spruce).— Sirex edwardsii. Picea engelmanni Parry (Engelmann spruce).—Sirex cyaneus, Urocerus albi- cornis, U. gigas flavicornis, Xeris spec- trum. (American 19] Picea glauca (Moench) Voss (white spruce).—Sirex cyaneus. Picea mariana (Mill.) B.S.P. (black spruce).—Urocerus albicornis. Picea pungens Engelm. (blue spruce).— Xeris spectrum. Picea sitchensis (Bong.) Carr. (sitka spruce ).—Sirex cyaneus, Urocerus albi- cornis, U. gigas flavicornis. Picea sp. (spruce).—Sirex cyaneus, S. ju- vencus, S. nigricornis, S. noctilio, Uro- cerus cressoni, U. sah. Pinus clausa (Chapm. ex Engelm.) Vasey ex Sarg. (sand pine).—Sirex nigricornis. Pinus contorta Douglas ex Loudon (lod- gepole pine).—Sirex areolatus, S. cy- aneus, S. juvencus, Urocerus gigas flav- icornis, Xeris spectrum. Pinus echinata Mull. (shortleaf pine).—Sir- ex edwardsii, S. nigricornis. Pinus elliottii Engelm. (slash pine).—Sirex edwardsii. Pinus jeffreyi Grev. & Balf. (Jeffrey pine).—Sirex areolatus, S. behrensit. Pinus lambertiana Dougl. (sugar pine).— Sirex areolatus. Pinus palustris Mill. Gongleaf pine).—Sir- ex edwardsii, S. nigricornis. Pinus ponderosa Dougl. ex Laws. (ponde- rosa pine).—Sirex behrensii, S. cyaneus, S. longicauda, Xeris spectrum. Pinus radiata D. Don (Monterrey pine).— Sirex areolatus, S. behrensti, S. noctilio. Pinus rigida Mill. (pitch pine).—Sirex ed- wardsil, S. nigricornis, Urocerus cres- SONL. Pinus strobus L. (eastern white pine).—Sir- ex edwardsii, S. nigricornis. Pinus taeda L. (loblolly pine).—Sirex ni- gricornis, Urocerus cressoni. Pinus virginiana Mill. (Virginia pine).— Sirex edwardsii, S. nigricornis. Pinus sp. (pine).—Sirex noctilio, Urocerus albicornis, U. sah, Eriotremex formosan- us’ GD): Platanus occidentalis L. (sycamore ).—Tre- mex columba. Pseudotsuga menziesii (Mirb.) Franco (Douglas-fir).—Sirex areolatus, S. cy- 192 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON is, S. longicauda, S. noctilio, Uroce- us albicornis, U. gigas flavicornis, Xeris spectrum. Pyrus sp. (apple, pear).—Tremex columba. Quercus alba L. (white oak).—Eriotremex formosanus. Quercus laurifolia Michx. (laurel oak).— Eriotremex formosanus. Quercus nigra L. (water oak).—Eriotremex formosanus. . Quercus phellos L. (willow oak).—Eriotre- mex formosanus. Quercus sp. (oak).—Tremex columba, Eri- otremex formosanus. Sequoia sempervirens (D. Don) Engl. (red- wood).—Sirex areolatus. Taxodium distichum (L.) Rich. (baldcy- press).—Sirex areolatus, Urocerus taxo- dii. Thuja plicata Donn ex D. Don (western redcedar).—Urocerus albicornis. Thuja sp. (cedar).—Sirex areolatus. Tsuga heterophylla (Raf.) Sarg. (western hemlock).—Sirex cyaneus, Urocerus al- bicornis. Ulmus spp. (elm).—Tremex columba. ACKNOWLEDGMENTS Most of the specimens on which this study is based are in the National Museum of Natural History, Smithsonian Institution, Washington, DC. We thank the curators of the following other collections examined: University of Arkansas, Fayetteville; Ari- zona State University, Tempe; University of Georgia, Athens; Iowa State University, Ames; University of Kansas, Lawrence; Mississippi State University, Mississippi State; University of Missouri, Columbia; Montana State University, Bozeman; Ne- braska State Museum, Lincoln; University of New Hampshire, Durham; New Mexico State University, Las Cruces; Ohio State University, Columbus; Oregon Department of Agriculture, Salem; Oregon State Uni- versity, Corvallis; Texas A&M University, College Station; Virginia Museum of Nat- ural History, Martinsville; Virginia Poly- technic Institute and State University, Blacksburg; West Virginia University, Mor- gantown; University of Wyoming, Laramie. We thank the following for allowing col- lections on their property: Mr. and Mrs. J. G. Kloke, Louisa and Essex counties, VA; T. J. Henry and D. R. Miller, Hardy Co., WV: R. Turner, Loudoun Co., VA; and M. E. Bowers, University of Virginia Blandy Experimental Farm and State Arboretum of Virginia, Clarke Co., VA. E. M. Barrows allowed study of his collections from Tuck- er Co., WV, and Allegheny and Garrett counties, MD. Cathy Anderson, Systematic Entomology Laboratory, USDA, took the scanning electron micrographs and _ ar- ranged the plates. Thanks are extended to the following for reviewing the manuscript: H. Goulet, Ag- riculture and Agri-Food Canada, Ottawa, and R. Ochoa and M. Pogue, Systematic Entmology Laboratory, USDA, Beltsville, MD, and Washington, DC, respectively. LITERATURE CITED Amman, G. D. 1969. Annotated list of insects infesting bark and wood of Fraser fir. Journal of Economic Entomology 62: 249-250. Bedding, R. A. and R. J. Akhurst. 1974. Use of the nematode Deladenus siridicola in the biological control of Sirex noctilio in Australia. Journal of the Australian Entomological Society 13: 129— 1376 Belyea, R. M. 1952. Death and deterioration of balsam fir weakened by spruce budworm defoliation in Ontario. Canadian Entomologist 84: 325-335. Benson, R. B. 1962. Holarctic sawflies (Hymenoptera: Symphyta). Bulletin of the British Museum (Nat- ural History) Entomology 12(8): 381—409. Blackman, M. W. and H. H. Stage. 1918. Notes on insects bred from the bark and wood of the Amer- ican larch—Larix laricina (Du Roc) Koch. Tech- nical Publication No. 10 of the New York State College of Forestry at Syracuse University, 115 Pp- Bradley, J. D. 1913. The Siricidae of North America. Journal of Entomology and Zoology 5: 1—36, 5 plates. Buchner, P. 1928. Holznahrung und Symbiose. J. Springer. Berlin. . 1965. Siricids, pp. 83-92. Jn Endosymbiosis of Animals with Plant Microorganisms. J. Wiley, N.Y. 909 pp. Revised English version. Cameron, E. A. 1967. Notes on Sirex juvencus cali- fornicus (Hymenoptera: Siricidae), with a descrip- VOLUME 104, NUMBER 1 tion of the male and a key to the California spe- cies of Sirex. 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Smithsonian Institution Press, Wash- ington, D.C., 1198 pp. 1987. Urocerus sah (Mocsary) (Hymenop- 194 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tera: Siricidae) new to North America and key to North American species of Urocerus. Proceedings of the Entomological Society of Washington 89: 834-835. . 1993. Systematics, life history, and diversity, pp. 3-32. In Wagner, M. and K. E Raffa, eds. Sawfly Life History Adaptations to Woody Plants. Academic Press, 581 pp. 1996. Discovery and spread of the Asian horntail, Eriotremex formosanus (Matsumura) (Hymenoptera: Siricidae), in. the United States. Journal of Entomological Science 31: 166-171. Stange, L. A. 1996. The horntails of Florida (Hyme- noptera: Siricidae). Florida Department of Agri- culture and Consumer Services, Division of Plant Industry, Entomology Circular No. 376, 3 pp. Stillwell, M. A. 1964. The fungus associated with woodwasps occurring in beech in New Bruns- wick. Canadian Journal of Botany 42: 495—496. . 1965. Hypopleural organs of the woodwasp larva Tremex columba (L.) containing the fungus Daedalea unicolor Bull. ex Fries. Canadian En- tomologist 97: 783-784. . 1967. The pigeon tremex, Tremex columba (Hymenoptera: Siricidae), in New Brunswick. Ca- nadian Entomologist 99: 685-689. Stivaly€. Es i ledes EB: SchaitzaySs Re -S; Ren- teado, and S. M. S. da Silva, eds. 1993. Confer- encia Regional da Vespa da Madeira, Sirex noc- tilio, Na América do Sul, 1992, Florian6polis. EMBRAPA, Centro National de Pesquisa de Flo- restas, Curitiba, Brazil. 278 pp. Tabata, M. and Y. Abe. 1995. Cerrena unicolor iso- lated from the mycangia of a horntail, Tremex lon- gicollis, in Kochi Prefecture, Japan. Mycoscience 36: 447-450. Tribe, G. D. 1995. The woodwasp Sirex noctilio Fa- bricius (Hymenoptera: Siricidae), a pest of Pinus species, now established in South Africa. Journal of the Entomological Society of Southern Africa 3: 215-217. 1997. Sirex woodwasp expands its range. Plant Protection News, Bulletin of the Plant Pro- tection Institute (Plantbeskermingsnuss) 49: 9-10. Viitasaari, M. 1984. Sahapistidiset 3. Siricicoidea, Orussoidea, ja Cephoidea. University of Helsinki, Department of Agricultural and Forest Zoology Reports 6, 66 pp. Viitasaari, M. and E Midtgaard. 1989. A contribution to the taxonomy of horntails with notes on the genus Sirex Linnaeus (Hymenoptera, Siricidae). Annales Entomologici Fennici 55: 103-110. Webster, E M. 1895. The black horned horntail, Uro- cerus Nigricornis. The Ohio Farmer, August 29, ItsXay, jos 7/7 Westcott, R. L. 1971. New host and distribution re- cords for three western wood-boring Hymenoptera (Syntexidae, Siricidae). Pan-Pacific Entomologist 47: 310. PROC. ENTOMOL. SOC. WASH. 104(1), 2002, pp. 195-210 NEW ALISOTRICHIA (TRICHOPTERA: HYDROPTILIDAE) FROM CENTRAL AND SOUTH AMERICA AND THE GREATER ANTILLES STEVEN C. HARRIS AND OLIVER S. FLINT, JR. (SCH) Department of Biology, Clarion University, Clarion, PA 16214, U.S.A. (e-mail: harris @ mail.clarion.edu); (OSF) Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560-0169, U.S.A. Abstract.—Twelve new species of Alisotrichia, A. panamensis, A. woldai, A. linterna, A. asta, and A. cuernita from Panama, A. neblina and A. viuda from Venezuela, A. muellita from Peru, A. kanukua from Guyana, A. cacaulandia from Brazil, and A. math- isi and A. paxilla from Jamaica are described and illustrated. A new record of A. fi- mouchela Botosaneanu from Venezuela is also provided. Key Words: The genus Alisotrichia is widespread throughout Mexico, Central America, northern South America and the West In- dies. There are currently 33 species and subspecies, including one fossil species from Dominican amber, assigned to the ge- nus (Flint et al. 1999). This paper describes 12 additional species, primarily from Pan- ama, but also from Venezuela, Peru, Guy- ana, Brazil, and Jamaica. Harris and Holzenthal (1993) divided the 32 then known species of Alisotrichia into eight monophyletic species groups. Since then, the three basal groups have been ele- vated to full generic status: the former A. blantoni Group as Mejicanotrichia Harris and Holzenthal, the former A. dominicensis Group as Cerasmatrichia Flint, Harris and Botosaneanu, and the former A. quemada Group as Scelobotrichia Harris and Bueno. Nine of the new species herein described are being placed in the A. orophila Group. Alisotrichia woldai, A. cacaulandia, A. ka- nukua, A. mathisi, A. paxilla, and A. neb- lina agreeing in all characteristics of that group. Alisotrichia panamensis and A. muellita although lacking the dorsal process Trichoptera, Hydroptilidae, Alisotrichia, new species, Neotropics and macroseta of segment VIII, agree with several other species in the group in the possession of the spring-like process of the venter of segment X and together with A. linterna in the ventral process of segment VII and internal structure of the phallus. One species, A. viuda, is tentatively placed in the A. hirudopsis Group based on the two pairs of apical processes from segment VIII. The final two species, A. asta and A. cuernita, based on the presence of elaborate lobes from the dorsolateral margin of seg- ment VIII are placed in the A. chorra Group. Morphological terminology follows that of Marshall (1979). Length is measured from the front of the head to the wing tip and is given as a range if more than one specimen is known. Type material is de- posited in the National Museum of Natural History, Smithsonian Institution, Washing- ton; DC. Alisotrichia panamensis Harris and Flint, new species (Figs. 1-5) In many respects this species is very sim- ilar to A. muellita, n.sp., from Peru and A. 196 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-10. Male genitalia. 1-5, Alisotrichia panamensis. 1, Lateral. 2, Dorsal. 3, Ventral. 4, Phallus, ventral. 5, Phallus, lateral. 6-10, A. muellita. 6, Lateral. 7, Dorsal. 8, Ventral. 9, Phallus, ventral. 10, Phallus, lateral. VOLUME 104, NUMBER 1 woldai, n.sp., also from Panama. In com- mon with these species, as well as A. cir- cinata Flint from Puerto Rico, A. panamen- sis has a single spring-like structure anter- oventrally from segment X and lying within the eighth sternite. Alisotrichia panamensis is distinguished from these other species on the basis of the presence of a elongate ap- icodorsal process from the eighth segment and from A. woldai and A. circinata in lack- ing the dorsal setate process from the eighth sternum. Male.—Length 1.8—1.9 mm. Antenna cream colored, black at tip with 18 seg- ments, scape greatly enlarged and setose. Head and thorax with central mass of white hair dorsally. Forewing with extensive white hairs, black on margin, with diffuse black bands or spots. Abdominal segment VII annular, with bifid ventromesal process. Segment VIII roughly rectangular in lateral view, truncate posteriorly with elongate dorsolateral process, and short, lobate ven- trolateral process; in ventral view longer than wide, with shallow mesal incision dis- tally. Segment IX short, incomplete ven- trally, apparently fused with X dorsally. Tergum X concave posteriorly in dorsal as- pect, anteriorly with pair of slender apode- mes extending through segment VII; highly modified tubular structure attached to ven- ter of these apodemes and elaborately coiled anteriorly (this structure appears to be loosely attached to venter of X, as it will move anteriorly and posteriorly). Phallus wide basally, tubular sheath posteriorly en- closing two pairs of slender lateral rods which originate at the same level, lateral- most rods longer than inner rods; slender ejaculatory duct protruding tongue-like be- yond sheath. Female.—Unknown. Type material—Holotype, ¢: Panama, Barro Colorado Island, Canal Zone, July 1967, W. W. Wirth, Malaise trap. Paratypes: same data as holotype, | d; same, but Sny- der-Molino trail, marker 3, light trap III, 19-25 August 1987, H. Wolda, | d; same, but 25 November—1 December 1987, 1 6; 197 same, but 9-15 December 1987, 1 3; same, but 2—8 November 1988, 1 ¢; same, but 23-29 November 1988, 1 6; same, but | 1— 17 January 1989, 2 3; same, but 29 No- vember—5 December 1989, | 3; same, but 2-8 January, 1991, | 3d; same, but light trap I, 4-10 November 1987, 1 5; same, but 2— 8 January 1988, 1 3d; same, but 28 Decem- ber 1988-3 January 1989, 1 d; same, but 6—12 December 1989, 3 3; same, but 13— 19 December 1989, 2 3d; same, but 3—9 Jan- uary 1990, 1 6. Etymology.—Named for the country of Panama. Alisotrichia muellita Harris and Flint, new species (Figs. 6-10) This is another species of the group pos- sessing a spring-like structure from the ven- ter of segment X. Alisotrichia muellita is very similar to A. panamensis differing pri- marily in the structure of the phallic rods which are united basally into an enlarged plate and the shape of the eighth segment that is conical apically with an elongate process projecting from the inner dorsal margin. Male.—Length 1.7—1.9 mm. Brown in alcohol with no distinguishing patterns. An- tenna with 18 segments, scape greatly en- larged and setose. Abdominal segment VII annular, with bifid ventromesal process. Segment VIII narrowing to rounded apex in lateral view, with elongate process project- ing from inner posterodorsal margin; in ventral view longer than wide, shallow me- sal incision distally. Segment IX short, in- complete ventrally, apparently fused with X dorsally. Tergum X rounded posteriorly, an- teriorly with pair of slender apodemes ex- tending through segment VII; | highly modified tubular structure attached to ven- ter of these apodemes and elaborately coiled anteriorly. Phallus wide basally, tu- bular sheath posteriorly enclosing two pairs of thin lateral rods, lateralmost rods about twice length inner rods, inner rods origi- nating from mesal margins of lateral rods; 198 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON slender ejaculatory duct protruding tongue- like beyond sheath. Female.—Unknown. Type material.—Holotype, ¢: Peru, Ma- dre de Dios, Manu, Pakitza, 11°56’S, ILLS W, Yel..9250.%m,! 12=18\ e September 1989, N. Adams et al., kitchen stream, Mal- aise trap, night collection. Paratypes: same data as holotype, 2 6. Etymology.—Spanish, “‘small spring,” referring to the structure within segment VI. Alisotrichia woldai Harris and Flint, new species (Figs. 11-15) Although similar to A. muellita in the structure of the phallus, and A. panamensis in the shape of segment VIII, A. woldai ap- pears to be more closely related to A. cir- cinata in the presence of seta-bearing pro- cesses from the dorsal margin of segment VIII. In common with all of these species, A. woldai has a spring-like structure within sternum VIII. Alisotrichia woldai is distin- guished from these other species on the ba- sis of the lateralmost pair of phallic rods which arise from a large, rectangular base. Male.—Length 1.4—1.7 mm. Brown in alcohol with no distinguishing patterns. An- tenna with 18 segments, scape greatly en- larged and setose. Abdominal segment VII annular, with bifid ventromesal process. Segment VIII tapering posteriorly to trun- cate apex in lateral view, anterodorsally with seta-bearing process on each side; in ventral view rectanguloid with wide mesal incision distally, forming inwardly hooked lobes laterally. Segment IX elongate dor- sally and apparently fused with X, incom- plete ventrally. Tergum X thin with mesal incision posteriorly, anteriorly with pair of slender apodemes originating from ventro- lateral margin IX + X and extending an- teriorly through segment VII; highly mod- ified spring-like structure attached to the venter of these apodemes. Phallus wide ba- sally, tubular sheath posteriorly enclosing two pairs of sclerotized rods, lateralmost rods longer than inner rods and widened at base, inner rods thin over length and orig- inating from base of lateral rods; slender ejaculatory duct protruding tongue-like be- yond sheath. Female.—Unknown. Type material——Holotype, d: Panama, Barro Colorado Island, Snyder-Molino trail, marker 3, light trap I, 7-13 December 1988, H. Wolda. Paratypes: same data as holo- type, but 21-27 December 1988, 1 ) Nn Chapius, FE 1876. Histoire naturelle des insectes. Gen- era des Coléopteres, Paris 12: 1—424. Crotch, G. R. 1871. List of Coccinellidae. Cambridge, 8 pp. . 1873a. Revision of the Coccinellidae of the United States. 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WASH. 104(1), 2002, pp. 237-239 NOTE New Data on the Structure of the Female Genitalia of Flea Beetles (Coleoptera: Chrysomelidae) Female genitalia of flea beetles have re- ceived much attention during last two de- cades and have become an important source of valuable diagnostic and phylogenetic characters (Cox 1997; Doguet 1994; Duck- ett 1999; Kangas and Rutanen 1993; Kon- stantinov 1987, 1998a, 1998c; Konstanti- nov and Vandenberg 1996; Lingafelter and Konstantinov 2000). They also became a subject of comparative morphological stud- ies (Duckett 1995; Konstantinov 1994, 1998b; Konstantinov and Rusakov 1993). As a result of these studies the structure and variability of the spermatheca, vaginal palpi and tignum are relatively well understood, however the relative position of these struc- tures, especially in regards to the gut, ter- gite 9 and vagina, and the way in which they are connected to each other remained unknown. This note intends to answer the afore- mentioned questions. In addition to stan- dard dissecting techniques (Konstantinov 1998c, Lingafelter and Konstantinov 2000), “Chlorazol Black’ was used to stain the membranes of the genitalia. That procedure revealed that each sclerite (tergite and ster- nite) consists of two membranous layers (Figs. 1A, B): an external layer (Fig. 1B thicker lines) usually more sclerotized in both tergites and sternites; and an internal layer usually much less sclerotized and ful- ly membranous (Fig. 1B thinner lines). This latter layer usually is very poorly visible without staining. In tergites the external, dorsal layer is more sclerotized than the ventral one, whereas in sternites ventral lay- er is external and more sclerotized than the internal, dorsal layer. The ventral layer of sternite 8 also contains a long sclerotized projection called the tignum. The internal membranous layer connects the distal part of the more external sclerite with the prox- imal part of the following sclerite, thus en- abling the genitalia to protrude telescopi- cally out of the abdomen. The maximum distance to which genitalia can be protruded depends upon the length of the internal lay- ers of segment 7, the external and internal layers of segments 8 and 9, and the distance between the base of the vaginal palpi and apex of tergite 9. The internal layer of ter- gite 8 forms at least the dorsal side of the gut. It also connects with the dorsal layer of tergite 9 which is situated only slightly below the dorsal side of the gut and which lateral sclerotizations are situated laterally of anus. The internal membrane of tergite 9 also extends into the ventral and _ lateral walls of the gut. The membranous ventral wall of the gut bends anteriorly and contin- ues on the dorsal side of the vagina. The vaginal palpi are formed by the invagina- tion of the dorsal side of the vagina, so their cavity opens in the body cavity. The ventral side of the vagina is a continuation of the internal layer of sternite 8. In addition the external and internal layers of tergite and sternite 8 are laterally connected to each other. The connections of the internal layers of segment 8 form lateral folds which con- tinue on the side of the vagina. This observation yields the rather unex- pected result that the vagina is formed by membranous parts of both segments 8 and 9. In order to avoid this interpretation, we would have to assume that tergite 9 1s ab- sent in flea beetles and everything between the external layer of tergite 8 and the vagina belongs to the internal layer of tergite 8. Although this latter explanation is the most parsimonious it should be tested by embry- ological or comparative morphological studies. Until then the question of the ho- 238 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON B Fig. 1. Female genitalia of Pseudodera xanthospila Baly. A, Three dimensional drawing with right side of several sclerites removed. B, Diagrammatic sagittal section. Abbreviations: an = anus; est7 = external layer of sternite 7; est8 = external layer of sternite 8; et7 = external layer of tergite 7; et8 = external layer of tergite 8; et9 = external layer of tergite 9; ist? = internal layer of sternite 7; ist8 = internal layer of sternite 8; it7 = internal layer of tergite 7; it8 = internal layer of tergite 8; it? = internal layer of tergite 9; gt = gut; spd = spermathecal duct; tg = tignum: vg = vagina; vp = vaginal palpus. VOLUME 104, NUMBER 1 mology of the vaginal palpi remains unan- swered. I am grateful to A. Norrbom (Systematic Entomology Laboratory, USDA, ARS, Washington, DC), A. Tishechkin (Depart- ment of Entomology, Louisiana State Uni- versity, Baton Rouge, LA), and anonymous reviewer for comments on the earlier ver- sions of the manuscript. LITERATURE CITED Cox, M. 1997. Homichloda barkeri (Jacoby) (Cole- optera: Chrysomelidae: Alticinae) a candidate agent for the biological control of prickly acacia, Acacia nilotica (Mimosaceae) in Australia. Jour- nal of Natural History 31: 935-964. Doguet, S. 1994. Coleopteres Chrysomelidae, Vol. 2. Alticinae. Faune de France, Paris, 681 pp. Duckett, C. N. 1995. Variation in reproductive struc- tures. Chrysomela Newsletter 29: 4—5. . 1999. A preliminary cladistic analysis of the subtribe Disonychina with special emphasis on the series Paralactica (Chysomelidae: Galerucinae: Alticini), pp. 105-136. In Cox, M. L., ed. Ad- vances in Chrysomelidae Biology 1. Backhuys Publishers, Leiden, 671 pp. Kangas, H. and I. Rutanen. 1993. Identification of fe- male of the Finnish species of A/tica Miller (Co- leoptera, Chrysomelidae). Entomologica Fennica 42(2): 115-129. Konstantinov, A. S. 1987. On the morphological struc- tures used for identification of females of Altica (Coleoptera, Chrysomelidae). Zoologicheski Zhurnal 54(1): 42—SO [in Russian]. 1994. Comparative morphology and some evolutionary trends in flea-beetles (Alticinae), pp. 383-391. In Jolivet, P. H., M. L. Cox, and E. Pe- titpierre, eds. Novel Aspects of the Biology of the Chrysomelidae. Kluwer Academic Publishers. Dordrecht/Boston/London, 582 pp. 239 . 1996. Review of Palearctic species of Crepi- dodera Chevrolat (Coleoptera: Chrysomelidae: Alticinae). Spixiana 19(1): 21—37. . 1998a. Revision of the Aphthona crypta group of species and a key to species groups in Aphthona Chevrolat (Coleoptera: Chrysomelidae: Alticinae). Coleopterists Bulletin 52(2): 134-146. . 1998b. On the structure and function of the female genitalia in flea beetles (Coleoptera: Chry- somelidae: Alticinae). Proceedings of the Ento- mological Society of Washington 100(2): 353— 360. . 1998c. Revision of the Palearctic species of Aphthona Chevrolat and cladistic classification of the Aphthonini (Coleoptera: Chrysomelidae: AI- ticinae). Memoirs on Entomology, International 11: 429 pp. Konstantinov, A. S. and A. M. Rusakov. 1993. Com- parative morphological study of the female geni- talia of Chrysomelinae (Coleoptera, Chrysomeli- dae). Vestnik BGU, Biology 2: 18-21 [in Rus- sian]. Konstantinov, A. S. and N. J. Vandenberg. 1996. Handbook of Palearctic flea beetles (Coleoptera: Chrysomelidae: Alticinae). Contributions on En- tomology, International 1(3): 237—439. Lingafelter, S. W. and A. S. Konstantinov. 2000. The monophyly and relative rank of Alticine and Gal- erucine leaf beetles: A cladistic analysis using adult morphological characters (Coleoptera: Chry- somelidae). Entomologica Scandinavica 30(4): 397-416. Alexander S. Konstantinov, Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Ag- riculture, Yo U.S. National Museum of Nat- ural History, Smithsonian Institution, Washington, D.C. 20560-0168, U.S.A. (e-mail: akonstan@ sel.barc.usda. gov) PROC. ENTOMOL. SOC. WASH. 104(1), 2002, pp. 240-241 NOTE Reinstatement of Rhithrogena manifesta Eaton (Ephemeroptera: Heptageniidae) The name Rhithrogena manifesta Eaton, 1885, was based on certain adult heptagen- iid mayfly specimens collected from Rock Island, Illinois in the early 1860s by B. D. Walsh. Walsh (1862) misidentified those specimens as Baetis debilis Walker (a spe- cies now in the genus Paraleptophlebia Lestage). Eaton (1885), as was his practice at the time, did not designate type speci- mens for his species, and any de facto syn- types of R. manifesta have been heretofore undocumented. Walsh’s collections, in gen- eral, were deposited in the Chicago Muse- um (subsequently destroyed in the Great Chicago fire of 1871) or with H. Hagen in Prussia (subsequently placed in the Muse- um of Comparative Zoology, Harvard Uni- versity [MCZ]) (see Burks 1953). Rhithro- gena manifesta was placed with species in the genus Heptagenia Walsh by Mc- Dunnough (1924), who originally believed the genus Rhithrogena Eaton was equiva- lent with Heptagenia. Although Mc- Dunnough (1926) later recognized some species in Rhithrogena, Eaton’s manifesta has continued to be regarded as Heptagenia manifesta. This is because McDunnough (1924) suggested that H. manifesta was re- lated to particular species of Heptagenia, which more recently have been transferred to the genus Nixe Flowers. Flowers (1980) chose not to place H. manifesta in Nixe un- til more was known about the species. The only other report of the occurrence of the species was from Quebec by Walley (1927). McCafferty (1996) treated H. manifesta as a nomen dubium because of the many un- certainties surrounding it. Recently, we located and studied four adult specimens in the MCZ that Walsh had collected from Rock Island in 1860 and 1863 and identified as B. debilis. These specimens were consistent with Eaton’s de- scription of manifesta. We also discovered and studied a series of 15 adult specimens from New Brunswick and Quebec in the Canadian National Collection of Insects, Agriculture and Agri-Food Canada (CNC) that had been identified as H. manifesta by McDunnough. We found the CNC speci- mens to be the same species as those in the MCZ and thus correctly identified by McDunnough. Our study of these two series of specimens also revealed important addi- tional data regarding the species. It was clear that the species belongs in the genus Rhithrogena, as Eaton (1885) had originally surmised, not in Heptagenia. Also, whereas the species fits the rather incomplete de- scription given by Eaton (1885), the species also fits the more completely defined and relatively well-known species R. pellucida Daggy (Daggy 1945, Burks 1953, Leonard and Leonard 1962, McShaffrey and Mc- Cafferty 1988, Durfee and Kondratieff 1994). Rhithrogena pellucida is a relatively widespread North American species that also is known from the vicinity of the type locality of Eaton’s species (Randolph and McCafferty 1998, Durfee and Kondratieff 1994). Our study of comparative material showed it to be the same as H. manifesta. Based on the above observations, four steps are now required to remediate the tax- onomic and nomenclatural problems asso- ciated with H. manifesta. First, we replace the species to its proper genus, as Rhithro- gena manifesta Eaton (recombination). Sec- ond, we place R. pellucida as a subjective junior synonym of R. manifesta (new syn- onym). Third, we designate a lectotype from the Rock Island material collected by Walsh (see material examined below) for the purpose of fixing the identity of R. man- ifesta. Fourth, we remove R. manifesta from nomen dubium status. It should be noted VOLUME 104, NUMBER 1 that although Eaton (1885) listed R. mani- festa as a renaming, R. manifesta was tech- nically a new name. Walsh (1862) did not describe B. debilis as new. Thus, no sec- ondary homonym of that name should be construed, as might be incorrectly inter- preted from the synonymy given by Eaton (1885) for R. manifesta. Material examined.—Rhithrogena mani- festa: lectotype, 1 d adult, Hlinois, Rock Island, 1863, Walsh [MCZ]. Other material: 1 3 adult, Hlinois, Rock Island, Walsh [MCZ]: 1 2 adult, Ilinois, Rock Island, 1860, Walsh [MCZ]; 1 @ adult, Illnois, Rock Island, Walsh [MCZ]; 1 6 adult, 6 @ adults, New Brunswick, Fredericton, VII- 1928, W. J. Brown [CNC]; 1 @ adult, Que- bec, Cascades Point, 30-VIII-1930, L. J. Milne [CNC]; 2 ¢ adults, same data but 26- VII-1930 (one set genitalia on slide) [CNC]; 1 2 adult, Quebec, Kirk’s Ferry, 4- VII-1925, G. S. Walley [CNC]; 1 @ adult, Quebec, Lachine, 6-VIII-1924, G. S. Wal- ley [CNC]; 2 6 adults, Quebec, Laprarie, PINVMEeN925, iP ide [CNC]; a2 tadult, Quebec, Richelieu, 5-VIH-1927, G. S. Wal- ley [CNC]. Rhithrogena pellucida: 1 ¢& adult, 1 & adult, Indiana, West Lafayette, 13-VII- 1974, H. R. Lawson [Purdue Entomological Research Collection, West Lafayette, Indi- ana]. We thank R. D. Waltz (Indianapolis, In- diana) and R. W. Flowers (Tallahassee, Florida) for comments on an early stage of the manuscript, and we thank A. V. Pro- vonsha (West Lafayette, Indiana) for hand- delivering specimens from the MCZ. This study has been funded in part by CanaColl Foundation grant 178 to LMJ and NSF grant DEB-9901577 to WPM. This paper has been assigned Purdue ARP Journal No. 16505. LITERATURE CITED Burks, B. D. 1953. The mayflies, or Ephemeroptera, of Illinois. Illinois Natural History Survey Divi- sion Bulletin 26: 1-216. 241 Daggy, R. H. 1945. New species and previously un- described naiads of some Minnesota mayflies (Ephemeroptera). Annals Entomological Society of America 38: 373-396. Durfee, R. S. and B. C. Kondratieff. 1994. New ad- ditions to the inventory of Colorado mayflies (Ephemeroptera). Entomological News 105: 222— 22k Eaton, A. E. 1883-88. A revisional monograph of re- cent Ephemeridae or mayflies. Transactions of the Linnean Society of London, Second Series Zool- ogy 3: 1-346. Flowers, R. W. 1980. Two new genera of Nearctic Heptageniidae (Ephemeroptera). The Florida En- tomologist 63: 296-307. Leonard, J. W. and F A. Leonard. 1962. Mayflies of Michigan trout streams. Cranbrook Institute of Science Number 43. Bloomfield Hills, Michigan. 139 pp. McCafferty, W. P. 1996. The Ephemeroptera species of North America and index to their complete no- menclature. Transactions of the American Ento- mological Society 122: 1—54. McDunnough, J. 1924. New Canadian Ephemeridae with notes, II. The Canadian Entomologist 56: 90-98, 113-122, 128-133. . 1926. Notes on North American Ephemerop- tera with descriptions of new species. The Cana- dian Entomologist 58: 184—196. McShaffrey, D. and W. P. McCafferty. 1988. Feeding behavior of Rhithrogena pellucida (Ephemerop- tera: Heptageniidae). Journal of the North Amer- ican Benthological Society 7: 87-99. Randolph, R. P. and W. P. McCafferty. 1998. Diversity and distribution of the mayfies (Ephemeroptera) of Illinois, Indiana, Kentucky, Michigan, Ohio, and Wisconsin. Ohio Biological Survey Bulletin New Series 13: i—vii, 1-188. Walsh, B. D. 1862. List of the Pseudoneuroptera on Illinois contained in the cabinet of the writer, with descriptions of over forty new species, and notes on their structural affinities. Proceedings of the Academy of Natural Sciences of Philadelphia 8: 361—402. Walley, G. S. 1927. Ephemeroptera. Annual Report of the Entomological Society of Ontario 57: 59—61. Luke M. Jacobus and W. P. McCafferty, Department of Entomology, Purdue Uni- West Lafayette, IN 47907-1158, U.S.A. (e-mail: luke-jacobus @ entm.purdue. edu) versity, PROC. ENTOMOL. SOC. WASH. 104(1), 2002, pp. 242-243 NOTE Variation in Abdominal sa2 and sa3 Setation in Larvae of Arctopsyche grandis (Banks) (Trichoptera: Hydropsychidae) Arctopsyche grandis (Banks) is the most abundant and widespread of the western Nearctic arctopsychine caddisflies, and the larvae are often collected in streams from 45—2,150 m in elevation (Givens and Smith 1980). Wiggins (1996) separated known larvae of Arctopsyche from larvae of the genus Parapsyche by the setation of the ab- dominal sa2 and sa3 positions. Parapsyche species bear a tuft of several long setae or scale hairs in each position, whereas Arc- topsyche species bear a single, long, hair- like seta in each position, occasionally ac- companied by one or two shorter setae. To quantify the extent of variation in the abdominal setation in A. grandis, I exam- ined larvae from four Rocky Mountain riv- ers: the Clark Fork River in Montana, the Arkansas and Eagle rivers in Colorado, and the Red River in New Mexico. Benthic ma- croinvertebrate samples for ecological stud- ies had been taken from 1994 to 1996 at 8— 14 individual sites in each river. A total of 660 A. grandis larvae were examined, and the number of setae in each of the sa2 and sa3 positions of abdominal segments I-VII was counted. The geometric mean number of setae per position was calculated, and these data were analyzed by general linear methods analysis of variance (GLM-ANOVA) using NCSS (Number Cruncher Statistical Systems 1997) to determine if statistical differences existed in setation between abdominal seg- ments, setal positions, or river systems. A 95% significance level (a = 0.05) was used for all tests. HZ 3 setae __| 2setae Mi 1 seta 0 setae Abdominal Segment 100 80 72) S c2000) eee een aie lane | eet ss ee eee INSTRUCTIONS FORJAUTHORS feigns ena ee NU Se raer ee mage te ease fora tre tar ety aera SH | 201 106 The: 51 91 164 160 174 221 24 138 244 242 240 VOL. 104 APRIL 2002 NO. 2 Ol (ISSN 0013-8797) a PROCEEDINGS of the PUBLISHED QUARTERLY ; CONTENTS BEAULIEU, FREDERIC and TERRY A. WHEELER—Insects (Diptera, Coleoptera, Lepidoptera) reared from wetland monocots (Cyperaceae, Poaceae, Typhaceae) in southern (OSE OSC aU has SALA as ORD Ri a SEE AIA aaa A ASRS AT Weg en ae Ye PU RS 300 BROWN, JOHN W., JORGE PENA, THERESA VASQUEZ, and JOAQUIN BAIXERAS— Description of a new tortricid pest (Lepidoptera: Tortricidae: Olethreutinae) of litchi (Litchi chinensis) in Florida, with a review of tortricid pests of litchi worldwide .................... 318 CANO, ENIO and MIGUEL ANGEL MORON—Adéditions to Phyllophaga subgenus Chlaenobia (Coleoptera: Melolonthidae: Melolonthinae) from Guatemala ......................0.020 ese 352 CARVALHO, CLAUDIO JOSE BARROS de and MARCIA SOUTO COURI—Cladistic and bio- geographic analyses of Apsil Malloch and Reynoldsia Malloch (Diptera: Muscidae) of south- GruVS OUUNVATMOnI Cal Malesia tite ena Um Roh rata rtiare ie mere easins leet tie heitte aa eine IMS bate cits, kha tevare 309 DIKOW, TORSTEN and WAYNE N. MATHIS—A revision and phylogenetic study of Actocetor Becker (Diptera: EH phy drmdacy isis. iver ee ein Guay eRe GIM Ny cae ia ee atte wales acho bel eeerats 249 EISNER, THOMAS, JAMES E. CARREL, EILEEN VAN TASSELL, E. RICHARD HOEBEKE, and MARIA EISNER—Construction of a defensive trash packet from sycamore leaf tri- chomes by a chrysopid larva (Neuroptera: Chrysopidae) ................ sec cece cece eee eee eee 437 GOEDEN, RICHARD D.—Description of immature stages of Tephritis stigmatica (Coquillett) (Diptera dephritidae) een) yo Nae, bal arh sey enti atSeis yeters ape RNE plese eib dim slat anette ayes aicte eseenl a eerete ccolaial et ec S55 GOEDEN, RICHARD D.—Life history and description of immature stages of Oxyna aterrima (Doane) (Diptera: Tephritidae) on Artemisia tridentata Nuttall (Asteraceae) in southern (CeINATOTigeWREY Tope Semis Oe apie n see Ml Re Mtb seep esteC OBL, 315 ANTI han AA ee EN Aba) Or Be tate ya 510 HALL, JASON P. W—A review of Chalodeta Stichel with a revision of the chelonis group Mbericl option: A CIMIGAS Los AN ee aah Pepin atiniah Re atta enn eftnin nes! Me hls gaan a Binet Sos alt 376 HARBACH, RALPH E. and THERESA M. HOWARD—Sabethes (Peytonulus) paradoxus, a new Species on sabethini (Diptera: Culicidae) from Pandas 2.5 .\, jens -)arcle sole esalele acl eleis.« njel-ieiel siecle 363 KEIPER, J. B. and S. C. HARRIS—Biology and immature stages of Ochrotrichia footei (Trichoptera: Hydroptilidae), a new microcaddisfly from a torrential mountain stream ...... 2911 (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON OFFICERS FOR 2002 GABRIELA CHAVARRIA, President MiIcHAEL G. PoGug, 7reasurer JONATHAN R. MAwps Ley, President-Elect RonALD A. OcHoa, Program Chair Stuart H. McKamey, Recording Secretary STEVEN W. LINGAFELTER, Membership Chair Ho..is B. WiLLiaMs, Corresponding Secretary Joun W. Brown, Past President Jon A. Lewis, Custodian Davip R. Smitu, Editor Publications Committee RAYMOND J. GAGNE THOMAS J. HENRY Wayne N. MartuHis Honorary President Louise M. RUSSELL Honorary Members Kart V. KROMBEIN RONALD W. HopGEs DonaLp M. ANDERSON WILLIAM E. BICKLEY All correspondence concerning Society business should be mailed to the appropriate officer at the following address: Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Wash- ington, D.C. 20560-0168. MEETINGS.— Regular meetings of the Society are held in the Natural History Building, Smithsonian Institu- tion, on the first Thursday of each month from October to June, inclusive, at 7:00 P.M. Minutes of meetings are published regularly in the Proceedings. MEMBERSHIP.—Members shall be persons who have demonstrated interest in the science of entomology. Annual dues for members are $25.00 (U.S. currency). PROCEEDINGS.—The Proceedings of the Entomological Society of Washington (ISSN 0013-8797) are pub- lished quarterly beginning in January by The Entomological Society of Washington. POSTMASTER: Send address changes to the Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Washington, D.C. 20560-0168. Members in good standing receive the Proceedings of the Entomo- logical Society of Washington. Nonmember U.S. subscriptions are $60.00 per year and foreign subscriptions are $70.00 per year, payable (U.S. currency) in advance. Foreign delivery cannot be guaranteed. All remittances should be made payable to The Entomological Society of Washington. The Society does not exchange its publications for those of other societies. PLEASE SEE PP. 247-248 OF THE JANUARY 2002 ISSUE FOR INFORMATION REGARDING PREPARATION OF MANUSCRIPTS. STATEMENT OF OWNERSHIP Title of Publication: Proceedings of the Entomological Society of Washington. Frequency of Issue: Quarterly (January, April, July, October). Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash- ington, D.C. 20560-0168. Editor: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Books for Review: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 9 April 2002 Periodicals Postage Paid at Washington, D.C. and additional mailing office. PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PROC. ENTOMOL. SOC. WASH. 104(2), 2002, pp. 249-290 A REVISION AND PHYLOGENETIC STUDY OF ACTOCETOR BECKER (DIPTERA: EPHYDRIDAE) TORSTEN DIKOW AND WAYNE N. MATHIS (TD) Universitat Rostock, Germany; National Museum of Natural History, Research Training Program, Smithsonian Institution, Washington, D.C. 20560-0169, USA (e-mail: torsten.dikow @stud.uni-rostock.de); (WNM) Department of Systematic Biology, Ento- mology Section, NHB 169, Smithsonian Institution, Washington, D.C. 20560-0169, USA (e-mail: mathis. wayne @nmnh.si.edu) Abstract.—The genus Actocetor Becker is revised, including a phylogenetic analysis of the seven recognized species. Two new species, Actocetor afrus (Liberia and Senegal) and A. yaromi (Ethiopia), are described. Actocetor hendeli de Meijere, A. margaritatus (Wiedemann), and A. panelii Frey are new junior synonyms of A. indicus (Wiedemann), and A. beckeri de Meijere and A. elegans Hendel are new junior synonyms of A. nigrifinis (Walker). Lectotypes, all females, are designated for the following species: Notiphila indica Wiedemann, Ephydra margaritata Wiedemann, Opomyza nigrifinis Walker, and Actocetor beckeri de Meijere. The cladistic analysis is based on 29 morphological char- acters and resulted in 10 equally most parsimonious trees (length of 60 steps and consis- tency and retention indices of 0.83 and 0.83 respectively). Three trees then resulted from application of successive weighting, and from these a strict concensus tree was derived that is typologically identical to one of the original 10 trees. From the consensus clado- gram, the following hypotheses can be made: (1) Actocetor is monophyletic and (2) the subgenera Actocetor Becker (4 species) and Poecilostenia Bezzi (3 species) are each monophyletic. Although we suggest an Afrotropical origin for Actocetor, the genus could have resulted from speciation events in the Oriental Region. Keys for the identification of all genera of Discomyzini and the species of Actocetor are provided. The distribution of all species of Actocetor and the biology of A. indicus from a coastal site in Israel are discussed. Key Words: Actocetor, Discomyzini, Ephydridae, phylogeny Actocetor Becker, which comprises species with spotted wings and beautifully colored Shore flies are attractive to study because they are “‘... a family of flies in the full flower of its evolution” (Oldroyd 1964: 189). Although appealing for research, many nonspecialists consider the adults to be rather drab, being nondescript, usually dark colored, and tiny. There are a few large shore-fly species (>5 mm), and others are comparatively pale colored, but with few exceptions, the drab generalization holds. One notable exception is the genus bodies. Like most shore-fly species, how- ever, specimens of Actocetor are generally small, with body lengths of usually less than 3.5 mm. This paper is a revision of this distinctive genus. Although specimens of Actocetor are easily recognized by their unique coloration and spotted wings, the species have never been treated comprehensively, and no key 250 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON levels is available for all included species. Lack- ing adequate identification tools, some spe- cies were described more than once, some- times from single females. The objectives of this revision are: (1) to clarify and re- describe all known species, (2) to investi- gate the phylogenetic relationships among them, (3) to summarize what we know about their biology, distribution, and bio- geography, and finally (4) to present a key and illustrations for their identification. A key to the genera of the tribe Discomyzini Acloque is also presented. Actocetor is known only from the Old World where the genus occurs primarily in the tropical zones of the Afrotropical and Oriental Regions. The most widely occur- ring species, A. indicus (Wiedemann; senior synonym of A. margaritatus (Wiedemann)), which is found in the Afrotropical, Oriental, and southern Palearctic Regions, is relative- ly well known because of its beautiful col- By ea Mling lon Color habitus of Actocetor indicus (Wiedemann). oration and spotted wings. Not surprisingly, this species has been reported relatively of- ten in the literature (see species’ synony- my). The literature on the other species is meager at best, and virtually nothing about their natural history is recorded in the lit- erature. The nomenclatural history of Actocetor as a genus 1s mostly confined to the 20th Century when Becker (1903) first proposed the genus with A. margaritatus (= A. in- dicus) as its type species. The only generic synonym is Poecilostenia Bezzi (1908), which has been given generic or subgeneric status with the latter being more common in recent treatments and catalogs (Wirth 1955, Cogan 1980, Mathis and Zatwarnicki NSIS). The nomenclatural history at the species level, however, is more convoluted, with the type species being an example. Wiede- mann (1830) named the type species in the VOLUME 104, NUMBER 2 genus Notiphila, apparently without realiz- ing that six years earlier (1824) he had de- scribed the same species as Ephydra indica. This same species was described a third time (Hendel 1917) but using a preoccupied name, A. beckeri. De Meijere (1924) rec- ognized the preoccupation and proposed A. hendeli as a replacement name. Frey (1958b) described this species yet again, but as a subspecies, A. margaritatus panellti, from specimens collected on the Cape Verde Islands. Two other species, A. nigri- finis (Walker) and A. decemguttatus (Bez- zi), have similar histories, each having been described three times. Specimens of these three species are the most common in col- lections. The nomenclatural histories of the remaining four species of Actocefor are rel- atively uncommon, and correspondingly, not encumbered with synonyms. METHODS AND MATERIALS The descriptive terminology, with the ex- ceptions noted in Mathis (1986) and Mathis and Zatwarnicki (1990a), follows that pub- lished in the Manual of Nearctic Diptera (McAlpine 1981). Because specimens are small, usually less than 3.5 mm in length, study and illustration of the male terminalia required use of a compound microscope. We have followed the terminology for most structures of the male terminalia that other workers in Ephydridae have used (see ref- erences in Mathis 1986, and Mathis and Za- twarnicki 1990a, 1990b), such as surstylus, which in Dryxini is divided into a presur- stylus (surstylus) and postsurstylus (clasp- er). Zatwarnicki (1996) has suggested that the pre- and postsurstylus correspond with the pre- and postgonostylus and that the su- bepandrial plate is the same as the medan- drium. The terminology for structures of the male terminalia is provided directly on Figs. 30-32 and 37-38 and is not repeated for comparable illustrations of other spe- cies. Species’ descriptions are composite and not based solely on the holotypes. One head and two venational ratios that are used in the descriptions are defined below (all 251 ratios are based on three specimens: the largest, smallest, and one other. Gena-to- eye ratio is the genal height measured at the maximum eye height/eye height. Costal vein ratio is the straight line distance be- tween the apices of R,,, and R,,./distance between the apices of R, and R,,;. M vein ratio is the straight line distance along vein M between crossveins dm-cu and r-m/dis- tance apicad of dm-cu. The phylogenetic analysis was performed using Hennig86©, a computerized algo- rithm that produces cladograms by parsi- mony. Character data were polarized using outgroup procedures. Although autapomor- phies were not included in the cladistic analysis (they were made inactive), which would skew the consistency and retention indices, we listed them on the cladogram and included them as part of generic treat- ments and phylogenetic considerations to document the monophyly of the lineages, particularly at the generic level. Although most specimens are in the Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. (USNM), we also borrowed and studied nu- merous specimens from the following mu- seums: AMNH- American Museum of Natural History, New York, USA. The Natural History Museum (formerly the British Museum (Natural History)), London, Eng- land, United Kingdom. Personal collection of Silvano Canzoneri, Venezia, Italy, now in Museo Regionale di Scienze Na- turali (MRSN), Torino, Italy. Canadian National Collection, Ottawa, Canada. Muséum National d’ Histoire Na- turelle, Paris, France. Musée Royal de l’Afrique Cen- trale (Koninklijk Museum voor Midden Afrika), Tervuren, Bel- gium. BMNH CANZ CNC MNHN MRAC 252 MRSN_ Museo Regionale di Scienze Na- turali, Torino, Italy. MZH Museum Zoological Helsinki, Helsinki, Finland. NMSA _ Natal Museum, Pietermaritzburg, South Africa. NMW _ Naturhistorisches Museum, Wien, Austria. NMWL._ National Museum of Wales, Car- diff, Wales, United Kingdom. SMN Staatliches Museum fiir Natur- kunde, Stuttgart, Germany. TAU Tel-Aviv University, Tel-Aviv, Israel. UZMC _Zoologisk Museum, Copenhagen, Denmark. ZX. Zoological Institute, Lund Uni- versity, Lund, Sweden. ZMAN Instituut voor Taxonomische Zoologie, ZoGlogisch Museum, Universiteit van Amsterdam, The Netherlands. ZMHB Zoologisches Museum, Hum- boldt Universitat, Berlin, Germa- ny. SYSTEMATICS TRIBE DISCOMYZINI Discomyzini Acloque 1897: 486. Type ge- i) . Vein R,,, lacking a subapical stump vein nus: Discomyza Meigen 1830. KEY TO GENERA OF DISCOMYZINI N Ww Vein R,,, bearing a subapical stump vein . . . Vein R,,; basad of crossvein r-m bearing at most 1—2 tiny, black setulae; wing with irreg- ular, transverse, wide white and brown bands; tibiae yellowish with brown rings; abdomen with maculate pattern of gray and brown mi- crotomentum Eremomusca Mathis Vein R,,; basad of crossvein r-m bearing 5— 6 setae; wing generally dark with pattern of white spots; tibiae and femora unicolorous, lacking bands; abdomen mostly shiny, lacking maculate pattern Trypetomima de Meijere . Wing hyaline or with anterior margin dark- Enedsbut nOMsSpoOtted ye: aa eee ee 5 Wing generally dark with a pattern of white SPOES eerie are tay machete tao s. aie on ae eee eae 4 . Facial setae 3—5 but usually 4, all well de- veloped; Ist flagellomere short, length sube- qual to height of pedicel; pseudopostocellar . Alula weakly developed, very narrow . Pseudopostocellar PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON setae as well developed as large, proclinate, fronto-orbital seta; legs unicolorous Se CRO Ce ME. ene oie Actocetor Becker Facial setae 2, only dorsal seta well devel- oped; Ist flagellomere longer than height of pedicel; pseudopostocellar setulae distinctly smaller than large proclinate fronto-orbital seta; tibia or tarsus generally paler colored than femur . Guttipsilopa Wirth, subgenus Guttipsilopa Alula well developed, earlike . Intrafrontal seta present; fronto-orbital setae 4 (anterior 2 setae proclinate, 3rd lateroclinate, 4th lateroreclinate) Paratissa Coquillett Intrafrontal seta absent; fronto-orbital setae 2— . Supra-alar seta well developed, length sub- equal to postalar seta .... Rhysophora Cresson Supra-alar seta moderately well developed, length about half postalar seta . Arista bearing 3—5 dorsal rays; facial setae 2, both inserted on ventral 3rd of face; vein Ry, ; basad of crossvein r-m bearing 3—4 black se- tulae; legs unicolorous Hostis Cresson Arista bearing 7—9 dorsal rays; facial setae sometimes 2 but usually 4 (becoming larger dorsally; sometimes with larger gap between dorsalmost seta and those ventrad) in a ver- tical row that extends dorsad to midfacial height; vein R,,; basad of crossvein r-m bare; legs dark with yellow tarsus, apical tarsomere dark Guttipsilopa Wirth, subgenus Nesopsilopa Mathis and Wirth . Prescutellar acrostichal seta absent; katepis- ternal seta 1; facial setae 3—5, all small and poorly developed; face conspicuously and deeply, transversely rugose; only the reclinate fronto-orbital seta well developed METRE NOT ETOROER TS ROTA 1G be Discomyza Meigen Prescutellar acrostichal seta present; katepis- ternal setae 2, dorsal seta larger; facial setae 2, dorsal pair cruciate, ventral pair poorly de- veloped; face at most with shallowly im- pressed, transverse striae; at least 1 proclinate fronto-orbital seta in addition to reclinate seta well developed +, ,., with costa: celll n.-. with subcircular, medial spot, in line with crossvein dm-cu, and an apical spot (some specimens with a trace of a 3rd sub-basal spot); discal cell with a wide, U-shaped, basal spot (apical arm sometimes constrict- ed) and a subapical, transversely oval-rect- angular spot, sometimes divided into 2 cir- cular spots; cell m with a transversely oval- rectangular spot near middle; cell cua, with a large, basal rectangular white and a sub- apical, transversely trapezoidal spot, vein CuA, straight; costal vein ratio 0.54—0.72; M vein ratio 0.84—1.0. Halter stem yellow; knob whitish yellow and distinctly clubbed. Legs generally whitish yellow to yellow; forecoxa and base of femora sometimes with whitish gray to gray microtomentum laterally. Abdomen (Fig. 28-38): Mostly shiny, gray microtomentum almost always present and as follows: base of Ist tergite with thin band; 4th tergite with wide, anterior band; sometimes with thin, lateral bands on 3rd and 5th tergites or with medial spot on 3rd tergite; color generally metallic, dark blue to black, often with yellow to yellowish red PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON at anterior and/or posterior margins or sometimes entirely yellowish red. 3rd and 4th sternites rectangular, broad, 1-2 lon- ger than wide; 5th sternite of male (Fig. 30) trapezoidal with medial weakness or slight indentation on concave posterior margin. Male terminalia (Figs. 31—38): epandrium (Figs. 31-32) higher than wide; presursty- lus (Fig. 33) large, ventromedial surface with a small emargination, bearing 2 round to oval lobes medially; postsurstylus (Fig. 34) much longer than wide, bearing 3—4, apical setulae, apex bilobed with a small, digitiform, medial, pointed process; post- surstylar process (Fig. 35) distinctly spatu- late, rounded process, shaped like an halter; aedeagus (Fig. 37) as a subrectangular ring basally from which a lateral phalange ex- tends apically; aedeagal apodeme (Fig. 38) in lateral view subtriangular with angle at attachment with base of aedeagus extended and acutely formed; postgonite in ventral view as a sickle shaped process, broader to- ward attachment with aedeagal apodeme; pregonite short, simple, bearing 2 setulae apically; hypandrium broadly and deeply pouchlike. Type material.—The lectotype 2 of No- tiphila indica Wiedemann, here designated to stabilize and make more universal the use of this name, is labeled *““TYPE [red]/ Notiphila indica Wied. Ind[ia]. orient. [handwritten]/Notiphila indica Wied. Det. B.H. Cogan 1976. [species name and “*76” handwritten]/Actocetor indicus (Wied.) Det. B.H. Cogan 1976. [species name and “76” handwritten|/LECTOTYPE @ Notiphila indica Wiedemann By Dikow & Mathis [all except “LECTOTYPE” and, (By, ghand- written; black submarginal border].’? The lectotype is directly pinned, is in fair con- dition (the specimen is greasy and the pin bears some verdigris), and is deposited in the UZMC. The lectotype ° of Ephydra margaritata Wiedemann, here designated to stabilize and make more universal the use of this name, is labeled ‘‘[a small pink square]/ Egypten [handwritten]/margaritata [hand- VOLUME 104, NUMBER 2 written] Coll. Winth. [black margin along longer sides of label|/Ephydra margaritata W Aegyptn. [handwritten]/Type [red ]/LEC- TOTYPE @ Ephydra margaritata Wiede- mann By Dikow & Mathis [all except “LECTOTYPE” and “By” handwritten; black submarginal border].’> The lectotype is pinned directly, is in fair condition (sev- eral setae missing), and is deposited in the NMW. A 4, presumably a syntype, is la- beled “‘Egypten [handwritten]/margaritata [handwritten] Coll. Winth./margaritata W Egyptn [handwritten].”” and is deposited in the NMW. A ¢ paratype of Actocetor margaritatus panelii Frey from the type locality was ex- amined, including structures of the dissect- ed male terminalia. The holotype ¢ of Actocetor beckeri Hendel (replaced with A. hendeli by de Meijere) is labeled “‘Algoa bay Capland 22 3 96 [22 Mar 1896; handwritten] Dr. Brauns./Actocetor Beckeri, n. sp. [2 black submarginal borders; handwritten]/viola- ceus nom. nov. det. Hendel [all except “‘det. Hendel”’ handwritten|/Coll. Hendel/HO- LOTYPE ¢ Actocetor beckeri Hendel [red submarginal border; species name, author, and gender handwritten]”’. The holotype is double mounted (minuten in a cardboard card), is in poor condition (both wings are missing and several setae are broken), and is deposited in the NMW. Other specimens examined.—AFRO- TROPICAL. ANGOLA. Bruco, 26 Feb—2 Mar 1972 (46, 42; BMNH); Cachoeiras (20 mi SW Gabela), 18-19 Mar 1972 (1°; BMNH); Rio Curoca (7 mi N Rio Alexan- dre), 25-26 Feb 1972 (26, 22; BMNH); Rio Giraul (10 mi NE Mocamedes), 27—29 Feb 1972 (12; BMNH); Santa Clara, 19 Mar 1972 (12; BMNH): Tundavala (8 mi NW Sa da Bandeira), 27—29 Mar 1972 (1°; BMNH). BOTSWANA. River Semowane, 23-24 Apr 1972 (16, 12; BMNH). CAMEROON. Kribi (Rt. N7; beach), 28-29 Nov 1987, A. Freidberg (29; USNM). 265 CAPE VERDE ISLANDS. Sao Vincen- te: Rib. Juliao, 26 Nov—2 Dec 1953, Lind- berg (1d; USNM). Sao Jorge dos Orgaos (alfalfa field), Jun 1986, A. van Harten (36, 12; NMWL). DEMOCRATIC REPUBLIC OF CON- GO. Rumonge (Urundi), 1932, A. Lestrade (12; MRAC). ERITREA. Ailet (from egg-pods), 22 Jul 1956, D. J. Greathead (36, 12; BMNH); Massawa (15 mi N, from egg-pods emerged), 20 Mar 1954, Natural Resources Institute Coll (1d, 12; BMNH). ETHIOPIA. Lake Langano, 1989. A: Freidberg, USNM). GAMBIA. Bakau, Cape St. Mary (at Sun Wing Hotel, swept in vegetation along beach), 5 Nov 1977, L. Cederholm, R. Dan- ielsson, O. Hammerstedt, K-J. Hedqvist, G. Samuelson (16, 22; BMNH, ZIL); Bakau at Tropic Bungalow (swept in meadow, rich in flowers, at beach), 4 Nov 1977, L. Ced- erholm, R. Danielsson, O. Hammerstedt, K- J. Hedqvist, G. Samuelson (1¢; BMNH); Jul-Sep 1927 (on ground-nut plots) (1d; BMNH). Central Banjol (3 km NW; gar- den), 21 Feb 1977, L. Cederholm, R. Dan- ielsson, O. Hammerstedt, K-J. Hedqvist, G. Samuelson (1°; ZIL). Central Banjol (3 km NW; in vegetation along mangrove), 21—22 Feb 1977, L. Cederholm, R. Danielsson, O. Hammerstedt, K-J. Hedqvist, G. Samuelson (12; ZIL). Kotu Stream (3 km SW Bakau; in vegetation), 23 Nov 1977, L. Cederholm, R. Danielsson, O. Hammerstedt, K-J. Hedqvist, G. Samuelson (2¢; ZIL). IVORY COAST. Abidjan, Riviera near Golf Club (05°10'N, 04°00'W; scrubland near edge Ebrie lagoon), 21 Apr 1989, J. G. H. Londt (16, 42; NMSA). Banco Na- tional Park, N Abidjan (05°22’N, 04°03'W; edge of wide track in forest), 23-27 Apr 1989, J. G. H. Londt (16; NMSA). Fresco (15 km W; 05°06'N, 05°43'W; airfield/for- est edge, near Palmindustrie), 24 Apr 1989, J. G. H. Londt (16; NMSA). KENYA. Biretwo (40 km E Eldoret), 12 May 1991, A. Freidberg, F Kaplan (1°; 3) Dee FE Kaplan (id: 266 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON hypandrium Gara. 1 wai ih 32 | postsurstylus 31 / presurstylus postsurstylar process 5" sternite aedeagal apodeme postgonite 38 — aedeagus 25; oF Figs. 30-38. Structures of the male preabdomen and terminalia of Actocetor indicus. 30, Fifth sternite and hypandrium, ventral view (only left side showing setae). 31, Epandrium, cerci, presurstylus, postsurstylus, gonite, and hypandrium, lateral view. 32, Same, posterior view. 33, Presurstylus, posterior view. 34, Postsurstylus, lateral view. 35, Postsurstylar process, posterior view. 36, Same, posteroventral view. 37, Aedeagus, aedeagal apodeme, dorsal view. 38, Aedeagal apodeme, lateral view. Scale bar equals 0.3 mm for Figs. 30-32 and 0.1 mm for Figs. 33-38. VOLUME 104, NUMBER 2 USNM). Bungoma, 6—7 Nov 1983, A. Freidberg (12; USNM). Isolo (5 km S), 30 Nov 1986, A. Freidberg (1 2; TAU). Mom- basa (beach) 5 Dec 1989, A. Freidberg, EF Kaplan (22; TAU, USNM). Mombasa (10 km N), 18 Aug 1996, A. Freidberg (1d; USNM). Mombasa (20 mi S; Malaise trap), 23-25 “Jan: 1968, .K..-V. ‘Krombeinjf Pes: Spangler (1 2; USNM). Nairobi (50 km SE; Rt. A109), 30 Apr 1991, A. Freidberg, F Kaplan (12; TAU). Rift Valley, Ol Arabe Gorge, 11 Nov 1988, R. K. Butlin (1d, 19; NMWL). Tsavo West, Ngulia Lodge, 16— 17 Aug 1983, A. Freidberg (1d, 39; USNM). Tseikuru, Jan 1957 (12; USNM). Mombasa, 9 Dec 1951, Deutche Ost Afrika Exps(lO3SMN). MALAWI. Monkey Bay, Lake Malawi, 15 Aug 1943, R. C. Wood (12; BMNH). NAMIBIA. Gross Barmen_ Resort (22°07'S, 16°42’E; roadside grass and dry river bed at camp), 29 Mar 1984, J. G. H. Londt, B. Stuckenberg (16; NMSA). NIGERIA. Ile-Ife, 25 Apr 1969, J. T. Medler (12; USNM). Lagos (shore), 15 Dee 1987, ‘A. Freidberg (2d, 22;.TAU): Lagos, Ikoyi, 14 Jan 1966, J. C. Deeming (12; CNC). Lagos, Victoria Island (shore), 15 Dec 1987, A. Freidberg (36, 12; TAU). Niger State, Mariga River (80 km W Mina), 11 Dec 1987, E Kaplan (1 2; USNM). Yan- kori, 8-14 Nov 1987, P. Neuenschwander (12; TAU). Zaria, Samaru, May—Sep 1979, J. C. Demming (26, 42; NMWL). SENEGAL. Dakar, 4—5 Oct 1978, J. For- tin, G. Hevel (1d; USNM). Brin (3 km SSE), Iguinchor (11 km SW), 9 Nov 1977, L. Cederholm, R. Danielsson, O. Hammer- stedt, K-J. Hedqvist, G. Samuelson (1°; ZX): SEYCHELLES. La Digue> La Passe (4220. 84S;55"49:8'E), 14° May * 1997). V. Hollmann (22; ZMHB); La Réunion G22 887Sh8S5°4916'E),. 14 May. 1997, We Hollmann, W. N. Mathis (56, 102; USNM, ZMHB). Mahé: Airport, 7-8 Apr 1986, W. N. Mathis (26, 22; USNM); Anse aux Pins (4°41.4’S, 55°31.7'E), 2 Apr-3 May 1986, 1997, V. Hollmann, W. N. Mathis (14, 1°; 267 USNM, ZMHB): Anse Soleil (4°44.8’S, 55°27.9'E), 19 May 1997, W. N. Mathis (16, 12; USNM); Beau Vallon, 24 Mar 1965, W.T. Tams, I.B. Nye (1d; BMNH); Police Bay (4°48.0'S, 55°31.3’E), 16 May 1997, V. Hollmann, W. N. Mathis (8d, 59; USNMr-ZMEHB). (Praslin: tAnsesEazio (4°17.6'S, 55°42.1'E), 8-13 May 1997, W. N. Mathis (12¢, 62; USNM); Anse Kerlan (4°18.3'S, 55°41.1'E), 9 May 1997, W. N: Mathis (12; USNM); Anse Kerlan Farm G218:5'S,4 55°42" E) 13 eMay 1997 7a: Hollmann (12°; ZMHB); Anse Lazio (4°17.6'S, 55°42.1’E), 8-13 May 1997, V. Hollmann, W. N. Mathis (206, 998; USNM); Baie Ste. Anne, Anse Takamaka (4°19.6’S, 55°46.3'E), 10-13 May 1997, V. Hollmann (2¢; ZMHB); Fond de L Anse (42047 S555"°43!5'E) 711 May 1997. W. Ne Mathis (16; USNM). SIERRA LEONE. Freetown (6 km SW; 9°26'N, 13°16’W; in garden), 21 Nov 1993, L. Cederholm, R. Danielsson (14; ZIL). SOKOTRA ISLAND. Hadibu Plains, 14 Dec 1898, W. R. O. Grant (1¢; BMNH). SOUTH AFRICA. Eastern Cape: Algoa Bay, 22 Mar 1896, Brauns (1d holotype of A. hendeli; NMW); East London (3227Dd; 5 m; coastal dunes), 16 Mar 1972, M. E. and B. J. Irwin (12; NMSA); The Haven (3228Bb; coastal dunes), 24—28 Jun 1979, R. Miller, P Stabbins (1d; NMSA). KwaZulu-Natal: Dukuduku Forest (4 mi W St. Lucia; 2832Ad; 3 m), 26 Nov 1971, M. E. and B. J. Irwin (22; NMSA); Dukuduku Forest (E Mtubatuba; 2832Ad), 21 Jul 1973, M. E. Irwin (12; NMSA); Dukuduku (between St Lucia and Matubatuba), 7—8 Apr 1960, B. and P Stuckenberg (2; NMSA, USNM); Durban, 27 Feb 1927, L. Bevis (12; BMNH); Gillitts (Pinetown Dis- trict), 28 Dec 1961, B. and P. Stuckenberg (12; NMSA); Jamisons Drift (Tugela Riv- ef), 120Apr 1974; Me E: Irwin (1d; NMSA); Kosi Bay Estuary (2632Dd; indig- enous bush area), 16—19 Mar 1982, D. A. Barraclough (5d, 12; NMSA); Lebombo Hills, H. A. Junod (10, 12, 1EX; BMNH); Mtubatuba, 24—25 Mar 1968, P. J. Spangler 268 (23, 12; USNM); Ndumu Reserve (Ing- wavuma District), Tongaland, 1-10 Dec 1963, B. and P. Stuckenberg (12; NMSA); Near Lilani (Ahrens District), Apr 1962, B. and P. Stuckenberg (1d; NMSA); Oribi Gorge Reserve, Umzinkulwana Valley, 21— 28 Nov 1960, B. and P. Stuckenberg (1d; USNM); Port St. Johns, 20—25 Nov 1961, B. and P. Stuckenberg (1d; USNM); Salt Rock (28°29'S, 31°15'E; 10 m; caravan park), 5=127Ochil9ol. J. (GH Londti(lex; NMSA); Scottsburgh, 15 Nov 1963, B. and P. Stuckenberg (16, 32; NMSA); St. Lucia Park, 7-8 Oct 1983, A. Feidberg (1°; USNM); Tugela Ferry (20 km W; 2830Ga; Malaise trap), 26—27 Feb 1977, R. Miller (12; NMSA); Umkomaas (NE; 3030Bb; along sand dunes), 21 Jun 1980, R. Miller, R. Stabbins (1d; NMSA); Umkomaas, South Coast, 11 Oct 1983, A. Freidberg (45, 82; USNM); Umlalazi Nature Re- serve (2831Dd, dune forest and edges), 2— 10 Oct 1982, J. G. H. Londt (12°; NMSA); Umlalazi Nature Reserve (28°57’S, 31°40'E; 20 m; dune forest), 28-29 Jan L9Ssh IMG. EeswondtsGes 7/25 tNMSA): Umlalazi Nature Reserve (1.5 km E Mtun- zini; 1831Dd; coastal dune vegetation), 30 Dec 1978, R. Miller (12; NMSA); Umla- lazi Nature Reserve (1.5 km E Mtunzini; 1831Dd; coastal indigenous vegetation), 27 Jan 1979, R. Miller (1d; NMSA); Umlalazi Nature Reserve (1.5 km E Mtunzini; 1831Dd; coastal indigenous vegetation), 24-25 Mar 1979, R. Miller (16; NMSA); Widenham Umbilo, 16 Dec 1914, L. Bevis (1d, 12; BMNH). Mpumalanga: Ofcolaco, Selati River (2430Ab), 7-8 Dec 1976, R. Miller (12; NMSA). Northern Province: Entabeni For. Station, Zoutpansberg Range (2230Cc; grassland), Jan 1975, B. Stuck- enberg (12; NMSA). SUDAN. Um Baghot (among hantot 7), 22 Aue 93 |. RECA ME Darling) (ice BMNH). TANZANIA. Tarangire National Park (36°10’E, 03°50'S), Dec 1994, D. Grimaldi (1d; AMNH). UGANDA. Kilembe, Ruwenzori Range PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (4500 ft), BMNH). ZIMBABWE. Gwaai, 14 Oct 1926, R. H. R. Stevenson (12; NMSA). Harare, 9 Jun 1938, A. Cuthbertson (1°; BMNH). Country unknown. Torina (Deutsch Ost Afrika Expedition), 4-18 Mar 1952 (19; USNM). ORIENTAL. INDIA. Karnataka: Mudi- gere, 6 Apr 1980, W. N. Mathis, A. Freid- berg (12: USNM); Tamil Nadu: Palni Hills (Kodaikanal; 6500 ft), Mar-Apr 1953, P. S. Nathan (1d; USNM). MALAYSIA. Penang (at light trap), 16 Jul 1957, H. T. Pagden (12; BMNH). SRI LANKA. Central Province: Nuwara Eliva: Horton Plains, 23 Apr 1980, W. N. Mathis, T. Wijeshine, L. Jayawickrema (22; USNM):; Southern Province: Ham- bantota: Palatupana Tank, 6 Feb 1975, K. V. Krombein (12: USNM). PALEARCTIC. UNITED ARAB EMIR- ATES. Abu Dhabi: Ruwais, 6 Nov 1985, M. J. Ebejer (12; NMWL). CANARY ISLANDS. Gran = Canaria: Las Plamas (on garden wall), 12 Nov 1927, Kisluik (36, 12; USNM); Las Palmas, R. Stora (lex; BMNH); 1-15 May 1901 (16; ZMHB). Tenerife: Laguna (6—700 m), 8—30 Jun 1904 (12; ZMHB); Puerto de la Cruz (dry sand on shore), 21—26 Apr 1979, C. E. Dyte (26, 82; BMNH); Puerto Orotava (low herba and brass near beach), 20 Dec— 2 Jan 1901 (60, 62; ZMHB); Jul 1907, Ca- brera (12; ZMHB). EGYPT. Heliopolis, 20—22 Oct 1964, E. Hargreaves (ex; BMNH). Egypten (12 ho- lotype, 1d; NMW). Sinai: Bir Zrir, 2 Sep 1970, J. Kugler (12; USNM); El-Arba’in (Sinai Mountains), 14 Jul 1974, E Kaplan (12; USNM); Danav, 23 May 1981, W. N. Mathis (12; USNM); El ’Arish, 9 Jan 1917, E. E. Austen (19°; BMNH); Ein Furtaga, 7 Jul 1969, Kugler (26, 32; USNM); Ein Hudra, 9 Jul 1969, A. Freidberg (22, lex; USNM); Et-Tur, 25 Jun 1968 (46, 198; TAU); Feiran, 9 Apr 1973, D. Furth (1d; TAU); Nuweiba, 14 May 1981, T. Furman (42, 56; USNM); Ofira (sewage), 21 May Dec 1934-Jan 1935 (lex: VOLUME 104, NUMBER 2 1981, A. Freidberg, W. N. Mathis (11d, 82, lex; TAU, USNM); Ofira, 22 Mar 1981, A. Freidberg (1 29; TAU); Ophira Air- ports) 22isApr: 1980; 9B: UR. - Pitkin (19; BMNH); Ras Mamlach, 11-15 Apr 1980, B. R. Pitkin (66, 42; BMNH); St. Kathar- ina, 12 Jul 1969, J. Kugler (12; USNM); Taba (1 km S), 10 Apr 1980, B. R. Pitkin (12; BMNH); Wadi Feiran, 25 May 1971, A. Freidberg (26, 22; TAU, USNM); Wadi Hibran, 11 May 1973, A. Freidberg, E Kap- lan (1d, 12; TAU); Wadi Watir, 5 Aug 1975, A. Freidberg (16; TAU). ISRAEL. Arava Valley: Hazeva Field School (30°46.8'N, 35°14.6'E; 2 km N; Na- hal Gidron; 110 m), 11 Mar—7 Jun 1995, M.E. Irwin (1d, 142; TAU); Hazeva, She- zaf Nature Reserve (30°46'’N, 35°15.4’E; north of water treatment plant, low sandy hummocks in small wadi; 80 m), 24 Mar— 20 May 1995, MLE. Irwin (36, 42; TAU); Hazeva (30°46.3'N, 35°16.3'E; Hahal Sha- haq, between agricultural fields; Malaise trap; 110 m), 15 Mar—16 Apr 1995, M.E. Irwin (106, 122; TAU); ?Iddan (30°48.9'N, 35°16.8'E; nahal running east of date palm orchard; 110 m; malaise trap), 13 Mar—5 Jun 1995, MLE. Irwin (216, 222; TAU); ’Iddan Springs (30°49'N, 35°17’E; in damp seep east of spring; 116 m), 13-17 Mar 1995, MLE. Irwin (32; TAU); ’Ir Ovot (30°56.8'N, 35°04.4'E; 3 km NW; water tank on hill- side), 6 Apr 1995, MLE. Irwin (26, 12; TAU). Ashgelon, 5 Sep 1960, Fatal (2d, 52; TAU, USNM). Biq’at Bet Zayda, 3 May-5 Aug 1973, 1986, A. Freidberg, I. Nussbaum (1d, 12; TAU). ’En Yahav, 20 Sep 1995, A. Freidberg (10d, 22; TAU, USNM). ’Enot Qane [’En Turaba], 30 Apr 1973p. Shurth (Gexs- TAU). “Herzliyya (beach; Malaise trap), 13 Jan—9 Nov 1975, 1981, 1982, 1994, A. Freidberg, FE Kaplan, D. Simon, I. Susman (31¢, 569; TAU, USNM). Mash’abbé Sade, 21 Aug 1986, A. Freidberg (1¢; TAU). Nahal Hatira, 19 Oct 1983, I. Nussbaum (12; TAU). Ne’ot ha- Kikkar, 20 May 1974, A. Freidberg (39, 13; TAU). Ne’ot Semadar, 21 Jun—29 Aug 1995, A. Freidberg (46,42; TAU, USNM). 269 Paran, 19 Sep 1977, A. Freidberg (26; TAU). Park HaYarden, 30 Sep 1982, FE Kap- lan (1d; TAU). Qalya, 20 Mar 1980, J. Ku- slere(. ose TAU) = GezironIZ8mAig1986n Nussbaum (12, 16; TAU). Ramat-ha- Sharon, 18 Aug 1975, D. Simon (1d; TAU). Rehovot, 30 Oct 1931, 1938, J. Aharoni (36; TAU, ZMHB). Tel Aviv, 22 Aug—27 Nov 1969, 1971, J. Kugler (132; TAU, USNM). Tel Aviv, Abu-Kabir, 13 Sep 1953, L. Fishelsohn (1d; TAU). Upper Galilee, Park HaYarden, 5 Aug 1986, W. N. Mathis (26; USNM). Yotvata, 23 Sep 1962, J. Ku- gler (1d, 42; TAU). Ze’elim, 6 Dec 1976, A. Freidberg (12; TAU). Zin Wilderness, Nahal Zin at ’En ’Aqrabbim (35°09'N, 39°E; cane-covered, sandy wadi; 61 m; Malaise trap), 27 Mar—15 May 1995, M.E. Irwin (QG222.2 TAD). OMAN. As-Seefa (coastal sand dunes), 4 Dec 1987, J. J. Ebejer (16, 12; NMWL). Ghuzayn (date culture), 10 Apr 1985, Pad (1d; ZIL). Hazm (date palm grove beside Fort on maize, sorghum, and grasses), 19 Oct 1990, M. D. Gallaghler, J. C. Deeming (32; NMWL). Mazara (date cult), 6 Apr 1985, Pad (1d, 52; ZIL). Muscat, 10 Apr 1985, BP Ard6é (26, 22; ZIL). Qurm (hotel garden), 5 Apr 1985, Pa6 (1d, 52; ZIL). Ruwi, Wattayeh, 1-7 Apr 1988, M. D. Gal- lagher (66, 62; NMWL). Viti Wadi, 7 Apr 1985, Padé (10, 12; ZIL). Wadi Bani Kha- rus (foothill of Jebel Akhdar Lilyah; 810 m; 2331 IUINGS 7-401 E) eliSeOctelO9O0s IME: Gallaghar, J. C. Deeming (1d; NMWL). SPAIN. Almeria, Oct 1973, Boness (10d, 62; BMNH). Distribution (Fig. 14).—Afrotropical: Angola, Botswana, Cameroon, Cape Verde Islands (Sao Vincente), Democratic Repub- lic of Congo, Eritrea, Ethiopia, Gambia, Ivory Coast, Kenya, Namibia, Nigeria, Rwanda (literature), Senegal, Seychelles (La Digue, Mahé, Praslin), Sierra Leone, Sokotra Island, South Africa (Eastern Cape, KwaZulu-Natal, Mpumalanga, Northern Province), Sudan, Tanzania, Uganda, Zim- babwe. Oriental: India (Karnataka, Tamil Nadu), Malaysia, Sri Lanka. Palearctic: Ca- 270 nary Islands (Gran Canaria, Tenerife), Egypt, Israel, Oman, Spain, United Arab Emirates (Abu Dhabi). This is the most widespread species of the genus, and its oc- currence at some sites may represent intro- ductions. We suspect this to be the case for the Seychelles and perhaps elsewhere, such as the Indian Subcontinent and Malaysia. Natural history.—On islands of the Sey- chelles, the majority of specimens collected by Mathis were associated with sandy areas with sparse to no vegetation that were sur- rounded by or in close proximity to grass covered habitats. Occasionally this species was found associated with organic debris and other garbage that had accumulated at the high tide mark on sandy beaches. Larvae of this species apparently feed on varied sources of highly organic debris, probably as saprophages. Dr. Amnon Freid- berg (personal communication) successfully reared this species on a medium of domes- tic cat dung in Israel, and Stower et al. (1958) and Greathead (1963) reared adults in Eritrea from damaged egg pods of desert locusts (Schistocerca gregaria Forskal). Observations on mating behavior.—Date: 16 Aug 1986 (Saturday). Time: 10:30 am. Weather: Temperature about 30°C, very slightly breezy, sunny. Location: Beach at Herzliyya, Israel. Shaded area at base of hill, mostly a dune, immediately adjacent to beach (spray belt). Hill covered with Oen- othera sp. (introduced; Onagraceae) and Pancratium maritimum L. (Amaryllida- ceae). Observers: A. Freidberg and W. N. Mathis. Several specimens were observed run- ning over the sandy substrate, primarily in the shade. Their wings were parted at about 80° and occasionally the wings were quick- ly moved in a scissoring motion. A few fe- males were observed to tap the substrate with their abdomens, perhaps searching for Ovipositional sites. When a male came within 5—7 cm of a female, he would im- mediately pursue her and attempted to mount, apparently without any precopula- tory behavior. Mounting was brief, usually PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 5 seconds or less (range 3-15 seconds based on 4 observations). For one pair we observed the tips of both abdomens touch- ing but could not see if the genitalia of the male engaged those of the female (we as- sume that they did). After dismounting, the pair would face each other and touch their extended probosces intermittently for 10— 20 seconds. While thus touching, the male and/or female would scissor its wings, the opposite sex would remain essentially sta- tionary, with its wings parted at nearly 150°. We could not determine whether trophallaxis occurred with the touching of probosces. While touching probosces there was apparently some foreleg-to-head touch- ing. Remarks.—Wiedemann described this species twice and in two different genera. As species of Actocetor are striking and un- likely to be overlooked, we suspect that Wiedemann failed to examine specimens in his own collection (he may have returned some specimens and did not have ready ac- cess to them, did not consult his previously published description, or his memory fal- tered). Regardless, our study of Wiede- mann’s primary types clearly revealed that the syntypes of both names are conspecific, with A. indicus having priority as the senior synonym. In view of the widespread distri- bution of this species, particularly in the Afrotropical Region, it is unfortunate that the senior synonym, A. indicus, alludes to an area on the periphery of this species’ distribution where it may represent an in- troduction. The comparison of the male terminalia of the paratype of A. margaritatus panelii with those from India and Sri Lanka indicates that both are conspecific and thus the names are synonyms despite the lack of character- istic microtomentum on tergites 3—5. The microtomentum on the abdomen is also var- iable in other specimens, especially on the 4th tergite. This was particularly evident in specimens from a single site in Nigeria where a majority of the specimens have a wide band of microtomentum and a few, VOLUME 104, NUMBER 2 271 AK, Zaft / LY 46 Figs. 39-46. Structures of the male preabdomen and terminalia of Actocetor nigrifinis. 39, Fifth sternite and hypandrium, ventral view (only left side showing setae). 40, Presurstylus, posterior view. 41, Postsurstylus, lateral view (Sri Lanka. Padaviya). 42, Same (Malaysia. Sabah: Kota Kinabalu). 43, Postsurstylar process, posterior view (Sri Lanka. Padaviya). 44, Same (Malaysia. Sabah: Kota Kinabalu). 45, Aedeagus, aedeagal apodeme, dorsal view. 46, Same, lateral view. Scale bar equals 0.3 mm for Fig. 39 and 0.1 mm for Figs. 40-46. including females, have sparse to no micro- = Actocetor (Actocetor) nigrifinis (Walker) tomentum. The structures of the male ter- (Figs. 39-47, 50) minalia were compared directly in detail Opompza aieriins walker S60 m6e and a the saute, and we ee thus ,oti,the Actocetor nigrifinis: Cogan and Wirth Bhilommiat they.,are \copspecmic: 1977: 328 [generic combination]. Tone Win @snbo (os 71 eeee a C2 terete Actocetor beckeri de Meijere 1916: 264.— semis onl) a Vey aetaner Seem) mcrolo” Cogan and Wirth 1977: 328 [Oriental cat- mentum on the 4th tergite and is otherwise alog]_—Mathis and Zatwarnicki 1995: 22 metallic, bluish black. Although an entirely world Gatnige = ZaMbu Sood sse (ae bluish black abdomen also occurs else- Fad = a where, specimens often have some yellow- ish orange to red color on at least the basal tus of syntypes]. New synonym. Actocetor elegans Hendel 1917: 41.—Cres- son 1929: 171 [comparison with A. mar- 2-3 tergites. The structures of the male ter- garitatus|.—Cogan and Wirth 1977: 328 minalia of the holotype are virtually iden- [synonymy with A. beckeri; Oriental cat- tical to those of typical A. indicus, and we alog]. New synonym. consider these specimens to be conspecific. Thus, A. hendeli is also a synonym of A. Diagnosis.—This species is distinguished indicus. from congeners by the following combina- 272 tion of characters: moderately small to me- dium-sized shore flies, body length 2.40— 3.70 mm. Description.—Head: Arista with 10-13 dorsal rays. Gena high, higher than height of pedicel, gena-to-eye ratio 0.22—0.30. Thorax: Mesonotum densely microto- mentose, reddish gray to whitish gray, pleu- ra whitish to silvery gray; 2 more or less short, brown bars may be present anteriorly, separated by width equal to that separating prescutellar acrostichal setae; rings at bases of larger setae (prescutellar acrostichal, pos- terior dorsocentral, and basal scutellar se- tae) usually not dark brown; scutellum with apical %—%4 dark brown, basal portion con- colorous with posterior portion of scutum. Wing (Fig. 50) normally developed with wide anal angle; alula narrow, bandlike, bearing long setulae (length twice height of alula) along posterior margin; vein R,,; bearing 5—6 setae basad of crossvein r-m; wing pattern as follows: cell c with basal % white; cell r, with | sub-basal rectangular spot; cell r,,,; with 2 spots, basal spot in line with spot in cell r,, apical spot, transversely oval/rectangular, immediately apicad of merger of vein R,,, with costa; cell r,,; with sub-basal rectangular spot in line with anterior spot in cell r,, a subcircular, medial spot, in line with crossvein dm-cu, and an apical spot; discal cell with a wide, U- shaped, basal spot (apical arm sometimes constricted or appears to be separated) and a subapical, transversely oval-rectangular spot; cell m with a transversely oval-rect- angular spot near middle; cell cua, with a large, basal rectangular white and a subapi- cal, transversely trapezoidal spot, vein CuA, straight; costal vein ratio 0.65—0.87; M vein ratio 0.94—1.17. Halter stem yellow; knob whitish yellow and distinctly clubbed. Legs generally whitish yellow to yellow; forecoxa and base of femora sometimes with whitish gray to gray microtomentum laterally. Abdomen: Entirely shiny but with gray microtomentum on base of Ist tergite; color of tergites 1-3 more or less yellowish red, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON remaining tergites metallic black; 5th ster- nite of male as in Fig. 39. Male terminalia (Figs. 40—46): epandrium higher than wide, somewhat narrowed dorsally; presurstylus (Fig. 40) large, ventromedial surface with a small emargination, bearing | round to oval lobe medially; postsurstylus (Figs. 41—42) much longer than wide, bearing 3—4, apical setulae, apex simple, narrowed, but not bi- lobed, widest subapically, then tapered to point; postsurstylar process (Fig. 43) dis- tinctly expanded apically, with anterior por- tion extended to acutely narrowed process, posterior margin narrowly rounded; aedea- gus in lateral view (Fig. 46) highest at base, tapered to narrowly rounded apex, in dorsal view (Figs. 45), with apical % triangular from a wide base to pointed apex; aedeagal apodeme (Fig. 46) in lateral view subtrian- gular with angle at attachment with base of aedeagus extended and acutely formed; postgonite in ventral view as a sickle shaped process, relatively narrow through length; pregonite short, simple, bearing 2 setulae apically; hypandrium broadly and deeply pouchlike. Type material.—The lectotype 2 of Opo- myza nigrifinis Walker, here designated to preserve stability and make more universal the use of this name, is labeled ““SYN-Type [circular label, green submarginal border, SYN handwritten]/Mak [circular label, gray |/nigrifinis [handwritten ]/684/Celebes: Makessar [handwritten ]/Actocetor nigrifinis Walk. SYNTYPE 1860 Proc. Linn. Soc. Lond. IV p. 168 (Opomyza) det. J.C. Deeming 1964 [handwritten except for “det. J.C. Deeming 196’? )/LECTOTYPE ? Opomyza nigrifinis Walker By Dikow & Mathis [all except “LECTOTYPE” and “By” handwritten; black submarginal bor- der].”’ The lectotype is double mounted (minuten in rectangular block of foam), is in good condition (left midleg and wing missing), and is deposited in the BMNH. A paralectotype @ is labeled ““SYN-Type [cir- cular label, green submarginal border, SYN handwritten]|/Celebes [circular label, gray] 58 142 X [on underside of label, means that VOLUME 104, NUMBER 2 specimens were purchased in 1858 and col- lected by Wallace]/nigrifinis [handwritten, label folded]/Notiphilinae 3/8.01. Czerny [brownish label with squares, handwritten ]/ Celebes: Makessar [handwritten ]/Actocetor nigrifinis Walk. SYNTYPE 1860 Proc. Linn. Soc. Lond. IV p. 168 (Opomyza) det. J.C. Deeming 1964 [handwritten except for “det. -€; Deeming 196’): The paralec- totype is double mounted (minuten in rect- angular block of foam; head, left fore- and midlegs missing), and is deposited in the BMNH. The lectotype 2 of Actocetor beckeri de Meijere, here designated to preserve stabil- ity and make more universal the use of this name, is labeled “‘Batavia [Djakarta] (Moara Antjol) [= Moaraantjol] XII. 07 [Dec 1907] Jacobson [handwritten]/Acto- cetor Beckeri det. de Meijere. type [species name and “‘type’’ handwritten; black sub- marginal border]J/HOLOTY PE Actocetor beckeri de Meij det. B.H. Cogan 1971 [handwritten except for “det. B.H. Cogan 197” |/Actocetor hendeli de Meij. Det. B.H. Cogan 1971. [handwritten except for “det. B.H. Cogan 197” |/Actocetor Beckeri de Meijere, 1916 ZMAN type DIPT.0437.1 [red]/LECTOTYPE & Actocetor beckeri de Meijere By Dikow & Mathis [all except “PECTORYPE >and By handwritten; black submarginal border].’’ The lectotype is double mounted (minuten in rectangular block of foam), is in good condition (min- uten has verdigris on it), and is deposited in the ZMAN. In the original description, de Meijere mentioned ““Sumpf” (swamp) as part of the type locality. There are also five 2 paralectotypes that bear the same lo- cality data as the lectotype with the addition of “by Bembex nest.” Although Cogan wrote that this specimen is a holotype, de Meijere’s original description noted “einige Exemplare’’ (some specimens) and thus a lectotype designation is needed. Cogan also attached a second label on which *“Acto- cetor hendeli de Meij.”” was written. Cogan apparently confused the name of this spe- cies with A. beckeri Hendel, which is pre- 273 occupied and needed a replacement name, A. hendeli, which de Meijere (1924) pro- vided. The holotype 2 of Actocetor elegans Hendel is labeled ““Ceylon, Colombo./6/VI 02. [6 Jun 1902] Dr. Uzel./Actocetor ele- gans H. det. Hendel [species name and au- thor handwritten]/Coll. Hendel/HOLO- TYPE & Actocetor elegans Hendel [red submarginal boarder; name, author, and gender handwritten].’” The holotype is dou- ble mounted (minuten in a_ rectangular block of foam), is in very good condition, and is deposited in the NMW. Other specimens examined.—ORIEN- TAL. AUSTRALIA. Cocos-Keeling Is- lands. Direction Island, 4 Jun 1952, T. G. Campbell (12; BMNH). MALAYSIA: Sabah: Kota Kinabalu (17 km S), 19. Awe i983 "Ger Hever aw:, E- Steiner (1d, 52; USNM); Tanjung Aru Beach,.(29 Aug | 1983; G. FicHevel iw. E- Steiner (12; USNM); Tomani, | Sep 1983, G. E Hevel, W. E. Steiner (5¢; USNM). PHILIPPINES. Palawan Brookes Point, Uring Uring, 25 Aug 1961, Noona Dan Ex- pedition (12; UZMC). Batangas: Bo. Cale (14°07'N, 121°06’E; 7 km NW Tanauan; ex. Zea mays L.), 16 Jul 1977, A. T. Barrion (ditENE): SRI LANKA. Central Province. Kandy: Mahiyangana (51 mi NW; Malaise trap at Hasalaka Irrigation Bungalow), 30 Mar—9 Apr 1971, P. and P. Spangler (16; USNM); Peradeniya Botanical Gardens, 25 Jan 1977, W. N. Mathis, T. Wijesinhe, L. Jayawickri- ma (2d, 82; USNM); Peradeniya (Malaise trap at Botanical Gardens), 26-28 Mar 197 igse. and PRP) Spaneler"“@2>)USNM): Northern Province. Jaffna: Klinochchi (80 ft), 25 Jan 1977, W. N. Mathis, T. Wijes- inhe, L. Jayawickrima (12; USNM). North Central Province. Anuradhapura: Padaviya (180 ft), 2-8 Nov 1970, O. S. Flint, Jr. (12; USNM); Padaviya (180 ft; Irrigation Bun- galow), 27 Feb—9 Mar 1970, D. R. Davis, W. Rowe (12; USNM). Polonnaruwa: Pimburettawa (13 mi S Mannampitiya; SSOMf)Ie9=12) Octr1970; -Onse Flint Ir 274 (36; USNM). Province of Sabaragamuwa. Ratnapura: Panamure (500 ft), 15-21 Oct 1970, O. S. Flint, Jr. 1d, 12; USNM); Ug- galkaltota (350 ft; Irrigation Bungalow), 31 Jan—8 Feb 1970, D. R. Davis, W. Rowe (22: USNM). Southern Province. Galle: Kanneliya, 27 Apr 1980, W. N. Mathis, T. Wijesinhe, L. Jayawickrima (2d, 1°; USNM). Hambantota: Kirinda, 25 Apr 1980, W. N. Mathis, T. Wijesinhe, L. Jay- awickrima (16; USNM); Palatupana, 3—6 Feb 1975, K. V. Krombein, P. B. Karunar- atne, P. Fernando, E. G. Dabrera (12; USNM); Palatupana Tank, 6 Feb 1975, K. V. Krombein (12; USNM). Province of Uva. Badulla: Digaluma Falls (1550 ft; black light), 17 Aug 1973, G. Ekis (14, 12; USNM); Girandurakotte Circ. Bunga- low (10 mi NNE Mahiyangana; UV trap), 4-7 Sep 1980, K. V. Krombein, P. B. Ka- runaratne, T. Wijesinhe, L. Jayawickrema, V. Gunawardane (1d, 22; USNM). Mon- aragala: Agunakolapelessa (on or in leaf litter), 8-9 Oct 1980, K. V. Krombein, P. B. Karunaratne, T. Wijesinhe, L. Jayawickre- ma, V. Gunawardane (1d, 102; USNM); Agunakolapelessa (Malaise trap), 27—28 Mar 1981, K. V. Krombein, T. Wijesinhe, L. Weeratunge (23, 92; USNM). North Province. Kudattanai (6 mi SE Point Pe- dro), 13 Feb 1962, H. Andersson, P. Brink, L. Cederholm (12; ZIL). Northwest Prov- ince. Kadalmparu (15 mi N Negombo; at shore of lagoon), 31 Jan 1962, H. Anders- son, P. Brink, L. Cederholm (1¢, 12; ZIL). Western Province. Colombo: Colombo, 18 Apr 1980, W. N. Mathis (82; USNM); Co- lombo, Museum Garden, 28-31 Jan 1975, 6 Jul 1976, K. V. Krombein, P. B. Karunar- atne, P- Fernando (22; USNM); Colombo, 6 Jun 1902, Dr. Uzel (12; NMW); Colom- bo (swept waste ground), 17 Sep 1963 (1°; BMNH); Katunayaka (near airport), 16 Jan 1977, K. V. Krombein (12; USNM); Rat- malana (9 mi S Colombo; dry grass on sandy beach), 7-13 Jan 1963, H. Anders- son, P. Brink, L. Cederholm (2¢, 12; ZIL). THAILAND. Chaing Mai, 5 Dec 1985, K. A. Spencer (12; NMWL). Hot (20 km PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON W; 200 m), 17 Oct 1993, FE Kaplan, A. Freidberg (1d; USNM); KhaoLak Beach (100 km N Phuket), 19 Oct 1993, E Kaplan, A. Freidberg (1d; USNM). Distribution (Fig. 47).—Oriental: Cocos- Keeling Islands, Indonesia (Java, Sulawesi), Malaysia, Philippines, Sri Lanka, Thailand. Natural history.—Like A. indicus, this species apparently also occurs around sandy habitats, at least in part. This suggestion is based on the observation that the paralec- totypes of A. beckeri (see label data noted above) were collected by a nest of Bembex, a genus of solitary wasps known to con- struct nests in sand. Remarks.—This is a widespread species in the Orient and is likely to occur in more countries than available locality records in- dicate. The lack of records from most of the countries of Southeast Asia is particularly noteworthy and probably reflects sampling error rather than the species’ absence. Cogan and Wirth (1977) determined pre- viously that A. elegans was a junior syno- nym of A. beckeri, and here we have con- cluded that both names are junior synonyms of A. nigrifinis, a species name that was generally overlooked until Cogan and Wirth (1977) correctly assigned this species to Ac- tocetor from Opomyza (family Opomyzi- dae). Subgenus Poecilostenia Bezzi Poecilostenia Bezzi 1908: 195 [as a genus; type species: Poecilostenia decemguttata Bezzi 1908, monotypy].—Wirth 1955: 51 [subgeneric status]. Diagnosis.—This subgenus is distin- guished from Actocetor sensu stricto by the following combination of characters: Description.—Thorax: Mesonotum gen- erally yellow microtomentose. Wing alula completely reduced, bearing few or no se- tulae along posterior margin; anal angle straight, cell cua, narrower than discal cell; vein R,,; bearing 5—6 setulae on dorsum, basad of crossvein r-m and 5—10 setulae ap- icad of crossvein r-m; vein CuA, conspic- VOLUME 104, NUMBER 2 NO | Nn Fig. 47. Distribution map for Actocetor nigrifinis. uously sinuous along posterior margin of discal cell; discal cell with at most 1 white spot. Knob of halter dark brown to black, not distinctly clubbed. Abdomen: 5th tergite of male bearing 4 well-developed setae along posterior mar- gin; epandrium lacking large setae along dorsum; other structures as in the species descriptions of A. decemguttatus and A. lindneri, the only species of Poecilostenia for which males are known. Discussion.—Specimens of Poecilosten- ia are uncommon generally and more spe- cifically when compared with Actocetor sensu stricto. Although the species can be determined fairly easily, some are repre- sented by a single female. We suspect that additional species will be found in this sub- genus as the Afrotropical Region is better sampled. The subgenus Poecilostenia is monophy- letic, as substantiated by the following syn- apomorphies (characters marked with an * are unambiguous): 6*. Color of mesonotal microtomentum yellowish; 11*. Alula com- pletely reduced and bearing few or no se- tulae along posterior margin; 12*. Cell cua, generally narrower than discal cell, and anal margin nearly straight; 13*. Vein R,,; bear- ing setulae basad and apicad of crossvein r- m; 14*. Anterior margin of vein CuA, ba- sad of crossvein dm-cu slightly concave; 20*. Knob of halter dark brown to black; 21*. Knob of halter not distinctly clubbed (only slightly expanded from diameter of stem); 22. Microtomentum on anterior mar- gins of 3rd and 4th tergites with narrow bands (secondarily modified in A. lindner?). Actocetor (Poecilostenia) decemguttatus (Bezzi) (Figs. 52, 55-59) Poecilostenia decemguttata Bezzi 1908: 195 [Botswana. Kang-Khakhea, Kalaha- ri; ST 6 2, deposition unknown].—Cres- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 48-54. Wings. 48, A. hovus. 49, A. indicus. 50, A. nigrifinis. 51, A. afrus. 52, A. decemguttatus. 53, A. lindneri. 54, A. yaromi. son 1946: 256 [review, list, South Afri- ca]. Actocetor (Poecilostenia) decemguttatus: Wirth 1955: 51 [generic combination].— Cogan 1980: 659 [Afrotropical catalog]. Actocetor anormalipennis Séguy 1933: 40.—Cresson 1946: 256 [synonymy with P. decemguttata Bezzi|].—Wirth 1955: 51 [key]|—Canzoneri and Rampini 1995: 254 [list, Sierra Leone].—Mathis and Za- twarnicki 1995: 23 [world catalog]. New synonym. Actocetor abnormalipennis: Cresson 1946: 256 [lapsus, synonymy with Poecilosten- ia decemguttata Bezzi|.—Cogan 1980: 659 [Afrotropical catalog]. Actocetor toniatabae Canzoneri 1981: 209 [habitus figure].—Mathis and Zatwarni- cki 1995: 22 [world catalog]. New syn- onym. Diagnosis.—This species is distinguished from congeners by the following combina- tion of characters: moderately small shore flies, body length 2.30—2.90 mm. Description.—Head: Arista with 8—9 dorsal rays. Gena-to-eye ratio 0.20—0.21. Thorax: Mesonotum densely microto- mentose, yellowish brown, pleura concolo- rous; 2 pale brown bars anteriorly, separat- ed by width equal to that separating pre- scutellar acrostichal setae; scutellum with VOLUME 104, NUMBER 2 Figs. 55-59. 277 Uh mi yy) YD, My S).2) lateral view. 56, Seneeatite posterior view. 57, at ere lateral view. 58, Aedeasus, aedeagal apodewie! dorsal view. 59, Same, lateral view. Scale bar equals 0.1 mm. apical % brown, basal portion concolorous with posterior portion of scutum. Wing (Fig. 52) very narrow, cell cua, narrower than discal cell, anal margin nearly straight; vein R,,; bearing 5 setulae basad and 5 se- tulae apicad of crossvein r-m; wing pattern as follows: cell c with rectangular white spot basad of subcostal break; cell r, with rectangular, medial spot; cell r,,; with 2 spots, basal rectangular/trapezoidal spot in line with spot in cell r,, apical, transversely oval/rectangular spot, immediately apicad of merger of vein R,,, with costa; cell r,,5 with a circular medial spot in line with api- cal spot in cell r,,,, and an apical spot; dis- cal cell with a circular, subapical spot; cell m with transversely oval-rectangular spot; cell cua, with 2 spots, rectangular sub-basal spot, and transversely trapezoidal spot in line with spot in discal cell, vein CuA, slightly concave between these spots; costal vein ratio 0.70—0.71; M vein ratio 1.17— 1.18. Halter stem at base yellowish white, apical part of stem and knob dark brown to black; knob only slightly to moderately ex- panded from diameter of stem, not distinct- ly spatulate. Legs generally whitish yellow to yellow. Abdomen: Mostly shiny, base of Ist ter- gite with gray microtomentum; base of 3rd and 4th tergites with tiny bands of gray mi- crotomentum; generally metallic black but sometimes with reddish anterior and/or pos- terior margins of tergites; 3rd and 4th ster- nites of male rectangular, moderately nar- row, 2—3X longer than wide; 5th sternite of 278 male trapezoidal, bearing numerous, mostly evenly scattered setulae, posterior margin evenly concave, lacking a less sclerotized area medially. Male terminalia (Figs. 56— 59): epandrium wider than high, bearing numerous setulae, none, especially along dorsum, enlarged; presurstylus (Fig. 56) large, similar to moose antlers, expanded ventral margin bearing 4 prongs, medial prong short, wide, apex curved dorsally as a small point; submedial prong narrow, dig- itiform, tapered to point; 2 lateral prongs wide basally, tapered to rounded point; postsurstylus (Fig. 57) very robust through- out length, bilobed apically, both lobes well developed, subequal, anterior lobe bearing a well-developed, stout, spinelike seta that extends into hypandrium; postsurstylar pro- cess (Fig. 57) short, robust, apex spatulate, round; aedeagus in lateral view (Fig. 59) shoelike, apex pointed, in dorsal view with base wider and lateral extension; aedeagal apodeme in lateral view (Fig. 59) broadly sickle shaped with a moderately prominent, raised keel; postgonite a curved sclerite be- tween juncture of hypandrium and aedeagal apodeme and postsurstylus; postsurstylar process with short stalk, apex somewhat spatulate; pregonite not distinct, perhaps greatly reduced or fused indistinguishably with other structures; hypandrium (Fig. 55) greatly modified, narrow but extended an- teriorly like a large, narrow, keel, pouch narrow but deep, bearing pronglike pro- cesses that are opposable to spinelike seta of postsurstylus. Type material.—The syntypes, apparent- ly 1 d and 1 &, of Poecilostenia decem- guttata Bezzi were not located despite searches in Italy (Museo Civico di Storia Naturale, Milano; Dr. Carlo Pesarini) and Germany (Zoologisches Museum, Hum- boldt Universitat, Berlin; Dr. Hella Wendt), and our determination of this species is based on the original description, especially the illustration of the wing. Bezzi (1908) published the following locality data about the syntypes and type locality: “‘Ein gut er- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON haltenes Parchen aus der Kalahari: Kang- Khakhea, Dezember 1904, No. 968.” The holotype 2 of Actocetor anormali- pennis Séguy is labeled ““MUSEUM PARIS MOZAMBIQUE ENV. VILA PERV. [Sé- guy wrote “Vila-Pery” in the original pub- lication; last line of label scratched out; blue]/P. LESNE 1929 [blue]/ouzinai 2 se- tebre [Séguy wrote ‘‘2.X” in the original publication as the day and month]/TYPE [red ink]/Actocetor anormalipennis TYPE E. SEGUY det 1933 [handwritten except for ““E. SEGUY det 19’’].” The holotype is double mounted, is in good condition, and is deposited in the MNHN. The information on this holotype was kindly provided by Dr. Tadeusz Zatwarnicki. The holotype @ of Actocetor toniatabiae Canzoneri is labeled ““GAMBIA. Toniata- ba[,] margini mangrovia veget[azione]. alof[ila]. 30.vii [30 Jul] Giordani Soika leg. 1973 [white label glued to a green backing; handwritten except for “Giordani Soika leg. 1973’ ]/HOLOTYPUS @& Actocetor toniatabae m. det. Canzoneri S[ilvano] [red; gender symbol, species name, and “m.” handwritten]/Actocetor decemguttatus (Bezzi) det. WNMathis 2000 [species name and year handwritten].””> The holotype is double mounted (minuten in a rectangular card), is in excellent condition, and is de- posited in the CANZ collection that is now in the MRSN. Other specimens examined.—AFRO- TROPICAL. BOTSWANA. Kuke Pan, 21— 30 Mar 1930, V.-L. Kal. Exp. (12; NMSA). Serowe (Malaise trap), 16-30 Apr 1985, P. Forchhammer (1 2°; BMNH). Tlokweng, 15 Mar 1990, J. M. Mashonja, (60, 8°; NMWL, USNM). GAMBIA. Western: Bakau, 15 Nov 1993, J. C. Deeming (12; NMWL). KENYA. Matembur (01°22'N, 35°02’E; 1550 m; Malaise trap), 26—27 May 1980, Lamoral (12; NMSA). Nairobi (50 km SE; Rt. A109), 30 Apr 1991, A. Freidberg, F Kaplan (12; USNM). Rift Valley: Nguru- VOLUME 104, NUMBER 2 Fig. 60. Distribution map for Actocetor decemguttatus (filled circles), A. lindneri (filled square), and A. yaromi (filled triangle). man, Oloibortoto River (1°48’S, 36°04’E; Malaise trap), 17-20 May 1999, R. Cope- land (12; USNM). NAMIBIA. Outjo (52 km W road 65; 20°14'S, 15°40'E; Mopane woodland on a rocky hillside), 24 Mar 1984, B. Stucken- berg, J. G. H. Londt (16; NMSA). Oka- handja, 2—4 Feb 1972 (12; BMNH); NIGERIA. Zaria, Samaru, Jun 1979, J. C. Demming (3d, 62; NMWL, USNM). SOUTH AFRICA. KwaZulu-Natal: Mhlopeni Nature Reserve (15 km S Muden; 2930Ab), 22 Dec 1983, J. G. H. Londt (12; NMSA). Weenen, Nov 1929, H. P. Tho- masset (1d, 12; BMNH, NMSA). Mpu- malanga: Shingwedzi, 9 Feb 1988, D. Si- mon (22; USNM). TANZANIA. Tarangire National Park (36°10’'E, 03°50'S), Dec 1994, D. Grimaldi (25, 42; AMNH). ZAMBIA. Livingstone, 1958, E. Lindner (12; SMN). Distribution (Fig. 60).—Afrotropical: Botswana, Gambia, Kenya, Mozambique, Namibia, Nigeria, South Africa, Tanzania, and Zambia. Remarks.—This is the only species of the subgenus Poecilostenia that is somewhat 16-20 Nov 280 common, and its distribution in the Afro- tropical Region is relatively widespread. This species was described three times with two of the descriptions being based on single females. Although males were un- available for two of the type series, we are confident of the conspecificity of the three names based on the stable and consistent wing pattern in the type series of all three names. Our assessment of the stability in the wing pattern is based on study of nu- merous specimens, including males, in ad- dition to the type series. Actocetor (Poecilostenia) lindneri Wirth (Figs. 53, 61-64) Actocetor (Poecilostenia) lindneri Wirth 1955: 52.—Cogan 1980: 659 [Afrotrop- ical catalog]|—Mathis and Zatwarnicki 1995: 23 [world catalog]. Diagnosis.—This species is distinguished from congeners by the following combina- tion of characters: moderately small shore flies, body length 2.90—3.00 mm. Description.—Head: Arista with 10-11 dorsal rays. Gena-to-eye ratio 0.18. Thorax: Mesonotum densely microto- mentose, yellowish brown, pleura concolo- rous; 2 pale brown bars anteriorly, separat- ed by width equal to that separating pres- cutellar acrostichal setae; scutellum with apical % pale brown, basal portion conco- lorous with posterior portion of scutum. Wing (Fig. 53) very narrow, cell cua, nar- rower than discal cell, anal margin nearly straight; vein R,,; bearing 7 setulae basad and 7 setulae apicad of crossvein r-m; wing pattern as follows: cell c with rectangular white spot basad of subcostal break; cell r, with a circular, sub-basal spot and a rect- angular, medial spot; cell r,,, with 2 spots, basal, transversely narrow, rectangular spot in line with spot in cell r,, apical, trans- versely oval/rectangular spot, immediately apicad of merger of vein R,,, with costa; cell r,,; with a tiny circular, sub-apical spot, and an apical spot; discal cell without spots but with a paler-colored, crescent-shaped PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON bulla on apical half; cell m with narrow, transversely trapezoidal spot; cell cua, with 2 spots, 1 sub-basal and | subapical narrow rectangular spot, vein CuA, concave be- tween these spots; costal vein ratio 0.53— 0.72; M vein ratio 1.12—1.15. Halter stem yellowish white basally, apical part of stem and knob dark brown to black; knob only slightly expanded from diameter of stem, not distinctly clubbed. Legs generally whit- ish yellow to yellow. Abdomen: Mostly covered by microto- mentum: Ist tergite entirely gray microto- mentose; 2nd tergite shiny, lacking micro- tomentum, following tergites entirely mi- crotomentose, basal part gray and apical part brownish; metallic black. Male 3rd and 4th sternites rectangular, moderately nar- row, 2—3X longer than wide; 5th sternite of male trapezoidal, bearing numerous, mostly evenly scattered setulae, posterior margin evenly concave, lacking a less sclerotized area medially. Male terminalia (Figs. 61— 64): epandrium wider than high, bearing numerous setulae, none, especially along dorsum, enlarged; presurstylus (Fig. 62) large, similar to moose antlers, expanded ventral margin bearing 4 prongs, medial prong short, wide, apex curved dorsally as a small point; submedial prong narrow, dig- itiform, tapered to point; 2 lateral prongs wide basally, tapered to rounded point; postsurstylus (Fig. 63) very robust through- out length, bilobed apically, both lobes well developed, subequal, anterior lobe bearing a well-developed, stout, spinelike seta that extends into hypandrium; postsurstylar pro- cess (Fig. 63) short, robust, apex spatulate, round; aedeagus in lateral view (Fig. 64) shoelike, apex pointed, in dorsal view with base wider and lateral extension; aedeagal apodeme in lateral view (Fig. 64) broadly sickle shaped with a moderately prominent, raised keel; postgonite a curved sclerite be- tween juncture of hypandrium and aedeagal apodeme and postsurstylus; postsurstylar process with short stalk, apex somewhat spatulate; pregonite not distinct, perhaps greatly reduced or fused indistinguishably VOLUME 104, NUMBER 2 281 et | ey: TE lr | | ty | jae 64 Figs. 61-64. Structures of the male preabdomen and terminalia of Actocetor lindneri. 61, Hypandrium, lateral view. 62, Presurstylus, posterior view. 63, Postsurstylus, lateral view. 64, Aedeagus, aedeagal apodeme, lateral view. Scale bar equals 0.1 mm. with other structures; hypandrium (Fig. 61) Type material—The holotype ¢ is la- greatly modified, narrow but extended an- beled “[Tanzania] Jipe-See 20.—23. V. 1952 teriorly like a large, narrow, keel, pouch D. O. Afrika Exp. [blue]/Jipe 21. V 52 narrow but deep, bearing pronglike pro- [handwritten]J/HOLOTYPE 2 Actocetor lin- cesses that are opposable to spinelike seta derni W. W. Wirth [red; species name hand- of postsurstylus. written]/Actocetor (Poecilostenia) lindneri 282 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Wirth det. WWirth [19]54 [species name, author, and ‘‘54”’ handwritten].’’ The holo- type is double mounted (minuten in a card- board card), is in poor condition (head, left wing, and right fore- and hindlegs are miss- ing), and is deposited in the SMN. Other specimens examined.—AFRO- TROPICAL. KENYA. Coast: Muhaka For- est (4°19.47'S, 39°31.45'E; Malaise trap), 2-9 Apr 2000, R. Copeland (16; USNM). Eastern: (At » Athi SRiver 9(2°3855 1S, 38°21.98'E; Malaise trap), 30 Aug—6 Sep 2000, R. Copeland (12; USNM). Distribution (Fig. 60).—Afrotropical: Kenya, Tanzania. Remarks.—This species was described from a single female that is in poor condi- tion. We examined an additional female and one male, both from Kenya. Although sim- ilar to A. decemguttatus, especially struc- tures of the male terminalia, this species is distinguished by characters of the wing. Better sampling of this species may reveal that the wing characters used to distinguish this species are clinal and thus, that this species may be conspecific with A. decem- guttatus. Actocetor (Poecilostenia) yaromi Dikow and Mathis, new species (Fig. 54) Diagnosis.—This species is distinguished from congeners by the following combina- tion of characters: moderately small shore flies, body length 2.70 mm. Description.—Head: Arista with 9 dor- sal rays. Gena-to-eye ratio 0.15. Thorax: Mesonotum densely microto- mentose, yellowish brown, pleura concolo- rous; 2 pale brown bars anteriorly, separat- ed by width equal to that separating pres- cutellar acrostichal setae; scutellum entirely concolorous with yellowish to slightly brownish yellow scutum. Wing (Fig. 54) very narrow, but cell cua, wider than discal cell, anal margin nearly straight; vein R,,; bearing 6 setulae basad and 5 setulae apicad of crossvein r-m; wing pattern as follows: cell c with rectangular white spot medially; cell r,; with rectangular, medial spot; cell r,; with 2 spots, basal rectangular spot in line with spot in cell r,, apical, transversely oval/rectangular spot, immediately apicad of merger of vein R;., with costa;(cell rt. with a circular spot in line with apical spot in cell r,,3, and an apical spot; discal cell without white spot; cell m with transversely oval-rectangular spot; cell cua, with 2 spots, a rectangular sub-basal spot, and a trans- versely trapezoidal spot in line with spot in discal cell; vein CuA, concave between these 2 spots; costal vein ratio 0.66; M vein ratio 1.05. Halter stem yellowish white ba- sally, apical part of stem and knob dark brown to black; knob only slightly expand- ed from diameter of stem, not distinctly clubbed. Legs generally whitish yellow to yellow. Abdomen: Mostly shiny, base of Ist ter- gite with gray microtomentum; base of 3rd and 4th tergites with bands of gray micro- tomentum; generally metallic black but 2nd and 3rd tergites with reddish anterior. Male unavailable. Type material—The holotype @ is la- beled “ETHIOPIA: GAMO GOFA, Arba Minch Springs, 1300m[,] 8.11.2000 [8 Feb 2000], I. YAROM & A. FREIDBERG/ wing slide [handwritten/HOLOTYPE & Actocetor yaromi Dikow & W.N. Mathis USNM [red; species’ name and gender and “Dikow &’” handwritten].’’ The holotype is double mounted (minuten in a block of plastic), is in good condition (left wing re- moved and slide mounted), and is deposited in the USNM. Distribution.—Afrotropical: Ethiopia. Etymology.—The species epithet, ya- romi, 18 a genitive patronym to honor and recognize the contributions of Dr. Ilan Ya- rom to the study of acalyptrate Diptera. Dr. Yarom collected the holotype. Remarks.—We are describing this spe- cies now despite having only a single fe- male specimen available to us. The species is distinctive, especially the characters of the wing, and confusion with congeners is unlikely. VOLUME 104, NUMBER 2 PHYLOGENETIC CONSIDERATIONS In the presentation on species-level rela- tionships that follows, the characters used in the analysis are noted first. Each char- acter is immediately followed by a discus- sion to explain its states and to provide per- spective and any qualifying comments about that character. After presentation of the information on character evidence, an hypothesis of the cladistic relationships is presented and briefly discussed. The clad- ogram (Fig. 65) is the primary mode to con- vey relationships, and the discussion is to supplement the cladogram and is intended only to complement the latter. In the dis- cussion of character data, an “‘O”’ indicates the state of the outgroup; a “1” or “2” in- dicates the derived states. Head: 1. Length of Ist flagellomere: (O) relative- ly long (longer than width of pedicel); (1) short (length subequal to width of pedicel; an autapomorphy for Actoce- tor). 2. Position of antenna on head: (0) posi- tioned near midheight level of head (frons conspicuously longer than wide); (1) positioned high, above dor- sal % in lateral view (an autapomorphy for Actocetor). 3. Development of pseudopostocellar se- tae: (0) setae reduced (as in Mimapsi- lopa), (1) setae well developed, as large as proclinate setae (often or most- ly slightly reclinate; a synapomorphy for Actocetor, Eremomusca, Guttipsi- lopa, and Trypetomima). 4. Development of facial setae: (0) face bearing 2 well-developed setae; (1) face bearing 4 well-developed setae in a vertical row, dorsalmost 1—2 setae cruciate (a synapomorphy for Actocetor and Guttipsilopa). 5. Gena-to-eye ratio: (0) variable but gena generally short with ratios of 0.08— 0.15; (1) 0.21—0.36 (a synapomorphy for the subgenus Actocetor); (2) 0.14— 0.20 (a synapomorphy for the subgenus Poecilostenia). Thorax: 6. Color of mesonotal microtomentum: (O) blackish; (1) grayish (a synapomor- phy for the subgenus Actocetor); @) yellowish (a synapomorphy for the subgenus Poecilostenia). 7. Coloration of scutellum: (0) scutellum entirely concolorous with scutum (as in A. yaromi); (1) apicad %4—’% dark brown (a synapomorphy for A. afrus and A. indicus); (2) apicad %—%4 dark brown (a synapomorphy for A. decemguttatus, A. hovus, A. lindneri, and A. nigrifinis). 8. Color surrounding base of prescutellar acrostichal, posterior dorsocentral, and basal scutellar setae: (0) not distinctly dark brown (as in A. decemguttatus, A. lindneri, A. nigrifinis, and A. yaromi); (1) distinctly dark brown (a synapo- morphy for A. afrus, hovus, and A. in- dicus). 9. Development of wing pattern: (0) wing generally hyaline (the plesiomorphic state as in Mimapsilopa); (1) wing mostly whitish and conspicuously mac- ulate (an autapomorphy for Eremomus- ca); (2) wing mostly blackish brown with white spots (a synapomorphy for Actocetor, Guttipsilopa subgenus Gut- tipsilopa, and Trypetomima), (3) wing with anterior margin darkened (a syn- apomorphy for Guttipsilopa subgenus Nesopsilopa and some Discomyza). 10. Development of apex of vein R,,,;: (O) apex normally developed at merger with costa (as in Actocetor); (1) vein R,,; with a subapical stump vein (a synapomorphy for Eremomusca and Trypetomima). 11. Development of alula: (0) alula a well- developed lobe (as in Guttipsilopa); (1) reduced to a thin bandlike lobe bearing numerous short or long setulae along posterior margin (a synapomorphy for the subgenus Actocetor and the genera Eremomusca, Mimapsilopa, and Try- petomima); (2) completely reduced and 284 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 9 fon GUETIPSIOpa 3 3.9 26 Mimapsilopa 46 001 9 22 0 0 Eremotrichoma 10 15 1 13 14 fh Trypetomima 21 1113 U [ _—A. yaromi 1 1 0 5 6 111213 14 20 21 22 24 26 27 28 29 16 22 Tb a A. lindneri De dieo det) 2 4. 4 oteegie 14 15 12747 A. decemguttatus 1 AL ie 18 22 25 26 Lb | A. nigrifinis 1-2"3-5 De 7 1516 20 22 oe ee | HHH A. afrus 25 26 17 18 123 ne ; A. hovus 0 7 16 22 25 26 27 LH | [ = A. indicus Aad ale (eon o Fig. 65. Cladogram depicting hypothetical cladistic relationships among species of Actocetor. bearing few or no setulae along poste- rior margin (an autapomorphy for the subgenus Poecilostenia). . Development of cell cua,: (O) cell cua, broad, broader than discal cell, and anal angle moderately well developed (as in the subgenus Actocetor); (1) cell cua, generally narrower than discal cell, and anal margin nearly straight (an auta- pomorphy for the subgenus Poecilos- tenia). . Development of setulae on vein R,,;: (O) no setulae present (as in Guttipsi- lopa); (1) setulae only basad of cross- vein r-m (a synapomorphy for the sub- genus Actocetor and the genera Ere- momusca and Mimapsilopa); (2) setu- lae basad and apicad of crossvein r-m 14. 1S 16. (a synapomorphy for the subgenus Poecilostenia and the genus Trypetom- ima). Shape of anterior margin of vein CuA, basad of crossvein dm-cu: (QO) straight (as in the subgenus Actocetor); (1) slightly concave (a synapomorphy for A. decemguttatus, A. lindneri, A. ya- rom, and Trypetomima). Number of white spots in discal cell: (O) no spot (as in A. yaromi and A. lind- neri); (1) 1 spot (a synapomorphy for A. decemguttatus and Trypetomima); (2) 2 spots (a synapomorphy for A. ho- vus, A. indicus, and A. nigrifinis); (3) 3 spots (an autapomorphy for A. afrus). Number of white spots in cell r,: (O) 1 spot (as in A. decemguttatus, A. hovus, VOLUME 104, NUMBER 2 I aie 20. Piss A. nigrifinis, and A. yaromi); (1) 2 spots (a synapomorphy for A. afrus, A. indi- cus, and A. lindneri). Number of setulae on vein R,,; basad of crossvein r-m: (QO) no setulae present (as in Guttipsilopa); (1) 2—3 setulae (an autapomorphy for A. hovus); (2) 4 or more setulae (a synapomorphy for A. afrus, A. decemguttatus, A. indicus, A. lindneri, A. nigrifinis, and A. yaromi). . Costal vein ratio: (0) 0.53—0.73 (as in A. afrus, A. decemguttatus, A. indicus, A. lindneri, and A. yaromi); (1) 0.73— 0.87 (a synapomorphy for A. hovus and A. nigrifinis). . M vein ratio: (0) 0.84—1.00 (as in A. hovus and A. indicus); (1) 1.01—1.21 (a synapomorphy for A. afrus, A. decem- guttatus, A. lindneri, A. nigrifinis, and A. yaromi). Color of knob of halter: (0) white to yellow (as in the subgenus Actocetor); (1) dark brown to black (a synapomor- phy for A. afrus, A. decemguttatus, A. lindneri, and A. yaromi). Shape of knob of halter: (QO) distinctly clubbed (as in the subgenus Actocetor); (1) knob of halter not distinctly clubbed (only slightly expanded from diameter of stem; an autapomorphy for the subgenus Poecilostenia). Abdomen: Wika Development of microtomentum on ab- domen (in addition to microtomentose anterior margin of Ist tergite): (O) en- tirely bare (as in A. hovus and A. ni- grifinis); (1) anterior margin of 4th ter- gite with distinct band (an autapomor- phy for A. indicus); (2) anterior mar- gins of 3rd and 4th tergites with narrow bands (a synapomorphy for A. decem- guttatus and A. yaromi); (3) anterior margins of 3rd, 4th, and 5th tergites with distinct bands (a synapomorphy for A. afrus); (4) tergites 3—7 entirely gray and brown microtomentose (an autapomorphy for A. lindneri); (5) en- tirely microtomentose (genus Eremo- musCQ). De 285 Development of setae on dorsum of epandrium: (0) bearing small setae only (as in A. decemguttatus and A. lind- neri); (1) bearing well-developed, dor- sally erect setae (a synapomorphy for A. afrus, A. hovus, A. indicus, and A. nigrifinis; no male available for A. ya- rom). 24. Shape of male 3rd and 4th sternites: (0) N Nn 26. rectangular, broad, 1-2 longer than wide (as in A. afrus, A. hovus, A. in- dicus, and A. nigrifinis); (1) rectangu- lar, moderately narrow, 2—3 longer than wide (a synapomorphy for A. de- cemguttatus and A. lindneri; no male available for A. yaromi). . Shape of male 5th sternite: (0) trape- zoidal with slightly concave posterior margin (as in A. decemguttatus and A. lindneri); (1) trapezoidal with medial weakness or slight indentation on con- cave posterior margin (an autapomor- phy for A. indicus); (2) trapezoidal, di- vided medially into two sclerites (a synapomorphy for A. afrus and A. ho- vus); (3) trapezoidal with concave pos- terior margin with medial weakness and 4—5 setulae on either side of weak- ness (a synapomorphy for A. nigrifinis; no male available for A. yaromi). Postsurstylar process: (O) absent; (1) present, cylindrical (an autapomorphy for Mimapsilopa); (2) present, clubbed with rounded knob (an autapomorphy for A. indicus); (3) present, expanded ventrally and projected to a point pos- teriorly (a synapomorphy for A. afrus and A. hovus); (4) present, clubbed but not stalked (a synapomorphy for A. de- cemguttatus and A. lindneri); (5) pres- ent, distinctly expanded apically, with anterior portion extended to narrow process, posterior margin narrowly rounded (an autapomorphy for A. ni- grifinis; no male available A. yaromi). 27. Postsurstylus: (0) bilobed subapically with a small, digitiform, medial, point- ed process (as in A. indicus); (1) bi- lobed apically, both lobes well devel- 286 Table | correspond with those used in the text). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Matrix of characters and taxa used in the cladistic analysis of Actocetor (numbers for characters neem Characters ——-vtt E Taxa Guttipsilopa (Nesopsilopa) Mimapsilopa Eremomusca Trypetomima Actocetor (A.) hovus Actocetor (A.) indicus Actocetor (A.) nigrifinis Actocetor (A.) afrus Actocetor (P.) yaromi Actocetor (P.) lindneri Actocetor (P.) decemguttatus OOOO OCROO COTTA 22222222 12345 6789012345678901 23456789 00110 1003000000?70??00 00000000 00000 0000010100?0??00 00001000 00100 0001110100?70??00 50000000 00100 000211021000??00 00000000 1 I AV OPOMO ZV LO OOO O23 2210 LT AO MO MOA MAO OCOR On AAG MAL LAOACIMMLOAOZT WO) AaLO)3) 52510) MUNIN LMAACAOLOSUAOMO SLOZS2i0 HAL TM, ZOO AOA 2 AN 0) OVA (OME AL IE 2) SB) 11112 2202021210120111 40104131 VN DAO 20) 2a Io) ANOVA aL aE 27(0) EO) ak 3} 3 oped, subequal (a synapomorphy for A. decemguttatus and A. lindneri); (2) apex not bilobed, bluntly formed or an- gulate, tapered to a point toward one side (a synapomorphy for A. afrus, A. hovus, and A. nigrifinis; no male avail- able for A. yaromi). 28. Presurstylus: (0) variable in form, but usually rather simple structures, longer than wide; (1) bearing | round to oval lobe medially (a synapomorphy for A. afrus and A. nigrifinis); (2) bearing 2 lobes along ventral margin, medial lobe much larger, | smaller, thumblike pro- cess between medial lobe and expand- ed base (a synapomorphy for A. hovus and A. indicus); (3) bearing 4 lobes (a synapomorphy for A. decemguttatus and A. lindneri; no male available for A. yarom1). 29. Shape of hypandrium: (0) pouchlike, often moderately deep (as in the sub- genus Actocetor); (1) very deeply formed, with an extended keel-like pro- cess that extends into the abdomen (a synapomorphy for A. decemguttatus and A. lindneri; no male available for A. yaromi). ANALYSIS, RESULTS, AND CONCLUSIONS Multistate characters 5, 6, 7, 9, 13, 15, 17, 22, 25, 26, 27, and 28 were treated as nonadditive (—); character 11 is a multistate character that was intuitively kept additive; characters 24 and 29 are autapomorphies that were made inactive (]) for the analysis so that they did not contribute to the cal- culations of the consistency and retention indices. The autapomorphies were kept in the matrix (Table 1) so that they would ap- pear on the cladogram as evidence corrob- orating the monophyly of the lineages they support. Using an exhaustive search, the implicit enumeration (ie*) option of Hennig86, 10 most parsimonious tree (Fig. 65) were gen- erated from the analysis of the 29 charac- ters. These cladograms have a length of 60 steps and consistency and retention indices of 0.83 and 0.83 respectively. The basal nodes in each of the 10 cladograms are identical with variation only in the more de- rived species within the two subgenera of Actocetor. The matrix was then subjected iteratively to successive weighing (xs w, ie*, cc) to determine a character’s contribution or weight (Carpenter 1988, Dietrich and McKamey 1995). The successive weighing stabilized at 418 steps and produced three cladograms that are identical to three of the first 10. A consensus of these three trees resulted in a tree that is identical to one of VOLUME 104, NUMBER 2 Table 2. Analysis of characters based on the cladogram (Fig. 61). Characters 1 2} 3 4 5 6 Steps ] ] l 2 y) Con. Index 100 100 100 100 + ©100- 100 Ret. Index 100 100 100 100 = =100~ 100 Characters 16 iL7/ 18 19 20 21 Steps 3 2 2 ] 2 1 Con. Index 355 OOM 100 00 50 =100 Ret. Index O 100 100 100 66 100 the three weighed trees and to one of the original 10 trees. The consensus tree, al- though with one unresolved lineage (a tri- tomy), is our cladogram of choice (Fig. 65). The analysis of the characters for the clad- ogram is given in Table 2 and the weights of the various characters are given in Table 3. Given these character weights, the anal- ysis of the cladogram resulted in revised consistency and retention indices of 0.94 and 0.95 respectively. Two basal, monophyletic clades are im- mediately evident within Actocetor, and as genus-group names are available for these clades, we are recognizing them as subgen- era, Actocetor and Poecilostenia (other- wise, recognition of these clades as species- groups would have been adequate). Poecilostenia comprises three species that occur exclusively in the Afrotropics and form an unresolved tritomy. One of these species, A. yaromi, is represented by a single female, and as we noted previously, we suspect that additional Afrotropical spe- cies will yet be discovered in this subgenus. The most widespread of these three species is A. decemguttatus. We predict that the 287 7 8 y 10 11 12 13 14 15 4 I 3 1 2 | 3 2 3 50 100 100 100 100 100 66 50 100 50 100 100 100 100 100 66 66 100 22 23 24 PID) 26 Dif 28 29 6 I 3 5 3 3 Soe LOO LOO ROO R 100 66 100 100 50 LOO MOOR OO 00 66 100 100 character states found in A. decemguttatus and A. lindneri, such as the lack of well- developed setae on the dorsum of the epan- drium, will also apply to A. yaromi. The subgenus Actocetor includes four species, and most of these have widespread distributions. An exception is A. afrus, which is thus far only in Senegal and Li- beria and which has a partially attenuate wing. The placement of A. afrus within Ac- tocetor sensu stricto and as the sister spe- cies of the lineage giving rise to A. indicus and A. hovus indicates that an attenuate wing, as found in A. afrus and conspicu- ously so in species of Poecilostenia, arose twice. As there are specific and evident dif- ferences in the narrowing (see descriptions), we suggest that its independent occurrence in two separate lineages is quite plausible. A concluding discussion point concerns the geographic origins of Actocetor. With Poecilostenia being exclusively Afrotropi- cal in distribution, and with most species of the subgenus Actocetor also occurring pri- marily in the Afrotropical Region, we sug- gest that the genus arose there. We would advocate this more strongly if we had con- Table 3. Weights (varying between 1-10) and status (additive = +, nonadditive = —, active = [, inactive = ]) of characters after successive weighing. Characters | 2 3 4 5) 6 i 8 9 10 Weight, status 10+] 10+[ 10+[ 10+[ KO=(I NO=|I 2=[ 10+[ 10—[ 10+[ Characters 11 12 13 14 15 16 iL7/ 18 19 20 Weight, status 10+[ 10+[ 4—[ arr IKO=|I O+[ 10-[ O+[ 10+[ 3+[ Characters 21 PEP 23 24 Dis) 26 2 28 29 Weight, status 10+[ 4-[ 10+[ 10+[ KO=|| IKO=|I 4—[ 10+[ 10+[ 288 clusive evidence on the sister group of Ac- tocetor. We suggest that the occurrence of A. indicus in the southern Palearctic Region and perhaps in the Oriental Region may represent subsequent dispersion of this spe- cies from its Afrotropical origins. An alter- native hypothesis is suggested by the clad- ogram. If Trypetomima, which is Oriental and Oceanian in distribution, is the sister group to Actocetor and if A. nigrifinis, which also occurs in the Orient, is the bas- almost species, then the genus could have originated from speciation events in the Orient/Oceanian area with subsequent dis- persion to the Afrotropics. It is evident that numerous speciation events occurred in the Afrotropical Region, resulting in the pre- sent Afrotropical fauna being comparative- ly rich. ACKNOWLEDGMENTS We gratefully acknowledge the assistance and cooperation of many organizations and individuals who contributed to the field work and production of this paper. To Dr. David A. Grimaldi (AMNH), Mr. Nigel Wyatt (BMNH), Dr. J. R. Vockeroth (CNC), Dr. Pier Mauro Giachino (MRSN), Dr. Ru- dolf Meier (UZMC), Dr. Herman de Jong (ZMAN), Ms. Jenny Pohl (ZMHB), Dr. Ruth Contreras-Lichtenberg (NMW), Dr. J. C. Deeming (NMWL), Dr. David Barra- clough (NMSA), Dr. Hans-Peter Tschorsnig (SMN), Dr. Eliane De Coninck and Mr. Jos Debecker (MRAC), and Dr. Amnon Freid- berg (TAU) and their institutions, who loaned specimens, we express our sincere thanks. Dr. Tadeusz Zatwarnicki examined the holotype female of A. anormalipennis and kindly provided us with the data on the labels. The illustrations were carefully inked by Mr. Young T. Sohn. The color frontispiece was beautifully rendered by Sophie Alling- ton. Lucrecia H. Rodriguez and Hollis B. Williams provided technical support. For reviewing a draft of this paper, we thank Drs. James EK Edmiston, Amnon Freidberg, Stephen D. Gaimari, Richard S. Zack, and PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Tadeusz Zatwarnicki. We are very grateful to the Smithsonian Women’s Committee for providing a stipend for Torsten to partici- pate in the 10-week Research Training Pro- gram at the National Museum of Natural History. We are also grateful to Dr. David Challinor, former Assistant Secretary for Research, and to Dr. Anna K. Bernensmey- er, former Associate Director for Science, National Museum of Natural History, and Dr. David Pawson, Associate Director for Science, National Museum of Natural His- tory, Smithsonian Institution, for financial support to conduct research at The Natural History Museum (BMNH), London, Eng- land, through grants from the Research Op- portunity Fund. Field work on Aldabra and the granitic Seychelles was funded by a grant from the Aldabra Project, Smithsoni- an Institution (Dr. Brian E Kensley, direc- tor), the Research Opportunity Fund (Dr. David L. Pawson, Associate Director for Science), and was supported and facilitated by the Seychelles Islands Foundation (Lindsey Chong-Seng, Marc Nicette) and the Seychelles Bureau of Standards (Mr. Terence Coopoosamy). This paper resulted from the senior au- thor’s participation in the Research Training Program at the National Museum of Natural History (Smithsonian Institution) for a ten- week period during the summer of 2000. We also gratefully acknowledge the support and enthusiasm of Ms. Mary Sangrey, di- rector of the Research Training Program at the National Museum of Natural History (Smithsonian Institution). Her tireless ef- forts have made this program successful. LITERATURE CITED Acloque, A. 1897. Faune de France. Bailliere et Fils, Paris, 516 pp. Becker, Th. 1903. Agyptische Dipteren. Mitteilungen aus dem Zoologischen Museum in Berlin 2(3): 67-195. Bezzi, M. 1908. Simuliidae, Bombyliidae, Empididae, Syrphidae, Tachinidae, Muscidae, Phycodromi- dae, Borboridae, Trypetidae, Ephydridae, Droso- philidae, Geomyzidae, Agromyzidae, Conopidae. In Schultze, L., ed. Zoologische und anthropolo- VOLUME 104, NUMBER 2 gische Ergebnisse einer Forschungsreise im wes- tlichen und zentralen Siidafrika ausgefiihrt in den Jahren 1903-1905. Erster Band: Systematik und Tiergeographie. IV. Insecta (Erste Serie. D. Dip- tera (1). Denkschriften der Medicinisch-naturwis- senschaftlichen Gesellschaft zu Jena, 13: 179— 201. 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Mitteilungen aus dem Zoologischen Mu- seum in Berlin 31(1—2): 48-58. Zatwarnicki, T. 1996. A new reconstruction of the or- igin of eremoneuran hypopygium and its impli- cations for classification (Insecta: Diptera). Genus 7(1): 103-175. PROC. ENTOMOL. SOC. WASH. 104(2), 2002, pp. 291-299 BIOLOGY AND IMMATURE STAGES OF OCHROTRICHIA FOOTEI (TRICHOPTERA: HYDROPTILIDAE), A NEW MICROCADDISFLY FROM A TORRENTIAL MOUNTAIN STREAM J. B. KEIPER AND S. C. HARRIS (JBK) Department of Invertebrate Zoology, Cleveland Museum of Natural History, 1 Wade Oval Drive, Cleveland, OH 44106, U.S.A. (e-mail: jkeiper@cmnh.org); (SCH) De- partment of Biology, Clarion University, Clarion, PA 16214, U.S.A. Abstract.—The egg, larvae, pupa, and adult male of Ochrotrichia footei, n. sp., taken from waterfalls in southern California are described. The species is univoltine, with early instars present during the summer months, and overwintering as fifth instars within their cases. First instars hatch from eggs deposited in abandoned pupal cases attached to vertical rock faces. Gut contents analysis and laboratory observations of living larvae indicate that larvae are scrapers of periphyton, including diatoms, green algae, cyanophytes, and fungal hyphae. The generalist trophic ecology of O. footei may be a preadaptation that allows this species to thrive in a harsh and variable environment. Key Words: Ochrotrichia footei, microcaddisflies, new species, streams, larvae, scrapers, diatoms, Bacillariophyta, Cyanobacteria, Chlorophyta The number of described species of mi- crocaddisflies (Trichoptera: Hydroptilidae) from North America has been growing steadily during recent years (e.g., Harris and Sykora 1996, Moulton et al. 1999, Har- ris and Huryn 2000). Hydroptila and Och- rotrichia are the two most speciose genera with more than 50 and 100 species, respec- tively (Morse 1993), and most of the newly described species are placed in these gen- era. Despite efforts to catalog the species richness of the Hydroptilidae, the small size and often cryptic habits of the immature stages have created a lag in studies of their natural history and life cycles. Indeed, of the 300+ species from North America, lar- va/adult associations are known for only approximately 10% (Wiggins 1996, Keiper 1998, Keiper and Walton 1999). Species of Ochrotrichia are found in lo- tic habitats (Wiggins 1996) where they are frequently associated with algal mats (Flint and Herrmann 1976; Keiper and Foote 1998, 1999, 2000; Keiper and Walton 2000) or surfaces of rocky substrate where they scrape periphyton (Vaillant 1984, English and Hamilton 1986). It appears that Och- rotrichia larvae are equipped with robustly cusped mandibles that allow them to either scrape relatively smooth substrates, macer- ate biofilms, or pierce cells within filaments of the green alga Cladophora (Vaillant 1984, English and Hamilton 1986, Keiper 1999, Keiper and Foote 2000, Keiper and Walton 2000). The only food sources re- ported for Ochrotrichia immatures are Cla- dophora and periphytic diatoms (Keiper and Foote 2000, Keiper and Walton 2000). Early instars do not build portable cases, whereas the fifth, and final, instar constructs either a bivalved purse-like case, or a dome- shaped case (Ross 1944, Wiggins 1996). During the summer of 1999, Ochrotri- chia adults of a new species were reared 292 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON from larvae collected from a southern Cal- ifornia stream. We describe the male, egg, larvae, and pupa, and give details on its nat- ural history and feeding habits. MATERIALS AND METHODS We collected larvae from Fullers Mill Creek in the San Bernardino National For- est of western Riverside County, near the city of Idyl Wild. Diatom-encrusted rock and algal mats were brought back to the laboratory with living larvae and placed in petri dishes or aerated rearing chambers (Keiper and Foote 1996). Periphyton was scraped with a knife and scooped into col- lecting vials to obtain early instars. Larval behavior and development was observed, and adults reared from pupae. All labora- tory observations of living material were made at 20°C, and a 12:12 light:dark pho- toperiod maintained with incandescent lights. Representative immature stages were fixed in Kahle’s solution (Wiggins 1996) for 24 h and then preserved in 70% ethanol. Adults were placed directly in ethanol. Im- matures were described following the meth- ods of Keiper and Foote (1999). Because the case size for fifth instars varied between dates of collection, larvae and cases were measured separately for each date. On 27 September 1999 and 18 May 2000, fifth instars were collected with for- ceps and placed directly into hot water car- ried in the field in an insulated thermos. After approximately 30 s, larvae were transferred to vials of 10% formalin for preservation. In the laboratory, formalin- preserved specimens were dissected, and the guts removed and smashed on a micro- scope slide with glycerin and a cover slip. The gut contents were described and enu- merated for 5 larvae from May and 9 larvae from September. Descriptions of adults and larvae followed the terminology of Mar- shall (1979) and Wiggins (1996), respec- tively. The collecting site, Fullers Mill Creek, is a low order, snow-melt stream. Due to the remoteness of the site and rugged terrain, the source was not found. The stream forms cobble-strewn channels and also cascades over bedrock, creating numerous waterfalls and madicoles. During the spring and early summer, the habitat is torrential and diffi- cult to access. As the summer progresses and less snow-melt is available, the water slows to a trickle, and many areas bordering the stream remain wet from capillary action only. The green alga Spirogyra and the cy- anophyte Cylindrospermum were visible in some areas, the red alga Lemanea was pre- sent at cascades, and rocky surfaces were frequently brownish-green with diatoms, unicellular green algae, and filamentous cy- anophytes. Mosses were abundant at pe- ripheral slash zones. Larvae were collected from three water- falls at approximately 1,800—1,900 m ASL, each with abundant mosses and diatoms. Each area had a central spill-way where most of the water ran, as well as lateral splash zones. Each area was open to the sunlight, and a variety of trees, mostly oaks (Quercus spp.) and pines (Pinus spp.), grew along the margins of the stream. RESULTS Ochrotrichia footei Keiper and Harris, new species (Figs. 1—4) Adult male.—Length 3.5 mm. 31 anten- nal segments. Brown in alcohol. Abdominal segment VII with short ventromesal pro- cess; segment VIII ventrally angular; seg- ment IX square in lateral view, reduced to narrow bridge dorsally (Fig. 1). Segment IX deeply incised mesally in dorsal view (Fig. 2), but rounded ventrally, with shallow lat- eral incisions posteriorly, a pair of lateral apodemes at outer juncture with inferior ap- pendages (Fig. 3). Tenth tergum bearing several elongate processes, distally with narrow mesal process tipped with heavy spine subapically, anteriorly with pair of acute lateral processes on right margin, an- teromost process short, posteromost process VOLUME 104, NUMBER 2 N \© 2 Figs. 1-4. elongate and curving inward (Fig. 2); in lat- eral view segment X thin and elongate, large spine basolaterally and ventrally, dor- sal process ending in downward projecting spine, remainder of segment rectangular, with narrow, downward curving posterior sclerite, tuft of fine hairs laterad (Fig. 1); in ventral view with basomesal process, thin anteriorly and rounded posteriorly (Fig. 3). Inferior appendages in lateral view trian- gular, distally tapering to thin, sclerotized apex (Fig. 1); in dorsal view rectangular, with series of short pegs on inner margin, heavier spine at apex, right inferior append- 4; Ealesy) ~ oe See oe Ochrotrichia footei, male terminalia. |, Lateral. 2, Dorsal. 3, Ventral. 4, Phallus. age with lobe near base bearing numerous short pegs, left inferior appendage with sev- eral short pegs on inner margin near mid- length (Fig. 2); in ventral view wide basal- ly, tapering to midlength, distally rounded on outer margins, with short pegs on inner margin (Fig. 3). Phallus tubular and thin, small triangle at apex (Fig. 4). Diagnosis.—Ochrotrichia footei appears to be most similar to O. salaris Blickle and Denning in the overall configuration of ter- gum X. Both species have an elongate me- sal process which terminates in a lateral spine, and both species have a pair of lateral 294 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Sees T <5 0:4 0.3 ab) . =! 0.2 IV £ . 0.1 0 0.09 0.11 0.13 0.15 0.17 0.19 Head capsule width (mm) Fig. 5. Histogram of head capsule width for field-collected early instar Ochrotrichia footei (n = 40). Num- bers placed over peaks indicate estimated instar. processes. However, the inferior append- ages of the new species are triangular in lateral aspect, more similar to those of O. mono (Ross) than to those of O. salaris, which are narrow at midlength and bear heavy spines from the ventral margin, both anteriorly and posteriorly. The features of the tenth tergum in combination with the shape of the inferior appendages will serve to readily identify O. footei. Type material.—Holotype, ¢: United States, California, Riverside County near Idyl Wild, Fullers Mill Creek, reared from larva collected 29 June 1999, emerged 7 July 1999, J. B. Keiper; deposited at the Cleveland Museum of Natural History, De- partment of Invertebrate Zoology. Para- types: | 6, 4 2, same location as holotype, except reared from larvae collected 27 June 1999, J. B. Keiper; 2 6 pupae, collected 3 July 2000, J. B. Keiper. Paratypes deposited with holotype male, except for | male and 2 females (27 June 1999) deposited at the National Museum of Natural History, Smithsonian Institution, Washington, DC. Egg.—Round, chorion colorless, no sur- face markings evident; embryo bright yel- low, developing dark brown to black eye spot with age. Maximum diameter 0.20— 0:23mmi (x = (0.21) mm) ae =e10): Larvae.—Except for field-collected fifth instars and first instars obtained from eggs in the laboratory, determination of larval stadium was only possible by measuring the head capsule width of field material (n = 40 early instars). Using peaks in a histo- gram we approximated head capsule width (Fig. 5). Instars 1-4 without cases, as is typ- ical for Hydroptilidae (Wiggins 1996). First instar: Flattened dorsoventrally, non-sclerotized areas of body white, nearly transparent. Head: width 0.09—0.10 mm, pale brown, posterior margin with thin brown band, sutures evident, newly hatched larvae lacking coloration except for prima- ry setae and eye spot (Fig. 6). Thorax: sclerites pale brown, primary setae well de- veloped. Abdomen: dorsal sclerites rectan- gular, covering most of dorsum of each seg- ment; segment 10 bifurcate, bearing large anal claws. (n = 10). Second instar: Similar to first instar, just differing in size and proportions of body parts; sclerites darkening with age. (n = 10). Third instar: Sclerites darkening consid- erably. Head: Primary seta near eye spot long, 1.5 length of head capsule, other- wise similar to first instar. Thorax: One pair of long setae laterally on each notal sclerite, others small and weak. Abdomen: Three pairs of strong setae laterally; dorsal sclerite on segment 9 with one pair of long setae positioned mesad, subequal in length to pri- mary seta near eye spot. Otherwise similar to first instar except in size and proportion (Fig. 7). (n = 10). Fourth instar: Similar to third instar, dif- VOLUME 104, NUMBER 2 N \O Nn Figs. 6-12. Ochrotrichia footei, larvae. 6, Head capsule of first instar, right side only with setation. 7, Slide- mounted third instar, dorsal view, right side only with setation, left side only with thoracic legs; note damage to posterior of head capsule and flattening of body a result of slide preparation. 8, Head capsule of fifth instar, right side only with setation. 9, Fifth instar, mandibles, ventral view. 10, Fifth instar, prosternal sclerites, legs removed. 11, Fifth instar, abdominal segments 1 and 2, dorsal view. 12, Fifth instar, pronotal sclerite and fore leg, lateral view. fering only in size and proportions of body parts, sclerites uniformly dark brown. (n = 4). Fifth instar: Shape and coloration typical of Ochrotrichia, head and thoracic seg- ments mostly dark brown, grading to black or tan in some areas. Head: width 0.15—0.19 mm (x = 0.17 mm); eye spot round, black, surrounded by narrow pale ring ca. 0.015 mm wide; position of primary setae as in Fig. 8; longest seta near eye spot approxi- mately same length as head; anterior edge of labrum slightly concave in dorsal view; antenna short, with narrow base, bearing short seta approximately same length as base; right mandible bearing one seta on posterolateral corner, left mandible bearing two (Fig. 9); left mandible with row of hair- like setae on inner margin among dentition; mandibles asymmetrical, each robustly cusped. Thorax: prosternal sclerites dark brown, anterior sclerite small, rectangular, 296 and divided by thin membranous area, two posterior sclerites narrowing laterally, trun- cate medially (Fig. 10); each sclerite mostly dark brown, with anterior margin lined with dark setae curving anteroventrad; notum with black posterior and lateral margins, black area extending somewhat along an- terior margin in darker specimens, never wider than 0.05 mm; legs dark brown to black, foreleg 0.30 mm, midleg 0.36 mm, and hindleg 0.36 mm, leg ratio 0.83:1.00: 1.00; fore tibia and tarsus with anterior margin lined with short hairs, fore tibia with strong ventral projection bearing stout setae, tarsal claw curved with short basal seta (Fig. 12); mesothoracic sclerite with black anterolateral corner; metathoracic sclerite with anterolateral corner produced, tan laterally. Abdomen: milky white, ap- proximately 3.0—3.5X as wide as thorax in lateral view, approximately 2.0 as wide as thorax in dorsal view, somewhat com- pressed laterally; brown sclerotized rings dorsally on segments 2—8 (Fig. 11), rings on segments 2—4 oblate with lateral edges truncate, rings on segments 5—8 more rounded; segment | with dorsal sclerite di- vided by central membranous band, sclerite fading to pale color laterally; sclerites on abdominal segments 9—10 uniformly dark brown, anal claw somewhat lighter brown than other sclerites, approximately 0.01 mm long; case comprised of minute sand grains in two purse-shaped valves, typical of that for Ochrotrichia (Wiggins 1996). (n = 10, mature fifth instars only). Newly molted fifth instars bear sclerites that are somewhat lighter brown than ma- ture larvae. The dorsomedial sclerite of ab- dominal segment | is less extensive and dif- ficult to see. Otherwise, these young fifth instars are similar except that the abdomen is not swollen and approximately the same girth as the thoracic segments. Length of larvae taken in June were 2.47—4.34 (* = 3.31) mm, while those taken in September measured 1.46—3.07 (x = 2.12) mm; cases collected in June were 3.21—4.18 (* = 3.68) mm in length and contained mature larvae, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON whereas those cases collected in September contained smaller larvae, and were 1.10— 2.04 (x = 1.61) mm in length. Pupa.—Typical of Hydroptilidae; 2 d, length 2.7 and 2.9 mm. No distinguishing characters. Etymology.—Named for Dr. Benjamin A. Foote, Professor Emeritus, Kent State University, who introduced the senior au- thor to Hydroptilidae, in honor of his many contributions to aquatic entomology. Biology.—One female was observed to emerge on 27 September 1999 from the face of a waterfall. The pharate adult slid down a shallow film of water running over the rock face. After travelling about 20 cm, the adult broke free and flew to mosses ad- jacent to the water; the exuvia continued to move downward and was lost. The adult sat motionless for approximately one minute, took wing, and was collected with an aerial net. Three other adult females were ob- served on the same day walking over a wa- ter film, each dragging its ovipositor over the water. No oviposition was noted, nor were eggs located during a search of the rock face. Two of the females were col- lected, brought back to the laboratory alive, and placed in petri dishes with moist dia- tom-covered rocks. No oviposition oc- curred. On 6 July 1999, an empty pupal case was found to contain approximately 50 eggs. The eggs adhered to each other forming a mass within the pupal case. Four more cas- es with similar egg masses were collected on 27 September. The eggs collected 27 September were placed in a petri dish with stream water and small pieces of diatom- covered rock; the eggs hatched two days later. Although larvae appeared to feed by scraping the substrate, no growth occurred. All larvae died within 5 d. Because only field collected eggs have been studied, the incubation period remains unknown. In the field, active fifth instars were re- stricted to areas of noticeable water flow, rather than the wet margins of the water- falls. Early instars were too small for field VOLUME 104, NUMBER 2 Table 1. and September. Algal and cyanobacterial taxa found in the guts of Ochrotrichia footei fifth instars taken in May Month Collected Genus Division May September Cocconeis Bacillariophyta xX xX Cymbella Bacillariophyta x x Diatoma Bacillariophyta x Epithemia Bacillariophyta x Gomphonema Bacillariophyta 4 Surirella Bacillariophyta xX Synedra Bacillariophyta x x Chlamydomonas — Chlorophyta xX Oedogonium Chlorophyta x Anabaena Cyanobacteria x Chroococcus Cyanobacteria x Phromidium Cyanobacteria 4 Pseudanabaena Cyanobacteria x Trachelomonas Euglenophyta xX observations, but scrapings of algal films from areas of flowing water contained early instars while scrapings from wet margins lacked them. Only fifth instars that had at- tached their cases prior to pupation, or that had temporarily attached their cases to the rock to facilitate case-building efforts, were found in the thin water film at the margins of waterfalls. Fifth instars brought back to the labora- tory scraped diatom covered rocks placed in petri dishes and rearing chambers, but did not form pupae. Three field-collected pupae produced adults (2 d, 1 2) after =10 days. Larvae did not survive long enough in the laboratory for detailed observations of feeding habits. Gut contents analysis revealed that dia- toms were the most common item ingested, and ranged in size from 2.0—20.8 microns in length. Most diatoms were fragmented. Larvae collected in September never had more than 10 diatoms in their guts, whereas those collected in May contained 32—210 diatoms. Eleven algal and cyanophyte taxa were found in the guts of specimens from May, while only six were identified in Sep- tember samples (Table 1). Fungal hyphae were present in specimens only in May. 29 Pupae formed aggregations in the field, and were often too numerous to count. All pupal cases were oriented parallel to the di- rection of water flow. During October 1999 visits to the stream, the rock substrates re- mained moist only in limited areas and many pupal cases were dry. Collections showed that no cases supported eggs, lar- vae, or pupae. A number of cases exhibited holes in one lateral valve, possibly made by the predatory behavior of an undetermined species of Rhyacophila (Rhyacophilidae) that inhabited the waterfall areas. DISCUSSION Fifth instars are separated from all other known Ochrotrichia larvae by the combi- nation of uniformly dark brown sclerites, narrow ring around eye spots, left mandible with two posterolateral setae, right mandi- ble with one posterolateral seta, prosternal sclerites separated by a median gap, medi- odorsal abdominal sclerites of segment | with longitudinal membranous band, ab- dominal segments II—VIII bearing medi- odorsal ring sclerites with membranous centers, and cases purse-shaped bearing two lateral valves. Immatures are probably re- stricted to waterfalls or madicolous habi- tats, which is not uncommon for hydroptilid species (e.g., Ross 1944, Vaillant 1984, Wells 1985, Keiper and Walton 2000). Much of the life history appears similar to that described for other Ochrotrichia spp. (Flint and Herrmann 1976, Vaillant 1984, English and Hamilton 1986, Keiper and Walton 2000). Certain species of Hydrop- tilidae have been reported to be intimately associated with filamentous algae as pierc- er/herbivores (Nielsen 1948; Keiper and Foote 1998, 1999, 2000), but O. footei nev- er exhibited the algal piercing habit. Water- falls that range from torrential to virtually dry may preclude populations of Hydrop- tilidae that exhibit specific trophic habits (e.g., Resh and Houp 1986; Keiper et al. 1998; Keiper and Foote 1998, 2000), thus the relatively generalized feeding habits may have preadapted O. footei for existence 298 at Fullers Mill Creek. The mandibles appear well-suited for attacking rock substrates, and allow larvae to consume a variety of materials, similar to that of O. quadrispina (Keiper and Walton 2000). There appeared to be a dietary shift from May to September as more diatoms were consumed and a greater number of algal/cyanophyte taxa were present in guts during that period of greatest water flow. During May, three cy- anophyte genera were present in guts, whereas Anabaena was the only cyano- phyte taxon present in September. The green alga Oedogonium was the only chlo- rophyte consumed during late summer, and has been reported as a food source of the hydroptilid Oxyethira arizona Ross (Keiper and Walton 1999). The dietary shift was probably not behavioral, but resulted be- cause of a change in the periphyton com- munity as the habitat dried. These obser- vations suggest that categorizing macroin- vertebrates into functional feeding groups based on gut contents analysis may lead to spurious results if such analyses are taken from specimens collected from only one time period. We can infer certain biological details based on laboratory and field observations. Ochrotrichia footei is probably univoltine and overwinters as fifth instars. Adults are active during summer months and multiple cohorts of immatures coexist. Small fifth in- stars with short cases are present in the au- tumn, and overwinter inside their cases. Larvae seal their cases in preparation for pupation after case building and growth are concluded in the spring. Due to the remote- ness of the area, difficulty associated with field observations in torrential streams, and difficulty with rearings, many biological de- tails remain unknown. A particularly fascinating aspect of the biology of O. footei is that females place their eggs in one mass inside of empty pu- pal cases. Based on laboratory and field ob- servations, it is evident that females leave their cases; parthenogenesis has never been reported in Trichoptera to our knowledge. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON After mating, females then must find a suit- able case for egg deposition. All six cases with eggs were undamaged, and open to the environment through the apical slit in the case formed by escaping pharate adults. Few observations of hydroptilid oviposition have been recorded (but see Nielsen 1948, Resh and Houp 1986, Ito 1997), but some species do create egg masses. The behav- ioral adaptation of inserting eggs into at- tached cases may provide certain benefits, such as protection from desiccation, ultra- violet light, predators, or torrential waters. ACKNOWLEDGMENTS We thank Michelle Sanford, Joshua Jian- nino (University of California, Riverside), Parker Workman (Harvard University), and Pamela Keiper (Cleveland Museum of Nat- ural History) for assisting the senior author on collecting trips to Fullers Mill Creek. Dale Casamatta (Ohio University) kindly identified algal and cyanophyte specimens. We extend our appreciation to Joseph Han- nibal (CMNH), Oliver Flint (Smithsonian Institution), and Benjamin Foote (Kent State University) for reviewing the manu- script. LITERATURE CITED English, W. R. and S. W. Hamilton. 1986. The larva of Ochrotrichia arizonica (Trichoptera: Hydrop- tilidae) with notes on distribution and geographic variation. Journal of the Kansas Entomological Society 59: 474-479. Flint, O. S. and S. J. Herrmann. 1976. The description of, and environmental characterization for, a new species of Ochrotrichia trom Colorado (Trichop- tera: Hydroptilidae). Annals of the Entomological Society of America 69: 894-898. Harris, S. C. and A. D. Huryn. 2000. New and rare microcaddisflies (Trichoptera: Hydroptilidae) from the eastern United States. Entomological News 111: 77-83. Harris, S. C. and J. L. Sykora. 1996. New species of microcaddisflies from the eastern United States (Insecta: Trichoptera: Hydroptildiae). Annals of the Carnegie Museum 65: 17-25. Ito, T. 1997. Oviposition preference and behavior of hatched larvae of an oligophagous caddisfly, Pa- laeagapetus ovatus (Hydroptilidae: Ptilocolepi- nae), pp. 177-181. Jn Holzenthal, R. W. and O. S. VOLUME 104, NUMBER 2 Flint, eds. Proceedings of the 7th International Symposium on Trichoptera. Ohio Biological Sur- vey, Columbus. Keiper, J. B. 1998. Biology, larval feeding habits, and resource partitioning by microcaddisflies (Tri- choptera: Hydroptilidae). Ph.D. Thesis, Kent State University, Kent, Ohio. 146 pp. . 1999. Morphology of final instar Ochrotrichia xena (Trichoptera: Hydroptilidae). Entomological News 110: 231-235. Keiper, J. B., D. A. Casamatta, and B. A. Foote. 1998. Use of algal monocultures by larvae of Hydroptila waubesiana and Oxyethira pallida (Trichoptera: Hydroptilidae). Hydrobiologia 380: 87-91. Keiper, J. B. and B. A. Foote. 1996. A simple rearing chamber for lotic insect larvae. Hydrobiologia 339: 137-139. . 1998. Biological Notes on Ochrotrichia xena (Ross) (Trichoptera: Hydroptilidae), a species newly recorded for Ohio. Proceedings of the En- tomological Society of Washington 100: 594—595. . 1999. Biology and immature stages of two species of Hydroptila Dalman (Trichoptera: Hy- droptilidae) which consume Cladophora (Chlo- rophyta). Proceedings of the Entomological So- ciety of Washington 101: 514-521. . 2000. Biology and larval feeding habits of coexisting Hydroptilidae (Trichoptera) from a small woodland stream in northeastern Ohio. An- nals of the Entomological Society of America 93: 225-234. Keiper, J. B. and W. E. Walton. 1999. Biology and morphology of the mature larva of Oxyethira ar- izona Ross (Trichoptera: Hydroptilidae). Pan-Pa- cific Entomologist 75: 212-220. . 2000. Biology and immature stages of Och- rotrichia quadrispina Denning and Blickle (Tri- 299 choptera: Hydroptilidae), a spring-inhabiting scraper. Proceedings of the Entomological Society of Washington 102: 183-187. Marshall, J. E. 1979. A review of the genera of the Hydroptilidae (Trichoptera). Bulletin of the British Museum of Natural History 39: 135-239. Morse, J. C. 1993. A checklist of the Trichoptera of North America, including Greenland and Mexico. Transactions of the American Entomological So- ciety 119: 47-93. Moulton, S. R., S. C. Harris, and J. P. Slusark. 1999. The microcaddisfly genus /thytrichia Eaton (Tri- choptera: Hydroptilidae) in North America. Pro- ceedings of the Entomological Society of Wash- ington 101: 233-241. Nielsen, A. 1948. Postembryonic development and bi- ology of the Hydroptilidae. Det Kongelige Danske videnskabernes selskabs Biologiske Skrifter 5: 1— 200. Resh, V. H. and R. E. Houp. 1986. Life history of the caddisfly Dibusa angata and its association with the red alga Lemanea australis. Journal of the North American Benthological Society 5: 28—40. Ross, H. H. 1944. The caddisflies, or Trichoptera, of Illinois. Bulletin of the Illinois Natural History Survey 23: 1-326. Vaillant, E 1984. The hydroptilid larvae of dripping rocks, pp. 407—412. In Morse, J. C. ed., Proceed- ings of the 4th International Symposium on Tri- choptera. Series Entomologica 30, Dr. W. Junk Publ., Boston. Wells, A. 1985. Larvae and pupae of Australian Hy- droptilidae (Trichoptera), with observations on general biology and relationships. Australian Jour- nal of Zoology, Supplementary Series 113: 1—69. Wiggins, G. B. 1996. Larvae of the North American caddisfly genera (Trichoptera), 2nd Ed. University of Toronto Press, Toronto. 457 pp. PROC. ENTOMOL. SOC. WASH. 104(2), 2002, pp. 300-308 INSECTS (DIPTERA, COLEOPTERA, LEPIDOPTERA) REARED FROM WETLAND MONOCOTS (CYPERACEAE, POACEAE, TYPHACEAE) IN SOUTHERN QUEBEC FREDERIC BEAULIEU AND TERRY A. WHEELER Department of Natural Resource Sciences, McGill University, Macdonald Campus, Ste- Anne-de-Bellevue, QC, H9X 3V9 Canada (e-mail: wheeler@nrs.mcgill.ca) Abstract.—Information about feeding habits and host-plants is given for 24 species of insects reared from the stem or leaves of wetland herbaceous monocots, primarily Carex lacustris Willd. and Phalaris arundinacea L., in southern Quebec. Nineteen species of Diptera in eight families (1 Dolichopodidae, 2 Otitidae, | Anthomyzidae, | Agromyzidae, 1 Chamaemyiidae, 1 Sphaeroceridae, 11 Chloropidae, 1 Scathophagidae), three Lepidop- tera in the families Cosmopterigidae, Elachistidae, and Noctuidae, and two weevils (Co- leoptera: Curculionidae) were reared from four species of host plants. Seven species are phytophagous primary invaders, thirteen are secondary invaders (mostly saprophagous) inside the burrows of primary invaders, two are predators of aphids and other invertebrates living on the leaves of the plants, and two have unknown feeding habits. Key Words: Diptera, Coleoptera, Lepidoptera, ecology, stem-borers, secondary invaders, host plants, wetlands, Carex, Phalaris Emergent wetland plants provide a mi- crohabitat and food source for many in- sects, especially Diptera (Gaevskaya 1969, Ferrar 1987). The higher Diptera fauna in wetlands is diverse and many species, par- ticularly in the family Chloropidae, are as- sociated with emergent plants such as sedg- es (Cyperaceae), rushes (Juncaceae) and grasses (Poaceae) (Davis and Gray 1966, Todd and Foote 1987a, 1987b, Pollet 1992). Although there have been several studies on the biology of selected Diptera feeding on wetland monocots (e.g., Valley et al. 1969b, Rogers et al. 1991, Allen and Foote 1992, Wearsch and Foote 1994, Valley and Foote 1996), the host plants and feeding habits are unknown for many species. Many flies associated with emergent plants are saprophagous secondary invaders of plant tissues damaged by other insects. In many cases, the feeding habits of pri- mary invaders other than Diptera are also poorly known. This study provided the op- portunity to obtain biological information on some species of phytophagous Lepidop- tera and Coleoptera associated with wetland plants. This paper documents the results of rearing studies on Diptera and other phytophagous insects associated with emergent wetland monocots in southern Quebec. Host records and ecological information are given for 19 species of Diptera, three species of Lepidop- tera, and two species of weevils (Coleoptera: Curculionidae) reared from Cyperaceae, Po- aceae and Typhaceae. MATERIALS AND METHODS Infested plants were collected from May— October 1999 in sedge meadows of the Lac St. Francois National Wildlife Area in southern Quebec, and May 2000 at Stoney- VOLUME 104, NUMBER 2 Table 1. 301 Insects reared from Carex lacustris and Phalaris arundinacea in southern Quebec. COL = Cole- optera; LEP = Lepidoptera; all other species are Diptera. Trophic Group Carex lacustris Phalaris arundinacea primary invaders Chlorops seminiger Cosmopterix fernaldella (LEP) Elachista sp. CLEP) Chromatomyia fuscula Apamea ophiogramma (LEP) Sphenophora costipennis (COL) secondary invaders Chaetopsis massyla Lasiosina canadensis Rhopalopterum atriceps Orthacheta hirtipes predators Plunomia elegans Thaumatomyia obtusa unknown Medetera aberrans Anthomyza sp. Chaetopsis ?fulvifrons Conioscinella zetterstedti Oscinisoma alienum Elachiptera penita Elachiptera angustifrons Eribolus longulus Eribolus nana Eribolus nearcticus croft Pond, adjacent to McGill University’s Morgan Arboretum in Ste-Anne-de-Belle- vue, Quebec. Infested stems of Carex, Scir- pus (Cyperaceae) and Typha latifolia L. (Typhaceae) were identified by the presence of curled, decumbent, yellowish-brown in- ner leaves, contrasting with green, straight, healthy outer leaves (Neff and Wallace 1969). Infested stems of Phalaris arundi- nacea L. (Poaceae) were detected by the presence of yellowish terminal leaves and shorter shoots than intact plants. Infested leaves were distinguishable by the presence of mines visible on the leaf surface. Infested plants were transported to the lab in plastic bags and either examined im- mediately or refrigerated for later exami- nation. Stems were dissected to locate lar- vae and any observed feeding behavior was noted. The lower end of each infested leaf was placed in a | cm X 5 cm vial filled with water. The vials containing leaves, or small cut sections of stems containing lar- vae, were placed in a plastic container 4.5 cm in diameter and 7.5 cm deep, covered with fine mesh. These were kept at 22°C in a sealed plastic bag containing moist paper towel in order to keep humidity high. Con- tainers were inspected every 2—3 days for emerged adults. Plants collected late in the season (late August to October) were put in an incubator at 4°C for 1—2 months before dissection to simulate overwintering for the larvae. Additional collecting in the field using sweep nets and pan traps around host plants provided data on the adult activity of some insect species. All specimens collected and reared are deposited in the Lyman Ento- mological Museum, McGill University, Ste- Anne-de-Bellevue, QC, except Microlepi- doptera, which are in the Canadian National Collection of Insects, Ottawa, ON. RESULTS Infested plants were mostly Carex lacus- tris, collected in the Lac St. Frangois Na- tional Wildlife Area, and Phalaris arundi- nacea, collected at Stoneycroft Pond. In- sects reared from these host plants are listed in Table 1. Species are classified as phy- tophagous primary invaders, secondary in- vaders (saprophages and/or facultative predators), and predators of other insects feeding externally on the plant. Ecological observations and previous records of reared species are included in the annotated list be- low. 302 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Diptera Medetera aberrans Wheeler (Dolicho- podidae).—Two adults were reared from larvae found on the inner leaves of two shoots of C. lacustris at a height of 17 and 20 cm from the culm base (Lac St. Fran- cois, 1S.1x1999, emerged 0421-20007): same except 01.x.1999, emerged 02.1.2000 (1)). A narrow, brown feeding trail was ob- served on two leaves of one of the shoots, but no other arthropods were found in as- sociation with the M. aberrans larvae. The larvae of several Medetera species live un- der tree bark, and are predators of soft-bod- ied arthropods (Bickel 1985). However, Bickel (1985) noted that M. aberrans and related species are more closely related to Thrypticus Gerstiacker (a phytophagous ge- nus) and Dolichophorus Lichtwardt than they are to Medetera sensu stricto and sug- gested that the larvae of M. aberrans may be associated with wetland plants. Although the presence of a feeding trail and the ab- sence of other arthropods might suggest that M. aberrans larvae are phytophagous, the evidence is not conclusive and the lar- val habits remain unknown. Chaetopsis ?fulvifrons Macquart (Otiti- dae).—Three adults were reared from two stems of P. arundinacea attacked by Apamea ophiogramma (Noctuidae) (Stoneycroft Pond, 31.v.2000, emerged 27.vi.2000). Val- ley et al. (1969a) considered C. fulvifrons a secondary invader in the heads of Echinoch- loa crusgalli (L.) (Poaceae) attacked by the phytophagous otitid Eumetopiella rufipes (Macquart). Johnson (1921) stated that C. ful- vifrons injures corn, sugar-cane and onions, but did not indicate whether he considered the larvae phytophagous or saprophagous. Chaetopsis massyla (Walker) (Otitidae).— Adults were reared from three species of host plants. Most specimens were reared from multiple stems of Typha latifolia, usually fol- lowing primary infestation by a moth or wee- vil (Lac St. Francois, 23.vii.1999, emerged latevanat L999) .(2) eel 2vaniel/ 999 =56) aan 13.vii.1999 (1); same except 7.viii.1999, emerged 16:vi11.1999' -(1))"same. except 12.viii.1999, emerged 17.viii.1999 (1) and 30.vili.1999 (1)); one adult was reared from C. lacustris (Lac St. Frangois, 16.vii.1999, emerged 05.viii.1999); three adults emerged from a rotting shoot of Scirpus fluviatilis (Torr.) Gray (Cyperaceae) along with an adult of Pullimosina pullula (Zetterstedt) (Sphaer- oceridae) (Lac St. Francois, 7.viii.1999, emerged 7.1x.1999). Adults were abundant on C. lacustris at Lac St-Frangois. This species was previously reared as a secondary invader from T. latifolia, C. lacustris and Scirpus mi- crocarpus Presl. (Valley et al. 1969b, Allen and Foote 1992). This is the first record of C. massyla from S. fluviatilis. Plunomia elegans Curran (Chamaemyi- idae).—Several adults were reared from pu- paria found between the leaves of C. lacus- tris (Lac St. Frangois, 05.vii.1999, emerged 11.vii.1999 (1); same except 12.vii.1999, found dead 22.xi.1999 (1); same except 23.v1i.1999, emerged 02.viii.1999 (1); same except 01.x.1999, emerged 14.11.2000 (1); same except 27.x.1999, emerged 14.11.2000 (1)). There are no previous records of the ecology of Plunomia spp., but given that all Chamaemyiidae whose feeding habits are known are predators of Aphidoidea and Coccoidea (Ferrar 1987), the larvae proba- bly prey on one of these taxa. Specimens of Thripsaphis ballii (Gillette) (Drepanosi- phidae) were abundant on C. /acustris at the study site and are the likely prey of P. ele- gans. Adults of P. elegans were abundant in stands of C. lacustris and C. aquatilis Wahlenb. by 14 May 1999, peaked in July, and persisted until early September. Unpub- lished collection data on specimens in the Canadian National Collection of Insects in- dicate that P. elegans lives in wet habitats such as grassy marshes, sedge meadows and bogs. Chromatomyia fuscula (Zetterstedt) (Agromyzidae).—Four adults were reared from leaf mines in individual leaves of P. arundinacea (Stoneycroft Pond, 31.v.2000, emerged 11.vi.2000).This species has been reared from several cereals and wild grass- VOLUME 104, NUMBER 2 es, including P. arundinacea, in Europe (Griffiths 1980); this is the first North American record from this host. Previous Nearctic records were from the grasses Beckmannia erucaeformis L. and Cinna la- tifolia (Trev.) Griseb. in western North America (Griffiths 1980). Anthomyza sp. (Anthomyzidae).—One adult was reared from a larva that was found 10—15 cm from the culm base of C. lacustris (Lac St. Francois, 19.x.1999, emerged 09.11.2000).The shoot had signs of internal feeding damage but no exit holes or larvae of other species were seen; this suggests that the anthomyzid larva was a primary invader, although this is not cer- tain. The larval feeding habits of the An- thomyzidae are practically unknown (Foote 1991). Some Anthomyza spp. have been found between the sheathing leaves of grasses, Juncus and Typha (Nye 1958, Uff- en and Chandler 1978, Ferrar 1987), but it is not known whether the larvae were phy- tophagous or saprophagous. This specimen was not identified to species because most Nearctic species of Anthomyza are undes- cribed and there is no key to species. Chlorops seminiger Becker (Chloropi- dae).—This species was reared as a primary invader from ten C. lacustris stems (Lac St. Francois, 01.x.1999, emerged 28.x1i.1999 (1); same except emerged 02.i1.2000 (1); same ex- cept 19 x1999 -emerced) Silexn1999 (1); same except emerged 24.1.2000 (1); same ex- cept emerged 29.1.2000 (1); same except 27.x.1999, emerged 241.2000 (5 from 4 stems); same except emerged 29.1.2000 (1)). Emerging adults were always solitary, except in One case where two flies emerged from the same stem. The mature larvae were deep in the culm, very close to the base. Adults were collected at the study site from mid-May to late June. Sedges from which the adults were reared were collected in October 1999; the larvae are more mature late in the season and damage to Carex stems is more evident. The flies probably overwinter as second- or third- instar larvae because larvae collected in Oc- tober were large (over 0.6 cm long) and be- 303 cause most other chloropid stem-borers of Cyperaceae whose life cycle is known over- winter as second or third instars (Rogers et al. 1991, Wearsch and Foote 1994). This is the first host-plant record for C. seminiger. Chlorops certimus Adams and C. frontosa Meigen have been reared from Carex species (Nye 1958, Rogers et al. 1991) and Chlorops obscuricornis Loew is a primary invader of stems of Eleocharis (Cyperaceae) (Wearsch and Foote 1994). Most other rearing records of Chlorops species are from grasses (Nye 1958, Ferrar 1987). Conioscinella zetterstedti Andersson (Chloropidae).—Two brachypterous adults emerged from one stem of P. arundinacea infested by Apamea ophiogramma, along with Eribolus nearcticus and Oscinisoma alienum (Stoneycroft Pond, P. arundina- cea, 31.v.2000, emerged 05.vii.2000). This is the first host-plant record for C. zetter- stedti. Conioscinella zetterstedti occurs in a range of habitats including peatlands, sedge meadows, shoreline debris, among grass roots and leaf litter and in seashore dune grasses (Wheeler 1994), suggesting that it is a generalist scavenger on plant material. Other Conioscinella species have a wide range of habits but many are apparently sa- prophagous (Ferrar 1987). Seven species of Chloropidae (C. zetter- stedti, Elachiptera angustifrons, Elachip- tera penita, Eribolus longulus, Eribolus nana, Eribolus nearcticus, Oscinisoma al- lenum) were reared from a total of four stems of P. arundinacea infested by A. ophiogramma. The chloropid larvae were mostly in the lower part of the burrow ex- cavated by the noctuid larvae, which was usually higher in the burrow. Elachiptera angustifrons Sabrosky (Chloropidae).—One adult emerged from one stem of P. arundinacea infested by Apamea ophiogramma, along with an adult of E. penita and Eribolus spp (Stoneycroft Pond, 31.v.2000, emerged 21.vi.2000). This is the first host-plant record for E. angus- tifrons. Species of Elachiptera are usually considered secondary invaders of grasses 304 and plant scavengers in general, usually in wet habitats (Valley et al. 1969b, Ferrar 1987). Elachiptera penita (Adams) (Chloropi- dae).—One adult emerged from one stem of P. arundinacea infested by Apamea ophiogramma, along with an adult of E. an- gustifrons and Eribolus spp (Stoneycroft Pond, 31.v.2000, emerged 21.vi.2000). This is the first host-plant record for Elachiptera penita. Eribolus longulus (Loew) (Chloropi- dae).—Adults emerged from two stems of P. arundinacea infested by Apamea ophio- gramma, along with E. nana and E. nearc- ticus (Stoneycroft Pond; 31.v.2000, emerged 17.vi.2000 (3) and 19.vi.2000 (1)).This is the first record of this species from Phalaris. Three other larvae and 18 pupae, probably Eribolus species, died be- fore emergence in one of the stems. Like Elachiptera, species of Eribolus are usually considered secondary invaders of grasses and other plant material in wet habitats (Ferrar 1987, Valley and Foote 1996) and E. longulus is apparently confined to wet open habitats dominated by sedges or grass- es (Valley and Foote 1996). Eribolus lon- gulus has been reared from damaged stems of Carex and Scirpus, developing flowers of /ris, and several genera of grasses (Ave- na, Echinochloa, Glyceria, Triticum) (Val- ley et al. 1969b, Valley and Foote 1996). Eribolus nana (Zetterstedt) (Chloropi- dae).—Adults emerged from two stems of P. arundinacea infested by Apamea ophio- gramma, along with E. longulus and E. nearcticus (Stoneycroft Pond, 31.v.2000, emerged 17.vi.2000 (1), 19.vi.2000 (1), 21.vi.2000 (1), 22.vi.2000 (1), 27.vi.2000 (1) and 30.vi.2000 (1)). This is the first re- cord of this species from Phalaris. Eribolus nana has been reared from Carex, Scirpus and Sparganium L. (Sparganiaceae) (Valley et al. 1969b, Uffen and Chandler 1978). Eribolus nearcticus Sabrosky (Chloropi- dae).—Adults emerged from three stems of P. arundinacea infested by Apamea ophio- gramma, along with EF. longulus and E. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON nana (in two stems), and with Oscinisoma alienum and Conioscinella zetterstedti (in a third stem) (Stoneycroft Pond, 31.v.2000, emerged 17.vi.2000 (1), 19.vi.2000 (6), Awa 000")/(3)5 | 27 NReZOOOUNGI) arama 30.vi.2000 (1)). This is the first host-plant record for E. nearcticus. Lasiosina canadensis Aldrich (Chloropi- dae).—Two adults were reared from a culm of) C.. Jdacustris\; (Hac “St )-Franeors 16.vii.1999, emerged 17.vili.1999). Adults were swept from stands of C. lacustris, and of C. aquatilis-Calamagrostis canadensis (Michx.) Beauv. from mid-May to late Oc- tober. Its relatively high abundance in Oc- tober may indicate multivoltinism. This species has been reported as a secondary invader of Carex interior, C. hystricina Muhl. and Scirpus cyperinus (L.) Kunth, feeding on the decaying tissue damaged by the primary invader (Valley et al. 1969b). Oscinisoma alienum (Becker) (Chlorop- idae).—One adult emerged from a stem of P. arundinacea infested by Apamea ophio- gramma, along with Conioscinella zetter- stedti and Eribolus nearcticus (Stoneycroft Pond, 31.v.2000, emerged O5.vii.2000). This is the first record from Phalaris; this species has been reared as a secondary in- vader from Scirpus microcarpus Presl. (Valley et al. 1969b). Rhopalopterum atriceps (Loew) (Chlo- ropidae).—Adults were reared on two oc- casions from C. lacustris (Lac St. Frangois, 09.vi1.1999, emerged 28.vii.1999 (1); same except 18.1x.1999, emerged 04.1.2000 (6), and 10.1.2000 (1)). In one case, seven adults emerged from the same stem but ten larvae were previously observed feeding deep in the culm within 5 cm of the base on rotting tissues previously attacked by a phytopha- gous invader. This is the first record of this species from C. lacustris. This species was reported feeding in shoots of Carex comosa Boott infested by the scathophagid Cordi- lura varipes (Walker) (Valley et al. 1969b). Most specimens were collected in June. Its higher abundance in Carex aquatilis—Cal- amagrostis canadensis stands suggest that VOLUME 104, NUMBER 2 one or both of these plants is also a host for R. atriceps. Rearing records of Rhopal- opterum spp. suggest that they are all sec- ondary invaders of herbaceous plants (Val- ley et al. 1969b, Armstrong et al. 1996). Thaumatomyia obtusa (Malloch) (Chlo- ropidae).—One adult was reared from a C. lacustris shoot (Lac St. Francois, 01.x.1999, emerged 04.1.2000). The single larva was found under the second leaf sheath 6 cm above the roots. Feeding dam- age possibly caused by a leaf-mining moth such as Elachista sp. was observed on the inner leaves and frass was present inside two of the leaves. At least three species of Thaumatomyia are predators of Aphidoidea (Harper 1963, Alleyne and Morrison 1977, Raspi 1996) and, despite the apparent leaf- miner damage near the 7. obtusa larva, this species probably also preys on aphids, such as Thripsaphis ballii, which were abundant between the inner leaves of Carex at the study sites. No adults of 7. obtusa were col- lected from C. /acustris stands. Pullimosina pullula (Zetterstedt) (Sphaeroceridae).—One adult emerged from decaying shoots of Scirpus fluviatilis (Lac St. Francois, 07.vii.1999, emerged 01.1x.1999). Although in this case the spec- imen was a secondary invader and emerged along with adults of Chaetopsis massyla (Otitidae) from a previously attacked plant, larvae of P. pullula probably feed on vari- ous types of decaying vegetation. Pullimo- sina pullula is found in many types of wet decaying plant matter (Marshall 1986). Orthacheta hirtipes Johnson (Scatho- phagidae).—One adult was reared from a stem of C. lacustris that was previously in- vaded, possibly by Cordilura sp. (Scatho- phagidae) or by a weevil (Lac St. Francois, 0O7.vili.1999, emerged 24.1.2000). Orthach- eta hirtipes has been reported as a faculta- tive predator of five Cordilura species, which are stem-borers of Carex species (Neff and Wallace 1969). This is the first record of this species from C. lacustris. 305 Coleoptera Sphenophora costipennis (Horn) (Curcu- lionidae).—Three adults were reared from C. lacustris (Lace St. Francois; 12.vii.1999- emerced BOlexe 599Gb); samen eXCept 23.vili.1999, emerged 08.x.1999 (1); same except 11.ix.1999 (1)). Several larvae were found in July and a few in August, within the base of the culm. Burrows examined were usually 6—7.5 cm long (rarely up to 13 cm). Numerous stems of C. lacustris were ob- served during these months with empty bur- rows and large exit holes close to the base, probably caused by S. costipennis; this spe- cies may contribute most to feeding injury of C. lacustris at Lac St. Frangois and may play an important role in the colonization of Dip- tera secondary invaders in such sedge mead- ows. Previous records of this species are from Carex comosa EF Boott and Scirpus validus Vahl. (Vaurie 1951). Sphenophora australis (Vaurie) (Curcu- lionidae). One adult was reared from a stem of Typha latifolia (Lac St. Francois, 21 .vin 1999> emerced yi-x.1999). ‘his stem-boring species is widespread in North America. Vaurie (1951) cited 7. latifolia as the preferred host, but also mentioned Spar- ganium eurycarpum Engelm. as a host. Lepidoptera Cosmopterix fernaldella Walsingham (Cosmopterigidae).—Five adults were reared from several mined leaves of C. la- eustris Wlkdc SSta Francois, 227-1999), emerged 09.11.2000). Leaf-mines were con- spicuous and abundant from August to Oc- tober 1999. Usually several leaves on the same host plant were mined, which sug- gests that a single female lays eggs on many leaves of the same shoot. The larvae were sometimes observed outside of the mine, probably feeding externally on the leaves. Hodges (1978) reported that C. fer- naldella feeds on Carex but did not specify a host species. Cosmopterix clemensella Stainton also mines the leaves of Carex (Hodges 1978) and an unidentified Cos- 306 mopterix sp. was reared from Carex vesi- caria L. (Frohne 1938). Elachista sp. (Elachistidae).—Adults were reared from leaf mines on C. lacustris (Lac St\Francois) 97x, 1999" semergsed 14.11.2000 (1); same except 27.x.1999, emerged 21.111.2000 (1)). Infested sedges were most commonly seen in October, when larval feeding damage was more ad- vanced. Only the basal parts of the inner- most leaves were attacked, in contrast to Cosmopterix fernaldella and Agromyzidae that attack the upper parts of the leaves. The mines of mature larvae were usually 15—20 cm in length, usually starting within | cm of the culm base. The first few centimeters of the mine were narrow, brown and zigzag shaped. The upper portion was filled with white frass with the caterpillar always at the upper end of the mine. Occasionally, two mines were found in the same Carex stem, but on different leaves; and other shoots were simultaneously attacked by Chlorops seminiger. In the lab pupation occurred out- side the mine in a cocoon attached to the mesh cover or bottom of the container. The immature stages and food plant are un- known for many species of Elachista (Braun 1948). A similar leaf mine was de- scribed for a species attacking Diarrhena americana Beauv. (Poaceae); that species was said to undergo a long resting period during which the early mine turns brown (Braun 1948). At least three species are known to mine Carex, but our biological observations indicate that the reared species is distinct from those species. Many Elach- ista species start feeding in fall; some start in spring, are dormant for the summer and resume feeding in fall; others have contin- uous development throughout the season (Braun 1948). The species we reared is probably of the dormant type, with two sep- arate feeding periods. Apamea ophiogramma (Esper) (Noctui- dae).—Two adults emerged from two stems of P. arundinacea (Stoneycroft Pond, 31.v.2000, emerged 28.vi.2000). Larvae or pupae were seen in six shoots and several PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON others had similar damage but were empty. Infested shoots were shorter than intact ones, with yellowish terminal leaves. In- fested shoots had an entry hole 2—3 mm in diameter and 2—11 cm above the base. On two occasions, two holes were observed on the same shoot with a mass of green frass just above each hole, indicating that two caterpillars attacked the same shoot. Nu- merous Diptera larvae were found feeding on frass and decaying plant tissues in the six A. ophiogramma burrows examined; several of these were reared to adults (see above). This is the first North American rearing record of A. ophiogramma (J. D. Lafontaine, pers. comm.). This introduced Palearctic species was first recorded in Brit- ish Columbia in 1992 (Troubridge et al. 1992) and is now also established in south- ern Quebec (Handfield 1999). Our speci- mens belong to the dark form of the spe- cies. The larvae are stem-borers of P. arun- dinacea, Glyceria maxima Hartm., Phrag- mites communis Trin. (Poaceae) and Tris pseudacornis L. (Iridaceae). DISCUSSION This study established 15 new host-plants for three phytophagous species and 12 sec- ondary invaders, including the first host re- cords for five chloropid species. This re- flects the current lack of knowledge on the feeding habits of Diptera, particularly Chlo- ropidae, associated with Poaceae and Cy- peraceae (Ferrar 1987). Other than the rear- ing survey of Valley et al. (1969b) and a few studies on the life-cycle and immature stages of selected species (e.g., Rogers et al. 1991, Wearsch and Foote 1994, Valley and Foote 1996), the biology of Nearctic Chloropidae is largely unknown. Based on rearing data and observations of infested plants, the four primary invaders recorded on C. lacustris (Table 1) all seem to be well established on this host. This is probably also true for Chromatomyia fus- cula and Apamea ophiogramma on P. arun- dinacea; both species were repeatedly reared or observed from a sample of the VOLUME 104, NUMBER 2 host plant taken from one small stand on a single date. There were at least three species of chlo- ropids in each of the four stems of P. arun- dinacea examined, apparently feeding on the decaying matter inside A. ophiogramma burrows. This diversity of chloropid sec- ondary invaders was unexpected and sug- gests that these flies may frequently be pre- sent in mixed infestations, competing for ephemeral food resources in a limited space. A similar situation probably occurs in a wide range of host plants. One of the reasons that so little biological information is available for larval Brachy- cera is that emergence rates are often low when rearing flies. Many Diptera breeding in sedges and grasses probably overwinter as third-instar larva or pupa within young shoots (Nye 1958, Wallace and Neff 1971, Rogers et al. 1991). Our success in rearing flies was higher when the insects were col- lected in the field as a mature larva or pupa near the beginning of the warm season or at the end of it. The search for infested plants should be concentrated during these periods to increase the probability of suc- cessful rearing. ACKNOWLEDGMENTS We thank the Canadian Wildlife Service for permission to collect at Lac St. Fran- cois. Thanks to Yang J.-T. (National Chung- Hsing University, Taiwan) for advice about rearing methods, R. Anderson (Canadian Museum of Nature), J. D. Lafontaine, J.-F Landry, and E. Maw (Agriculture and Agri- Food Canada) for insect identifications. 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Foote. 1996. Biology and im- mature stages of Eribolus longulus, with notes on E. nanus (Diptera: Chloropidae), secondary invad- ers of herbaceous wetland plants. Memoirs of the Entomological Society of Washington 18: 273— 279. Valley, K., J. A. Novak, and B. A. Foote. 1969a. Bi- ology and immature stages of Eumetopiella rufi- pes. Annals of the Entomological Society of America 62: 227-234. Valley, K., T. Wearsch, and B. A. Foote. 1969b. Larval feeding habits of certain Chloropidae. Proceedings of the Entomological Society of Washington 71: 29-34. Vaurie, P. 1951. Revision of the genus Calendra (for- merly Sphenophorus) in the United States and Mexico (Coleoptera, Curculionidae). Bulletin of the American Museum of Natural History 98: 31— 186. Wallace, J. B. and S. E. Neff. 1971. Biology and im- mature stages of the genus Cordilura (Diptera: Scathophagidae) in the Eastern United States. An- nals of the Entomological Society of America 64: 1310-1330. Wearsch, T. K. and B. A. Foote. 1994. Biology and immature stages of Chloropidae (Insecta: Diptera) associated with spike-rushes (Cyperaceae: Eleo- charis) 1. Stem borers. Annals of the Carnegie Museum 63: 193-213. Wheeler, T. A. 1994. Conioscinella zetterstedti An- dersson (Diptera: Chloropidae), a chloropid fly with polymorphic wing reduction, new to the Ne- arctic and central Palearctic regions. The Cana- dian Entomologist 126: 1377-1381. PROC. ENTOMOL. SOC. WASH. 104(2), 2002, pp. 309-317 CLADISTIC AND BIOGEOGRAPHIC ANALYSES OF APSIL MALLOCH AND REYNOLDSIA MALLOCH (DIPTERA: MUSCIDAE) OF SOUTHERN SOUTH AMERICA CLAUDIO JOSE BARROS DE CARVALHO AND MARCIA SOUTO COURI (CJBC) Departamento de Zoologia, Universidade Federal do Parana, Curitiba, Parana, 8 1531-990, Brazil (e-mail: cjbcarva@bio.ufpr.br); (MSC) Departamento de Entomologia, Museu Nacional, Quinta da Boa Vista, 20940-040, Rio de Janeiro, RJ, Brazil (e-mail: mcouri @attglobal.net) Abstract.—Apsil Malloch and Reynoldsia Malloch are genera endemic to Patagonia and southern Chile. A cladistic analysis for each genus is presented. The analysis of Apsil supports its monophyly on the basis of two synapomorphies, flat head and high gena. The phylogenetic relationships of Apsil species found are: ((A. pennata (A. dilata, A. spatu- lata)) (A. maculiventris, A. apicata) (A. maculipennis, A. biseta)) A. atripes)). The anal- ysis of Reynoldsia supports its monophyly on the basis of very long and slender palpi. The phylogenetic relationships of Reynoldsia species found are: ((R. pectinata (R. pter- opleuraris, R. rufoapicata)) (R. coxata (R. brevitarsis (R. aurifera, R. scutellata)))). The cladistic biogeographic analysis of both genera shows a distribution pattern congruent with recent biogeographic reconstructions of southern South America. The distribution pattern of Apsil species is more restricted than that of Reynoldsia. By the distribution pattern found in Reynoldsia, the occurrence of Reynoldsia species on Malvinas Islands could be expected. Key Words: Apsil, Reynoldsia, Muscidae, cladistic, biogeography, South America Among the 29 recognized genera of the Coenosiini (Muscidae, Coenosiinae) (Couri and Pont 2000), Apsil and Reynoldsia, both described by Malloch (1929, 1934), are the only ones endemic to Patagonia and south- ern Chile. Both genera can be easily rec- ognized with the key to the world genera of the Coenosiini (Couri and Pont 1999). According to Hennig (1959), the cos- mopolitan range of many genera is regarded as indicator of their antiquity. Skidmore (1985) called attention to the predominance of the coenosiines over other muscids both in polar regions and in the high montane equatorial regions. All known adults of Coenosiinae are predators and are possibly so also in the larval stage. No second instar coenosiine larva has been described (except for certain Lispe) and probably they are all monomor- phic (Skidmore 1985). Few references to these taxa have been made in the literature since the original de- scriptions. Recently, Couri (1995, 1998) re- described Reynoldsia and Couri (2000) re- vised the eight known Apsil species. Our objectives are to perform a cladistic analysis of the species of Apsil and Rey- noldsia and to analyze the geographic dis- tribution patterns of the species of these genera. MATERIAL AND METHODS The cladistic analyses of Apsil and Rey- noldsia, considered by Couri and Pont 310 Table 1. of Apsil used in the analysis. 0 = plesiomorphic char- acter states; | and 2 = apomorphic character states. Data matrix and characters of the species Outgroup 00000 00000 00000 00000 00 A. pennata 10011 00000 01000 01010 20 A. dilata ETA OOMOO OMOMOTO G2 HORA0 1000 00000 01010 11210 12 EOP O20) OLA OM O OMe: A. spatulata A. maculiventris A. apicata db AL (OO) 2, aRAL(OVAL(O) (O)ab aL Lak aly O}abal, (oye A. maculipennis 11010 10001 10101 10100 01 A. biseta iAMO)ALAL AL(OOVOVAL aL aa aLaL IL O/aLoVf0) (O)al A. atripes 10111 11010 00011 10000 11 1. Shape of head. [0] round, not flat: [1] flat [ci:100: ri: 100; weight:10; steps:1]. 2. Male: color of the pollinosity at face. [0] silver; [1] golden [ci:50; ri:50; weight:2; steps:2]. 3. Shape of eyes. [0] not elongate; [1] elongate [C.I.: 33; ri:0; weight:0; steps:3]. 4. Width of gena. [0] high; [1] very high [ci:33; ri: O: weight:0; steps:3]. 5. Number of frontal setae. [0] 5; [1] 4; [2] 3 [ci:40; ri:0; weight:0; steps:5]. 6. Hairs at arista. [0] almost absent; [1] short hairs present [ci:100; ri:100; weight:10; steps:1]. 7. Length of antennal flagellum related to pedicel. [O] twice longer or more; [1] less then twice longer [ci:50; 11:0; weight:0; steps:2] (unordered). 8. Enlargement at apex of palpi: [0] absent; [1] pres- ent. [ci:50; ri:0; weight:0; steps:2). 9. Acrostichals presutural hairs. [0] developed: [1] not developed; [2] one cilia [ci:66; 11:50; weight:3; steps:3]. 10. Brown cloud at stigma in wind. [0] absent; [1] present [ci:100; ri: 100; weight:10; steps:1]. 11. Small dark mark in first posterior cell almost below apex of second vein. [0] absent; [1] present [ci: 100; ri: 100; weight:10; steps:1]. 12. Length of hind tarsus related to hind tibia. [0] much over % as long as tibia; [1] not over % as long as tibia [ci:33; ri:0; weight:0; steps:3]. 13. Color of tibia. [0] brown; [1] yellow or yellowish [ci: 100; ri:100; weight:10; steps:1]. 14. Size of claws and pulvilli. [0] developed; [1] very reduced [ci:33; ri:O; weight:0; steps:3]. 15. Width of frons at level of anterior ocellus. [0] about %; [1] larger than % [ci:100; ri:100; weight:10; steps: 1]. 16. Antennal insertion. [0] not projected; [1] slightly projected [ci:33; ri:33; weight:1; steps:3]. 17. Insertion of antenna relative to the transverse mid-line of head (head viewed from in front): [0] near: [1] a little far above. [ci:33; ri:33; weight: 1; steps:3]. 18. Length of lower calypter related to upper one. [O] twice longer; [1] almost the same size; [2] 1.5 times longer [ci:100; ri:100; weight:10; steps:2] (unordered). 19. Posteroventral series of bristles at fore femur re- lated to posterodorsal series. [0] not stouter than those of posterodorsal series; [1] stouter [ci:50; ri:66; weight: 3; steps:2]. 20. Brown marks at abdominal tergites 1+2, 3 and PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (2000) as sister-groups, were made sepa- rately and were carried out using Hennig&86 version 1.5 (Farris 1988), and ‘‘Tree Gar- dener’’ version 2.2 (Ramos 1997), a pro- gram designed for running Hennig86 under Windows environment. Minimum-length trees were calculated using options “‘ie’’ as- sociated with “‘successive weighting.”’ The data matrix and the characters are shown in Tables 1 and 2. Eight species of Apsil were analyzed us- ing on 22 characters, and Reynoldsia with its seven species was analyzed using 25 characters. Characters were polarized by the outgroup method (Watrous and Wheeler 1981, Maddison et al. 1984). The outgroups were represented by three species: Schoen- omyza armipes Malloch 1934, Notoshoen- omyza costata Snyder 1957 and Spathi- Ppheromyia guttipennis Thomson 1869. These three genera compose, together with Apsil, Reynoldsia, and Schoenomyzina Mal- loch 1934, a larger monophyletic group of Coenosiini, according to Couri and Pont (2000) mainly distributed in southern South America. Characters were coded as binary and multistate, the latter were considered as additive or nonadditive (characters 18 and 22 in Apsil, and 7 and 20 in Reynoldsia), depending on the availability of informa- tion on contiguity of states in the outgroups. Cladograms were made using WINCLADA (Nixon 1999, version 0.9.9 beta). Cladistic biogeographic methods (see Morrone and Crisci 1995 for review) were used to construct taxon-area cladograms from the different taxon cladograms. A comparison of the general area cladogram was made based in some published papers. a 4 abdomen. [0] two marks; [1] a unique large mark [ci: 100; r1:100; weight:10; steps:1] (unordered). 21. Color of abdominal tergite 5. [0] all same color; [1] with 2 longitudinal brown marks; [2] with 2 round brown marks [c1:33; ri:20; weight:0; steps:6]. 22. Marks at scutellum. [0] all same colour; [1] with 2 lateral brown lines reaching the apex; [2] with 2 lateral brown lines reaching the apex [ci:66; 11:66; weight:4; steps:3] (unordered). VOLUME 104, NUMBER 2 Table 2. Data matrix and characters of the species of Reynoldsia used in the analysis. 0 = plesiomorphic character states; 1 and 2 = apomorphic character states; ? = missing data. 000 00000 00000 00000 00000 R. pectinata TOMOKO aL Aaa” (ONO O)aE IE Lak WO\(O) alae"? R. pteropleuralis 10111 10111 01100 01011 ??010 R. rufoapicata I O)aL TAL ak? aL aL (OG) ALLO) (OIE A ae O)al(O) abst R. coxata 11100 00001 10100 00002 10101 R. brevitarsis 11100 00001 11200 00000 100?? R. aurifera 11100 01000 10200 01110 ???10 R . scutellata 11100 00000 10201 10010 ???01 1. Shape of palpi. [0] not very long and slender; [1] very long and slender. [ci:100; ri:100; weight:10; steps: 1]. 2. Setulae on fronto-orbital plate. [0] numerous; [1] few and weak [ci:100; ri:100; weight:10; steps:1]. 3. Setulae on parafrontale. [0] numerous; [1] few and weak [ci:50; ri:0; weight:0; steps:2]. 4. Length of antennal flagellum related to pedicel. [O] twice longer or less; [1] more than twice longer [ci: 100; ri:100; weight:10; steps:1]. 5. Antennal insertion. [0] not projected; [1] slightly projected [ci:100; ri: 100; weight:10; steps:1]. 6. Width of gena. [0] not very high; [1] very high [ci:100; ri:100; weight:10; steps:1]. 7. Color of pruinosity at face. [0] grey; [1] golden [2] silver. [ci:66; ri1:0; weight:0; steps:3] (unordered). 8. Color of frontal triangle. [0] dark, contrasting with the ground colour; [1] not contrasting with the ground colour [ci:100; ri:100; weight:10; steps:1]. 9. Length of frontal triangle. [0] short; [1] long [ci: 100; ri:100; weight:10; steps:1]. 10. Marks at scutellum. [0] all same colour; [1] with 2 lateral brown lines reaching the apex. [ci:50; ri:50; weight:2; steps:2]. 11. Color of knob: [0] totally brown; [1] yellow and brown. [ci:100; ri: 100; weight:10; steps:1]. 12. Acrostichals presutural hairs. [0] developed; [1] not developed [ci:50; ri:0; weight:0; steps:2]. 13. Disc of katepisternum: [0] with many ground- setulae; [1] with few ground-setulae; [2] almost bare [ci:66; 11:66; weight:4; steps:3]. 14. Number of humeral bristles. [0] 2 or 3; [1] more than 3 [ci:50; 11:0; weight:0; steps:2]. 15. Number of setae on median third of anteroventral surface of hind tibia: [0] 2; [1] 3 [ci:50; 1:0; weight: O; steps:2]. 16. Number of setae on median third of anterodorsal surface of hind tibia: [0] 2 or 3; [1] 4 [ci:50; 11:0; weight:0; steps:2]. 17. Number of setae on median third of anterodorsal surface of mid tibia: [0] 2; [1] 3 [ci:50; r1:66; weight:3; steps:2]. 18. Number of supramedian setae on posterior sur- face of mid tibia: [0] 1; [1] 2 [ci:33; ri:0; weight:0; steps:3]. 19. Color of fore femur. [0] all same colour, dark brown; [1] dark brown, yellow at apex [ci:50; 11:66; weight:3; steps:2]. 311 The distribution data of Apsil and Reynold- sia were taken from Malloch (1929, 1934) and (Couri 1995, 1998, 2000) (Appendices ira?) RESULTS AND DISCUSSION Phylogenetic analysis——Only one tree was found for each analysis, with a length of 137 and 162, a consistency index (C.I.) of 88 and 91 and a retention index (R.I.) of 90 and 93, respectively for Apsil (Fig. 1) and Reynoldsia (Fig. 2). The analysis of Apsil supports its mono- phyly on the basis of the following shared synapomorphies: flat head, high gena and four frontal setae. The most basal dichoto- my divides the genus into two groups: (A. pennata + A. dilata + A. spatulata) and (A. maculiventris + A. apicata + A. maculi- pennis + A. biseta + A. atripes), the latter defined by two synapomorphies; presence of short hairs at arista and width of the frons larger than % of head. Apsil biseta and A. maculipennis are the only Apsil species that show a brown cloud at the stigma in the wing, a small dark mark in the first pos- terior cell almost below the apex of the sec- ond vein, and the calypters almost of same size. Malloch (1934) mentioned the remark- able morphological similarity of these two species. The analysis of Reynoldsia supports its monophyly on the basis of very long and slender palpi. The most basal dichotomy di- oe 20. Color of hind femur. [0] dark brown, yellow at apex; [1] dark brown on basal half and yellow at apical half; [2] totally dark brown [ci:100; ri:100; weight: 10; steps:2] (unordered). 21. Sternite 5 arms. [0] large [1] medium [ci:100; ri: 100; weight:10; steps:1]. 22. Width of cercal plate. [0] large; [1] thin [ci: 100; ri: 100; weight:10; steps:1]. 23. Aedeagus apodem. [0] shorter and larger; [1] lon- ger and thinner [ci:50; ri:0; weight:0; steps:2]. 24. Bristles on sternite 8 of ovipositor. [0] long and fine; [1] short and strong [ci:50; ri:50; weight:2; steps: Pil. 25. Shape of epiproct. [0] not divided; [1] divided [ci:33; ri:0; weight:0; steps:3]. ws) Nm PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON A pennata 12 17 19 21 Neat Imac as ®-e—O-—@ dilata O—O 4 42 “\ BY A 22 O-O-O-O spatulata O70 ee2 3) 5.1809.) 46.21 , 1 4 5 O-O-0-@-0-0 maculiventris —@-@_@—_ 17 2 A. OM dh 32. 02 14 4 > ae 45722 ; @ O—O—O 2 2 2 "be a ba ba apicata ©O—-©O 4 14 5 12 14 O-O-O—— maculipennis 6 9 4 15 16 2 py i han ONTO Cane OF vena tote ct. dees A biseta Ca aes : @-2-@ atripes er 1 Fig. 1. Cladogram depicting the phylogenetic relationships among the species of Apsi/. Length: 137; C.L.: 88; ri: 90. Black boxes, synapomorphies; gray boxes, homoplasies; clear boxes, reversals. 1 8 28 147 17 18 22) 24 brevitarsis ei 18) A425 aurifera 2 Fig. 2. Cladogram depicting the phylogenetic relationships among the species of Reynoldsia. Length: 162; C.I.: 91; ri: 93. Black boxes, synapomorphies; gray boxes, homoplasies; clear boxes, reversals. VOLUME 104, NUMBER 2 313 Central Chile Central Argentina Chaco a) Morrone (1993) a sal SE PO pow i ay Maule Valdivian ain I Malvinas M See coe M forest Sa b) Morrone et al. (1994) N Chile ARG SBrazil CChile Malvinas I ir were NGSoas c) Amorim & Pires (1996) [based on Morrone (1993) and Morrone (1994)]. Central Chile Coquinbo Santiago Curicd Nuble << (d) Morrone et al. 1997 3 Fig. 3. Biological area cladogram for southern South America. C = Central; I = Isles; M = Magellanic; N = North; PAT = Patagonian province; S = South; SUB = Subantarctic province; T = Tierra del. 314 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON pennata dilata spatulata maculiventris Subantarctic apicata maculipennis \ Curico + Subantarctic Fig. 4. Taxon-area cladogram of species of Apsil. vides the genus into two groups: (R. pectin- ata + R. pteropleuralis + R. rufoapicata) and (R. coxata + R. brevitarsis + R. auri- fera + R. scutellata). The first group is de- fined by five synapomorphies (the slightly projected antennal insertion, the very high width of the gena, the color of the frontal triangle not contrasting with the ground col- or, the long frontal triangle, and a thin cer- cal plate) and the second one by three (pres- ence of few and weak setulae on the fronto- orbital plate, the brown and yellow knob, and sternite 5 arms medium in width). Biogeographic analysis.—The biogeo- graphical patterns of distribution of Mus- cidae are scarcely known in the world. For the Neotropical Region, only Hennig (1965) and more recently Carvalho (1999) analyzed patterns of distribution of Musci- dae species in South America with a dis- persalist and a cladistic view, respectively; however, in the southern part of this conti- nent, below 30°S latitude but including the Andean highland northern of this latitude (Kuschel 1969, Cabrera and Willink 1973, Crisci et al. 1991, Morrone 1993, Morrone et al. 1994, Morrone and Lopretto 1994, Amorim and Pires 1996), there are no his- torical biogeographic studies with Musci- dae. The biogeographical patterns of some en- demic taxa in southern South America have been recently studied with cladistic biogeo- graphic methods (Crisci et al. 1991; Mor- rone 1993, 1994; Morrone et al. 1994, 1997). Selected biological area cladograms proposed for southern South America are shown in Fig. 3. The taxon-area cladograms of Apsil (Fig. 4) and Reynoldsia (Fig. 5) were compared with those patterns found (Fig. 3) and shown to be congruent with those biogeo- graphic reconstructions. The distribution pattern of Apsil species is more restricted than Reynoldsia species. Apsil atripes is the only species (Fig. 4) ex- clusively occurring to the north of the sub- antarctic area corresponding broadly to the Curic6 area of central Chile of Morrone et al. (1997). They tried to explained the pat- terns of central Chile and their study sup- ports a vicariant explanation of the patterns analyzed. All Reynoldsia species are spread in the VOLUME 104, NUMBER 2 IPS, Se subantarctic area, with only Reynoldsia pectinata occurring in Tierra del Fuego (Appendix 2). By the distribution pattern found in Reynoldsia, compared with the bi- ological reconstruction of southern Chile by Morrone et al. 1994 (Fig. 3b), it could be expected that undescribed species of Rey- nodsia occur in Malvinas Islands, which are part of the subantarctic area, here under- stood as a major monophyletic area (see Amorim and Pires 1996). ACKNOWLEDGMENTS We thank Juan J. Morrone (Museo de Zoologia, Universidad Nacional Aut6noma de México, México), Dalton de Souza Amorim (Universidade de Sao Paulo, R1- beirao Preto, Brazil), Adrian Charles Pont (University Museum of Natural History, Oxford, U.K.), and a anonymous referee for their critical rewiews of the manuscript. We are also grateful to Sionei R. Bonatto for technical assistance in drawing the clado- grams. We are grateful to the ““Conselho Nacional de Desenvolvimento Cientifico e Tecnolégico,” an agency of the Brazilian government, for scientific and technological Oo — Nn pectinata pteropleuralis rufoapicata coxata Subantarctic brevitarsis aurifera scutellata Taxon-area cladogram of species of Reynoldsia. development, and the support provided by a grant to CJBC (Proc. Nr. 300043/86-4) and to MSC (Proc. Nr. 300386/80-0). MSC is grateful to Fundagao Universitaria José Bonifacio for financial support (Proc. 8700- OQ). LITERATURE CITED Amorim, D. S. and M. R. S. Pires. 1996. Neotropical biogeography and a method for maximum biodi- versity estimation, pp. 183-219. /n Bicudo, C. E. M. and N. A. Menezes, eds. Biodiversity in Brazil, a first approach. Sao Paulo, CNPq, 326 pp. Cabrera, A. L. and A. Willink. 1973. Biogeografia de América Latina. Washington, D.C., Organizacion de los Estados Americanos, 117 pp. Carvalho, C. J. B. de. 1999. Revision, Cladistics and Biogeography of the Neotropical genus Souzalo- pesmyia Albuquerque (Diptera, Muscidae). Pro- ceedings of the Entomological Society of Wash- ington 101: 123-137. Crisci, J. V.. M. M. Cigliano, J. J. Morrone, and S. Roig Junent. 1991. Historical biogeography of southern South America. Systematic Zoology 40: 152-171. Couri, M. S. 1995. Contribution to the Knowledge of Reynoldsia Malloch (Diptera: Muscidae). Pro- ceedings of the Biological Society of Washington 108: 281-29. . 1998. Redescriptions of two species of Rey- 316 noldsia Malloch (Diptera, Muscidae). Revista Brasileira de Entomologia 42: 101—103. . 2000. Revision of Apsil Malloch (Diptera, Muscidae, Coenosiinae, Coenosiini). Studia Dip- terologica 7: 45—57. Couri, M. S. and A. C. Pont. 1999. A key to the world genera of the Coenosiini (Diptera, Muscidae, Coe- nosiinae). Studia Dipterologica 6: 93-102. . 2000. Cladistic analysis of Coenosiini (Dip- tera: Muscidae: Coenosiinae). Systematic Ento- mology 25: 373-392. Farris, J. S. 1988. Hennig 86, version 1.5. Published by the author, Port Jefferson, New York, 18 pp. Hennig, W. 1959. Muscidae (Part, Lieferung 204). Jn Lindner, E., ed. Fliegen der Palaearctic Region, 63b: 233-288. . 1965. Vorarbeiten zu einem phylogenetischen System der Muscidae (Diptera: Cyclorrhapha). Stuttgarter Beitrage zur Naturkunde 141, 100 pp. Kuschel, G. 1969. Biogeography and ecology of South America Coleoptera, pp. 709-722. Jn Fittkau, E. J., J. Ilies, J. Klinge, G. H. Schwabe, and H. Sioli. Biogeography and Ecology in South America, Volume II. Maddison, W. P., M. J. Donoghue, and D. R. Maddi- son. 1984. Outgroup analysis and parsimony. Sys- tematic Zoology 33: 83-103. Malloch, J. R. 1929. Exotic Muscaridae (Diptera).— XXVI. Annals and Magazine of Natural History 10(4): 97-120. . 1934. Muscidae. /n Diptera of Patagonia and South Chile 7: 171-346. London, British Muse- um. Morrone, J. J. 1993. Cladistic and biogeographic anal- yses of the weevil genus Listroderes Schoenber (Coleoptera: Curculionidae). Cladistics 9: 397— 411. 1994. Distributional patterns of species of Rhytirrhinini (Coleoptera, Curculionidae) and the historical relationships of the Andean provinces. Global Ecology and Biogeography Letters 4: 188— 194. Morrone, J. J. and J. V. Crisci. 1995. Historical Bio- geography: Introduction to methods. Annual Re- view Ecological Systematic 26: 373-401. Morrone, J. J., S. R. Junent, and J. V. Crisci. 1994. South American Beetles. National Geographical Research and Exploration 10: 104—115. Morrone, J. J. and E. C. Lopretto. 1994. Distributional pattern of freshwater Decapoda (Crustacea: Ma- lacostraca) in southern South America: A panbi- ogeographic approach. Journal of Biogeography 21: 91-109. Morrone, J. J., L. Katinas, and J. V. Crisci. 1997. A cladistic biogeographic analysis of Central Chile. Journal of Comparative Biology 2: 25—42. Nixon, K. C. 1999. WINCLADE, Beta version 0.9.9. Published by the author: L.H. Bailey Hortorium, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Cornell University, Ithaca, New York. Available as shareware via (www.Cladistics.com/). Ramos, T. C. 1997. Tree Gardner 2.2. Privately dis- tributed by Thiago Courrol Ramos, Museu de Zoologia, Universidade de Sao Paulo, Brazil. Skidmore, P. 1985. The biology of the Muscidae of the world. Series Entomologica 29, 550 pp. Watrous, L. E. and Q. D. Wheeler. 1981. The outgroup comparison method of character analysis. System- atic Zoology 30: 1-11. APPENDIX 1 Localities of species of Apsil in South America including latitude and longitude. Abbreviations: E = East, m = meters, k = kilometers, v. valley. Apsil apicata Malloch 1934.—ARGEN- TINA: Lake Correntoso (40°44'S, 71°40’W); San Carlos de Bariloche (41°11'S, 71°23’W). CHILE: Casa Pangue (41°03’S, 71°52’W). Apsil atripes Malloch 1934.—CHILE: Rio Colorado (33°04'S, 71°39'W); Curic6 (G4°59'S, 71°14'W);:, Talca, 800 m' Gs 20's; 71°46'W); Concepcion (San Rosendo) Gi dGS. 72 49. Wo): Apsil biseta Malloch 1934.—ARGENTI- NA: Volcan (36°26'S, 67 °09'W): San Carlos de Bariloche (41°11’S, 71°23'W). CHILE: Las Cabras, 1480 m (34°18'S, 71°19’W); An- gol (37°47'S, 72 °45'W); Malalcahuello (4— 14k E., 1080-1570 m.) (38°27'S, 71°35’W); Curacautin .(G8°28'S, 71°52’ W): Villanicas 1250 m (39°15'S, 72°30'W); Coihaique (v. of Simpson river) (45°35'S, 72°08'W). Apsil dilata Malloch 1934.—ARGENTI- NA: San Carlos de Bariloche (41°11'S, 71°23’ W))... CHULE: , Ancud, , Givaass: 73°50'W); Puntra (42°07'S, 73°49W); Castro (42°30'S, 73°46’W). Apsil maculipennis Malloch 1934.—AR- GENTINA: Volcan (36°26'S, 67°09'W); Lake Correntoso (40°44'S, 71°40'W); Puerto Blest (41°02'S, 71°59'W). CHILE: Malalca- huello (12 k E, 1080 m) (38°27'S, 71°35'’W); Curacautin (38°28'S, 71°52’W); Villarica (39°15'S, 72°30'W); Osorno (40°35’S, 73°14'W); Parque Nacional Puyehue (40°40'S, 72°37'W); Coihaique (v. of Simp- son river) (45°35'S, 72°08'W). VOLUME 104, NUMBER 2 Apsil maculiventris Malloch 1929,— CHILE: Perales (36°40’S, 72°39’W). Apsil pennata Malloch 1934.—ARGEN- TINA: San Carlos de Bariloche (41°11'S, 71°23'W). CHILE: Casa Pangue (41°03’S, JA 32 W): Apsil spatulata Malloch 1934.—ARGEN- TINA: San Carlos de Bariloche (41°11'S, fal23eW ))°> CHIE E:*|, Peulilas(41°06'S, 72°02'W); Puntra (42°07'S, 73°49’W). APPENDIX 2 Localities of species of Reynoldsia in South America including latitude and longi- tude. Abbreviation: m = meters. Reynolsia aurifera Bigot 1885.—ARGEN- TINA: Lolog (40°05’S, 71°19’W). CHILE: Casa Pangue (41°03’S, 71°52’W); Ancud (41°53’S, 73°50'W). Reynolsia brevitarsis Malloch 1934.—AR- GENTINA: San Carlos de Bariloche (4°11'S, Mia Whe Wake ‘(Gutterrez. (41° 11'S, 72°23'W). S117) Reynolsia coxata Malloch 1934.—AR- GENTINA: Lolog (40°05’S, 71°19’W); San Carlos de Bariloche (41°11'S, 72°23’W). Reynolsia pectinata Malloch 1934.—AR- GENTINA: Rio Grande (53°47'S, 67°42'W); Estancia Viamonte (54°02’S, 67°22’W). CHILE: Punta Arenas (53°09’S, 70°55’W). Reynolsia pteropleuralis Malloch 1934.— ARGENTINA: San Martin de los Andes (40°10'S, 71°21'W); Nahuel Huapi (41°03’S, 71°12’W); CHILE: Perales (36°40’S, 7239 W). Reynolsia rufoapicata Malloch 1934.— ARGENTINA: San Martin de los Andes, 1500 m (40°10’S, 71°21'W); Lake Corrento- so (40°44’'S, 71°40’W): San Carlos de Baril- oche (41°11’S, 71°23’W): Puerto Blest (41°02’S, 71°50’W). Reynolsia scutellata Malloch 1934.— CHILE: Angol (37°47'S, 72°45'W); Cerro Nahuelbuta, 650 m (37°48'S, 73°04'’W); Gal- varino (38°24’S, 72°47'W). PROC. ENTOMOL. SOC. WASH. 104(2), 2002, pp. 318-329 DESCRIPTION OF A NEW TORTRICID PEST (LEPIDOPTERA: TORTRICIDAE: OLETHREUTINAE) OF LITCHI (LITCHI CHINENSIS) IN FLORIDA, WITH A REVIEW OF TORTRICID PESTS OF LITCHI WORLDWIDE JOHN W. BROWN, JORGE PENA, THERESA VAZQUEZ, AND JOAQUIN BAIXERAS (JWB) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, c/o National Museum of Natural History, Smithsonian Insti- tution, Washington, DC 20560-0168, U.S.A. (e-mail: jbrown @sel.barc.usda.gov); (JP, TV) University of Florida, Institute of Food and Agricultural Services, Tropical Research and Education Center, 18905 SW 280 Street, Homestead, FL 33031, U.S.A.; (JB) Instituto Cavanilles Biodiversidad y Biologia Evolutiva, Universidad de Valencia, Apartado oficial de correos 22085, 46071 Valencia, Spain Abstract.—-The adult, larva, and pupa of Crocidosema litchivora Baixeras, new spe- cies, are described and illustrated; details of the life history are presented. Since 1993, an increase in the abundance of C. litchivora in southern Florida has resulted in significant damage to commercially grown litchi (Litchi chinensis Sonnerat; Sapindaceae). The larvae feed on the inflorescences and buds of the host plant, significantly reducing crop yield. Owing to the absence of records of C. litchivora prior to 1992, it is likely that this Neotropical species is a relatively recent arrival in southern Florida. A list of tortricid pests of litchi worldwide is presented. Key Words: tion, Caribbean Litchi chinensis Sonnerat (Sapindaceae) is a large evergreen tree native to China. The fruit, referred to as lychee, leechee, or litchi, has long been a favorite in south- eastern Asia; over the last decade it has gained popularity in the United States. In the contiguous states, L. chinensis is grown commercially only in tropical southern Florida, where growers harvested 1.37 mil- lion pounds (0.63 million kg) in 1996, val- ued at $2.75 million (Stanley 1998). The small but growing litchi industry in southern Florida was nearly destroyed in 1992 by hurricane Andrew. Subsequently, the acreage planted in litchi increased sig- nificantly; some of the new plant material was imported from Hawaii. Starting in Crocidosema, Litchi chinensis, pest species, new record, species introduc- about 1993, larvae of a tortricid moth be- came a pest in Florida orchards, and in 1996-1997, this species inflicted damage to litchi resulting in as high as 40% crop loss in some places. In 1997, examples of the moth were sent to the Systematic Entomol- ogy Laboratory, USDA, for identification. Although morphologically similar to the lantana flower-cluster moth (Crocidosema lantana Busck), which is reported to feed on litchi in the Hawaiian Islands (Zimmer- man 1978), detailed studies revealed that the Florida specimens represented an un- described species. The purposes of this pa- per are to present descriptions and illustra- tions of the new species, provide details on its biology, and briefly review tortricid pests of litchi worldwide. VOLUME 104, NUMBER 2 MATERIALS AND METHODS Systematics.—Specimens of the new Crocidosema species were collected as lar- vae from infested litchi trees (described be- low) and as adults by black-light collecting in southern Florida. Depositories are abbre- viated as follows: BMNH, The Natural His- tory Museum, London, England; IFAS, University of Florida, Institute of Food and Agricultural Services, Gainesville, Florida, USA; INBio, Instituto Nacional de Biodiv- ersidad, Santo Domingo de Heredia, Costa Rica; MEM, Mississippi Entomological Museum, Mississippi State, Mississippi, USA; USNM, National Museum of Natural History, Smithsonian Institution, Washing- ton, iC. USA: VBC,. Vitor Becker per- sonal collection, Planaltina, Brazil. Dissec- tion methodology follows that summarized in Brown and Powell (1991). Forewing measurements were made with an ocular micrometer mounted in a dissecting micro- scope. Terminology for wing venation and genitalic structures follows Horak (1984); terminology for larval features follows Brown (1987). Abbreviations and symbols are as follows: FW = forewing; HW = hindwing; DC = discal cell; n = number examined; ca. = circa (approximately); * = mean; bl. = blacklight. Biology.—Newly formed litchi leaf flushes with no evidence of Lepidoptera lar- vae or eggs were marked and checked daily at a single site in Dade County, Florida, from 30 March to 15 April 1998. Upon dis- covery of newly oviposited eggs, apical and lateral buds were collected and brought into the laboratory for further inspection. The infested buds were placed individually in 3- ml vials held at 22 + 3°C and 75% relative humidity and observed daily until the egg chorion began to darken, indicating the de- velopment of the embryo to an advanced stage. The infested buds then were trans- ferred to 30-ml plastic cups containing syn- thetic diet (Robinson et al. 1977). Eggs were observed daily for eclosion. A sub- sample of developing larvae were checked 319 daily for pupation, and pupae were checked daily for adult emergence and sex deter- mination. Representative examples of lar- vae were collected daily and preserved in 75% ethanol. Width of larval head capsules of all instars was measured for 106 individ- uals. Three newly emerged females and two males were held at 22 + 3°C, 80% relative humidity, and 12:12 L:D in a 30 X 30 X 30 cm plexiglass cage with water and honey and litchi leaf flushes as a potential ovipo- sition substrate. Leaf flushes were exam- ined daily for evidence of eggs. This ex- periment was repeated 5 times. RESULTS Systematics Crocidosema litchivora Baixeras, new species (Figs. 1—6) Diagnosis.—Crocidosema litchivora is similar to C. lantana Busck 1910, C. lon- gipalpana (Moschler 1890), C. calvifrons (Walsingham 1892), and several undescribed species in the shared possession by males of strongly upturned labial palpi, with an ex- tremely long second segment, and modified scaling on the convex frons. A distinctive male hairpencil extending from the poste- rior notal hindwing process to the abdomen is shared by C. litchivora and C. lantana. Although forewing pattern is similar in the latter two species, C. lantana always has reduced green suffusion and hence a more defined pattern, allowing superficial sepa- ration of the two in well preserved speci- mens. The layered sex scales beneath the costal fold of the forewing are dark brown in C. litchivora and silvery in C. lantana. The scales of the hairpencil from the pos- terior notal hindwing process are slightly different between the two: in C. lantana they are clubbed and thickened distally, whereas in C. litchivora they are piliform. The lateral pockets in the abdomen that re- ceive the scales also are different between the two: in C. lantana the sclerotized areas are complex, surrounded by an extensive 320 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-2. membranous region, and the pocket is cov- ered by broad, non-deciduous scales; in C. litchivora the structure is comparatively re- duced, with no membranous region, and lacks broad scales. Male and female geni- talia are distinct as well. Crocidosema lan- tana lacks the pointed process on the distal ventral part of the sacculus, and the entire cucullus is curved upward. In the female genitalia the sterigma is pentagonal in C. lantana, with a small but evident dorsal mi- crosclerite in the junction between the ste- rigma and the ductus bursae. This sclerite, which strongly stains by fuchsin acid and is 70—80p in length, also is present in C. litchivora, but much more reduced (25—50 w) and ill defined. The pair of irregularly rounded, densely scaled pockets on the sev- enth abdominal sternite are opened poste- riorly in C. /antana and relatively closed in C. litchivora; in the latter there is a lateral tooth in each pocket that is lacking in C. lantana. Description.—Male: Head: Antennal cil- ia approximately 0.5 times width of flagel- lomere. Labial palpus conspicuously elon- gate, strongly upturned, extending beyond top of head in resting position, tan, with brown and orange scales. First segment Adults of Crocidosema litchivora. 1, Male paratype. 2, Female paratype. with basal third having dorsal margin curved and ventral margin forming approx- imate right angle, apical two-thirds extend- ing forward in parallel axis of body, apex articulating with second segment at 90° an- gle; second segment long, ca. 2 times that of first, and 3 times that of third, basal third slightly curved, remainder straight and di- rected dorsally; third segment straight, short. Length of organ of vom Rath equal to approximately 0.25 length of segment; opening of organ in apical position. Pro- boscis developed, unscaled. Frons concave, clothed by small, charcoal black to oliva- ceous, appressed scales; scales aligned from eyes to a narrow vertical ridge at middle of frontoclypeus; ridge protruding ventrally in a group of erect longer scales and dorsally between antennae in a broad row of scales. Vertex anteriorly with overhanging tuft of blackish brown and gray scales between compound eyes; ocelli and chaetosemata well developed; tuft of erect dark brown to whitish long scales from lateral posterior part of vertex to occiput surrounding eyes. Thorax: Smooth-scaled, dark brown to mostly black dorsally; tegula covered by dull dark brown, sometimes yellowish, sim- ple scales; mesonotum and metanotum cov- VOLUME 104, NUMBER 2 ered by elongate glossy dark brown scales; scales erect on metascutum; pleural and sternal portions of thorax light brown to yellowish. Legs unmodified. Forewing (Fig. 1): Length 4.5-5.2 mm (* = 4.9; n = 5). Costal fold large, well defined, extending at least 0.45 length of FW and 0.66 distance between base of FW and point where R, meets costa; basal portion of R-stem obso- lete; R, and R; stalked for more than half of their length with R, extending to costa and R,; to termen; M-stem and chorda ill- defined if present; discal cell at least 0.7 times length of wing; M, and M, straight; M,, M,;, and CuA, approximate at base with M, and CuA, parallel and curved; CuP at- rophied except in distal portion; anal loop equal to 0.33 length of 1!A+2A. Upperside mostly mottled black, gray, and reddish brown, with large areas of green; space be- tween Sc and basal portion of Cu-stem cov- ered by a layer of smooth, small dark brown scales within costal fold; costal fold enclos- ing a strongly developed row of white pet- iolate sex scales along Sc; ground color brown to reddish brown; costa with two ill defined strigulae between base of wing and Sc on distal half of costal fold; two strigu- lae between Sc and R,, more basal less dis- tinguishable; three distinct strigulae be- tween veins R,—R;, R,—R;, and R;-R, re- spectively; three silvery striae extending from strigulae, confluent to termen between R, and M,; termen without strigulae; fasciae poorly defined; basal and subfasal fasciae fused, dark brown from costa to Al+A2, yellowish white from A2 to dorsum; me- dian fascia dark brown, extending from cos- ta to CuA,, broader and darker on discal cell; postmedian fascia disjunct, with a con- spicuous dark spot between R; and M, im- mediately anterior to ocellus, extending to termen in a narrow line from M, to inter- vening space between CuA, and CuA,; sub- terminal fascia represented by a spot on costa; apical spot between R, and R; as a small, but distinctive group of dark brown to black scales; ocellus present, green to yellowish green with strong silvery suffu- ee) i) sion and four small, sometimes indistinct, fascial dark brown spots; green to yellow- ish-green area from outer margin of sub- basal fascia to tornus on dorsum. Fringe mostly green; brown in apical region. Un- derside brown, with four white strigulae from Sc to R,; area of wing overlap whitish. Hindwing: Costa with basal 0.66 expanded to form broadly rounded lobe underlying forewing, narrowed before Sc+R,; Rs and M, stalked; Rs extending to costa near apex, M, to termen; M, and M,+CuA, aris- ing from same point; M, and CuA, stalked; upperside uniformly grayish brown, except area of forewing overlap anterior to Sc+R, densely covered by silvery scales; 3 rows of modified sex scales on wing: (1) cubital pecten slightly developed consisting of a sparse row of slender elongate, brownish scales along CuP; (2) a denser row of stiff, somewhat erect tan scales along 2A in anal loop that presumably connects with erect scales on metascutum; (3) a rather large patch of long, silky scales between inner margin and 3A, denser at wing base and progressively less dense in distal half to wing margin. Fringe concolorous with re- mainder of hindwing. Underside of hind- wing uniformly light brown. A dense hair- pencil of long pale orange scales arising from posterior notal wing process, extend- ing into a pocket of specialized scales on dorsolateral portion of second abdominal segment, emerging again from abdominal pocket on third abdominal segment. Abdo- men: First, second, and third pleural mem- branes strongly modified into a series of three extended sclerotized plates; plates slightly concave and corrugated, forming depressions on first, second, and third seg- ments, concealed by large, flat, somewhat erect, specialized scales; first three abdom- inal tergites covered with small dark brown sex scales; remainder light brown to yel- lowish. Genitalia (Fig. 3): Uncus short, wider at base, subtriangular, ca. 0.5 length of socius, finely setate dorsally, rounded apically. Socius well developed, crescent- shaped. Anal tube with two longitudinal 322 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 3. ventrolateral sclerotized bands that basally connect with the two lateral arms of anel- lus. Valva upturned; costa with conspicuous notch approximately 0.5 distance from base to apex. Sacculus short with distinct, swol- len, densely setose pointed ventral process approximately 0.20 distance from base to apex. Cucullus densely setose from neck of valva to apex. Ventral edge slightly thick- ened before apex. Aedeagus simple, nearly straight, with dense fascicle of 10—15 slen- der cornuti; attached to tegumen at base of gnathos and anal tube by a pair of slender lateral arms. Female: FW length 4.7-5.6 mm (x = 5.0; n = 10) (Fig. 2). Similar to male, with following exceptions: antennal cilia less than 0.1 times width of flagellomere; labial palpus weakly upturned, unmodified; frons brown, convex. Mesonotum concolorous with basal and subbasal fasciae in forewing; metanotum concolorous with hindwing up- perside; no erect scales on metascutum. Forewing without costal fold; four evident strigulae and striae basal to Sc; basal and subbasal fasciae poorly developed; median and postmedian fasciae slightly more de- veloped than in male; no specialized sex scaling or hairpencil on wings or posterior \ WE SY Male genitalia of Crocidosema litchivora, valvae spread, aedeagus in situ. notal hindwing process. First three abdom- inal segments unmodified, lacking sex scales. Genitalia (Fig. 4): Papillae anales simple, slender. Sterigma forming a slightly sclerotized subtriangular plate projected lat- erally and posteriorly; bearing 10—15 long setae arranged symmetrically and oblique- ly; sterigma forming an anterior funnel- shaped region leading to ostium. Sternite of sixth segment with a pair of irregularly rounded, densely scaled pockets with one lateral conspicuous tooth in each one. Junc- tion of ostium and ductus bursae with small, irregularly shaped dorsal sclerite, most easily seen if ductus removed from sterigma. Ductus bursae moderately long, broadened near junction with corpus; mem- branous in anterior 0.33, with strongly sclerotized plate in following 0.33, and membranous in posterior 0.33. Ductus sem- inalis arising from dorsal position in middle of sclerotized plate. Corpus bursae pyri- form, with a pair of long, curved, bladelike signa; a curved patch of short spines near junction of corpus and ductus that sur- rounds anteriorly a slightly developed lobe. Larva (Fig. 5). Based on 3 last instars. General: Length 11 mm. Head pale orange yellow, with a dark brown genal bar and VOLUME 104, NUMBER 2 Fig. 4. ra. Female genitalia of Crocidosema litchivo- dark patch in stemmatal region; prothoracic and anal shields yellow to tan; body white to dingy grayish white, with well-defined, pale grayish brown pinacula contrasting with body color (absence of spiculae on pinacula enhances contrast); body covered o>) i) ie) with short, fine spiculae; spiracles on T1 and A8 nearly equal in size, approximately 1.3 times diameter of those on Al—7. Head: Frons extending ca. 0.7 to occipital fora- men; stemmata arranged in a semicircle ex- cept for S5; S2 slightly smaller in diameter. Thorax: Prothorax with trisetose L-group on well defined pinaculum, LI slightly closer to L2 than to L3; meso- and meta- thorax with Dls and D2s on same pinacu- lum, arranged nearly vertically. Abdomen: SD2 either without distinct pinaculum or on narrow extension of large SD1 pinaculum; spiracles round, larger than base (or socket) of SD1 setae on Al—7; SD1 pinaclum on A8 large, curved along posterior edge to accommodate spiracle; SV group on 1, 2, 7, 8, 9 always 373:2:2:27 Digand SD! on same pinaculum on A9; L3 on same pina- culum as LI and L2 on AQ; distance be- tween Vs on AQ slightly greater than dis- tance between Vs on A8 and A7; anal fork well developed, with 5—6 elongate teeth. Crochets 35—38 in a biorinal circle; anal crochets 21—24 in a continuous band. Pupa (Fig. 6). Based on a single female. Typically olethreutine; head without projec- tion; no conspicuous sculpturing; dorsum of thorax with posterior-projecting, mesal lobe; abdomen with 2 rows of spines dor- sally on A2-7, spines of posterior row smaller, more numerous; anterior row on A4-7 extends from spiracle to spiracle; sin- gle row of strong spines on A8—9. Cremas- ter poorly developed, extremely short, blunt, ending in row of strong spines. Type material.—Holotype 6: USA: FLORIDA: Dade Co. 6 km NW _ Home- stead, 20 March 1998, bl. (J. Brown & J. Pena, USNM). Paratypes: 1d, same data as holotype (genitalia slide number 89828); 13, Florida, Homestead, November 1997, ex-litchi (genitalia slide number 88652); 72, same data as holotype; 2°, same data as holotype, but 20/23 March 1998. All par- atypes deposited in USNM. Additional material examined.—USA: Florida, Dade Co., Homestead, 36, 2°, 15- I-1992, as larvae on Litchi chinensis, 324 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fee OY, ean me, a ae & L3 | ay ae gee galsiael |. ay, ie) pad , L2gp ee +) ie a) L3 gees THe © SV iS ain ea ) KW svi» : a it ee KA Y “rr S) So a) eS ie} | ° L2 ey Ny cf | | C QL & io D2 v4 \SV1 fae or Sa D1 \p Se ea Abe ‘ © sD\_ % D2 ma rat al Po a PI D1 d li | aN “ imal iW Spe Be ee 2S Se a em ea Pa RS MwA SV aga SY2GN* : AP\( =< ae ae ~$ ) ae aN aR oNGt vi WS Fig. 5. Larva of Crocidosema litchivora. a, Head, T1—3. b, mandible. c, Al—2. d, A6—7. e, A8-10. VOLUME 104, NUMBER 2 6 A a ie : : “= <« rt 2 Ere ~ a —r~e Ec SS |S2 s fe = ELE - | © ES . ae < _ re onare B < = <= see SS ee yea | Fy < —e aime sre -° Z SS, pal Ze 5 St eee sn wee” (rears “ = SS \t + «] Est ¢ < < . = i705 Pee col 5 <_< os = Ss ee oo eae <8 < ea, E Seat Bd —d ee Ie == = «€ od =< f a en = oa = =< | ns = “See oe F 7 te ae Z aS = |: v, | b Ls Ga 4a 4 ae aA Fig. 6. emerged 21-H/9-III-92 (J. Pena, IFAS), 1d, 25-II-1992, 1¢, 25-II-1992, both as larvae on Eriobotrya japonica (R. Duncan, IFAS), 12, 21-I-1992, on Clausena lansium (R. Duncan, IFAS), 56, 10%, 5-XII-1996 as larvae on Litchi chinensis buds (A. Cati- neiras, IFAS), 2d, 39, XI-1997, ex-Litchi (J. Pena, USNM); Homestead, 146, 32, No- vember 1997, as larvae on Litchi chinensis; 6 km NW Homestead, | 2, 20 March 1998; 12, 20/23 March 1998; 4d, 39, October 1998 (USNM). BRAZIL: Planaltina, Distri- to Federal, 1000 m, 1d, 15 September 1984 (V. Becker, VBC). COSTA RICA: Province of San José, PN. Braulio Carrillo, Estacion Carrillo, 700 m, 1d, July 1990 (INBio); Province of Limon, P. N. Tortuguero, Cerro Tortuguero, 0-120 m, 12 April 1991, 1d June 1991 (INBio); Province of Limon, Sector Cerro Cocori, Fea. de E. Rojas, 150 m, 12, November 1990, 16 December Pupa of Crocidosema litchivora. a, Dorsum. b, Venter. 1991 (INBio); Province of Guanacaste, P. N. Guanacaste, 9 km S Santa Cecilia, Es- tacion Pitilla, 7OO m, 1d, 23—26 June 1993 (INBio). DOMINICAN REPUBLIC: Prov- ince of Dajabon, 13 km S Loma de Cabrera, 400 m, 2d, 20-22 May 1973 (USNM); Province of La Vega, Hotel Montana, 10 km NE Jarabacoa, 520 m, 1d, 28 May 1973 (USNM). JAMAICA: Kingston, St. An- drew, 1d, 12 March 1966 (S. S. & W. D. Duckworth, USNM). PUERTO RICO: Centro Vacacional Monte del Estado near Maricao, 650 m, 12, 1-9 March 1971 (USNM). VENEZUELA: Auyan Tepui, Ca- marata, 1d, 4-17 August 1974 (B. V. Ri- dout, BMNH). Etymology.—The species name is de- rived from the host plant genus. Comments.—Crocidosema litchivora be- longs to a group of Neotropical species that includes C. lantana, C. longipalpana, C. 326 Table 1. Duration of immature stages of Crocido- sema litchivora. n Days (x + SD) Egg 40 Al a= (0)! 1st instar 26 p) Se (Ve) 2™4 instar 7/ Syrse) 1128) 3 instar 19 pas (0) 4" instar Pal es 1X0) 5" instar 13 2-0-5 6" instar 10 Dt OL9 Larva total 40 (fre), = RA Pupa 40 lil 2= (09) calvifrons, and several undescribed species with Caribbean distributions. Secondary sexual characters are highly developed in males of this group. Sex scales are present on the head, thorax, wings, and abdomen of various species. Males of this group are dis- tinguished easily from other Crocidosema by the strongly upturned labial palpi and modified scaling on the convex frons, men- tioned above. Larvae of C. litchivora will key convinc- ingly to couplet 25 in the “key to some common tortricid larvae on apple, peach and generically related fruits’? (Brown 1987: 425). However, they do not fit either couplet of 25, 1.e., D1l’s on A8 are only slightly closer together than D2’s, and the pinacula are conspicuous. They can be dis- tinguished from the closely related C. lan- tana by their greater number of crochets: C. lantana has 20—30 crochets on prolegs A3— 6 and 15-17 on A10 (see MacKay 1959); in contrast, C. litchivora has 35—38 on pro- legs A3—6 and 20—23 on A10. Distribution of relatives —Crocidosema lantana was described from Hawaii (Busck 1910) based on specimens imported from Mexico for biological control of weedy lan- tana (Lantana camara L.; Verbenaceae) (Perkins and Swezey 1924, Zimmerman 1978). It subsequently has been introduced into Australia (Tyron 1914, Common 1957) and Micronesia (Zimmerman 1978) for bio- control. Crocidosema longipalpana was de- scribed from Puerto Rico (Méschler 1890) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 2. litchivora. Head capsule widths of Crocidosema Width in mm Stadium n Ge =SD) 1s* instar 26 O21 == 10/05 24 instar 17 0322-="0:08 3" instar 19 056) == ONG 4" instar 21 (O)9/S) Ze (114! 5" instar 13 0.97 + 0.09 6" instar 10 OUP se (OO! and has not been reported from the United States. Material examined of C. litchivora indicates that it may be restricted to Central America, Florida, and the Caribbean (in- cluding Puerto Rico). It probably is native to southern Florida, southern Texas, Mexi- co, and the Caribbean. The complex also includes several undescribed species from the Caribbean and Central America. Based on specimen evidence (IFAS), C. litchivora has been in the Homestead area at least since 1992, but in low density. Its increase in abundance in southern Florida since 1992 may be attributable to the in- crease in acreage of litchi plantings follow- ing hurricane Andrew. Owing to the ab- sence of records prior to 1992, it is likely that this Caribbean species is a fairly recent arrival to southern Florida. Food plants.—In addition to Litchi chi- nensis, C. litchivora has been reared from wampi (Clausena lansium (Lour.) Skeels; Rutaceae) (n = 1) and loquat (Eriobotyra Japonica (Thunb.) Lindl.; Rosaceae) (n = 2) in southern Florida. Biology Based on laboratory rearings, the total duration of the immature stages from egg Table 3. Commercial importation of Litchi into the UES: in! 1998" Country of Origin # Shipments Weight China 17 456,418 kg Israel 4 16,789 kg Mexico 83 204,533 kg Taiwan 14 295,486 kg 327 VOLUME 104, NUMBER 2 0661 asneID 2 UOIMEN “SZ6I “TE 19 FOTIND C661 PRAY 2 Pyesemvy ‘nseN ZR6I JJOUOYLIG ‘961 PAID suatutoads SH] pue WNSN C66] YOIURAMA'] 2 OYOINY C661 YOtURAMET I OYOINY *[ 86] AUUIAA 29 297] Sho] PUT] RISO 9L61 AUPID ‘T86I “0961 JJoUOyLIG 0661 asneID 2 UOIMEN ‘9/61 OULID P66] souor ‘8/6 URWAWILUIZ “9/61 OP[D “8961 “0961 OUOyeIG +66 SOUOL ‘8/6 URWIOUIUUTZ toded jyuasaid SLé6l UBULIOUIUUTZ €661 YOIURAMIT WZ OYOINY “TRG ‘O96I “6E6I JJouoyeiqd €661 YOLURAMAT 2 OYOINY *[ 6] AUUIAA 2 99] €661 YOUURAMAT 2 OYOINY “O661 AOL [861 AOUUIAA 27 99] T861 Jfouoyriq ‘OL61 MUeID €661 NSeN ROLY BISY }SeoyINOS vIsy JsvoyINos epHoly puepreyL vISY [izelg vISY JSeoyINOS PISQUOIOYAY BISOUOIOTAY ‘Te MEH TeMeHy BpHoly neMeHy vISV pueliey BIpuy eISVY AS suoy SuOH vIsSy JseayINoOS uevderp ‘wreujar, (IMAI) VlasJoona] viquojpuNDy J, (‘SOY 2% ‘plaq) Vuposip syosayINIS (oA) sidvona] syosaysjnis YOMUIaH VDI du sp1ajoajold (ADM) VUDJOnNpa DUOWOP (1OUIDIN) Diuvaffod DUOWOH (RUITT BISOD) WwnuUDYUDIND DULOSOAPUDUUKD (OLA) Djogosdy vnpng (oA) VIUSVyad DigajydojdKyD (1I9MOTJ) DIJaposquo vigajydojdi14y (tong) vpidazji vigajydojdiiy DAOAIYIJ] DWASOPIIOAD yosng pupjup] vulasop1g0sZD (YOLAI) V4981]]99 Dpogajsauy (Jaye AA) VUDaIvIIU SdiyI4V (OLAV) Ss7doyoou sdiyosy (‘9[SOY “A JAYOSt4) Vud10 sakydoxopy (Taye) VUBINIIISD{ sakydoxopy AOSJOUZNY VUDUIAOLSIU DINIOAIV SIDUDIOJOY uolsoy o1ydeis0aH so1sadg ‘OPIMPPIOM 1/9177 WO] PAPIOIII ePIOLNIOT, “pf [qe] 328 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON through pupa is 34 + 1 days. Eggs hatch 3—4 days after oviposition, and there are six larval instars. Duration of all stages is listed in Table 1. Larval head capsule width in- creases from 0.21 mm in first instars to 1.01 mm in final instars (Table 2). Females begin to oviposit 5 + 1 days after emergence, with the last oviposition bout coming 15 + 3 days after emergence. Adults lived 11 + 7 days. The number of eggs obtained per female (n = 6) was 33 + 5. Eggs of C. litchivora were found singly or in groups of no more than 3 eggs on a bud. First instars emerge by chewing through the chorion in the region of the mi- cropyle, leaving the empty eggshell in place. At first they feed externally on the bud, but eventually bore into it. Feeding damage is characterized by a small, nearly elliptical hole in the bud, usually accom- panied by light brown frass protruding from the hole. First instars feed mostly in buds; later instars bore into young stems, but also feed on flowers, predominantly in the ova- ry, or bore into developing fruits. When flowers are present, the larvae tie the pollen and petals together with silk and use the inflorescence as a pupation site. If only small fruits are present, larvae chew into the lateral part of the fruit, often leaving it hollow. Once fruits are sufficiently mature, larvae do not attack them; only rarely did we encounter mature fruit with larval feed- ing damage. Infrequently, larvae leave the bud, inflorescence, or fruit, and roll leaves of the food plant, and pupate in this shelter. At least one species of Tachinidae emerged from late instars (n = 20) collected in the field. Larval feeding can result in signifi- cant economic loss as a result of lowered fruit production. Other Tortricid Pests of Litchi In 1998, approximately 973,226 kilo- grams (over 1,000 tons) of litchi were im- ported into the United States, primarily from China, Taiwan, Mexico, and Israel (Table 3). Imported litchi are treated rou- tinely with methyl bromide to prevent the entry into the United States of exotic pest species (J. Cavey, pers. comm.). However, based on inspections of personal baggage and other cargo, it is likely that hundreds of kilograms of Litchi enter this country il- legally, undetected, and untreated every year, primarily from the Orient. The larvae of a large number of Tortrici- dae have been recorded feeding on litchi in the tropical regions of the world (see Table 4). Although most are reported infrequently, Cryptophlebia illepida (Butler), C. ombro- delta (Lower), and C. peltastica (Meyrick) are major pests of this crop (e.g., Newton and Crause 1990, Jones 1994), particularly in Hawaii and the Far East. Of the 19 spe- cies listed in Table 4, three are known from the contiguous United States (1.e., Florida)— Crocidosema_ litchivora, C. lantana, and Proteoteras implicata Heinrich. These three appear to be Neotropical, for the most part, ranging into Florida from either Central America or the Caribbean. Crocidosema lit- chivora and P. implicata at times have in- flicted considerable damage in commercial litchi orchards in Florida, but C. /antana ap- parently has not. The importation of Litchi plant stock into Florida from Hawaii creates a potential for the Hawaiian Litchi pests, C. illepida and C. ombrodelta, to inadvertently be brought into Florida. Hence caution must be exercised in the movement of Litchi plant material. Because larvae of many of these pests feed primarily in the fruit, the unde- tected, illegal import of Litchi fruit from the Far East creates a much greater potential for the introduction of additional Litchi pests from that geographic area. ACKNOWLEDGMENTS We thank the University of Florida Insti- tute of Food and Agricultural Sciences for facilitating a visit by JWB to litchi orchards near Homestead, Florida. We thank Ron Sponaugle, USDA, APHIS-PPQ, Riverdale, Maryland, for information on the importa- tion of Litchi into the U.S. We are grateful to Richard Brown, Mississippi State Uni- versity, for information on the identity of VOLUME 104, NUMBER 2 this species. We acknowledge Eugenie Phil- lips (INBio) and Kevin Tuck (BMNH) for allowing us to examine specimens in their care. We thank the following for helpful comments on the manuscript: Richard Brown, Mississippi State University, Mis- sissippi, USA; FE Christian Thompson and David Smith, USDA, Systematic Entomol- ogy Laboratory, National Museum of Nat- ural History, Washington, D.C., USA. Illus- trations of the male and female genitalia, larva, and pupa were prepared by David Adamski, USDA, Systematic Entomology Laboratory, National Museum of Natural History, Washington, D.C., USA. Finally, we thank the Bioresource LSF Project of the Natural History Museum London (EU- DGXII TMR Programme) which contrib- uted funds allowing J. Baixeras to visit that institution. LITERATURE CITED Brown, J. W. and J. A. Powell. 1991. Systematics of the Chrysoxena group of genera (Lepidoptera: Tortricidae: Euliini). University of California Pub- lications in Entomology 111, 87 pp. Brown, R. L. 1987. Tortricidae (Tortricoidea), pp. 419—433. In Stehr, E, ed. Immature insects. Ken- dall/Hunt Publishing Company, Dubuque, Iowa, 754 pp. Busck, A. 1910. New Central American Microlepidop- tera introduced into the Hawaiian Islands. Pro- ceedings of the Entomological Society of Wash- ington 12: 132-135. Clarke, J. E G. 1976. Microlepidoptera: Tortricoidea. Insects of Micronesia 9, no. |. Bernice P. Bishop Museum, Honolulu, 144 pp. Common, I. E B. 1957. The occurrence of Epinotia lantana (Busck) (Lepidoptera: Olethreutidae) in Australia. Proceedings of the Linnean Society of New South Wales 82: 230—232. Costa Lima, A. 1945. Insectos do Brasil 5 (28), Lep- id6pteros, parte 1. Escola Nacional Agronomia, Rio de Janeiro, Brazil, 379 pp. Diakonoff, A. 1939. The genera of Indo-Malayan and Papuan Tortricidae. Zoologische Mededeelingen van het Rijksmuseum van Natuurlijke Historie 21: 111-240. 1960. A second note on Microlpeidoptera from South China (Lepidoptera: Tortricidae). Bei- triage zur Entomologie 10: 132-133. . 1968. Microlepidoptera of the Philippine Is- lands. United States National Museum Bulletin 257, 483 pp. B29 . 1982. On a collection of some families of Mircolepidoptera from Sri Lanka (Ceylon). Zool- ogische Verhandelingen, Leiden 193, 124 pp. + figs. Horak, M. 1984. Assessment of taxonomically signif- icant structures in Tortricinae (Lep., Tortricidae). Bulletin de la Société Entomologique Suisse 57: 3-64. Jones, V. P. 1994. Oviposition patterns of koa seed- worm and litchi fruit moth (Lepidoptera: Tortri- cidae) on macadamia and litchi in Hawaii. Journal of Economic Entomology 87: 1278-1284. Kuroko, H. and A. Lewvanich. 1993. Lepidopterous pests of tropical fruit trees in Thailand. Japan In- ternational Co-operation Agency, Tokyo, 133 pp. Lee, L. H. Y. and R. Winney. 1981. Check list of ag- ricultural insects of Hong Kong 1981. Hong Kong Agriculture and Fisheries Department Bulletin 2: 1-164. MacKay, M. 1959. Larvae of North American Oleth- reutidae (Lepidoptera). Canadian Entomologist, supplement 10, 338 pp. Moschler, H. A. 1890. Die Lepidopteren der Insel Por- to Rico. Abhandingen nerasgeschen von der Senckenbergischen Naturforsden Gesellschaft Frankfort am Main 16: 330—346. Nasu, Y. 1993. New record of Acroclita nigrovenana (Lepidoptera, Tortricidae) from Japan. Japanese Journal of Entomology 61: 216. Nasu, Y., K. Kawasaki, and N. Arakaki. 1993. Stath- erotis discana (Felder et Rogenhofer) (Lepidop- tera: Tortricidae) injurious to litchi from Japan. Applied Entomology and Zoology 28: 97-101. Newton, P. J. and C. Crause. 1990. Oviposition on Li- tchi chinensis by Cryptophlebia species (Lepidop- tera: Tortricidae). Phytophylactica 22: 365-367. Perkins, R. C. L. and O. H. Swezey. 1924. The intro- duction into Hawaii of insects that attack lantana. Bulletin of the Hawaiian Sugar Planters’ Associ- ation, Entomology, Ser. 16, 83 pp. Quilici, S., B. Verbizier, B. Trahais, and R. Manikom. 1988. Note sur les ravageurs du litchi a la Re- union. Fruits (Paris) 43: 459—464. Robinson, J. E, A. Day, R. Cathbert, and E. V. Wann. 1977. The pickleworm: Laboratory rearing and ar- tificial infestation on cucumber. Journal of Eco- nomic Entomology 72: 305-307. Stanley, D. 1998. Florida growers like lychees & lon- gans. Agricultural Research 46: 8. Tuck, K. 1990. A taxonomic revision of the Malaysian and Indonesian species of Archips Hiibner (Lepi- doptera: Tortricidae). Entomologica Scandinavica PANE WID=MNLIO. Tyron, H. 1914. Report of the entomologist and veg- etable pathologist. Annual Report of the Depart- ment of Agriculture, Stock, Queensland, 1913-14: 114-120. Zimmerman, E. C. 1978. Insects of Hawaii, Volume 9, Microlepidoptera, Part 1. University of Hawaii Press, Honolulu, 881 pp. PROC. ENTOMOL. SOC. WASH. 104(2), 2002, pp. 330-334 FURTHER CHARACTERIZATION OF PALEOMASTACRIS AMBARINUS PEREZ ET AL. (ORTHOPTERA: EUMASTACIDAE) FROM DOMINICAN AMBER DANIEL E. PEREZ-GELABERT Research Associate, Department of Systematic Biology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0105, U.S.A. (e-mail: perez.daniel @nmnh.si.edu) Abstract.—Three new specimens of fossil Eumastacidae in Dominican amber are char- acterized and identified as the previously described Paleomastacris ambarinus Perez et al. 1997. The better preservation of these specimens allows examination of features of the head not visible in the holotype. Paleomastacris ambarinus lacks the protruding fas- tigium typical of extant Hispaniolan Eumastacidae and in this respect seems to resemble Central American Episactinae more closely than those of Hispaniola. This fossil species likely represents an early link between these two extant groups of eumastacids. Resumen.—Se estudian tres nuevos especimenes de Eumastacidae foésiles en ambar do- minicano y se identifican como el previamente descrito Paleomastacris ambarinus Perez et al. 1997. La mejor preservaciOn de estos especimenes permite examinar rasgos de la cabeza no visibles en el holotipo. Paleomastacris ambarinus carece del fastigio protuberante tipico de los Eumastacidae vivientes en la Hispaniola y en este respecto se asemeja mas a los Episactinae centroamericanos que a los de la Hispaniola. Esta especie fosil podria repre- sentar una conecciOn temprana entre estos dos grupos vivientes de eumastacidos. Key Words: Fossil eumastacid grasshoppers were known until recently only as compression fossils from North American and Asian lo- cations, varying in age from Miocene (15— 30 mya) to lower Jurassic (190 mya) (Zeu- ner 1944; Sharov 1968; Lewis 1974, 1976; Lin 1980). More recently, Martins-Neto (1991) reported fossil eumastacoids from the lower Cretaceous of the Santana for- mation in Araripe, Brazil. Paleomastacris ambarinus Perez et al. 1997, from the Mio- cene Dominican amber, is the first eumas- tacid and only acridomorph grasshopper re- ported so far from amber. The type speci- men (American Museum of Natural History no. DR-10-1795) is a juvenile male with several morphological features that distin- amber fossils, Dominican Republic, Eumastacidae, grasshoppers guish it from extant Eumastacidae. The ex- tant fauna of eumastacid grasshoppers in Hispaniola comprises at least 5 genera and 10 species (Perez-Gelabert 1999). Although the taxonomic characterization of the group on the island is still incipient, this is prob- ably a nearly complete representation of the lineages found there today. Geologically the amber mines found north of Santiago are considered Miocene in age (15—20 million years old) (Iturralde-Vinent and MacPhee 1996) although others (Lambert et al. 1985) have previously argued that they could be as much as 33—40 million years old, based on studies of nuclear magnetic resonance spectroscopy. In this paper, I report on three new fossil VOLUME 104, NUMBER 2 specimens of Eumastacidae in Dominican amber and assign them to the previously de- scribed species P. ambarinus. Since these specimens are preserved better than the ho- lotype, I describe features of the head not previously visible and discuss its relation- ship to extant eumastacids. SPECIMENS Through the kind assistance of Robert E. Woodruff and George O. Poinar, Jr., I had the opportunity to examine three eumasta- cid specimens in Dominican amber. One of these specimens appeared in a photograph together with a hypothetical illustration of its ancient environment in the book “The Amber Forest” by Poinar and Poinar (7999): Specimen MNHNSD (Figs. 4, 6).—Male juvenile, embedded diagonally in an oval- shaped flat piece, approximately 23 x 16 X 6 mm in size and 1.8 grams in weight. The fossil has the dorsal and ventral portions of the head obscured by products of decom- position and detritus. This specimen was donated by Dr. R. E. Woodruff to the en- tomological collection of the Museo Nacio- nal de Historia Natural, Santo Domingo (MNHNSD). Specimen O-2-I15A (Fig. 5).—Male ju- venile, positioned flat and centrally on a golden flattened piece approximately 12 x 6 mm that allows very good visibility of most external features on both sides. Its preservation condition is very good. De- posited in the Poinar collection. Specimen O-2-15C (Figs. 1, 2, 3).—Male juvenile, included near the top of a 5 mm cubic piece that is polished flat at its base. Also in good preservation condition. De- posited in the Poinar collection. METHODS Although measured and studied at differ- ent times, the same 7—30X stereoscope fit- ted with an ocular micrometer with preci- sion to 0.01 mm was used to examine the holotype and the new specimens. The spec- imens were examined directly without us- S3il ing immersion techniques. Photographs of the included eumastacids were taken with a Sony DKC-5000® digital photo camera mounted on a Leica MZ APO® stereoscope. The fossils were compared to adult and ju- venile specimens of several extant Hispa- niolan and Central American species. Paleomastacris ambarinus Perez et al. Me Paleomastacris ambarinus Perez et al. 1997: 150 (3; from Dominican amber; photo of complete specimen, figs. of head, pronotum, legs, hind tibia, abdom- inal end); Eumastacidae in Poinar and Poinar (1999: 53, photo 49; 57, habitus illustration). The unique combination of characters that allows the identification of these three new specimens as P. ambarinus are the short and flattened pronotum with anterior and posterior margins quadrate; genicular lobe of hind femur forming well-defined spine; triangular epiproct slightly constrict- ed in its middle and elongated just past ab- dominal end; and cerci simple, slightly glo- bose at base, then finely elongated, slightly projecting at sides of abdominal end. The split male subgenital plate described from the holotype as a unique feature, is some- what visible in the MNHNSD specimen (Fig. 4), but not in the other two. This is apparently due to the positioning of the specimens in the amber pieces and not to any obscuring debris or imperfection of the matrix. The approximate dimensions of the new specimens are given in Table |. As in the holotype, all three specimens represent male juvenile individuals of moderately well-defined morphology. Based on com- parisons to nymphs of several extant spe- cies, I believe that P. ambarinus was prob- ably smaller than all known extant Hispa- niolan eumastacids. Both specimens from the Poinar collection show the head intact; the head was incompletely characterized in the original description. In neither specimen eS) oS) i) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-6. Photos of Paleomastacris ambarinus. 1, Specimen O-2-15C. 2, Detail of head in specimen O-2- 15C, showing the non-elongated fastigium of vertex (fv). 3, Abdominal end of specimen O-2-15C showing finely pointed cerci (c) and elongated epiproct (ep). 4, Abdominal end of specimen MNHNSD showing split subgenital plate (sgp). 5, Specimen O-2-15A. 6, Specimen MNHNSD. In all figures scale bar represents 0.5 mm. Table 1. Some approximate dimensions (mm) of the three new specimens of P. ambarinus and the previously described holotype. Holotype MNHNSD O-2-15A O-2-15C ne ee ee ee Body length 5.50 7.10 7.47 6.32 Hind femur length 3.00 4.74 2.89 DAS) Pronotum length 1.00 — 0.74 0.74 VOLUME 104, NUMBER 2 does the fastigium of vertex obviously pro- trude beyond the eyes (Figs. 1, 2, 5). Extant Dominican eumastacids have a distinctively large and protruding fastigium of vertex even as small nymphs. The condition in the amber specimens is rather reminiscent of the Central American Episactinae (Episac- tus Burr, Gymnotettix Bruner and Maya- mastax Uvarov), which also lack this pro- truding fastigium. DISCUSSION Living Hispaniolan eumastacids charac- teristically occupy defined altitudinal strata of various mountain forests and apparently have rather limited distribution ranges. Be- cause of these characteristics, it is not sur- prising that the three additional fossil spec- imens which probably originated in the same area as the type (amber mines of Cordillera Central north of Santiago), represent the de- scribed species. The lack of a protruding fas- tigium in P. ambarinus resembles what is found today in extant Central American Epi- sactinae suggesting a closer affinity to them than to extant Hispaniolan species. This in- terpretation fits models of Caribbean histor- ical biogeography (e.g., Buskirk 1985) that indicate a closer proximity between the Greater Antilles and Central America during the Miocene than today. The phylogenetic relationships within and between the His- paniolan and Central American Episactinae are not known. The only phylogenetic data available on this group of eumastacids are from the analyses of Rowell and Flook (1998), in which phylogenetic trees based on 12S and 16S mtRNA genes, show the His- paniolan genera Antillacris Rehn & Rehn, Espagnoleta Perez-Gelabert and Espagno- lopsis Perez et al. on one branch of the Epi- sactinae clade somewhat separated from the branch containing the Central American ge- nus Episactus, which may be their closest relatives. Paleomastacris ambarinus likely represents an early link between these groups that have subsequently evolved, geo- graphically separated from each other for at least 20—30 million years. eS) Oo i>) ACKNOWLEDGMENTS I thank Robert E. Woodruff (Center for Systematic Entomology, Gainesville, FL) for allowing examination of his amber fos- sil collection, cordial hospitality during vis- its to Gainesville, and for the donation of his specimen of P. ambarinus to the MNHN, Santo Domingo. I am also indebt- ed to George O. Poinar, Jr. (Oregon State University, Corvallis, OR), for the loan of amber specimens, thus providing the nec- essary incentive to complete this paper. Do- nald Azuma facilitated examination of specimens of Central American eumasta- cids from the Academy of Natural Sciences, Philadelphia, PA, collection. C. H. E Ro- well (University of Basel, Switzerland) and Scott E. Miller (Department of Systematic Biology, National Museum of Natural His- tory, Smithsonian Institution, Washington, DC) read the manuscript and provided use- ful suggestions. LITERATURE CITED Buskirk, R. E. 1985. Zoogeographic patterns and tec- tonic history of Jamaica and the northern Carib- bean. Journal of Biogeography 12: 445—462. Iturralde-Vinent, M. A. and R. D. E. MacPhee. 1996. Age and paleogeographical origin of Dominican amber. Science 273: 1850-1852. Lambert, J. B., J. S. Frye, and G. O. Poinar, Jr. 1985. Amber from the Dominican Republic: Analysis of nuclear magnetic resonance spectroscopy. Ar- chaeometry 27: 443-51. Lewis, S. E. 1974. Four specimens of fossil grasshop- pers (Orthoptera: Caelifera) from the Ruby River Basin (Oligocene) of southwestern Montana. An- nals of the Entomological Society of America 67: 523-524. Lewis, S. E. 1976. A new specimen of fossil grass- hopper (Orthoptera: Caelifera) from the Ruby Riv- er Basin (Oligocene) of southwestern Montana. Annals of the Entomological Society of America 69-5120! Lin, Q.-b. 1980. Fossils of Mesozoic insects from Zhe- jian and Anhui provinces, pp. 211—234. Jn Nan- jing Institute of Geology and Paleontology, Aca- demia Sinica, eds. Division and Correlation of Mesozoic Deposits of Volcanic Origin in Zhejian and Anhui Provinces. Scientific Publishing House, Beijing. Martins-Neto, R. G. 1991. Primeiro registro de Eu- mastacoidea (Insecta, Caelifera) da formagao San- 334 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tana, Cretadceo Inferior do nordeste do Brasil. Anais de la Academia Brasileira de Ciencias 63: 91-92. Perez, D. E., B. Hierro, G. Dominici, and D. Otte. 1997. New eumastacid grasshopper taxa (Orthop- tera: Eumastacidae: Episactinae) from Hispaniola, including a fossil new genus and species from Do- minican amber. Journal of Orthoptera Research 6: 139-151. Perez-Gelabert, D. E. 1999. Saltamontes eumastacidos de la Reptiblica Dominicana. Novitates Caribaea 1: 53-57. Poinar, G. O., Jr. and R. Poinar. 1999. The Amber For- est—A Reconstruction of a Vanished World. Princeton University Press, Princeton, 239 pp. Rowell, C. H. FE and P. Flook. 1998. Phylogeny of the Caelifera and the Orthoptera as derived from ri- bosomal gene sequences. Journal of Orthoptera Research 7: 147-156. Sharoy, A. G. 1968. Phylogeny of the Orthopteroidea. Transactions of the Institute of Paleontology of the Academy of Sciences U.S.S.R 118: 1-251. Zeuner, E E. 1944. The fossil Acrididae (Orth. Salt.) Part IV. Acrididae incertae sedis and addendum to Catantopinae. Annals and Magazine of Natural History (11) 78: 359-383. PROC. ENTOMOL. SOC. WASH. 104(2), 2002, pp. 335-347 DESCRIPTION OF IMMATURE STAGES OF TEPHRITIS STIGMATICA (COQUILLETT) (DIPTERA: TEPHRITIDAE) RICHARD D. GOEDEN Department of Entomology, University of California, Riverside, CA 92521, U.S.A. (e-mail: richard.goeden @ucr.edu) Abstract.—The pre-imaginal stages of Tephritis stigmatica (Coquillett) are described. Tephritis stigmatica is an oligophagous, multivoltine, facultatively axillary-bud-gall-form- ing and florivorous tephritid associated with Senecio spp. (Asteraceae) in southern Cali- fornia. The egg, first-, second-, and third-instar larvae and puparium are described and figured, and selected characteristics of these stages are compared with those of other southern California Tephritis. The egg of T. stigmatica is covered by a smooth, membra- neous sheath of unknown function. The descriptions of the first, second, and third instars are the most complete reported to date for the genus Tephritis. The integumental petals of all three instars are found in medial and lateral positions on the gnathocephalon and increase in numbers from the first to the last instar. The most lateral integumental petal is fused with the stomal sense organ in the first instar. The anterior spiracle has four or five papillae in the second instar, and four papillae in the third instar. The pairs of stelex sensilla surrounding the posterior spiracular plates increase in number from the first to the second instar, and retain the same number and locations, but include two pairs of verruciform sensilla dorsally and laterally in the third instar. The lateral spiracular com- plexes of the metathorax and first abdominal segment of the second and third instars are described and pictured. The lateral spiracles of the third instar uniquely are prominent on raised peritremes, unlike the second instar, in which they are difficult to locate, as in most other tephritine larvae. Key Words: Insecta, Tephritis, Senecio, Asteraceae, nonfrugivorous Tephritidae, taxon- omy of immature stages, egg, larvae, puparium, spiracles To date, the life histories and immature stages of five species of Tephritis have been described in detail from southern Califor- nia: 7. arizonaensis Quisenberry (Goeden et al. 1993), 7. baccharis (Coquillett) (Goe- den and Headrick 1991), T. joanae Goeden (Goeden 1993, 2001b), T. teerinki Goeden (Goeden 200Ic) and T. footei Goeden matica is relatively well known (Tauber and Toschi 1965; Goeden 1988, 1993). MATERIALS AND METHODS Monthly samples of overwintered, axil- lary bud galls containing eggs, larvae, pu- paria of 7. stigmatica were collected in February, March and April, 2000, from Se- (Goeden 2002). The immature stages of a sixth species, 7. stigmatica (Coquillett), are described in this paper to facilitate their comparison with the immature stages of the other five species. The biology of T. stig- necio flaccidus Lessing var. douglassii (deCandolle) B. Turner and T. Barkley at the same two southern California sites used in my 1984-85 field study of the life his- tory of this tephritid (Goeden 1988): (1) 336 south of Lamont Peak at Spanish Needle Creek; Sequoia National Forest (north sec- tion); (T)ownship 22 (S)outh, R(ange) 36 E(ast); Kern County; and (2) 2 km south of Pearblossom; T5 North), R10 W(est); Los Angeles County. Samples of galls were transported in cold-chests in an air-condi- tioned vehicle to the laboratory and stored under refrigeration for subsequent dissec- tion, description, photography, and mea- surement. Three eggs, 18 first-, 16 second-, and 17 third-instar larvae, and three puparia dissected from galls were preserved in 70% EtOH for scanning electron microscopy (SEM). Specimens for SEM were hydrated to distilled water in a decreasing series of acidulated EtOH. They were osmicated for 24 h, dehydrated through an increasing se- ries of acidulated EtOH and two, I-h im- mersions in hexamethyldisilazane (HMDS), mounted on stubs, sputter-coated with a gold-palladium alloy, and studied and dig- itally photographed with a Philips XL-30 scanning electron microscope in the Central Facility for Advanced Microscopy and Mi- croanalysis, University of California, Riv- erside. Plant names used in this paper follow Hickman (1993); tephritid names follow Foote et al. (1993). Terminology and tele- graphic format used to describe the imma- ture stages follow Goeden (2001a, b, c, d), Goeden et al. (1993), Goeden and Headrick (1991), Goeden and Teerink (1999), Teer- ink and Goeden (1999), and our earlier works cited therein. Voucher specimens of immature stages of 7. stigmatica reside in my research collection. Digitized photo- graphs used to construct text figures were processed with Adobe Photoshop® Version 6. RESULTS AND DISCUSSION first-, sec- ond-, and third-instar larvae, and puparium of Tephritis stigmatica are described below. Egg. Four eggs measured in situ in field- collected, immature, overwintered galls were white, opaque, smooth, elongate-ellip- soidal, 0.8 mm long by 0.2 mm wide, Immature stages.—The egg, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 20 um Fig. 1. anterior to left; (B) pedicel showing pattern and shapes Egg of Tephritis stigmatica: (A) habitus, of aeropyles; (C) pedicel of a different egg, for com- parison. smoothly rounded at tapered basal end (Fig. 1A); pedicel button-like, 0.02 mm long, cir- cumscribed apically by different-sized, ir- regularly-, semi-rectangular, or semicircu- lar-shaped aeropyles arranged singly or in VOLUME 104, NUMBER 2 rows of two parallel to the long axis of the ess (Riss: 1B, C): The egg of T. stigmatica (Fig. 1A), like those of 7. joanae (Goeden 2001b), T. teer- inki (Goeden 2001c), and 7. footei (Goeden 2002) differs from eggs of T. baccharis (Goeden and Headrick 1991) and T. arizo- naensis (Goeden et al. 1993) by lacking prominent polygonal reticulation of the chorion. Also, the egg of 7. stigmatica, like those of 7. teerinki, T. joanae, and T. footei, apparently is covered by a smooth, mem- braneous sheath (Figs. 1A, B; Goeden 2001b, c, 2002), which remains intact and is not partly shed and peeled back during oviposition as in 7. baccharis (Goeden and Headrick 1991) and JT. arizonaensis (Goe- den et al. 1993). The function of this mem- braneous sheath remains unknown. It was first reported for 7. arizonaensis by Goeden et al. (1993), who then belatedly recognized it in 7. baccharis, and apparently only has been reported to date from the eggs of these six species of Tephritis. In T. arizonaensis (Goeden et al. 1993), this membraneous Sheath also is prominently, longitudinally striated. Weak longitudinal striations are present at the anterior, pedicellar end of the egg of T. footei, but otherwise are not seen on the rest of the egg body (Goeden 2002). First instar larva: White, cylindrical (Fig. 2A); gnathocephalon conical; dorsal sensory organ well-defined, domed (Fig. 2B-1); anterior sensory lobe (Fig. 2B-2) with terminal sensory organ (Fig. 2B-3), lateral sensory organ (Fig. 2B-4), suprala- teral sensory organ (Fig. 2B-5), and pit sen- sory organ (Fig. 2B-6); stomal sense organ (Figs. 2B-7, C-1) ventrolaterad of anterior sensory lobe and fused with flattened, pro- trudent, lateral integumental petal (Figs. 2B-8, C-2) dorsad of each mouthhook (Figs. 2B-9, C-3), two medial integumental petals between anterior sensory lobes, up- per one bulbous, and lower one, less prom- inent (Fig. 2B-10); mouthhook (Figs. 2B-9, C-3) bidentate; median oral lobe laterally compressed, apically rounded, ventrally flattened (Figs. 2B-11, C-4), basally at- 337 tached to labial lobe (Fig. 2B-12) bearing two, prominent apical pores; posterior spi- racular plate bears two ovoid rimae (Fig. 2D-1), ca. 0.01 mm long, and four inter- spiracular processes (Fig. 2D-2), each with one or two, foliose branches with one to three, apical teeth, longest branch measur- ing 0.01 mm; stelex sensillum dorsolaterad (Fig 2E) of posterior spiracular plate, ven- trolateral stelex sensillum not seen; inter- mediate sensory complexes (Figs. 2D-3, F) each with a stelex sensillum (Fig. 2F-1) and a medusoid sensillum (Fig. 2F-2). Among southern California Tephritis spp., the first instar previously was de- scribed in detail only for 7. teerinki (Goe- den 2001c), which along with the partial description for the first instar of T. footei (Goeden 2002), provide at least some basis for comparison. For example, the integu- mental petal fused with the stomal sense or- gan in T. stigmatica (Figs. 2B-7, 8, C-1, 2) also distinguishes the first instars of 7. teer- inki (Goeden 2001c) and 7. footei (Goeden 2002), as well as the first instars of at least five species of Neaspilota (Goeden 2001a), from subsequent instars. This character was first reported for the first instar of Trupanea vicina (Wulp) (Goeden and Teerink 1999). The integumental petals of the first instars of 7. footei (Goeden 2002) and T. stigma- tica (Figs. 2B-8, 10, C-2) are similarly shaped and separable into lateral and me- dial positions. The mouthhooks of the first instars of Tephritis teerinki (Goeden 2001c) and 7. stigmatica (Figs. 2B-9, C-3) are bi- dentate. The intermediate sensory complex- es ventral to the posterior spiracular plates of the first instars of 7. footei (Goeden 2002) and T. stigmatica (Figs. 2D-3, F) are similarly composed of a stelex sensillum and a medusoid sensillum, and the posterior spiracular plates are flanked, at least dor- solaterally in both species, by stelex sensilla (Goeden 2002, Fig. 2E) Second instar larva: White, cylindrical, rounded anteriorly, truncated posteriorly, body segments well-defined (Fig. 3A); gnathocephalon conical, anteriorly flat- 338 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 2. First instar of Tephritis stigmatica: (A) habitus, anterior to left: (B) gnathocephalon, ventrofrontal view, 1—dorsal sensory organ, 2—anterior sensory lobe, 3—terminal sensory organ, 4— lateral sensory organ, 5—supralateral sensory organ, 6—pit sensory organ, 7—stomal sense organ, 8—lateral integumental petal, 9— mouthhook, 10—median integumental petal, 11—median oral lobe, 12 labial lobe; (C) gnathocephalon, dor- solateral view, 1—stomal sense organ, 2—lateral integumental petals, 3—mouthhooks, 4—median oral lobe: (D) caudal segment, 1—rimae, 2—interspiracular processes, 3—intermediate sensory complexes; (E) dorsolateral stelex sensillum; (F) intermediate sensory complex, (1) stelex sensillum and (2) medusoid sensilllum. VOLUME 104, NUMBER 2 339 Fig. 3. Second instar of Tephritis stigmatica: (A) habitus, anterior to left; (B) prothorax, lateral view, 1— median vertical suture on gnathocephalon, 2—minute acanthae, 3—verruciform sensilla, 4—anterior spiracle; (C) gnathocephalon, dorsolateral view, 1—dorsal sensory organ, 2—anterior sensory lobe, 3—medial integu- mental petals, 4—lateral integumental petals, 5—mouthhook, 6—stomal sense organ, 7—oral ridges, 8—median oral lobe, 9—minute acanthae; (D) close-up of gnathocephalon, 1—dorsal sensory organ, 2—anterior sensory lateral sensory organ, 5—supralateral sensory organ, 6—pit sensory organ, lateral integumental petals, 9—stomal sense organ, 10—oral ridges; (E) lobe, 3—terminal sensory organ, 4 7—medial integumental petals, 8 gnathocephalon, ventrolateral view, 1—lateral integumental petals, 2—stomal sense organ, 3—oral ridges, 4— mouthhook, 5—median oral lobe, 6—minute acanthae; (F) anterior spiracle with middle papilla partially dis- placed. 340 tened, and medially cleaved by a vertical suture (Fig. 3B-1); posteriorly directed, spi- nose, minute acanthae incompletely circum- scribe gnathocephalon anteriorly (Figs. 3B- 2, C-9, E-6), minute acanthae also circum- scribe meso- and metathorax, and abdomi- nal segments Al—A2 anteriorly, with bands of minute acanthae widening ventrally and laterally on A3—A5, but narrowing again on A6—A8, prothorax also circumscribed dor- so- and lateromedially by verruciform sen- silla (Fig. 3B-3); dorsal sensory organ well- defined, rounded dome (Figs. 3C-1, D-1); anterior sensory lobe (Figs. 3C-2, D-2) bears terminal sensory organ (Fig. 3D-3), lateral sensory organ (Fig. 3D-4), suprala- teral sensory organ (Fig. 3D-5), and pit sen- sory organ (Fig. 3D-6); three or four, me- dial, integumental petals in vertical row be- tween anterior sensory lobes to each side of medial crease (Figs. 3C-3, D-7) and two each, lateral, spatulate or papillate, integu- mental petals (Figs. 3C-4, D-8, E-1) in two rows above each mouthhook (Fig. 3C-5), and separate from stomal sense organ (Figs. 3C-6, D-9, E-2) ventrolaterad of anterior sensory lobe; four or five, complete oral ridges (Figs. 3C-7, D-10, E-3) laterad of each anterior sensory lobe; mouthhook tri- dentate (Figs. 3C-5, E-4); median oral lobe laterally compressed, apically pointed (Figs. 3C-8, E-5); anterior spiracle with to- tal of four (Fig. 3B-4) or five (Fig. 4A-1), doliform or subquadrate papillae; lateral spiracular complexes of metathorax (Fig. 4A) and first abdominal segment (A1) (Fig. 4C) each consist of a spiracle (Figs. 4A-2, B, C-1, D) and two, verruciform sensilla (Figs. 4A-3, C-2), one sensillum vertical to the other; posterior spiracular plate (Figs. 4E-1, F) bears three ovoid rimae (Fig. 4F- 1), ca. 0.02 mm long, and four interspira- cular processes (Fig. 4F-2), each with two to four foliose branches, each with one or two apical teeth, longest branch measuring 0.01 mm; stelex sensilla dorsad (Fig. 4E-2), dorsolaterad (Fig. 4E-3), laterad (Fig. 4E- 4), and ventrolaterad (Fig. 4E-5) of poste- rior spiracular plate; intermediate sensory PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON complexes (Figs. 4E-6, F-3) each with a stelex sensillum and a medusoid sensillum. The habitus of the second instar of T. stigmatica (Fig. 3A) approximates those of T. baccharis (Goeden and Headrick 1991), T. arizonaensis (Goeden et al. 1993), T. Joanae (Goeden 2001b), 7. teerinki (Goe- den 200Ic), and 7. footei (Goeden 2002). The anterior spiracle of the second instar of T. stigmatica bears four (Fig. 3B-4) or five (Fig. 4A-1), doliform or subquadrate papil- lae, not five subquadrate or subglobose pa- pillae like those of 7. joanae (Goeden 2001b) and T. footei (Goeden 2002), nor four, doliform papillae like 7. teerinki (Goe- den 2001c). The lateral spiracular complex- es of the meso- and metathorax (Fig. 4A) and Al (Fig. 4C) are the first described to date for a second instar Tephritis. Likewise, the position and number of stelex sensilla surrounding the posterior spiracular plate are recorded here for the first time for a second instar Tephritis. Noteworthy, too, is the pair of dorsal stelex sensilla (Fig. 4E- 2), not reported in any other tephritine sec- ond instar. These dorsal sensilla also are present in the third instar of 7. stigmatica (see below, Fig. 7A), in which they are ver- ruciform, not stelex sensilla. Only minor differences in the number of branches on the interspiracular processes, 1.e., one to two for 7. footei (Goeden 2002), one to three for 7. teerinki (Goeden 2001c), versus two to four for 7. stigmatica (Fig. 4F-2) and T. joanae (Goeden 2001b), were noted. Also, the branches of the interspiracular processes of 7. stigmatica are foliose (Fig. 4F-2), like those of the second instars of T. Joanae (Goeden 2001b) and 7. teerinki (Goeden 2001c), not lanceolate, like those of T. footei (Goeden 2002). Third instar larva: White, ellipsoidal, distinctly segmented, tapered anteriorly, truncated posteriorly (Fig. 5A); gnathoce- phalon conical, anteriorly flattened, and me- dially cleaved by a vertical suture (Fig. 5B- 1); posteriorly directed, spinose, minute acanthae sparse on dorsal margin and ven- tral margin of gnathocephalon (Figs. 5B-2, VOLUME 104, NUMBER 2 Fig. 4. Second instar of Tephritis stigmatica, continued: (A) lateral spiracular complex of metathorax, 1— anterior spiracle, 2—metathoracic lateral spiracle, 3—verruciform sensilla; (B) metathoracic lateral spiracle; (C) lateral spiracular complex of first abdominal segment, anterior to right, 1—spiracle, 2—-verruciform sensilla; (D) lateral spiracle on first abdominal segment; (E) caudal segment, 1—posterior spiracular plate, 2—dorsal stelex sensilla, 3—dorsolateral stelex sensillum, 4—lateral stelex sensillum, 5—ventrolateral stelex sensilla, 6—1inter- mediate sensory complex; (F) posterior spiracular plate, |—rimae, 2—interspiracular processes, 3—intermediate complexes. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Third instar of Tephritis stigmatica: (A) habitus, anterior to left; (B) gnathocephalon, frontolateral view, |—-vertical, medial suture of gnathocephalon, 2—minute acanthae, 3—verruciform sensilla, 4—median oral lobe; (C) gnathocephalon, frontolateral view, 1—minute acanthae, 2—dorsal sensory organ, 3—anterior sensory lobe, 4—terminal sensory organ, 5—lateral sensory organ, 6—supralateral sensory organ, 7—pit sensory organ, 8—medial integumental petals, 9—lateral, spatulate, integumental petals, 10—lateral, papillate, integu- mental petals, |1—central, papilliate, integumental petal, 12—oral ridges, 13—mouthhook, 14—median oral lobe; (D) 1—dorsal sensory organ, 2—anterior sensory lobe, 3—terminal sensory organ, 4—lateral sensory organ, 5—supralateral sensory organ, 6—pit sensory organ, 7—lateral, spatulate, integumental petals, 8—oral ridges; (E) anterior spiracle; (F) mesothoracic and metathoracic, lateral spiracular complexes, 1—anterior spi- racle, 2—verruciform sensilla on mesothorax, 3—metathoracic spiracle, 4—verruciform sensilla on metathorax. VOLUME 104, NUMBER 2 C-1) and in bands that circumscribe inter- segmental areas of thorax and abdomen, bands widest ventrally and laterally be- tween segments A3 and A6; verruciform sensilla circumscribe the prothorax medi- ally (Fig. 5B-3); dorsal sensory organ well- defined, domed (Figs. 5C-2, D-1); anterior sensory lobe (Figs. 5C-3, D-2) bears ter- minal sensory organ (Figs. 5C-4, D-3), lat- eral sensory organ (Figs. 5C-5, D-4), su- pralateral sensory organ (Figs. 5C-6, D-5), and pit sensory organ (Figs. 5C-7, D-6); three or four, medial integumental petals (Fig. 5C-8) and two, lateral, spatulate (Figs. 5C-9, D-7) and two, lateral, papillate, in- tegumental petals (Fig. 5C-10) in two rows of two above each mouthhook, plus a cen- tral pair of papillate petals (Fig. 5—11), the lower, lateral, spatulate petal (Fig. 5C-9) separate from stomal sense organ (present, but not shown), ventrolaterad of anterior sensory lobe; at least three complete oral ridges (Figs. 5C-12, D-8) laterad of each anterior sensory lobe; mouthhook (Fig. 5C- 13) at least bidentate; median oral lobe (Figs. S5B-4, C-14) laterally compressed, apically pointed; anterior thoracic spiracle (Figs. SE, F-1) on posterior margin of pro- thorax bears four subglobose or subquad- rate papillae; mesothoracic, lateral spiracu- lar complexes with three verruciform sen- silla (Fig. 5F-2) in vertical row, mesotho- racic spiracle not seen; metathoracic lateral spiracular complex with well-defined spi- racle, on an oval, raised peritreme (Figs. 5F-3, 6A-1, B-1, C) and three verruciform sensilla (Figs. 5F-4, 6A-2) in vertical series posterior of spiracle; lateral spiracular com- plex of first abdominal segment (A1) (Fig. 6D) with well-defined spiracle on a circular, raised peritreme (Figs. 6A-3, D-1, E-1, F) anteriorad of two verruciform sensilla (Fig. 6E-2), one vertical to the other; caudal seg- ment (Fig. 7A) bears posterior spiracular plates (Figs. 7A-1; B) surrounded by pair of dorsal verruciform sensilla (Figs. 7A-2, C), pair of dorsolateral stelex sensilla (Figs. 7A-3, D), pair of lateral verruciform sen- silla (Figs. 7A-4, E), and ventrolateral pair 343 of stelex sensilla (Fig. 7A-5); each posterior spiracular plate bears three ovoid rimae (Fig. 7B-1), ca. 0.06 mm in length, and four, five to six-branched, single-, bi-, or trifurcately-tipped, interspiracular process- es, the longest ca. 0.03 mm long (Fig. 7B- 2); intermediate sensory complex (Fig. 7A- 6, F) with a stelex sensillum (Fig. 7F-1) and a medusoid sensillum (Fig. 7F-2). The habitus of the third instar of 7. stig- matica differs from those of four other de- scribed congeners in at least two ways. The ellipsoidal shape of the third instar (Fig. 5A) is closest to the elongate-ellipsoidal shape of the third instar of 7. footei (Goe- den 2002), both of which appear interme- diate to the ovoidal shape of the third in- stars of 7. joanae (Goeden 2001b) and T. teerinki (Goeden 2001c) and the cylindrical shape ascribed to third instars of 7. bac- charis (Goeden and Headrick 1991) and 7. arizonaensis (Goeden et al. 1993). The pro- thorax (Fig. 5B) is much smoother than that of 7. footei (Goeden 2002), that uniquely is circumscribed by many integumental petals, and thus, is much more like the prothoracic segments of the four other species of Te- phritis examined to date (Goeden and Headrick 1991; Goeden et al. 1993; Goeden 2001b, c). Few minute acanthae anteriorly circumscribe the prothorax of the third in- stars of 7. stigmatica (Figs. 5B-2, C-1) like that of 7. footei (Goeden 2002), rather than those of 7. baccharis (Goeden and Head- rick 1991), 7. arizonaensis (Goeden et al. 1993)1) joanae (Gocden 2001b) and i teerinki (Goeden 200Ic). On the other hand, the gnathocephalon, or at least the an- terior sensory lobes of all six species are separated by a vertical medial suture (Goe- den and Headrick 1991; Goeden et al. 1993; Goeden 2001b, c, 2002). The integumental petals in the third in- stars of all six congeners examined to date are arranged in a double row above each mouthhook, but those of TJ. stigmatica (Figs. 5C-9, 10) occur in two rows of two like 7. teerinki (Goeden 200Ic) and 7. foot- ei (Goeden 2002), with an additional, me- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Third instar of Tephritis stigmatica, continued: (A) lateral spiracular complex of metathorax, 1— metathoracic, lateral spiracle, 2—verruciform sensilla, 3—lateral spiracle on Al; (B) 1—metathoracic, lateral spiracle, 2—verruciform sensillum; (C) metathoracic, lateral spiracle; (D) lateral spiracular complex of first abdominal segement, 1—lateral spiracle on Al, 2—verruciform sensilla; (E) 1—lateral spiracle on Al, 2— verruciform sensillum; (F) lateral spiracle on Al. dial pair papillate in shape (Fig. 5C-11). However, the integumental petals of T. teer- inki (Goeden 2001c), 7. footei (Goeden 2002), and 7. stigmatica (Fig. 5C-8) are still fewer in number than those of 7. bac- charis (Goeden and Headrick 1991), 7. ar- izonaensis (Goeden et al. 1993), and T. joanae (Goeden 2001b). Only 7. footei and VOLUME 104, NUMBER 2 ee) a Nn Fig. 7. Third instar of Tephritis stigmatica, continued: (A) caudal segment, 1—posterior spiracular plate, 2—dorsal, verruciform sensilla, 3—dorsolateral, stelex sensillum, 4—lateral, verruciform sensillum, 5—dorso- ventral, stelex sensillum, 6—intermediate sensory complexes; (B) posterior spiracular plate, 1—rimae, 2—in- terspiracular processes; (C) dorsal, verruciform sensillum; (D) broken, dorsolateral, stelex sensillum; (E) lateral, verruciform sensillum; (F) intermediate sensory complex, 1—stelex sensillum, 2—medusoid sensillum. T. teerinki apparently lack the additional, sion separating the anterior sensory lobes medial, integumental petals found in a ver- (Fig. S5C-8; Goeden and Headrick 1991; tical double row above these lateral papil- Goeden et al. 1993; Goeden 2001b, c, lae, one to each side of the medial depres- 2002). The integumental petals increase in 346 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 8. number between the last two instars of T. footei (Goeden 2002), T. joanae (Goeden 2001b), T. teerinki (Goeden 200Ic), and T. stigmatica (Figs. 3C-3, 4, D-7, 8, E-1; 5C- SaO 103 Lb). The anterior spiracle of the third instar of T. stigmatica (Figs. 5E, F-1) bore four pa- pillae, instead of four or five papillae in the second instar (Figs. 3B-4, 4A-1). In com- parison, the anterior spiracle of 7. footei (Goeden 2002) bears three or four papillae in the third instar, and one or two less than the second instar; whereas, T. joanae (Goe- den 2001b) has three papillae in the second instar, and three or four in the third instar; and T. teerinki (Goeden 2001c) has four in the second instar and five in the third instar. The lateral spiracular complexes on the mesothorax, metathorax, and A1 of third in- star 7. stigmatica (Figs. 5K 6A, D) are the most fully described among the six species of Tephritis reported to date. The lateral spiracles on the metathorax and Al are uniquely prominent in the third instar oc- curring as they do on raised peritremes (Figs. 6B-1, C, D-1, F), and thus, are not difficult to locate as they are in the second instar (Figs. 4B, D), and in most other Te- phritinae larvae examined by my coworkers and me to date. Goeden (2001c) discussed the sensilla surrounding the posterior spiracular plates Puparium of Tephritis stigmatica: (A) habitus, anterior to left, (B) anterior end, |—invagination scar, 2—anterior spiracle. of the third instars that apparently differ in number and kinds among the Tephritis spe- cies examined to date. These sensilla are unique in 7. stigmatica, which alone has a pair of dorsal verruciform sensilla (Fig. 7A- 2) and a pair of lateral stelex sensilla (Figs 7TA-4, D) in addition to dorsolateral and ventrolateral pairs of verruciform sensilla, like those that surround the caudal seg- ments of the third instars of 7. footei (Goe- den 2002), 7. joanae (Goeden 2001b), and T. teerinki (Goeden 2001c). In addition, each of these four species bears a ventral pair of intermediate spiracular complexes (Fig. 7A-6, F), each composed of a stelex sensillum (Fig. 7F-1) and a medusoid sen- sillum (Fig. 7F-2), which bears short apical papillae typical of this type of chemosen- sillum (Goeden 2001a, b, c, d; Goeden and Teerink 1999, and references therein). Thus, two pairs of dorsal sensilla surrounding the spiracular plate of the third instar are ver- ruciform (Fig. 7A-2), not all stelex sensilla, as in the second instar (Fig. 4E-2). Puparia: Dull black, ovoidal, and smoothly rounded at both ends (Fig. 8A); the anterior end bears the invagination scar (Fig. 8B-1) and anterior spiracles (Fig. 8B- 2). Twenty-one puparia averaged 4.22 + 0.06 (range, 3.65—4.6) mm in length; 1.93 + 0.025 (range, 1.75—2.15) mm in width. VOLUME 104, NUMBER 2 ACKNOWLEDGMENTS I thank Andrew C. Sanders, Curator of the Herbarium, Department of Botany and Plant Sciences, University of California, Riverside, for identifications of plants men- tioned in this paper. Krassimer Bozhilov in the Central Facility for Advanced Micros- copy and Microanalysis, University of Cal- ifornia, Riverside, greatly facilitated my scanning electron microscopy. I also am grateful to Jeff Teerink for his technical as- sistance and to David Headrick and Jeff Teerink for their helpful comments on ear- lier drafts of this paper. LITERATURE CITED Foote, R. H., EF L. Blanc, and A. L. Norrbom. 1993. Handbook of the Fruit Flies (Diptera: Tephritidae) of America North of Mexico. Cornell University Press, Ithaca, New York. Goeden, R. D. 1988. Gall formation by the capitulum- infesting fruit fly, Tephritis stigmatica (Diptera: Tephritidae). Proceedings of the Entomological Society of Washington 90: 37-43. . 1993. Analysis of known and new host re- cords for Tephritis from California, and descrip- tion of a new species, 7. joanae (Diptera: Tephri- tidae). Proceedings of the Entomological Society of Washington 95: 425—434. . 2001a. Life history and description of im- mature stages of Neaspilota footei Freidberg and Mathis (Diptera: Tephritidae) on Aster occidental- is (Nuttall) Torrey and A. Gray (Asteraceae) in southern California. Proceedings of the Entomo- logical Society of Washington 103: 191—206. . 2001b. Life history and description of im- mature stages of Tephritis joanae Goeden (Dip- tera: Tephritidae) on Ericameria pinifolia (A. Gray) H. M. Hall (Asteraceae) in southern Cali- fornia. Proceedings of the Entomological Society of Washington 103: 586—600. . 2001c. Life history and description of im- 347 mature stages of Tephritis teerinki Goeden (Dip- tera: Tephritidae) on Hulsea vestita A. Gray (As- teraceae) in southern California. Proceedings of the Entomological Society of Washington 103: 807-825. . 2002. Descriptions of Tephritis footei and T. headricki, new species (Diptera: Tephritidae), with notes on their life histories in southern California. Proceedings of the Entomological Society of Washington 104: 142-159. Goeden, R. D. and D. H. Headrick. 1991. Life history and descriptions of immature stages of Tephritis baccharis (Coquillett) on Baccharis salicifolia (Ruiz & Parvon) Persoon in southern California (Diptera: Tephritidae). Pan-Pacific Entomologist 67: 86-98. Goeden, R. D., D. H. Headrick, and J. A. Teerink. 1993. Life history and descriptions of immature stages of Tephritis arizonaensis Quisenberry (Diptera: Tephritidae) on Baccharis sarothroides Gray in southern California. Proceedings of the Entomological Society of Washington 95: 210— 222% Goeden, R. D. and J. E Teerink. 1999. Life history and description of immature stages of Trupanea vicina (Wulp) (Diptera: Tephritidae) on wild and culti- vated Asteraceae in southern California. Proceed- ings of the Entomological Society of Washington 101: 742-755. Hickman, J. C. (ed.) 1993. The Jepson Manual. Uni- versity of California Press, Berkeley and Los An- geles. Jenkins, J. and W. J. Turner. 1989. Revision of the Baccharis-infesting (Asteraceae) fruit flies of the genus Tephritis (Diptera: Tephritidae) in North America. Annals of the Entomological Society of America 82: 674-685. Tauber, M. J. and C. A. Toschi. 1965. Life history and mating behavior of Tephritis stigmatica (Coquil- lett) (Diptera: Tephritidae). Pan-Pacific Entomol- ogist 41: 73-79. Teerink, J. A. and R. D. Goeden. 1999. Description of the immature stages of Trupanea imperfecta (Co- quillett). Proceedings of the Entomological Soci- ety of Washington 101: 75-85. PROC. ENTOMOL. SOC. WASH. 104(2), 2002, pp. 348-351 THE STATUS OF SOME PLESIOMORPHIC BLISSID BUGS (HETEROPTERA: LYGAEOIDEA: BLISSIDAE) FROM THE ORIENT JAMES A. SLATER Department of Ecology & Evolutionary Biology, University of Connecticut, Storrs, CT 06269, U.S.A. (e-mail: lygslat@ galaxyinternet.net) Abstract.—Five species of Oriental Blissidae are removed from the genus /[schnodemus and placed in the genus Capodemus. New combinations are Capodemus ambiguus (Slater, Ashlock, and Wilcox 1969), C. fumidus (Slater, Ashlock, and Wilcox 1969), C. nigroce- phalus (Slater, Ashlock, and Wilcox 1969), C. sinuatus (Slater, Ashlock, and Wilcox 1969), and C. thoracicus (Distant 1909). Capodemus nepalensis, n. sp., is described from Nepal. A discussion of the zoogeographic and phylogenetic relationships is included as are two dorsal view illustrations. Key Words: For many years I have been perplexed by the relationships of a group of species found in the Orient. As early as 1969 in a joint paper, the late P. D. Ashlock, D. Wil- cox and I described several species from Thailand and “Indo-China” and_ placed them in the genus /schnodemus. We em- phasized that they were not closely related to other species of that genus and were placed there more for convenience until re- lationships could be better understood to avoid creating another generic entity. The chief problem presented by these species was that the condition of the fore coxal cavities, which Slater (1979) used ex- tensively in his phylogenetic analyses, ap- pears to break down among what otherwise seems to be a closely related group of spe- cies. Slater and Ashlock (1976) had previ- ously discussed this coxal cavity feature and pointed out that it was not correlated with body length, but that several condi- tions existed (see Slater 1979 for a discus- sion). The primary differentiating character was whether the fore coxal cavities were open (as they are in most lygaeoids) or closed as they are in many blissid genera. Blissidae, Lygaeoidea, Oriental, Ethiopian, Gondwanaland, plesiomorphic In the tiny Oriental species considered here and at present placed in /schnodemus Fieber [ambiguus Slater, Ashlock, and Wil- cox (Fig. 1); fumidus Slater, Ashlock, and Wilcox; nigrocephalus Slater, Ashlock, and Wilcox (Fig. 2); sinuatus Slater, Ashlock, and Wilcox and thoracicus (Distant)], the coxal cavities are open in ambiguus and fumidus but very narrowly closed in the other species. All of these species have the dorsal surface of the head and pronotum completely shining and non-pruinose (con- sidered to be an apomorphic condition), the scutellum completely pruinose as is the en- tire propleuron up to a line just below the lateral edge of the pronotum, and all have rounded metathoracic scent gland auricles. These species were retained in [schnodemus by Slater (1979) but with a footnote that at least ambiguus and fumidus did not belong belong there and might be more closely re- lated to the South African genus Capode- mus Slater and Sweet. In the 1980’s, the late Thomas Wood- ward sent me a series of remarkable mi- cropterous blissids from Mt. Gingera, ACT, VOLUME 104, NUMBER 2 ey E> —% Figs. 1-2. Australia. These specimens resembled Ca- podemus species in many ways, differing chiefly in having a completely pruinose dorsal body surface and especially a para- mere with a unique outer projection, which is greatly enlarged and curves across the base of the blade (see Slater 1986). Since the pruinose body is considered plesio- morphic in the blissids (Slater 1979), I de- scribed these specimens as representing a new genus, Archaedemus (Slater 1986), but noted close resemblance in many ways to Capodemus Slater and Sweet. In the same paper I described an additional genus (Pro- pinquedemus), based on a micropterous fe- male from the mountains of Burma that I had held for many years hoping for addi- tional specimens, especially a male and/or a macropter. This female was also noted as being similar to species of Capodemus, but like Archaedemus also had a completely pruinose dorsal surface of the pronotum and 349 1, Capodemus ambiguus, dorsal view. 2, C. nigrocephalus, dorsal view. scutellum, but that the head was shining and non-pruinose, except at the extreme base. This paper is an attempt to place the five Oriental species listed above relative to Ca- podemus, Archaedemus, and Propinquede- mus. | also describe an additional species of Capodemus from Nepal. Capodemus is represented in South Af- rica by a number of species. There is a marked radiation in the extreme southwest- ern Cape and the genus is not known in Africa other than from South Africa and Zimbabwe. The fore coxal cavities are nar- rowly open. Very importantly in members of this genus the pruinosity differs from species to species. The propleuron may be completely pruinose (variabilis Slater and Sweet and a number of other species), or confined to the pleural area anterior to the acetabulum. The dorsal surface of the head and pronotum are mostly shining and non- 350 pruinose but pruinosity is frequently present along the lateral area of the dorsal surface of the pronotum. Scutellar pruinosity is pre- sent, but the meson is always shining and the pruinosity is either confined to the an- terior half of the scutellum or to the lateral areas. The parameres are simple and blade- like without any enlargement or twisting of the outer lobe. Spines may be present or absent on the fore femur. See Slater and Sweet (1972) for an extended discussion of variability among the South African spe- cies. All of the Oriental species mentioned above, plus an additional undescribed spe- cies, fall within the defining characters of Capodemus, except those with very nar- rowly closed fore coxal cavities. These spe- cies thus should be removed from /schno- demus and placed in Capodemus. Such action has interesting zoogeograph- ic implications. It suggests that Capodemus as here defined is an old taxon probably of Gondwana origin. This conclusion is based upon the close relationship to Archaedemus in Australia and the essential absence of /s- chnodemus in the otherwise rich Oriental blissid fauna. In the Orient these small Ca- podemus species seem to be rare and large- ly replaced by such genera as Macropes, Cavelerius, Dimorphopterus, Iphicrates, and a number of bizarre highly modified and specialized taxa. The presence of these Oriental Capodemus gives the impression of a taxon largely supplanted by other taxa. By contrast [schnodemus is abundant and diverse in Africa with many species show- ing close relationship to Neotropical spe- cies. It is also noteworthy that African spe- cies of Capodemus are largely confined to the extreme southwestern Cape, known to be the refuge area for many otherwise re- placed taxa. Such a distribution suggests an old taxon, once more widely distributed in Africa but supplanted by other taxa and now confined largely to refugial cool areas of winter rainfall in the southwestern Cape Province. A distribution such as that of /schnode- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON mus, with strong Neotropical-Ethiopian re- lationships and only marginal Old World Tropical relationships (1.e., essential ab- sence from the Oriental Region), is not common in the Lygaeoidea and suggests not only the antiquity of the Capodemus fauna, but also the radiation of /schnodemus after the separation of western Gondwana- land. The following new combinations are thus established: Capodemus ambiguus (Slater, Ashlock, and Wilcox 1969), new combination from Ischnodemus. Capodemus fumidus (Slater, Ashlock, and Wilcox 1969), new combination from Ischnodemus. Capodemus nigrocephalus (Slater, Ashlock, and Wilcox 1969), new combination from Ischnodemus. Capodemus sinuatus (Slater, Ashlock, and Wilcox 1969), new combination from Ischnodemus. Capodemus thoracicus (Distant 1909), new combination from /schnodemus. All measurements are in millimeters. KEY TO ORIENTAL SPECIES OF CAPODEMUS 1. Fore femur with a bifid spine present .. sinuatus BEorevtemuTmMute we eee ieee eee 2, 2. Prosternal pruinosity confined to area anterior TOECORAS oe NAS Manet ee he ema Ren aw thoracicus ' Entire prosternum pruinose ............. 3 3. Scutellum shining distally on meson .. nepalensis —' Entire scutellum pruinose .............. 4 4. Head black, strongly contrasting with pale col- or of pronotum and rest of body (Fig. 2) ADS aia Real wet is Mita Leh art rstith ie eh nigrocephalus —' Head nearly unicolorous with dorsal surface of PLONOCUIMSs:.°3 (2 fe sli ihale A te Eee 2) 5. Membrane of forewing (except veins) com- pletely white or hyaline; second antennal seg- ment pale testaceous (Fig. 1) ....... ambiguus —' Membrane of forewing smoky brown with dif- fering white veins; second antennal segment darksbrown) to blacks.) ac eee eons fumidus Capodemus fumidus (Slater, Ashlock, and Wilcox) This species was described from Thai- land. I have examined three males and one VOLUME 104, NUMBER 2 female from SABAH: Tomani, 1.1X.1983 (G. E Hevel & E. Steiner) (National Mu- seum of Natural History and J. A. Slater collections). These specimens appear to be conspecific with fumidus but have the sec- ond antennal segment pale. Capodemus nepalensis Slater, new species Description.—Head, anterior two-thirds of pronotum, scutellum, pleuron and ster- num black. A conspicuous sub-apical corial macula (not reaching lateral corial margin) and a small trianguloid macula at distal end of radial vein near anterior end of apical corial margin and distal half of fourth an- tennal segment dark brown. Posterior one- third of pronotum and lateral margins of ab- dominal sterna dark yellowish tan. Legs and antennal segments pale yellow. Clavus, co- rium and membrane white, latter with veins slightly infuscated. Head and pronotum dorsally and median area distally on scu- tellum shining, non-pruinose, remainder of scutellum pruinose. Lateral surface of head and thorax (dorsally to lateral pronotal mar- gin) entirely pruinose. A pair of large ovoid black spots on mesosternum. Metathoracic scent gland dull reddish. Dorsal surface bearing numerous semi-decumbent silvery hairs. Head non-declivent, eyes not stalked but set well away from anterior pronotal mar- gin. Length head 0.46, width 0.60, intero- cular space 0.36. Lateral pronotal margins little narrowed from humeral angles to level of calli, then strongly curving mesad to an- terior margin. Length pronotum 0.62, width 0.90. Length scutellum 0.31, width 0.42. Length claval commissure 0.32. Metatho- racic scent gland auricle broadly rounded, ovoid. Fore femur moderately incrassate, mutic. Antenna slender, filiform. Length an- 351 tennal segments I 0.12, IJ 0.31, III 0.31, IV 0.42. Labium relatively short, barely reach- ing anterior area of mesosternum, latter with a distinct median furrow. Length labial segments 10:22; 10:28; 1h OsGF IV 0:22. Total body length 2.92. Type material—RHolotype: 6 NEPAL: Meglavly Terai. Chitwan Nat. Park 7.IV.86 H. Brailovsky. In National Museum of Nat- ural History, Smithsonian Institution, Washington, DC. Paratypes: 2 5, same data as holotype. In Instituto de Biologia, Univ- ersidad Nacional Autonoma de Mexico and J. A. Slater collections. ACKNOWLEDGMENTS I extend my appreciation to Dr. Harry Brailovsky (Instituto de Biologia Universi- dad Nacional Aut6noma México) for the opportunity to study material of the new species from Nepal and to Dr. Jane O’Donnell (University of Connecticut) for aid in reviewing the manuscript. LITERATURE CITED Slater, J. A. 1979. The systematics, phylogeny, and zoogeography of the Blissinae of the world (He- miptera, Lygaeidae). Bulletin of the American Museum of Natural History 165: 1—80. . 1986. Two new genera and species of ple- siomorphic Blissinae from Australia and South- east Asia (Hemiptera: Lygaeidae: Blissinae). Jour- nal of the Kansas Entomological Society 59: 628— 634. Slater, J. A. and P. D. Ashlock. 1976. The phylogenetic position of Praetorblissus Slater with the descrip- tion of two new species (Hemiptera: Lygaeidae). Journal of the Kansas Entomological Society 49: 567-579. Slater, J. A., P. D. Ashlock, and D. Wilcox. 1969. The Blissinae of Thailand and Indochina (Hemiptera: Lygaeidae). Pacific Insects 11: 671-733. Slater, J. A. and M. H. Sweet. 1972. Capodemus, a new genus of Blissinae from South Africa with the description of twelve new species (Hemiptera: Lygaeidae). Journal of the Entomological Society of Southern Africa 35: 211—234. PROC. ENTOMOL. SOC. WASH. 104(2), 2002, pp. 352-362 ADDITIONS TO PHYLLOPHAGA SUBGENUS CHLAENOBIA (COLEOPTERA: MELOLONTHIDAE: MELOLONTHINAE) FROM GUATEMALA ENIO CANO AND MIGUEL ANGEL MORON (EC) Laboratorio de Entomologia Sistematica, Universidad del Valle de Guatemala, Apartado Postal 82, Guatemala 01901, Guatemala; (MAM) Departamento de Entomolo- gia, Instituto de Ecologia, A.C. (SEP-CONACYT) Apartado Postal 63, Xalapa, Veracruz 91000, México (e-mail: moron-_ma@ecologia.edu.mx) Abstract.—Three new species of Phyllophaga (Chlaenobia) are described from eight Guatemalan localities: P. fraternaria from cloud forests and coffee plantations at 1,320— 1,800 m in the departments of San Marcos and Quetzaltenango; P. schusteriana from pine-oak forests at 1,500 m in the departments of Alta and Baja Verapaz; and P. maril- ucasana from oak forests at 1,600—2,100 m in the departments of Santa Rosa, Sacate- pequez, and Solola. Drawings of male genital capsules, female genital plates, metatibiae, and tarsal claws are provided. Phyllophaga (Chlaenobia) aegrota (Bates) and P. (C.) scabripyga (Bates) are reported for the first time in Guatemala. Resumen.—Se describen tres especies nuevas de Phyllophaga (Chlaenobia) represen- tadas por 34 ejemplares procedentes de ocho localidades guatemaltecas: P. fraternaria de un bosque nebular y plantaciones de cafeto situados entre 1,320 y 1,800 m en los depar- tamentos de San Marcos y Quetzaltenango; P. schusteriana de bosques de pino-encino situados a 1,500 m en los departamentos de Alta y Baja Verapaz, y P. marilucasana de bosques de encinos ubicados entre 1,600—2,100 m en los departamentos de Santa Rosa, Sacatepequez y Solola. Se incluyen ilustraciones de las capsulas genitales masculinas, de las placas genitales femeninas, de las metatibias y de las unas tarsales. También se regis- tran por primera vez para Guatemala a P. (Chlaenobia) aegrota (Bates) and P. (C.) sca- bripyga (Bates). Key Words: Phyllophaga, May beetles, new species, montane forests, Guatemala Species of the Phyllophaga subgenus Chlaenobia have the following combination of characters: body elongate, slender; dorsal tegument shiny, yellowish, glabrous; head without transverse carina on the vertex; me- sotibiae with one complete oblique carina at the middle of external surface; tarsome- res depressed, elongate, usually with abun- dant setae on ventral side; tarsal claws bifid o narrowly bifurcated; sexual dimorphism usually noticeable in the form and surface of the pygidium, shape and lenght of tibial spurs and width of tarsomeres. At present, the subgenus consists of 18 species distrib- uted between Texas and Panama (Chapin 1935; Moron 1986, 1992). Blackwelder (1944) listed only Phyllo- phaga (Chlaenobia) rodriguezi (Bates) and P. (C.) tumulosa (Bates) from Guatemala. Field work between 1994—1997 led to the discovery of P. (C.) aequata chiapensis (Chapin), P. (C.) latipes (Bates), P. (C.) ve- xata (Horn), and two undescribed species of this subgenus in Guatemala (Cano and VOLUME 104, NUMBER 2 Mor6n 1998). Recently, E. Cano and J. Monz6n obtained samples of two species not known from Guatemala, another undes- cribed species from departments of Alta Ve- rapaz and Santa Rosa, and many new lo- cality records of all the other species except P. (C.) rodriguezi, confirming that the sub- genus is represented in Guatemala by at least ten species. Geographic distribution of six of these species range between southern United States, Mexico, Guatemala and Cos- ta Rica, but the other four appear to be re- stricted to the Guatemalan mountains. In support of current studies of Phyllophaga (Chlaenobia) by M. A. Moron, we describe three additional species from Guatemala. Two new country records are also present- ed. The terms used in the descriptions are those of Moré6n (1986, 1992). Drawings were made with the aid of a camera lucida and stereomicroscope; measurements were obtained with an ocular micrometer or cal- iper. Acronyms used in the text are as fol- lows: UVGC = Universidad del Valle de Guatemala; UNSM = University of Ne- braska State Museum, Lincoln; and MXAL = private collection, M. A. Moron, Xalapa, México. Phyllophaga (Chlaenobia) fraternaria Cano and Moré6on, new species (Figs. 1-7) Description.—Holotype male: Clypeus yellowish brown, disk reddish brown, an- terior border darker, frons reddish brown, vertex darker; pronotum shiny yellowish, borders reddish; elytra straw yellowish without macroscopic vestiture, glabrous; mouthparts, sterna, pygidium and legs shiny straw yellowish. Clypeus 2.94 wid- er than long, anterior border sinuate, with elevated margin, surface glabrous, slightly irregular, almost plain, with many uniform- ly distributed, deep, rugose punctures. Frontoclypeal suture sinuate and deeply im- pressed. Frons 1.92 wider than long, gla- brous, convex to plain, regularly and deep- ly, rugo-punctate. Vertex without punctures. ey) Nn o>) Antenna 10-segmented, with 3-segmented club, lamellae narrower than length of basal segments combined (0.86:1). Frons 2.94 wider than dorsal diameter of each eye. Eye canthus long and narrow, with 7-8 setae. Labrum bilobed, deeply sinuate, with scat- tered long, slender setae on borders. Men- tum slightly concave, impunctate, with scarce lateral setae, anterior border sinuate. Pronotum 1.74 wider than long and 2.35% wider than frons. Pronotal disk shiny, with round, moderately deep punc- tures separated by less of one diameter, forming rugo-punctate texture; lateral bor- ders widely angulated, lateral marginal bead with scattered, short or long, slender setae; basal bead strongly indicated, except in front of scutellum where it is indicated only by scarce punctures; anterior angles obtuse, not prominent; posterior angles not directed downward, obtuse. Scutellum 1.45 wider than long, with some minute punctures. Elytron 2.65 longer than wide, shiny, glabrous, densely rugo-punctate; punctures small, slightly deep, separated by 1—2 diameters; epipleural border very wide at base, narrow along complete margin, with some scattered, short, slender setae; humeral callus rounded, prominent; apical callus rounded. Metathoracic wings com- pletely developed. Propygidium shiny, yel- lowish with scattered, transversal punctua- tion and many short decumbent setae to- wards basal border. Pygidium convex, shiny, strongly rugose, but without visible punctures, with rounded-conical promi- nence located toward middle of preapical border, basal border without setae; apical margin with 7 long, slender setae; basal margin complete. Pterosternum with mod- erately dense, long yellowish setae. Visible abdominal sternites II to IV of similar length, slightly concave, with scattered, mi- nute setae at middle; sternite V with wide transverse concavity, almost plain toward midline, with 2 procumbent, short setae near middle; posterior border with marginal bead clearly impressed. Anal plate short, slightly concave at midline, posterior bor- 354 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-7. Phyllophaga (Chlaenobia) fraternaria. 1, Male protarsal claw, lateral view. 2, Male metatarsal claw, lateral view. 3, Genital capsule, lateral view. 4, Apex of aedeagus, dorsolateral view. 5, Parameres, distal view. 6, Female protarsal claw, lateral view. 7, Female genital plates, ventral view. Scale lines = 1 mm, except figs. 1, 2, 6 = 0.5 mm. der thickened, poorly crenulate at sides of midline, with 10 long and short erect setae along complete border. Protibia shorter than protarsus (0.56:1), with external border tri- dentate, proximal tooth much shortened, preapical spur narrow, nearly straight, apex acute, half as long as 2nd protarsomere. Mesotibia with one oblique, sharp, setife- rous carina on external side; upper apical spur straight, narrow, and 1.33 longer VOLUME 104, NUMBER 2 than lower spur. Metatibia shorter than metatarsus (0.63:1), with one oblique, sharp, setiferous carina on external side; up- per apical spur articulated, curved, apex acute, slightly shorter than basal metatar- somere, and 1.45 longer than lower spur; lower apical spur articulated with tibial bor- der, with acute apex. Tarsomeres depressed, elongate, with enlarged apices, with some setae apically and two lines of long, thick setae on flattened ventral side. Protarsal claws narrowly cleft, with lower tooth shorter than upper tooth; meso- and meta- tarsal claws with lower tooth slightly short- er than upper tooth (Figs. 1-2). Genital cap- sule with large parameres, dorsally and ven- trally fused, asymmetrical, apex of each elongated, acute, tooth-like and directed downward. Aedeagus with two internal pro- jections, one of them notably largest, and apex directed upward (Figs. 3—5). Tectum (= phallobase) shortened, with dorsal de- pression near middle. Length of genital cap- sule from apex of parameres to border of basal piece: 5.4 mm. Total body length: 17.0 mm. Humeral width: 7.3 mm. Allotype female: Similar to male except as follows: clypeus, frons and pronotum reddish brown; head with more fine punc- tures; antenna with lamellae of segments 8— 10 slightly shorter or equal to the length of six preceeding segments combined. Visible abdominal sternites II to V convex, with se- tiferous punctures near middle; anal plate convex, with 3 scattered setiferous punc- tures, and 10 slender setae at posterior bor- der. Pygidium very convex, without conical prominence, with 10 setae along apical bor- der. Protibia with teeth of external border wider and longer than in male. Both apical spurs of metatibia articulated, wide, lance- olate and curved. Protarsus longer than pro- tibia (1.17:1). Tarsal claws similar on all legs; lower tooth slightly shorter than upper (Fig. 6). Ventral genital plates moderately sclerotized, symmetrical, with scattered short setae, with shallow depression near apex, apical and lateral borders markedly serrate; dorsal genital plates with borders 359) markedly serrate, setae on distal border (Fig. 7). Total body length: 19.0 mm. Hu- meral width: 7.7 mm. Variation.—Male: Similar to holotype except as follows: Head and pronotum red- dish brown or yellowish brown; some spec- imens with reddish punctures on pronotum; mentum punctate and micropunctate or only densely micropuntate; pygidium with prea- pical process less, equal or more prominent, with more or less apical setae; abdominal sternites II-[V with more or less setae, ster- nite V with not so wide transverse concav- ity or without concavity, almost plain. Total body length: 17.2—-18.0 mm; humeral width: 7.0—7.9 mm. Female: similar to al- lotype except head and pronotum yellowish brown, disc and apical border of anal plate with more or less setae, lower tooth of tar- sal claws almost equal or slightly shorter than upper. Total body length: 17.2—18.2 mm; humeral width: 7.0—7.9 mm. Type material.—Described from 13 6, 6 2. Holotype ¢ UVGC: Guatemala: San Marcos, La Fraternidad, 1,800 m, 5/10-II- 1994, bosque nuboso, J. Monz6én’’. Allo- type 2 UVGC: Guatemala: Quetzaltenan- go, Viejo Palmar, Fca. San Juan Patzulin, 1,320 m, 21-III-1998, I. Chavez. Paratypes: same data as holotype (146) (MXAL); same data except 6-III-1994 (26) (UVGC); ex- cept 25/29-III-1994, 1850 m (16d, 22) (MXAL); except 15/17 IV 1994 (76, 3@) (UVGC); Guatemala: San Marcos, El Tum- bador, Fea. Australia, 1,510 m, 4-IV-1998, J. Amau (16) (UVGC). Type locality—La Fraternidad, depart- ment of San Marcos, Guatemala (aprox. 14°55'N; 91°50'W). Biological data.—Males and females of P. fraternaria were collected at UV and Hg lights in median altitude cloud forests and coffee plantations, at 1,320—1,950 m alti- tude. Phenology: February (2), March (6), April (11). Remarks.—Phyllophaga (Chlaenobia) fraternaria belong to the species group “vexata” (sensu Moron 1986). By body shape, size, color, general vestiture, punc- 356 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tuation on the clypeus, pronotum and elytra, as well as the general shape of the male genital capsule, it is similar to P. (Chiro- dines) zunilensis (Bates), P. (Chlaenobia) solanophaga Moron, and P. (Chlaenobia) ratcliffeiana Moron. However the detailed structure and proportions of the parameres and aedeagus are clearly different in all of those species. Also, the narrowly cleft hind tarsal claws separate P. fraternaria from P. zunilensis and the coarsely rugose pigydium and its preapical prominence separate it from P. solanophaga and P. ratcliffeiana. Etymology.—The species name is de- rived from the type locality, La Fraternidad, a town near the Mexican border. Phyllophaga (Chlaenobia) schusteriana Cano and Moron, new species (Figs. 8-14) Description.—Holotype male: Clypeus reddish brown, disk reddish brown, anterior border darker, frons and vertex reddish brown; pronotum shiny straw yellowish, borders reddish; elytra straw yellowish; mouthparts reddish brown, sterna, pygidi- um and legs shiny straw yellowish. Clypeus 3.04 wider than long, anterior border sin- uate, with elevated margin, surface with mi- croscopic vestiture, slightly convex, with many uniformly distributed, close, deep, al- most circular, rugose punctures. Frontocly- peal suture sinuate and deeply impressed. Frons 2.09 wider than long, convex, reg- ularly and deeply, rugo-punctate. Vertex with few small punctures. Antenna 10-seg- mented, with long 3-segmented club, la- mellae slightly short or equal to length of basal segments combined (0.99:1). Frons 3.61 wider than dorsal diameter of each eye, with microscopic vestiture. Eye can- thus long and narrow, with 12—13 setae. La- brum bilobed, deeply sinuate, with scattered long, slender setae on borders. Mentum concave, impunctate, with scarce lateral se- tae, anterior border sinuate. Pronotum 1.94 wider than long and 2.4 wider than frons. Pronotal disk shiny, with microscopic vestiture and round, slightly deep, rugose punctures regularly separated by 1-2 di- ameters, more dense and rugose toward lat- eral borders; lateral borders widely angu- lated, lateral marginal bead slightly crenu- late, with scattered, short or long, slender setae; basal bead complete, strongly indi- cated; anterior angles obtuse, not promi- nent; posterior angles not directed down- ward, obtuse. Scutellum 1.37 wider than long, with some minute punctures. Elytron 2.82 longer than wide, shiny, with micro- scopic vestiture, densely rugo-punctate; punctures small and shallow, separated by |—2 diameters; 2nd elytral stria prominent, well marked along it complete length, ex- cept at basis; epipleural border very wide at base, narrow along complete margin, with some scattered, long, slender setae; humeral callus rounded, prominent; apical callus rounded. Metathoracic wings com- pletely developed. Propygidium shiny, yel- lowish with scattered, minute punctuation and minute decumbent and procumbent se- tae. Pygidium triangular, convex, shiny, rugo-punctate, with shallow punctures reg- ularly distributed, separated by 1—2 diame- ters, basal border without setae; apical mar- gin almost straight with 16 long, slender se- tae, preceeding by transverse depression transversely micro-rugose; basal margin ef- faced medially. Pterosternum with moder- ately dense, long yellowish setae. Visible abdominal sternites II to V_ of similar length, very concave, with microscopic se- tae in middle and a few more long setae intermixed, more abundant on sternite II; posterior border with marginal bead clearly impressed. Anal plate long, slightly con- cave, posterior border thickened, notched at midline, with a few long, erect setae along complete border. Protibia shorter than pro- tarsus (0.49:1), with external border triden- tate, proximal tooth well marked but much shortened, preapical spur narrow, straight, apex acute, half as long as 2nd protarso- mere. Mesotibia with one oblique, sharp, setiferous carina on external side; upper apical spur straight, narrow, and 1.2 lon- ger than lower spur. Metatibia equal to the VOLUME 104, NUMBER 2 357 ae) Figs. 8-14. Phyllophaga (Chlaenobia) schusteriana. 8, Apex of male metatibia, latero-distal view. 9, Male protarsal claw, lateral view. 10, Genital capsule, lateral view. 11, Parameres, distal view. 12, Apex of aedeagus, dorsolateral view. 13, Apex of female metatibia, laterodistal view. 14, Female genital plates, ventral view. Scale lines = | mm, except fig. 9 = 0.5 mm. length of metatarsus, without carina on ex- ternal side, laterally flattened on ventral half and clearly widened toward the apex (Fig. 8); upper apical spur articulated, very curved, apex truncated and wide, slightly shorter than basal metatarsomere, and 1.14 longer than lower spur; lower apical spur articulated with tibial border, with bas- al %4 wide and apical % narrowed with apex truncated. Tarsomeres depressed, elongate, with enlarged apices, with some setae api- cally and two lines of long, thick setae on flattened ventral side. All tarsal claws sim- ilar, narrowly cleft, with lower tooth longer and wider than upper tooth (Fig. 9). Genital capsule with parameres dorsally fused, ven- trally notched, symmetrical, apex of each elongated, wide, and directed inward; mar- gin of apical area covered with abundant, golden short setae, posteriorly more long; dorsal area bare, ventral area with numer- ous golden, decumbent, short setae. Aedea- gus widely sclerotized forming a deeply channeled structure, with two dorsal, claw- shaped, laterally flattened, sclerotized pro- jections fused basally (Figs. 10-12). Tec- 358 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tum large, longitudinally sulcated in apical half. Length of genital capsule from apex of parameres to border of basal piece: 4.9 mm. Total body length: 23.0 mm. Humeral width: 9.3 mm. Allotype female: Similar to male except as follows: clypeus, frons and pronotum reddish brown; head with more fine punc- tures; antenna with lamellae of 8th to 10th segments slightly longer or equal to length of six preceeding segments com- bined. Visible abdominal sternites II to IV convex, V sternite large, transversely con- cave; anal plate short, convex, with various scattered setiferous punctures and few slen- der setae at posterior border. Pygidium con- vex, apical depression more deep and wide than male, apical border rounded, with 15 setae along apical border. Protibia with teeth of external border wider and longer than in male. Metatibia not flattened, with one oblique, sharp, setiferous carina on ex- ternal side; dorsal border serrated; apical spurs articulated, similar length, lanceolate, upper straight, lower slightly curved (Fig. 13). Protarsus longer than protibia (1.42:1). Ventral genital plates moderately sclero- tized, symmetrical, dorsal genital plates fused with central, longitudinal keel, and border bisinuated, centrally notched, with 8—10 setae on each side near border (Fig. 14). Total body length: 22.0 mm. Humeral width: 9.5 mm. Variation.—Male: Similar to holotype except with more setae on abdominal ster- nites II-V; pygidium more strongly longi- tudinally rugose; pronotum more or less punctured; elytra more or less densely punctured; total body length: 21.5—22.0 mm; humeral width: 9.0—9.3 mm. Type material.—Described from 7 ¢, 1 2. Holotype ¢ UVGC: Guatemala: Alta Verapaz, Purulha, camino a Pantin, V-1995, J. Monz6n. Allotype 2 UVGC: same data as holotype. Paratypes: same data as holo- type (1d) (MXAL); Guatemala: Baja Ve- rapaz, 19-24 km N Salama, 1,500 m, 25/ S1=V=1989,«J:, Ea Wappées Ged) /(UNGE MXAL, UNSM). Type locality—Old road between Sala- ma and Purulha, between departments of Alta Verapaz and Baja Verapaz, Guatemala (approx. 15°11'N; 90°18’W). Biological data.—Males and female of P. schusteriana were collected at UV and Hg lights in pine-oak forests with Liguidambar, at 1,500 m altitude. Phenology: May (8). Remarks.—Phyllophaga (Chlaenobia) schusteriana is tentatively placed in the species group “‘vexata’’? (sensu Moron 1986). By body size, general vestiture, punctuation on the clypeus, pronotum, and elytra, it differs from other species of Chlaenobia. The structure and proportions of the parameres and aedeagus are clearly different from other species in the subge- nus, and have some similarity with species in the subgenus Phytalus, such as P. gua- temala Saylor. Also, the flattened shape of male metatibia of P. schusteriana is unique among the species of Phyllophaga (s. lato) with cleft tarsal claws. Etymology.—This interesting species is named in honor of Dr. Jack C. Schuster, well known coleopterist, rock-folk musi- cian, and acclaimed professor of biology. Phyllophaga (Chlaenobia) marilucasana Cano and Moron, new species (Figs. 15-21) Description.—Holotype male: Clypeus yellowish testaceous, anterior border darker, frons and vertex reddish brown, frontocly- peal suture yellowish; pronotum shiny straw yellowish; elytra shiny straw yellow- ish without macroscopic vestiture; mouth- parts, sterna, pygidium and legs shiny yel- lowish. Clypeus 2.87 wider than long, an- terior border widely sinuate, with elevated margin, surface shiny, slightly irregular, slightly convex, with many uniformly dis- tributed, shallow, rugose punctures, sepa- rated by 1 or less than Idiameter, each puncture with microscopic seta. Frontocly- peal suture sinuate and deeply impressed. Frons 1.80 wider than long, convex, with punctuation similar to clypeus, but deeper and concentrated toward posterior border. VOLUME 104, NUMBER 2 20 Figs. 15-21. 359 21 Phyllophaga (Chlaenobia) marilucasana. 15, Male protarsal claw, lateral view. 16, Male meta- tarsal claw, lateral view. 17, Genital capsule, lateral view. 18, Parameres, distal view. 19, Apex of aedeagus, laterodistal view. 20, Female protarsal claw, lateral view. 21, Female genital plates, ventral view. Scale lines = 1 mm, except figs. 15-16, 20 = 0.5 mm. Vertex with small punctures on lateral bor- der. Antenna 10-segmented, with 3-seg- mented club, lamellae narrower than length of basal segments combined (0.77:1). Frons 3.4 wider than dorsal diameter of each eye. Eye canthus long and narrow, with 8— 9 setae. Labrum bilobed, deeply sinuate, with scattered long, slender setae on bor- ders and across antero-superior border. Mentum slightly concave, impunctate, with scarce lateral setae, anterior border sinuate. Pronotum 1.92 wider than long and 2.6 wider than frons. Pronotal disk shiny, with round, shallow punctures, each with micro- scopic setae, scarcely separated, more abun- dant and with rugose aspect towards lateral and anterior margins; lateral borders widely angulated, lateral marginal bead almost en- tire, with scattered, short or long, slender setae; basal bead strongly indicated, except in front of scutellum indicated only by scarce punctures; anterior angles obtuse, not prominent; posterior angles not directed downward, obtuse. Scutellum 1.3 wider than long, with some minute punctures and 5 long, procumbent apical setae. Elytron 3.17 longer than wide, shiny, under great magnification tegument micro-rugose, near- 360 ly chagrinate, under low magnification sur- face densely rugo-punctate; punctures small and shallow, each with minute seta, sepa- rated by 1—2 diameters; epipleural border very wide at base, narrow along complete margin, with some scattered, short, slender setae; humeral callus rounded, prominent; apical callus rounded. Metathoracic wings completely developed. Propygidium shiny, yellowish with scattered, transversal punc- tuation and many minute decumbent and erect setae towards basal border. Pygidium very convex, shiny, strongly rugose, with visible some scattered shallow punctures regularly distributed, with rounded-conical prominence located near middle of preapi- cal border, basal border without setae; api- cal margin with 13 long, slender setae; bas- al margin complete, preceeding by a shal- low, transverse sulcus. Pterosternum with moderately dense, long yellowish setae. Visible abdominal sternites II to IV of sim- ilar length, slightly concave, minutely se- tose at middle; sternite V with wide trans- verse posterior sulcus, anteriorly with 4 long, slender setae near the middle; poste- rior border with marginal bead effaced at middle. Anal plate short, slightly concave at midline, posterior border thickened, notched at midline, with a few long erect setae along complete border. Protibia short- er than protarsus (0.72:1), with external border tridentate, proximal tooth poorly de- veloped, almost imperceptible, preapical spur narrow, nearly straight, apex acute, half as long as 2nd protarsomere. Mesotibia with one oblique, sharp, setiferous carina on external side; upper apical spur straight, narrow, and 1.47 longer than lower spur. Metatibia shorter than metatarsus (0.59:1), with one oblique, sharp, setiferous carina on external side; upper apical spur articu- lated, curved, apex rounded to acute, slight- ly shorter than basal metatarsomere, and 1.47 longer than lower spur; lower apical spur articulated with tibial border, curved with rounded apex. Tarsomeres depressed, elongate, with enlarged apices, with some setae apically and two lines of long, thick PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON setae on flattened ventral side. Protarsal claws narrowly cleft, with lower tooth shorter than upper tooth; meso- and meta- tarsal claws with lower tooth slightly short- er than upper tooth (Figs. 15-16). Genital capsule with large parameres, dorsally and ventrally fused, almost symmetrical, apex of each elongated, acute, tooth-like and di- rected downward, each with one asymmet- rical laminar projection on internal face, di- rected upward. Aedeagus widely sclerotized with 2 lateral, narrow projections and lower apical, flattened, very sclerotized projec- tions (Figs. 17-19). Tectum shortened, with dorsal depression near distal third. Length of genital capsule from apex of parameres to border of basal piece: 5.5 mm. Total body length: 17.5 mm. Humeral width: 7.0 mm. Allotype female: Similar to male except as follows: antenna with lamellae of 8th to 10th segments equal to length of six pre- ceeding segments combined. Visible ab- dominal sternites IT to V convex, V sternite with setiferous punctures near middle; anal plate slightly convex, with many scattered setiferous punctures, and 11 slender setae at posterior border. Pygidium very convex, with 6 setae along apical border, without conical prominence, with 6 setae along api- cal border. Protibia with teeth of external border wider and longer than in male. Both apical spurs of metatibia articulated, wide, lanceolate and curved. Protarsus longer than protibia (1.26:1). Tarsal claws similar on all legs; lower tooth slightly longer than upper or equal in length (Fig. 20). Ventral genital plates sclerotized, nearly symmetri- cal, with complex combination of keels and sulci, surface with scattered minute setae; dorsal genital plates with a notable apical acute process with short setae on distal bor- der (Fig. 21). Total body length: 19 mm. Humeral width: 9 mm. Variation.—Male: similar to holotype except as follows: frontoclypeal suture red- dish brown, mentum with 6-8 punctures. Female: similar to allotype, except apical VOLUME 104, NUMBER 2 process of dorsal plates less acute and less notable. Type material.—Described from 3 6, 4 2. Holotype ¢ UVGC: Guatemala: Santa Rosa, Pueblo Nuevo Vifias, Finca Mira- mundo, 1,900 m, 10-VI-1999, J. Monz6n y E. Cano. Allotype 2 UVGC: same data as holotype. Paratypes: Guatemala: Sololda, Santa Maria Visitaci6n, Sierra Parraxquim, 2,100 m, 3-V-2000, bosque nuboso, J. Or- dofez (CDC-CECON), col. (2¢) (MXAL, UVGC); same data except 6-V-2000 (2°) (MXAL, UVGC); Guatemala: Sacatepe- quez, Antigua Guatemala, 1,600 m, 26-IV- 2000, A.C. Bailey y J. Monzon (12) (UVGC). Type locality—Cerro Miramundo, de- partment of Santa Rosa, Guatemala (approx. 14°12’N; 90°30'W), an extinct Quaternary strato-volcano. Biological data.—Males and females of P. marilucasana were collected at UV and Hg lights in a pine-oak forest near a coffee plantation and an oak forest with some Pi- nus, Ostrya, and Carpinus, at 1,600—2100 m altitude. Phenology: April (1), May (4), June (2). Remarks.—Phyllophaga (Chlaenobia) marilucasana belongs to the species group “vexata”’ (sensu Mor6n 1986). By body shape, size, color, general vestiture, punc- tuation on the clypeus, pronotum and elytra, as well as the general shape of male genital capsule, it is similar to P. (Chirodines) zu- nilensis, P. (Chlaenobia) solanophaga, P. (Chlaenobia) ratcliffeiana, and P. (Chlae- nobia) fraternaria, but the detailed struc- ture and proportions of the parameres and sclerotized accesories of the aedeagus are clearly different in all of these species. Also, the narrowly cleft hind tarsal claws separate it from P. zunilensis and _ the coarsely rugose pigydium and its preapical prominence separate it from P. solanopha- ga and P. ratcliffeiana. Phyllophaga (C.) marilucasana is allied to P. fraternaria, and it is difficult to separate them except by the male and female genitalia. Etymology.—This species is dedicated to 361 Maria Luisa Castillo, whose collecting ef- forts in many rotten logs have provided a large number of interesting samples of Me- lolonthidae and Passalidae. NEw COUNTRY RECORDS Phyllophaga (Chlaenobia) aegrota (Bates)—Guatemala: Huehuetenango, Ba- rillas, aldea Malpais, 1,200 m, VII-1998, bosque nuboso, E.Cano (6 specimens) (UVGC). Phyllophaga (Chlaenobia) scabripyga (Bates).—Guatemala: Sacatepequez, Es- cuintla, finca Eminencia, Volcan de Agua, 1,400 m, 4-[V-2000, bosque nuboso. J. Monzon (16 specimens) (UVGC). Guate- mala: Huehuetenango, Huehuetenango, al- dea Chivacabé, 7-VI-1997, P. Alarcén (12) (UVGC). ACKNOWLEDGMENTS Field work in Guatemala was possible by the support of project No. 2 FONACYT, Guatemala. Curatorial work in Xalapa was allowed by CONACYT, México. Both au- thors are indebted to José Monz6n (UVGC) and Brett C. Ratcliffe (Lincoln, Nebraska), who provided interesting specimens of Phyllophaga used in this study. This paper is a contribution to the project ““Sistematica y Biologia del género Phyllophaga en Méx- ico y América Central’’ (225260-5-25723- N), supported by CONACYT, México. LITERATURE CITED Blackwelder, R. E. 1944. Checklist of the Coleopter- ous insects of Mexico, Central America, the West Indies, and South America. Part 2. Smithsonian Institution United States. National Museum Bul- letin 185: 222-228. Cano, E. and M. A. Moron, 1998. Las especies de Phyllophaga (Coleoptera: Scarabaeidae: Melo- lonthinae) de Guatemala. Diversidad, distribucién e importancia, pp. 7-18. Jn Avances en el estudio de la diversidad, importancia y manejo de los co- ledpteros edaficolas americanos. Benemérita Uni- versidad Autonoma de Puebla y Sociedad Mexi- cana de Entomologia, México. Chapin, E. A. 1935. Review of the genus Chlaenobia Blanch. (Col. Scarabaeidae). Smithsonian Miscel- laneous Collection 94(9): 1—20. 362 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Moron, M. A. 1986. El género Phyllophaga en Méx- . 1992. Nuevas especies mexicanas del subgé- ico. Morfologia, Distribuci6n y Sistematica Su- nero Chlaenobia de Phyllophaga (Coleoptera: praespecifica (Insecta: Coleoptera). Publ. 20, Ins- Melolonthidae). Giornale italiano di Entomologia tituto de Ecologia, México. 341 pp. 6: 15-31. PROC. ENTOMOL. SOC. WASH. 104(2), 2002, pp. 363-372 SABETHES (PEYTONULUS) PARADOXUS, A NEW SPECIES OF SABETHINI (DIPTERA: CULICIDAE) FROM PANAMA RALPH E. HARBACH AND THERESA M. HOWARD Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, U.K. (e-mail: r.harbach@nhm.ac.uk; t.howard@nhm.ac.uk) Abstract.—The male, pupa and larva of Sabethes (Peytonulus) paradoxus Harbach, n. sp., are described and illustrated from Panama. The adult male lacks the brilliant metallic-colored scaling that is characteristic of all other Sabethes, and the paddle of the pupa is heterogeneous for the presence of a basal membranous area, a feature that has not been observed in other members of subgenus Peytonulus. Key Words: When Hall et al. (1999) described Sa- bethes luxodens, subgenus Peytonulus Harbach included 11 formally named species and at least four unnamed spe- cies. The species described in this paper is one of the four previously unnamed Species, the species which Harbach (1991) said “‘lacks metallic scutal scaling and bears certain other stasimorphic char- acters.” Species of subgenus Peytonulus are readily identified by unique apomorphies that include the remarkably modified seta 1-VII of larvae and the absence of its on- togenetic homolog in pupae. Neverthe- less, like all groups of organisms, mem- bers of the subgenus exhibit combinations of both ancestral and derived features, and the species described herein is a striking example of this. While the species bears the apomorphic characters diagnostic of the subgenus, it exhibits the plesiomorph- ic condition of the scutal scaling that is characteristic of genus Wyeomyia Theo- bald. The derived position of Sabethes Robineau-Desvoidy relative to Wyeomyia is supported by the cladistic analysis of Harbach and Kitching (1998). Diptera, Culicidae, Sabethes, Peytonulus, new species, mosquito, Panama MATERIALS AND METHODS This study is based on specimens bor- rowed from the National Museum of Nat- ural History (USNM), Smithsonian Insti- tution, Washington, DC. Observations of the adults were made under simulated nat- ural light. Larval and pupal chaetotaxy were studied using a combination of bright field and differential interference contrast microscopy. Measurements and counts were taken from all specimens of the type series where the structures in question were present. Numbers in parentheses rep- resent modes of the reported ranges unless indicated otherwise. The form of presen- tation, descriptive terminology and abbre- viations used in the description follow Harbach and Knight (1980, 1982) and re- cent papers published as part of an ongoing revision of genus Sabethes (Harbach and Peyton 1991; Harbach 1991, 1994, 1995a, 1995b; Harbach and Petersen 1992; Hall et al. 1999; Moses et al. 2000). Sabethes (Peytonulus) paradoxus Harbach, new species (Figs. 1-3) Sabethes (Sabethinus) sp 2 of Heinemann and Belkin 1978: 193. 364 This species is unique in lacking the bril- liant metallic-colored scaling that is char- acteristic of Sabethes (Harbach and Peyton 1991). The female is unknown. For the most part, the male resembles species of Wyeomyia in overall ornamentation, partic- ularly the relatively dull coloring of the scu- tal scaling. Male.—Relatively small species with broad, flat, distinctly metallic-colored scal- ing only on head, antepronota and thoracic pleura; scales of vertex with different com- binations of metallic blue, violet and green reflections depending on angle of light; scales of postgena, thoracic pleura and cox- ae silvery white; antepronotum ranging from brown to bright blue and violet de- pending on angle of view, lower margin with few silvery white scales; scutum and scutellum densely covered with moderately broad dark scales with dull bluish reflec- tions; mesopostnotum without scales; pro- boscis and legs predominantly dark-scaled with subdued blue and violet reflections when viewed from certain angles; wing with veins entirely dark-scaled, anterior veins darker with bluish sheen depending on angle of light; abdominal terga mainly dark-scaled with weak bluish, greenish and golden reflections, lateral margins with rather large rounded pale patches; sterna with yellowish-brown scaling. Head: Eyes joined above and below. Occiput with transverse row of short semi-erect scales at back of head. Ocular setae moderately long, dark, close to margin of eye; 2 long, bronzy, approximated interocular setae present. An- tenna dark; length about 1.8 mm, slightly shorter than proboscis; pedicel large, sur- face pubescent, with inconspicuous fine se- tae on dorsomesal area; flagellum rather strongly verticillate, proximal whorls with 14 setae, longest setae about 0.4 length of antenna. Clypeus and frons without setae and scales, clypeus with dense covering of silvery pubescence. Proboscis short, length about 2.0 mm, longer than antenna, about 0.8 length of forefemur; distal 0.2 of pre- mentum flattened, expanded laterally to 4 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON times width of proximal part and bent downward; ventral surface white-scaled be- ginning 0.2—0.4 from base and extending to expanded distal part, white scales grading to pale yellow at base of expanded part, dark scaling of expanded part distinctly darker (black), especially ventrally, than other dark scaling of prementum; labella short, very broad and dark-scaled proxi- mally. Maxillary palpus short, 0.12—0.15 length of proboscis; dark-scaled, ventral surface without scales. Thorax: Integument brown. Dorsum with dark setae on anterior promontory (9—12), antepronotum (10-14), supraalar area (22—24), scutellum (6,7) and mesopostnotum (9—14). Pleura with 2 or 3 prespiracular setae, 2—5 upper proepisternal setae, 6—8 long lower mesokatepisternal se- tae extending well above ventral margin of mesepimeron and 11—14 upper mesepimer- al setae; prespiracular setae dark, others yellow or golden. Lower part of proepister- num without scales, scales on upper part contiguous with scales on anteprocoxal membrane; scales absent from postprocoxal membrane; mesopleuron with scales except on lower anterior margin of mesokatepi- sternum, upper posterior margin of mese- pimeron and mesomeron; scales absent from metapleuron, metameron and _ post- metacoxal membrane. Wing: Length 3.5 mm; scales on veins moderately broad and slightly asymmetrical, smaller on cubitus and anal vein; alula with fine piliform scales on margin distally; calypters without setae. Halter: Scabellum without scales, in- tegument pale; pedicel and capitellum dark- scaled. Legs: Without paddles; coxae and trochanters with silvery-white scales, tro- chanters with some dark scales dorsally at apices; femora dark above and golden be- low; tibiae mainly dark with golden scaling proximally on ventral surface; tarsi entirely dark-scaled except ventral surface of hind- tarsomere 5 white-scaled. Forefemur about 1.3 length of proboscis, same length to slightly shorter than midfemur, about 1.3 length of hindfemur; hindtibia about as long as hindfemur, hindtarsomere | longer than VOLUME 104, NUMBER 2 iS lateral ] Howard Fig. 1. tergal 365 Male genitalia of Sabethes paradoxus, aspects as indicated. A, Gonocoxopodite. B, Gonostylus. C, Aedeagus, with parameres attached. D, Proctiger. E, Tergum IX. Abbreviations: A, E, C and M = gonostylar lobes; BML = basal mesal lobe; mp = membranous process; tms = tergomesal setae. Scale in mm. hindfemur. Ungues small, simple, black. Abdomen: Coloration as noted above; lat- eral pale areas of terga with silvery scales dorsally grading into yellow scales ventral- ly. Genitalia (Fig. 1): Tergum VIII (ventral in position; not figured) with posterolateral corners produced and broad V-shaped emargination at middle of posterior margin, border on either side of emargination with 4 or 5 irregular rows of long close-set setae extending to posterolateral corners, setae become longer and more numerous toward corners; scattered scales before and among setae; anterior 0.3—0.4 of tergum bare. Ter- gum and sternum IX fused laterally, form- ing a complete ring of sclerotization; ter- gum IX widely separated by relatively nar- row bridge, lobes slightly produced, each with 4—7(5) flattened setae with apices bent laterad. Gonocoxite elongate, tapered in PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 366 Fig. 2. Fourth-instar larva of Sabethes paradoxus. Reconstructed from exuviae of holotype; positions of setae may differ slightly from those shown. Setae missing from the right side of the exuviae (see Table 1) are drawn from those on the left side; an empty alveolus indicates that a seta was missing on both the right and left sides of the exuviae. A, Head, dorsal (left) and ventral (right) aspects of left side. B, Thorax and abdominal segments I-VI, dorsal (left) and ventral (right) aspects of left side. C, Abdominal segments VII—VIII and X, left VOLUME 104, NUMBER 2 distal half, tergomesal surface membranous, distal part of sternal surface covered with scales and setae, bearing one very long and 2 unequal shorter tergomesal setae below level of basal mesal lobe; basal mesal lobe irregular in shape, produced caudomesally, covered with small slender setae and bear- ing 2 large setae at caudolateral angle. Gon- ostylus about 0.75 length of gonocoxite; stem narrow and curved in lateral view, slightly longer than head; head with 3 lobes developed as figured; lobe A,E large, lat- erally flattened and roughly ellipsoidal in lateral view, with fringe of setae from ter- goapical angle to base of sternal side, setae on sternoapical edge noticeably flattened and expanded subapically; lobe M_ promi- nent, arising from sternolateral area of lobe A,E, tapered and bent apically, bearing 2 relatively large flattened setae on tergal margin and row of small stout setae on ster- nolateral surface; lobe C a relatively large stemmed process arising sternolaterally at base of head, bent mesad and bearing a globular head covered with rows of minute decumbent spicules; lateral side of head with a dangling bag-like membranous pro- cess (mp). Aedeagus longer than wide, broadly oval in tergal view; with submedian tergal arms joined at midline to form nar- row median tergal bridge; apical tergal arms broadly fused and bearing a median tubercle; median sternal plate membranous, apex flared and hood-like. Proctiger (lateral view) with very broad basal sclerotization (tergum X) narrowly fused with base of paraproct; paraproct narrow, slightly en- larged apically, apex bearing 3 or 4 small teeth and 3 subapical cercal setae; cercal sclerite distinct. Egg.—Unknown. Larva, fourth instar (Fig. 2).—Recon- structed from single exuviae (holotype), ae 367 character of setae as figured (positions may differ slightly from those shown), numbers of branches in Table 1. Exhibiting the sub- generic characters noted by Harbach (1991). Head: Slightly wider than long, widest in posterior half; length approxi- mately 1.0 mm; width approximately 1.1 mm; moderately tanned. Occipital foramen widely V-shaped with arms extending dor- solaterally to point laterad of level of seta 9-C, margins heavily tanned, ventrocaudal margin with collar-like edge. Anterior mar- gin of labiogula minutely denticulate; hy- postomal suture complete, gently curved. Dorsomentum short, roughly triangular, with 8 teeth on either side of median tooth, median tooth and most lateral tooth of ei- ther side larger than the others. Mandible without distinctive features. Maxilla rela- tively short, with 4 large lateral teeth of nearly equal size, about 0.25 length of api- cal tooth; apical tooth large, about half length of maxillary body, curved mesad. Setae 4—6-C single, simple; 7-C double; 11,13-C relatively short, about length of 12- C; 14-C much thicker and stiffer than other cranial setae, double; 15-C inserted cepha- lad of 14-C near anterior margin of labi- ogula. Antenna: Short, cylindrical, length 0.3 mm; moderately tanned, progressively lighter toward apex, surface smooth. Seta 1-A single, simple, borne dorsally about 0.8 from base, length about twice width of an- tenna at point of insertion. Thorax. Integ- ument hyaline, smooth. Setae 1-P,T, 4,5-T and 8,14-P weakly developed; 8-M much shorter than usual, slightly shorter than 7- M; 13-T shorter than thorax, with 3 thick aciculate branches. Abdomen: Integument hyaline, smooth. Setae 1-I weakly devel- oped, 1-II-VI relatively well developed, with stiff thickened branches, borne on bas- al plates; 2-I-VII weakly developed, small side. D, Comb scales. Abbreviations: A = antenna; C = cranium; CS = comb scale; P = prothorax; M = mesothorax; S = siphon; T = metathorax; I-VI, X = abdominal segments I-VIII, X; 0-15 = setal numbers for specified areas, e.g., seta 5-C. Scales in mm. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON = = == aa = = == = — = = 9/S CI = I/I 1/1 a = a _ = = = 8/01 -/7 T/T tl aa = rai vl/— b/P €/S S/P LIL C/S e/€ L/9 = S/S el = = —/— T/E 1/1 I/I 1/1 1/1 = 1/1 1/1 1/1 S/9 ral = = 1/1 8/11 b/€ c/E S/P 9/9 C/é TT 1/1 1/1 O/L a (Ge —/— —/T c/P 1/1 1/1 1/1 1/1 —/T 1/1 9/L E/E Ol E/E ‘S-B] a6 1/Z 1/1 1/1 1/1 1/1 1/1 11/6 S/S 9/9 p/P 6 TE ‘S-] —/O1 01/6 b/I C/E GS 1/1 = L/S T/T T/T c/€ 8 a= = 1/1 —/€1 81/61 L/9Té O@/1Z c/E C/E 6/8 CT ZI/ZI UT i. aa = —/r1 1/1 I/- 1/1 1/1 p/P b/P 9/9 1/1 -/€ 1/1 9 = ran 1/1 LI/L 9/8 b/L 9/S b/€ TT 1/1 1/1 TT 1/1 ¢ 8/8 —/I ra b/P LIL 1/1 1/1 o/t S/L TT 1/1 L/S 1/1 7 9/9 94/6 1/1 €/T I/I 7/7 I/I 1/1 1/1 b/S 1/1 T/T 1/1 € 9/9 T/T 1/1 1/1 1/1 1/1 1/1 1/1 1/1 1/1 1/1 1/1 = é b/P c/€ €/Z b/p C/E b/€ €/P €/Z TE 1/1 T/T i/I I/I I = I/1 1/1 1/1 I/I 1/1 1/1 1/1 = = = 9/S 1/1 0 ». IA WA © IA A AI Il II I at W d 2) PIOS SES —— ee ee ee eee ee sjusuisag yeurumopqy xXP1Oy ‘ysep Woys vB Aq payeoIpul aie sejas SUISSIPY “~poutadsIp aq jou prnoo Soyoursg JO Joquinu 9Y) yey) sayeorpul yeu uoNsonb vy ‘yseys ve Aq payesedas are Apoqg 24} JO Sapls IYSII pure Ia] ay) UO seIAS Jo sayoURIG JO JaquINN ‘snxopo.ind saylaqvgy Jo adAjopoy ay} JO avIANXO [RAIL] IeISUI-YLNOJ ay JO AxwIOJORYD “| BqRL VOLUME 104, NUMBER 2 and single, 2-I well laterad of 1-I, 2-II-VI well mesad and slightly anterior to seta 1; 6,7-I,1] similarly developed but seta 6 sig- nificantly longer than seta 7, both strongly aciculate; 6-III-VI long, single, strongly aciculate; 7-III small, resembling 7-IV—VI, small with numerous branches; 9-I—VII all of normal size and development; 13-I—V well developed, stellate, with stiff aciculate branches, borne on basal plates; 4-IIL1TV distinctly mesad of seta 1, more or less mid- way between setae | and 2; 12-VI shorter than usual, double or triple; 11-VII ventral to seta 13-VH; 13-VII not strongly devel- oped, double; punctures absent from seg- ments HI-V. Segment VIII: Comb an irreg- ular single row of 10,11 small thorn-like scales without lateral fringes of minute spicules. Siphon: Short, tapered, slightly flared at apex; moderately tanned, with dark plates laterally at base; lateral surfaces cov- ered with offset rows of blunt stud-like pro- jections that decrease in length posteriorly; length 0.90 mm, width at base 0.35 mm, index 2.57. Pecten of approximately 70 fil- aments staggered irregularly along posterior margin, filaments arise individually (about 25) and in clusters, 2 large clusters at mid- length arise from basal rings of paler integ- ument that resemble alveoli, proximal of these clusters with 12 filaments, distal one with 6 filaments. Seta 1-S inserted about 0.4 from base. Segment X: Saddle lightly tanned, with imperceptible rows of minute spicules mainly on sides; length 0.22 mm; siphon/saddle index 4.1. Setae 1—4-X equally well developed, shorter than usual, multiple with aciculate branches. Pupa (Fig. 3).—Character and positions of setae as illustrated, numbers of branches in Table 2. Cephalothorax: Lightly to mod- erately tanned, some darker mottling on scutum. Seta 1-CT strongly developed, double or triple, branches not noticeably sigmoidally curved, with hooked tips; 5-CT also well developed, single or double. Trumpet (Fig. 3A,C): Moderately and even- ly tanned, short, rather abruptly expanded at base; length 0.38—0.47 (x = 0.43 mm), 369 width at midlength 0.14—0.16 mm (* = 0.15), index 2.38-3.21 (« = 2.95); pinna short, length 0.06—0.11 mm (« = 0.09). Ab- domen: Lightly to moderately tanned, ster- na II-VII darker anteriorly; length 4.09— 5.20 mm (x = 4.45 mm). Seta 5-I laterad of seta 4-I; 7-I shorter than seta 6-I, with 2—4 branches; 2-II anterolateral to 1-II; 3- II on level anterior to level of 2-II; 5-H, much smaller than 5-IV—VI, 5-IV—VI lon- ger than length of following tergum; 7-II dorsal to 9-II; 8-II present or absent, alve- olus present when seta absent, 8-VI dorsal; 10-II present; punctures absent from seg- ments III—V. Genital lobe: Lightly tanned, length (male only) about 0.6 mm. Paddle: Lightly tanned, broadest at base, narrowed apically, outer part wider than inner part, minutely spiculate along margins, demar- cated basal membranous area present (Fig. 3D) or absent (Fig. 3B) on dorsal surface; length 0.59—0.70 mm (x = 0.66 mm), width at widest point 0.39-0.46 mm (* = 0.43 mm), index 1.46—1.65 (« = 1.52). Systematics.—The female of Sabethes paradoxus is unknown, but the male is eas- ily distinguished from that of all other spe- cies of the genus by the absence of brilliant metallic-colored scutal scaling, the greatly expanded apical portion of the proboscis, and features of the gonostylus. Assuming that the female of this species resembles the male in general habitus, it is possible, if not likely, that it will be confused for species of Wyeomyia. It will be necessary to note the absence of prealar setae to correctly identify the female as a species of Sabethes. The larva of Sa. paradoxus most closely resembles that of Sa. soperi Lane and Cer- queira, but is easily distinguished from this and all other Sabethes by the short, unique- ly ornamented siphon. Within subgenus Peytonulus, the robust development of seta 14-C is shared with Sa. soperi as well as Sa. hadrognathus Harbach and Sa. undosus (Coquillett). As far as known, the reduced seta 8-M of Sa. paradoxus is a unique fea- ture of this species. The pupae of Peytonulus are generally 370 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON / Howorcl Fig 3. Pupa of Sabethes paradoxus. A, Left side of cephalothorax, dorsal to right. B, Dorsal (left) and ventral (right) aspects of metathorax and abdomen. C, Trumpet. D, Paddle of paratype GG105-121 showing basal membranous area of dorsal surface. Abbreviations: CT = cephalothorax; Pa = paddle; I-VI = abdominal segments I-VI; 0-14 = setal numbers for specified areas, e.g. seta 3-I. Scales in mm. VOLUME 104, NUMBER 2 371 Table 2. Numbers of branches for setae of pupae of Sabethes paradoxus. Range (mode, when evident) of the type series (holotype and three paratypes). Cephalo Abdominal Segments thorax Paddle Seta (1h I I I IV Vv VI Vil Vit IX Pa O — — l 1 1 l 1,2(1) 1 1 — — i 2,3(2) 38-45 14 EZ CS) 1,2(1) 1,2(1) os — —— — 2 12) ] 1,2(1) 1 ] 1 —- — —— 3 52 ] | 2—5(3) | 1 1 4 | 1-3(3) = =24(3) 1—3(2) 1-3(2) 2-7(4) ] 1 1 —- — 3) eZ. 1—-3(1) 14(2) 2-403) ] 1 07/1(0?) = = = 6 1E2(2)) 1 1 1 XC!) 1 1,2() 7 1-3(2) 24(@) 2-5(3) 3-7(5) 1,2(1) 2—5(4) 1,2(1) 1 8 i — 0#,1(0*) 1,2 1—3(1) 141) 48(5) 3-7(4) — —- — 9 1 1,2(1) i i ] I 9-14 11-15 10 1,2(1) (OF 1 1 1,2(1) ! 1 1 — — — 11 1 | 0°/1(1) ] i ] 1 —— ae — 12 1 — = — = — — — — — — 13 — — — — — — — — —- = a 14 — — — — = — — — i = — 4 Alveolus present. very similar and more difficult to differen- tiate than the larvae. The pupa of Sa. par- adoxus resembles that of Sa. hadrognathus and differs from other species of the sub- genus in the dorsal placement of seta 8-VI. This seta is usually but not always dorsal in the latter species, and whether this is a fixed feature of Sa. paradoxus will remain uncertain until additional material becomes available for study. The three paratypes of Sa. paradoxus (see below) were reared from pupae col- lected from bamboo. The pupal exuviae of one of these specimens (GG105-121) bears a differentiated membranous area at the base of the paddle. This is paradoxical for two reasons: the holotype and other para- types have a normal paddle, and the feature is not known to occur in any other species of Peytonulus. A membranous area at the base of the paddle is characteristic of sub- genus Sabethoides and occurs in some spe- cies of subgenus Sabethes (Harbach 1991). The exuviae does not appear to be incor- rectly associated with the adult male be- cause seta 8-VI is dorsal in position and the specimen otherwise agrees with the holo- type and other paratypes in all other fea- tures. Seta 8-VI is ventral in species of sub- genus Sabethoides and those species of sub- genus Sabethes that have a membranous area at the base of the paddle. The genetics that control the expression of the character are unknown, but it seems plausible that this involves latent genes that may be influ- enced by environmental conditions. The function of the membranous area is like- wise unknown, but it could be a region of weakness to facilitate abscission of the pad- dle should it be grasped by a predator or entangled in debris. Etymology.—The specific name of par- adoxus is a Latin adjective (masculine) meaning strange or contrary to expectation. The name refers to the uncharacteristic col- oration of the adult male and the enigmatic presence of a delineated membranous area in one pupa of the type series. Bionomics.—The holotype was collected as a larva found in a terrestrial bromeliad; the paratypes were collected as pupae found in bamboo. In addition to the holotype, the bromeliad also contained larvae of the Cu- lex (Microculex) imitator group, Johnbelk- inia ulopus (Dyar and Knab), Limatus dur- hamii Theobald, a species of Toxorhynchi- 372 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tes (Lynchiella), a species of Wyeomyia (Hystatomyia), and two species of the Wy. (Wyeoymia) pertinans group. Species found in bamboo with the paratypes include the following (superscript | indicates collection GG105; 2 indicates collection GG109): Cu- lex (Carrollia) antunesit Lane and Whit- man!”, an unidentified species of Cx. (Cu- lex)', Limatus durhamii', Onirion sirivan- akarni (Duret)!?, Sa. (Peytonulus) identicus Dyar and Knab!?, Sa. (Pey.) hadrogna- thus'’, a species of Shannoniana'? (same species in both collections), Trichoproso- pon pallidiventer (Lutz), Tr. sp. near pal- lidiventer'’, Tr. castroi Lane and Cerquei- ra’, a species of Tx. (Lynchiella)'! (?same species found in bromeliad), and two spe- cies of Wyeomyia (Wyeomyia)'* (same two species in both collections). Nothing is known about the biology of the adults. Distribution.—Known only from the type locality in the Darien of Panama. The species probably occurs in neighboring countries in Central America and northern South America. Material examined.—Thirteen specimens (4 5,4 ¢ genitalia, | larval exuviae, 4 pu- pal exuviae) from | larval and 3 pupal rear- ings. Holotype: d (GG108-101, with asso- ciated larval and pupal exuviae and dis- sected genitalia on separate microscope slides), PANAMA: Darien, specific locality not known, 7 Jul 58, terrestrial bromeliad (Gorgas Memorial Lab. personnel). Para- types, same data as holotype except as fol- lows: | 6 (GG105-121, with associated pu- pal exuviae and dissected genitalia on sep- arate microscope slides), Pucro, ‘Paya Camp’, 50 m, 6 Jul 58, bamboo; 2 6 (GG109-112 and —122, both with associ- ated pupal exuviae and dissected genitalia on separate microscope slides), Pucro, 50 m, 8 Jul 58, bamboo. The holotype and par- atypes are deposited in the USNM. ACKNOWLEDGMENTS We thank personnel of the Walter Reed Biosystematics Unit, Smithsonian Institu- tion, Washington, DC, for the loan of spec- imens. We are grateful to John F Reinert (Center for Medical, Agricultural and Vet- erinary Entomology, United States Depart- ment of Agriculture, Agricultural Research Service, Gainesville, Florida) and Rory Post (The Natural History Museum, Lon- don) for commenting on the manuscript. LITERATURE CITED Hall, C. R., T. M. Howard, and R. E. Harbach. 1999. Sabethes (Peytonulus) luxodens, a new species of Sabethini (Diptera: Culicidae) from Ecuador. Me- morias do Instituto Oswaldo Cruz 94: 329-338. Harbach, R. E. 1991. A new subgenus of the genus Sabethes (Diptera: Culicidae). Mosquito System- atics 23: 1-9. . 1994. The subgenus Sabethinus of Sabethes (Diptera: Culicidae). Systematic Entomology 19: 207-234. . 1995a. A new Sabethes of the subgenus Pey- tonulus (Diptera: Culicidae) with an unusual fourth-instar larva. Entomologica Scandinavia 26: 87-96. . 1995b. Two new species of the subgenus Pey- tonulus of Sabethes (Diptera: Culicidae) from Co- lombia. Memorias do Instituto Oswaldo Cruz 90: 583-587. Harbach, R. E. and I. J. Kitching. 1998. Phylogeny and classification of the Culicidae (Diptera). System- atic Entomology 23: 327-370. Harbach, R. E. and K. L. Knight. 1980. Taxonomists’ Glossary of Mosquito Anatomy. Plexus Publish- ing, Inc., Marlton, NJ, xi + 415 pp. . 1982. Corrections and additions to Taxono- mists’ Glossary of Mosquito Anatomy. Mosquito Systematics 13: 201-217. Harbach, R. E. and J. L. Petersen. 1992. Two species previously confused under the concept of Sabeth- es tarsopus in Central America (Diptera: Culici- dae). Mosquito Systematics 24: 102—124. Harbach, R. E. and E. L. Peyton. 1991. Transfer of the subgenus Davismyia from Wyeomyia to Sabethes and description of the type species, Miamyia pe- trocchiae (Diptera: Culicidae). Mosquito System- atics 22: 149-159. Heinemann, S. J. and J. N. Belkin. 1978. Collection records of the project ““Mosquitoes of Middle America” 10. Panama, including Canal Zone (PA, GG). Mosquito Systematics 10: 119-196. Moses, D. A., T. M. Howard, and R. E. Harbach. 2000. A new species of the subgenus Sabethoides of Sa- bethes (Diptera: Culicidae) from Venezuela and Brazil. Proceedings of the Entomological Society of Washington 102: 991—1002. PROC. ENTOMOL. SOC. WASH. 104(2), 2002, pp. 373-375 DESCRIPTION OF A NEW SPECIES OF THE GENUS MASAAKIA TAKEUCHI (HYMENOPTERA: TENTHREDINIDAE) FROM JAPAN IcHII TOGASHI 1-chome, Honmachi, Tsurugi-machi, Ishikawa Prefecture 920-2121, Japan Abstract.—Masaakia katayamai, new species, from Honshu, Japan, is described and illustrated, and a key is provided for the three Japanese species of Masaakia. Key Words: Masaakia Takeuchi is a small genus of the subfamily Blennocampinae and is en- demic to Japan. It contains two described species, M. longivaginata Takeuchi 1950 and M. shinoharai Togashi 1998. Recently, I received two specimens of Masaakia through the courtesy of Mr. E. Katayama, Ohtawara City, Tochigi Prefecture, Honshu, Japan. These two specimens are distin- guished from the two described species by the yellow tegula, shape of the pedicel, shape of the sawsheath, and the serrulae of the lancet. Thus, I concluded that these specimens represent a new species. Here, I describe and illustrate this new species and give a revised key to the Japanese species of the genus. KEY TO THE JAPANESE SPECIES OF MASAAKIA 1. Apex of sawsheath pointed in lateral view (Fig. 11); tegula black; pedicel longer than wide stom) epee. Fiona o Goede eee erctee a icone Le 2 — Apex of sawsheath truncate in lateral view (Fig. 8); tegula yellow; pedicel shorter than wide (Fig. 4) 2. Antenna entirely black; radial crossvein of See robs doh eaeeth katayamai, 0. sp. FOKe wine CURVEC ss). a = longivaginata Takeuchi — Antenna black with undersides of 4th to 9th segments and apical half of underside of 3rd antennal segment reddish brown; radial cross- vein of forewing nearly straight (as in Fig. 5) | 2G) suche, a) clioun “Geen Gmeclod even ewe mean shinoharai Togashi Symphyta, Tenthredinidae, Blennocampinae, Masaakia, new species, Japan Masaakia katayamai Togashi, new species (Figs. 1-9, 12, 13) Female.—Length, 5 mm. Body including antenna black, but apical half of mandible reddish brown and tegula yellow. Wings hyaline; stigma and veins dark brown to black. Legs black with following parts milky white: tibiae except for apical portion and hind basitarsus. Head from above transverse (Fig. 1); postocellar area transverse, slightly convex: OOL:POL:OCL = 1.1:1.0:0.9; posterior half of circumocellar furrow distinct, ante- rior half indistinct; interocellar and posto- cellar furrows distinct; lateral furrows dis- tinct and deep, circular (Fig. 1); frontal area nearly flattened; median fovea distinct and deep, circular, with short longitudinal fur- row in middle; lateral fovea distinct and deep, circular, with a small conical-like pro- jection in middle; supra-antennal tubercles distinct; antenno-ocular distance about twice as long as distance between antennal sockets; front margin of clypeus emarginate (Fig. 3); malar space narrow, 0.2 as long as diameter of front ocellus; postorbital groove distinct (Fig. 2); postgenal carina distinct near mandible only (Fig. 2). Antenna (Fig. 4) shorter than costa of forewing; relative lengths of segments about 16:1:0:4.7:2.6:2.6:2.3: 0.62620; 374 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON =F Figs. 1-11. 4" J 14 1-9, Masaakia katayamai, holotype. 1, Head, dorsal view. 2, Head, lateral view. 3, Clypeus, front view. 4, Antenna, lateral view. 5, Forewing. 6, Fore inner tibial spur, lateral view. 7, Tarsal claw, lateral view. 8, Sawsheath, lateral view. 9, Sawsheath, dorsal view. 10-11, M. longivaginata. 10, Antenna, lateral view. 11, Sawsheath, lateral view. pedicel rectangular, length to width about 1.0:1.3 (Fig. 4). Thorax with mesoscutellum slightly con- vex; epicnemium absent. Forewing with ra- dial crossvein (2r) nearly straight (Fig. 5); vein 2A+3A straight. Hindwing with peti- ole of anal cell about twice as long as ner- LOvn kel Figs. 12-13. Masaakia katayamai, paratype. 12, Apical portion of lancet. 13, 9th to 11th serrulae of lancet. vulus; cell M absent. Legs with fore inner tibial spur furcate at apex (Fig. 6); hindbas- itarsus nearly as long as length of following 2 segments combined; tarsal claws with long inner tooth and basal lobe (Fig. 7). Abdomen with sawsheath rather long (Fig. 9) in dorsal view, apical portion trun- cate in lateral view (Fig. 8); cercus short, 0.4X< as long as sawsheath. Lancet with 16 serrulae, serrulae lobelike, rounded at api- ces; apical portion of lancet as in Fig. 12; Oth to 11th serrulae as in Fig. 13. Head and thorax covered with fine setig- erous punctures, those of clypeus closer than those on vertex; mesopleuron practi- cally impunctate, shining. Abdominal ter- gites nearly impunctate. Male.—Unknown. Food plant.—Unknown. Types.—Holotype 2, 12.1V.2000, Yoi- chi, Ohtawara City, Tochigi Prefecture, E. Katayama leg. Paratype, 1 2, 29.1V.2000, VOLUME 104, NUMBER 2 Shimoishigami, Ohtawara City, Tochigi Prefecture, E. Katayama leg. Both deposit- ed in the collection of the National Science Museum (Natural History), Tokyo. Remarks.—This new species comes out to Masaakia in the keys to genera by Tak- euchi (1952) and Okutani (1972). There are two minor discrepancies with the generic concept of Masaakia, the broad almost square pedicel and the shorter sawsheath with the apex truncated in lateral view. The typical species of Masaakia have the pedi- cel elongate, longer than broad, and the sawsheath longer and more pointed at its apex in lateral view. Other characters, how- ever, support the placement of M. kataya- mai in Masaakia, e.g., the bifid tarsal claws with a basal lobe, third antennal segment longer than the fourth segment, emarginate clypeus, narrow malar space, short genal carina, presence of a hind orbital groove, lack of an epicnemium, straight vein 2A+3A of the forewing, and petiolate anal cell and absence of cell M of the hindwing. Masaakia katayamai is similar to both M. longivaginata and M. shinoharai, but it is distinguished by the yellow tegula (in M. longivaginata and M. shinoharai, the tegula is black), by the broad pedicel (elongate in the other two species, compare Figs. 4, 10), by the truncate sawsheath (pointed at the apex in the other two species, compare Figs. 8, 11), and by the deeper, more round- ed serrulae of the lancet (shallower and flat- ter in the other two species, compare Figs. 12, 13 and Togashi 1998, figs. 15, 16). ACKNOWLEDGMENTS I thank Dr. David R. Smith, Systematic Entomology Laboratory, U.S. Department of Agriculture, Washington, DC, for re- viewing the manuscript. I also thank Mr. E. Katayama, Ohtawara City, Tochigi Prefec- ture, for lending me the valuable speci- mens. LITERATURE CITED Okutani, T. 1972. A new genus and a key to Japanese genera of the subfamily Blennocampinae (Hym. Tenth.). Entomological Review, Japan 24: 57-61. Takeuchi, K. 1952. A generic classification of the Jap- anese Tenthredinidae (Hymenoptera: Symphyta. Kyoto. 90 pp. Togashi, I. 1998. A new species of the sawfly genus Masaakia (Hymenoptera, Tenthredinidae) from Hokkaido, Japan. Bulletin of the National Science Museum, Tokyo, Series A 24(1): 27—30. PROC. ENTOMOL. SOC. WASH. 104(2), 2002, pp. 376-389 A REVIEW OF CHALODETA STICHEL WITH A REVISION OF THE CHELONIS GROUP (LEPIDOPTERA: RIODINIDAE) JASON P. W. HALL Department of Systematic Biology, Entomology Section, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0127, U.S.A. (e-mail: jpwhall @ hotmail.com) Abstract.—An overview of the Neotropical riodinid genus Chalodeta Stichel is pre- sented which defines the taxon, delineates its member species, and discusses its systematic position within the tribe Riodinini. A revision of the Chalodeta chelonis group includes notes on the taxonomy and biology of its species, and illustrations of the adults and male and female genitalia (where known) of all taxa. Four species are recognized, including two that are previously described, C. chelonis (Hewitson 1866) and C. chaonitis (Hewitson 1866), and two that are described here, C. chitinosa, n. sp., and C. chlosine, n. sp. The taxon stilbos Stichel 1910, is synonymized with C. theodora (C. and R. Felder 1862) (n. syn.). Key Words: The monophyly of most genera in the Riodinidae has never been critically as- sessed, but to do so is vitally important if the family’s classification is to become a predictive tool in broader evolutionary studies. The purpose of this paper is two- fold. The first is to provide an overview and diagnosis for the small riodinid genus Chal- odeta Stichel 1910, which has historically often been confused with other genera in the tribe Riodinini (sensu Harvey 1987), particularly Charis Hiibner [1819], delin- eate its constituent species, and discuss its systematic position, biogeography, and _ bi- ology. Chalodeta is hypothesized here to consist of two monophyletic groups, and the second purpose of this paper is to pre- sent a revision of one of these, the chelonis group, whose true species diversity has pre- viously gone undetected. It consists of two named species, C. chelonis (Hewitson 1866) and C. chaonitis (Hewitson 1866), and two additional species, widely sympat- Chalodeta, Charis, Neotropics, Riodinini, taxonomy ric with C. chaonitis, that are described here. All four chelonis group species are il- lustrated here, while adequate color figures of all theodora group species may be found in d’ Abrera (1994) (C. theodora (C. and R. Felder 1862) and C. lypera (Bates 1868)) and Hall and Willmott (1998) (C. pescada Hall and Willmott 1998 and C. panurga Sti- chel 1910). METHODS Dissections were made using standard techniques, abdomens being soaked in hot 10% potassium hydroxide solution for ap- proximately five minutes, and subsequently stored in glycerol. Specimens dissected are indicated in the material examined sections with an asterisk. Morphological terms for genitalia follow Klots (1956) and Eliot (1973), and the terminology for wing ve- nation follows Comstock and Needham (1918). The protocol for listing material ex- amined follows Hall (1999). VOLUME 104, NUMBER 2 Chalodeta chelonis group specimens have been examined and their locality data recorded in the following collections, whose acronyms are used throughout the text. Only locality data are given in the spe- cies accounts of described taxa, but full la- bel data are given for new species. AME Allyn Museum of Entomology, Florida Museum of Natural His- tory, Sarasota, FL, U.S.A. Collection of Boyce Drummond, Florissant, CO, U.S.A. The Natural History Museum, London, U.K. Collection of Jason Hall and Keith Willmott, Washington, DC, U.S.A. Museo de Historia Natural, Univ- ersidad Nacional Mayor de San Marcos, Lima, Peru Reading Public Museum, Read- ing, PA, U.S.A. Senckenberg Museum, Frankfurt, Germany Staatliches Museum fiir Tierkun- de, Dresden, Germany National Museum of Natural His- tory, Smithsonian Institution, Washington, DC, U.S.A. Zoologische Museum fiir Natur- kunde, Humboldt Universitat, Berlin, Germany Zoologische Staatssammlung, Munich, Germany BD BMNH JHKW MUSM RPM SMF SMTD USNM ZMHU ZSM REVIEW OF CHALODETA Chalodeta Stichel 1910 Chalodeta Stichel 1910b: 15. Type species by original designation: Charis theodora Cand R: Felder 1862: 72. Diagnosis and systematic position.— Chalodeta species are small to medium- sized riodinids (forewing length 10 mm [theodora]| to 16 mm [chelonis]) with com- pact wing shapes, often slightly falcate forewing apices, and rounded hindwings. The dorsal surface is typically brown with SY) one (all theodora group species except theodora) or two (all chelonis group species and theodora) submarginal blue or greenish silver lines, three dark brown markings in the discal cell, and discal, postdiscal and submarginal bands of dark brown spots. The ventral surface is typically iridescent blue or purple in males and brown in fe- males and has similar markings to the dor- sal surface except no silver submarginal lines are present (see Figs. 1A—H). The fringe of both wings is often entirely white. All members of the often confused genus Charis except the misplaced ocellata group (Hall and Harvey, in prep.) have two dorsal submarginal silver lines, and all except one have some ventral submarginal silver mark- ings. The exception is a recently described member of the Charis gynaea group (Hall and Harvey 2001), which was figured by DeVries (1997) as ““Chalodeta candiope” and presumably placed by him in Chalod- eta because of its lacks of ventral silver markings. The male genitalia of all Chalodeta spe- cies (see Figs. 2A—D, 3) possess the deep notch in the anterior margin of the tegumen characteristic of the tribe Riodinini. The un- cus is rectangular and in chelonis group species typically forms a small bifurcate posterior projection medially along the dor- sal margin. The falces and tegumen are of average size and shape for the tribe, and the vinculum is evenly narrow and somewhat arched medially. The aedeagus is character- istically short, narrow and straight, unlike that of Charis and most other riodinine spe- cies, in which it 1s long and variably asym- metrically curved; no cornuti are present. The structure of the posteriorly elongate pedicel is unique. It is tightly appressed to the aedeagus, its tip forms a ventrally di- rected plate, and its basal ventral margin is unsclerotized. The ventral tip contains elon- gate spines around its perimeter in chelonis group species, but only very small spines (lypera only) or no spines in theodora group species. The valvae of the two Chal- odeta species groups are very distinct. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VOLUME 104, NUMBER 2 Those of the chelonis group (Fig. 2) have a small narrow lower process that is un- sclerotized at its base and a_ posteriorly elongate upper process that typically has a medial dorsal projection and long broad spines at the tip of both upper projections (only chitinosa lacks a dorsal upper pro- cess). The transtilla is connected to the up- per valve process only at its base and is narrow and very posteriorly elongate with two upwardly curving brachia at its tip. Valvae of the theodora group (Fig. 3; see also male genital illustration of C. pescada in Hall and Willmott 1998) have an entirely sclerotized lower process and an upward and slightly outwardly directed upper pro- cess with no spines. The transtilla is con- nected to the upper valve process along its entire length, creating a broad concave dor- sal plate that narrows to a small bifurcate and grooved tip within which the tip of the aedeagus is confined. The female genitalia (see Figs. 4A—C) differ slightly between the two _ species groups. Those of the chelonis group have an elongate corpus bursae with elongate in- vaginated spine-like signa and a large area of sclerotization on the eighth abdominal sternite that is often medially desclerotized. The known females of the theodora group have a rounded corpus bursae with the sig- na either forming rectangular invaginations with an elongate base at the wall of the cor- pus (theodora) or two sclerotized bands at the wall of the corpus (/ypera), and a small area of sclerotization on the eighth abdom- inal sternite. In all species, the ductus bur- sae is relatively short and unusual in ex- tending to the dorsal tip of a posteriorly elongate ostium bursae. The position in all Chalodeta species of the small ventral sclerotized plate in the ductus bursae at the oe Fig. 1. 379 opening of the corpus bursae instead of im- mediately before the ostium (creating an elongate ductus seminalis parallel to the ductus bursae) is not known elsewhere in the tribe. As indicated above, despite a superficial external similarity (including the possession of hairy eyes), the genital morphology does not suggest a particularly close relationship between Chalodeta and Charis within the Riodinini, although the presence of most wing pattern elements in the species of these genera suggests they are both rela- tively basal within the tribe. Currently very little is known about relationships between genera in the Riodinini and what makes the pursuit of this knowledge all the more dif- ficult is the relative lack of conservative characters. While the tremendous interspe- cific variation in genitalia provides good di- agnostic characters at the species and spe- cies-group levels, it acts to confound the elucidation of relationships at the generic level and above. The other putatively basal riodinine genera which still possess most wing pattern elements are Metacharis But- ler 1867, Dachetola Hall 2001, Calephelis Grote and Robinson 1869, Caria Hiibner 1823, Amphiselenis Staudinger 1887, La- saia Bates 1868, and Exoplisia Godman and Salvin 1886. However, the male geni- talia of all but the first two of these genera possess pedicels tipped with the typical riodinine scobinate patch. Chalodeta may be most closely related to Metacharis and Dachetola, which possess a somewhat sim- ilar wing pattern devoid of ventral silver, and a posteriorly elongate ‘“‘rod’’-like ped- icel and a simple strap-like pedicel respec- tively. History of classification.—Stichel (1910a) described the genus Chalodeta to Chalodeta adults (dorsal surface on left, ventral surface on right, unless otherwise stated). A, d C. chelonis, Petropolis, S.E. Brazil (USNM). B, 2 C. chelonis, Petropolis, S.E. Brazil (USNM). C, Holotype ¢ C. chlosine, dorsal surface, Pakitza, Peru (USNM). D, Holotype d C. chlosine, ventral surface. E, Holotype 3 C. chitinosa, Tingo Maria, Peru (USNM). E Allotype 2 C. chitinosa, Pakitza, Peru (USNM). G, 3 C. chaonitis, Parque do Gama, S.E. Brazil (USNM). H, 2 C. chaonitis, Parque do Gama, S.E. Brazil (USNM). 380 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON include the species theodora, lypera, pan- urga, Chaonitis, and calagutis Hewitson 1871, designating the first of these as the type species, but subsequently added che- lonis, epijessa Prittwitz 1865, azora Godart [1824] (Stichel 1910b) and speusippa Schaus 1928 (Stichel 1930-31). The taxon virido Lathy 1958, was added to Chalodeta by Rebillard (1958) as a subspecies of che- lonis, but subsequently raised to species rank by Callaghan (1995). In his first riod- inid catalog, Bridges (1988) placed cala- gutis in Charis, presumably inadvertently, although calagutis and the remaining mem- bers of the ocellata group are actually ge- nerically distinct from Charis (Hall and Harvey, in prep.). d’Abrera (1994) con- fused the generic status of Chalodeta by combining it with Charis in his pictorial overview of the family, stating: ““Some workers follow Stichel (1910) in separating certain species in this group into the genus Chalodeta Stichel, 1910.... This writer considers this confusing and unnecessary and will retain them all in Charis Hiibner.” However, Bridges (1994) retained Chalod- eta distinct from Charis. Most recently, DeVries (1997) synonymized speusippa with /ypera, Hall and Willmott (1998) de- scribed pescada, and Hall (2001) trans- ferred virido and azora (with epijessa as a synonym) to Dachetola. Since two species are newly described here, I recognize eight species for Chalodeta in the systematic checklist below. Chalodeta theodora is highly variable even within populations, and as the name stilbos Stichel 1910 does not represent a geographically discrete phe- notype, it is synonymized with C. theodora. A dash ‘*—’’ indicates a synonym. Chalodeta Stichel 1910 chelonis group chaonitis (Hewitson 1866) chelonis (Hewitson 1866) chitinosa Hall, n. sp. chlosine Hall, n. sp. theodora group lypera (Bates 1868) —speusippa Schaus 1928 panurga Stichel 1910 pescada Hall and Willmott 1998 theodora (C. and R. Felder 1862) —stilbos Stichel 1910, n. syn. —calligramma (Rebillard 1958) KEY TO SPECIES OF CHALODETA (MALES) No key is given for females as those of C. chlosine, C. pescada and C. panurga are not known. ile Dorsal submargin with one silverish colored Jin: 3.6 ea. old Ach cetee eed on eee ene yD = Dorsal submargin with two silverish colored HIMES! po, dees nts NS 2 4 Spee eo eee 4 2(1). Ventral surface prominently iridescent blue i ahadkades aly Lars, oS SRN Oe ee 3 - Ventral surface brown with faint purple iri- GESCENCE™ 2. gh 0G 5 4 oth ee ee lypera 3(2). Dorsal surface with dark shades of irides- cént"blue «0.22.5 sede ae ee panurga Dorsal surface brown ........... pescada 4(1). Dorsal submarginal lines closely spaced, narrow and bluish silver - Dorsal submarginal lines distantly spaced, broad and greenish silver ........ theodora 5(4). Dorsal postdiscal line with no dark shading proximally v5 22 .P fen ee ee 6 - Dorsal postdiscal line with dark shading proximally 6. oc oss ciqsie co cyee | ey 7 . Forewing length typically 16 mm, forewing apex strongly falcate, ventral purple irides- Cenceistrons =... a.) 5. Cee Cee chelonis - Forewing length typically 13 mm, forewing apex weakly falcate, ventral purple irides- cence weak chlosine 7(5). Dark shading proximal to dorsal postdiscal line narrow, distal portion of dorsal wings brown. o2. ¢d.46y 28 32.3) expe chitinosa - Dark shading proximal to dorsal postdiscal line broad, distal portion of dorsal wings PATASINIS NN [KONA 4g canncocoscecoe chaonitis Biogeography.—Chalodeta species are distributed throughout the Neotropics, from Mexico to west Ecuador, throughout the Amazon basin and Guianas, and extend as far as southeastern Brazil (see Fig. 5). Two species, C. panurga and C. pescada exclu- sively inhabit lower premontane forest (Hall and Willmott 1998), while the re- mainder inhabit wet lowland rainforest and may also extend into lower premontane habitats (e.g., C. lypera, C. theodora and C. chelonis). The highest number of species occurs in the five northern and central An- dean countries, where all species but C. VOLUME 104, NUMBER 2 chelonis, a southeastern Brazilian endemic, should occur. Only C. lypera and C. chaon- itis are known to occur west of the Andes and throughout the Guianas. Biology.—All Chalodeta species except C. theodora are uncommon to very rare. Males are rarely encountered perching in small groups on hilltops, along streamsides or shaded forest paths at a variety of heights above the ground and usually in the early morning or early to late afternoon; they make rapid sorties and rest only briefly on the tops of leaves with their wings half open (Brévignon and Gallard 1998, Hall and Willmott, unpubl. data). Males are most frequently encountered in rotting fish baited canopy and subcanopy traps (Hall and Will- mott 2000), suggesting that males are infre- quently seen because they perch in the can- opy. Two species, C. lypera and C. chaon- itis have been recorded visiting flowers (Brévignon and Gallard 1998, Hall and Willmott 2000). The early stages are known for two species, C. lypera (incorrectly re- ferred to as C. chelonis by Kaye 1921) and C. chaonitis, which have been recorded feeding on young leaves and flowers of plants in the Melastomataceae, Passiflora- ceae and Sterculiaceae (Kaye 1921, Kirk- patrick 1954, DeVries et al. 1994). The lar- vae bear long tufts of lateral setae and the pupae are squat and bulbous with a broad cremaster (DeVries 1997). REVISION OF CHALODETA CHELONIS GROUP Chalodeta chelonis (Hewitson 1866) (Figs. 1A, B; 2A; 4A; 5) Charis chelonis Hewitson 1866: pl. 57, fig. 9. TL: Rio de Janeiro, S.E. Brazil. Syn- type 6 BMNH [examined]. Identification and taxonomy.—Typical forewing length: male 16 mm; female 15 mm. This species is readily distinguished by its large size, pointed wing shape, prom- inently falcate forewing apex and more uni- formly pale iridescent purple ventral sur- face with weakly defined markings. The lack of dark shading proximal to the dorsal 381 postdiscal bands occurs elsewhere only in C. chlosine (described below), which also has very similar male genitalia and appears to be its closest relative. The two species are distinguished in that species account. Biology.—Unknown. Distribution.—This species appears to be endemic to the northern states of south- eastern Brazil. Material examined.—BRAZIL: Espirito Santo, No specific locality 1 ¢, 1 2 BMNH; Minas Gerais, Campo Belo 3 @ ZMHU; Maromba 2 6 BMNH; 1 do SMF; Léopoldina ies 1 9 -ZMHU; 1 36, 1 ¢ SMTD,; Rio de Janeiro, Rio de Janeiro 1 6, 1 2 BMNH; 1 ¢ SMTD; 1 2 SMF; Nova Friburgo 1 ¢6 BMNH; Laguna de Sacuare- mai2 56-91? BMNEH-’ 18) W, 340. m,. 30) Sept 1991 (O. Mielke) (USNM). Paratypes: COLOMBIA: Putumayo, | 6: Rio Mulato, Mocoa, 29 Mar 1929 (ZMHU). Amazonas, 1 ¢: Florida, Sept 193 1G. ius) (BMNH). ECUADOR: Napo, | 3*: Tena, 1°01’S 77°49'W, 550 m, 6-10 Nov 1988 (R. Robbins) (USNM). 1 d*: Apuya, km 20 Tena-Puyo rd., 600 m, 6 Dec 1996 (K. Willmott) (JHKW). PERU: Loreto, 1 3: Rio Pacaya, Lower Rio Uca- yali, Aug 1912 (BMNH). Junin, | 5: Chan- chamayo (ZSM). Madre de Dios, Parque Nacional Pakitza, 11°55'48”S 71°15'18’W, 340 m (USNM), 1 6: 14 Oct 1991 (R. Rob- bims);16G215 Oct 1991 (G. Lamas), 1 6 *: 20 Oct 1991 (M. Casagrande). | 3d: Boca Iowa iorme: 20 Oct 1983°(G.. Lamas) (MUSM). Puno, 2 3: Yahuarmayo, 1,200 ft; Feb/Mar 1912 (H. & C. Watkins) (BMNH). BOLIVIA: La Paz, 1 3: Mapiri (SMTD). BRAZIL: Mato Grosso, 1 6*: Diamantino, Alto Rio Arinos, 14°13’S 56°12 W, 24 Sept 1989 (E. Furtado) (USNM). 3 6: Cuiabé (BMNH). 1 <¢: ‘““Mato Grosso” (Zobrys & Wolter) (BMNH). Etymology.—The species name is a eu- 383 Fig. 3. theodora. Male genitalia in lateral view of Chalodeta phonious anagram of “‘chelonis,”’ the name of its most closely related species. Diagnosis.—Chalodeta chlosine is super- ficially most similar to C. chaontits and C. chitinosa (described below), but lacks dark shading proximal to the dorsal postdiscal bands and has different genitalia (see those species accounts for further details). It ap- pears to be most closely related to the southeastern Brazilian endemic C. chelonis, but is smaller, has a more rounded wing shape, a less falcate forewing apex, and more prominent ventral markings. The male genitalia do not differ consistently. Biology.—Chalodeta chlosine is the rar- est of the Amazonian chelonis group spe- cies. The ratio of chlosine, chitinosa, and chaonitis specimens examined in collec- tions, respectively, is 2:7:10. An Ecuadori- an male was attracted to a canopy trap bait- ed with rotting fish. Distribution.—This species is currently known only from the western Amazon, from Ecuador to Bolivia and into south- western Brazil. Chalodeta chitinosa Hall, new species @igs: LE, Fy 2C:;,4B:,5) Description.—Male: Forewing length 14 mm. Forewing costal margin approximately 384 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON straight, distal margin slightly convex; hindwing rounded. Dorsal surface: Fore- wing ground color brown with subtle green- ish iridescence at oblique angle; three black marks in discal cell, one at base of cell Cul, two towards base of cell Cu2; a disjointed black postdiscal band extends from vein 2A to costa and is proximally kinked in cells Cul and M3, black scaling extends proxi- mally; two parallel submarginal silver lines encompass area of dark orange brown scal- ing containing a single black spot in each of cells Cul to R4+5 and two in cell Cu2, dark orange brown at distal margin; fringe VOLUME 104, NUMBER 2 brown with white scaling at distal tips of veins Cu2 to R4+5. Hindwing same as forewing except black scaling proximal to postdiscal band considerably less promi- nent. Ventral surface: differs from dorsal surface as follows: ground color pale gray brown overlaid with purple iridescence, dis- tal margins gray without iridescence (es- pecially hindwing) and with undulating proximal margin, silver submarginal lines absent. Head: Labial palpus brown. Eye brown and densely setose with brown scaling at margins. Frons brown. Antennal segments black with white scaling at base; club black, tip orange brown. Body: Dorsal surface of thorax and ab- domen dark brown, ventral surface pale brown; tegula brown. Legs brown with some iridescent purple setae. Genitalia (Fig. 2C): Uncus rectangular, posterior dorsal margin approximately straight; falces of average size and shape for family, small, deep semicircular notch in anterior margin of tegumen; vinculum narrow and ribbon-like, extends dorsally over anterior portion of tegumen; aedeagus relatively short, narrow and straight with pointed tip; pedicel extends from a point on aedeagus one-third distance from base to tip as narrow weakly sclerotized tube to form short posterior projection tipped with a heavily sclerotized oval plate with short spines around ventral perimeter, ventrally unsclerotized except at tip; valvae consist of a short, rounded lower process that is unsclerotized at its base and a tapering, rounded posteriorly projecting upper pro- cess with two long stout spines at tip, pos- teriorly elongate transtilla connected to dor- sal anterior portion of upper valvae process and forms two small, upwardly curving rounded posterior projections at tip. Female: Differs from male as follows: Distal margin of forewing more convex. Dorsal surface: Ground color paler. Ventral surface: Ground color brown with no pur- ple iridescence, distal margins and wing ba- ses pale orange brown. 385 Genitalia (Fig. 4B): Corpus bursae elon- gate, signa medium-sized spine-like invag- inations; ductus bursae narrow and mem- branous with small sclerotized ventral plate at anterior end, ductus seminalis connects to this sclerotized plate; eighth abdominal sternite a small sclerotized plate with me- dially and ventrally positioned ostium bur- sae forming a rounded posteriorly elongate projection, small round opening for ductus bursae at dorsal tip. Type material.—Holotype: d*, PERU: Hudnuco, Tingo Maria, 800 m, 24 June 1982 (S. Nicolay) (USNM),. Allotype: @, PERU: Madre de Dios, Parque Nacional Pakitza, 11°55'48"S 71°15'18"W, 340 m, 16 Sept 1989 (R. Rob- bins) (USNM). Paratypes: VENEZUELA: Amazonas, | 2: Yavita (Lichy) (AME). COLOMBIA: 1 6: >Gagueta, 1 Feb) (ZMHU): | ECUA- DOR: Sucumbios, Limoncocha (B. Drum- mond) i(BD)n hed 3 25cApr 19745: 8 Oct 1974. PERU: Loreto, 1 @: Castafia, 048228 75 1440 Wets Om; 17sOeu 19 ot (G. Lamas) (USNM). 1 ¢: Puerto Almen- dra, Rio Nanay, 03°50'S 73°23'W, 120 m, 3 Sept 1995 (R. Robbins) (USNM). Iquitos, 1 ¢: (H. Whitely) (BMNH), 1 6: (Hahnel) (ZMHU). 1 6: Pebas (Hahnel) (ZMHU). San Martin, Yurimaguas, 1898 (Michael) (ZMHU). 1 ¢: Juanjui (RPM). Pasco, 1 &: Monte Alegro, Rio Pachitea (G. Tessman) (ZMHU). 1 6: Pachitea (SMTD). Madre de Dios, Parque Nacional Pakitza, 11°55'48”S 71°15'18"W, 340 m (USNM), 2 6: 23 Sept I9890@D) Harvey), Ga27 "Sept 199s (©: Mielke)2: 9: 2:}Octd991s(Gatamas); dike: 6 Oct 1990 (R. Robbins), 1 d, 1 2*: 6 Oct 1991 (R. Robbins), 1 d: 10 Oct 1991 (O. Mielke), 1 ¢: 11 Oct 1991 (R. Robbins), 1 G2 Oct 199 (On Mielke) s2ndiysh@et 1991): (@iMielke)s ded #5 tr Oct 199 1n(@: Mielke), 1 d: 20 Oct 1991 (O. Mielke). 1 3: 30 km S.W. of Puerto Maldonado, 300 m, 22 Oct 1983 (S. Nicolay) (USNM). | 6: ‘Peru’? (SMF). Puno, 1F: La Union, Rio Huacamayo & Rio Carabaya, 2,000 ft, Nov 1904 (Ockenden) (BMNH). BOLIVIA: La 386 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Paz, 2 6: Mapiri (BMNH). 1 6, 1 2: Yun- gas (ZMHU). 1 ¢6*: “Peru” [= Bolivia] (coll. W. Schaus) (USNM). 2 ¢: “‘Bolivia”’ (SMF). BRAZIL: Amazonas, Sao Paulo de Olivenga, 6 6: Jan 1933 (S. Waehner) (BMNH), 1 2: (M. Moss) (BMNH), 1 &: (H. Bates) (BMNH), 2 3: Nov 1930 (S. Wucherpfennig) (SMF), 1 d: (Hahnel) (ZMHU), 3 6: (SMTD). 1 d: Tonantins (H. Bates) (BMNH). Manicoré, 1 od: 1887 (Hahnel) (ZMHU), 1 6: (SMTD). 1 2: Ma- naus, 1886 (Hahnel) (ZMHU). Rondénia, vicinity of Cacaulandia, 10°32'S 62°48’W, 160—350 m (J. Kemner) (USNM), | 6*: 23 Oct\1 991, Me 293@cr 1991. Paral &: km 1,666 Cuiaba-Santarém highway (C. Callaghan) (AME). | @: Itaituba, Rio Ta- pajos, 1890 (Michael) (ZMHU). Belém, 2 2: (H. Bates) (BMNH), 1 6: (ZMHU), 1 OA ZSMD: cAmazon*? (bed) Ca: Bates) (BMNH), 1 6, 1 2: (ZSM). GUYANA: 1 2: “Guyana” (Parish) (BMNH). Etymology.—The species name is a eu- phonious anagram of “‘chaonitis,”’ the name of a closely related species. Diagnosis.—Chalodeta chitinosa has long been confused with and is most similar to C. chaonitis and C. chlosine. It has some dark shading proximal to the dorsal post- discal bands, unlike C. chlosine, although this is less extensive than in C. chaonitis, lacks the greenish-brown distal coloration of C. chaonitis and typically has prominent alternating white and brown hindwing fringe elements. In C. chlosine, the hind- wing fringe is always entirely white and in C. chaonitis it is pale brown with only faint dirty white fringe elements. Each of these three species has a slightly different config- uration to the postdiscal forewing band. In C. chitinosa and C. chlosine, the postdiscal markings in cells Cul and Cu2 form an out- wardly directed semicircle while in C. chaonitis they form a stepped outwardly di- rected line. The postdiscal forewing spot in cell R4+5 is also more distally positioned in C. chlosine than it is in C. chaonitis and C. chitinosa. The genitalia of C. chitinosa are the most distinctive of all species in the group. The male genitalia have a straight instead of bi- fid posterior dorsal margin to the uncus, a broad posteriorly tapering upper valve pro- cess that has only two long spines at the tip and lacks a second spine-tipped dorsally elongate anterior process. The female gen- italia have a smaller eighth abdominal ster- nite that forms a single smoother sclero- tized plate with a more posteriorly elongate ostium bursae. Biology.— Unknown. Distribution.—Chalodeta chitinosa oc- curs from Venezuela to Bolivia and throughout Amazonian Brazil. It is unclear how widely it may be distributed in the Guianas, and only a single female in the BMNH is labeled from Guyana. Chalodeta chaonitis (Hewitson 1866) (PissalG, EoD; ACs) Charis chaonitis Hewitson 1866: pl. 57, figs. 7, 8. TL: “Amazon.” Lectotype male BMNH [designated]. Identification and taxonomy.—Typical forewing length: both sexes 13 mm. Chal- odeta chaonitis appears to have been de- scribed from two males and one female, all of which reside in the BMNH and bear Hewitson “‘Amazon” labels. However, al- though the two male syntypes represent C. chaonitis as treated here, the female syn- type represents C. chitinosa described above. Thus, the only labeled syntype male of C. chaonitis in the BMNH is designated as a lectotype. It bears the following labels: ‘‘Hewitson Coll./79-69./Charis/chaonitis. 1/ Amazon.” and ‘‘Type.” This lectotype is designated to fix and stabilize the identity of this species. Chalodeta chaonitis differs from C. chi- tinosa and C. chlosine, which have previ- ously been confused with it, by possessing distal greenish-brown dorsal coloration, ex- tensive dark shading proximal to the dorsal postdiscal bands, an outwardly diagonal in- stead of semicircular portion to the fore- wing postdiscal band in cells Cul and Cu2, VOLUME 104, NUMBER 2 387 © Chalodeta chelonis @ Chalodeta chlosine A Chalodeta chitinosa ®@ Chalodeta chaonitis Fig. 5. and a pale brown hindwing fringe with only weak dirty white elements. The upper valve process of the male genitalia possesses an elongate and upwardly curving posterior portion tipped with spines, and the eighth abdominal sternite of the female genitalia is divided into a heavily ribbed upper portion with the ostium bursae forming a separated ventral portion (unlike in C. chitinosa). Biology.—DeVries (1997) reported find- ing aggregations of males perching in Ec- uador along forest edges and in forest light gaps from 0.5 to | m above the ground be- tween 1500 and 1530 hrs, however, these individuals may also be referrable to C. 0 500 1000 Distributions of Chalodeta chelonis group species. chlosine and C. chitinosa. Bona fide Ecu- adorian males of C. chaonitis have been en- countered feeding on low weedy Astera- ceous flowers in secondary growth and at- tracted to canopy traps baited with rotting carrion in primary forest during the late af- ternoon (Hall and Willmott 2000). DeVries (1997) illustrated a mature larva and pupa of C. chaonitis reared from La Selva, Costa Rica. The larvae feed on flowers of Micon- ia longifolia (Melastomataceae) (DeVries et al. 1994). Distribution.—This is the most wide- spread chelonis group species and ranges from Mexico to western Ecuador, the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Guianas, Amazon basin and Trinidad, and as far south as southern Brazil. The follow- ing additional localities are listed by de la Maza and de la Maza (1993) for MEXICO: Chiapas, No specific locality; by DeVries (1997) for COSTA RICA: Heredia, La Sel- va; Puntarenas, Palmar Norte; and by Bar- cant (1970) for TRINIDAD: Fondes Aman- des. Since the report of this species in southern Mexico by de la Maza and de la Maza (1993) is not accompanied by an il- lustration, this record requires confirmation. Material examined.—COSTA RICA: Li- mon, Guapiles 2 ¢ USNM. PANAMA: Chiriqui, No specific locality 1 2 BMNH; 1 2 ZMHU; Canal Zone, Cocoli2 6,1 °* USNM,; Darién, Cana 2 6* USNM. VEN- EZUELA: Bolivar, La Vuelta, Rio Caura 1 2 BMNH. COLOMBIA: Auila, Neiva 1 @ RPM (possibly mislabelled). ECUADOR: El Oro, nr. Pasaje 1 ¢ JHKW; Sucumbios, Limoncocha 1 3d BD; Napo, Pano 1 3* JHKW; Pastaza, Sarayacu 1 ¢ ZMHU; Morona-Santiago, Bomboiza 1 6 JHKW; Zamora-Chinchipe, ““Loja” 1 ¢ RPM; No locality data | 6 BMNH. PERU: Loreto, Arcadia 2 6 USNM;; Iquitos 1 6 BMNH; 1 6 SMTD; Pebas 1 56 ZMHU; San Martin, Rio Chambirayacu, nr. Yurimaguas | 6 BMNH; Juanjui 4 d6 SMF; 1 ¢d RPM; Je- pelacio 1 ¢ RPM; Junin, La Merced 1 6 BMNH; Madre de Dios, 10 km N. of Puerto Maldonado | ¢ USNM; 30 km S.W. of Puerto Maldonado 2 6 USNM; Cuzco, No specific locality 1 d ZMHU. BOLIVIA: Lay Pag sRio) Songot3 id SBMNE 30 ZMEG: vives) SMT Ds, (Rigs Suapieilie G BMNH; Farinas | 6 BMNH; Mapiri 1 6 ZSM; Yungas 3 6 BMNH; No specific lo- cality 2 6 BMNH; No locality data 1 6 BMNH; 1 6 ZMHU; 1 3d ZSM; 1 6 SME BRAZIL: Amazonas, Sao Paulo de Oliven- ¢a-5' 6 eBMINEE ding), ZMBU aly do, SMTD; 8 3d SMF; Rio Tacana 1 ¢ SMTD; Fonte- boa | d6 BMNH; Tefé 3 6 BMNH; Mani- coré 1 d SMF; Parintins 1 6 BMNH; Up- per Amazon | 6 ZMHU; 3 6 BMNH; 1 ¢ SMTD; Pard, Belém 1 6 ZMHU: Mato Grosso, Cuiaba 4 6 BMNH; Melguira, 10 km S. of Diamantino 1 ¢d BMNH; No spe- cific locality 1 6 ZMHU; Distrito Federal, Planaltina 1 6 USNM; Parque do Gama 1 6*, 1F USNM; Goids, Vianépolis 1 6 SME GUYANA: No locality data 1 @ BMNH. SURINAM: No locality data 1 6 ZMHU. FRENCH GUIANA: Saint Laurent du Maroni, Saint Laurent du Maroni | 6 BMNH; Cayenne, Cayenne 2 6 BMNH; 1 36 RPM; Sinnamary 1 6* USNM; Saint Georges 2 6 BMNH; No locality data 3 6, 1 2 BMNH. TRINIDAD: St. Annes 1 @ BMNH. No locality data 1 d BMNH. ACKNOWLEDGMENTS I thank the following for giving me ac- cess to the riodinid collections in their care: P. Ackery (BMNH), B. Drummond (Flor- issant), M. Feyers (RPM), A. Hausmann (ZSM), W. Mey (ZMHU), L. and J. Miller (AME), W. Nassig (SMF), M. Nuss (SMTD), and R. Robbins (USNM). I thank the following for the financial assistance of field and museum research: Oxford Univer- sity (Poulton Fund), Cambridge University (Christ’s College), The Royal Entomologi- cal Society, Sigma Xi, Equafor, The Na- tional Geographic Society (Research and Exploration Grant #5751-96), the Smith- sonian Institution (two Postdoctoral Fellow- ships), and The National Science Founda- tion (Biotic Surveys and Inventories Grant). I thank the Pontificia Universidad Catdlica, the Museo Nacional de Ciencias Naturales and INEFAN, in Quito, for arranging the necessary permits for research in Ecuador. LITERATURE CITED Barcant, M. 1970. Butterflies of Trinidad and Tobago. Collins, London, 314 pp. Brévignon, C. and J.-Y. Gallard. 1998. Inventaire des Riodinidae de Guyane Frangaise III Riodininae: Riodinini. Description de nouveaux taxa. (Lepi- doptera). Lambillionea 98(1): 7—24. Bridges, C. A. 1988. Catalogue of Lycaenidae and Riodinidae (Lepidoptera: Rhopalocera). C. Bridg- es, Urbana, Illinois, 798 pp. . 1994. Catalogue of the Family-Group, Genus- Group and Species-Group Names of the Riodini- dae and Lycaenidae (Lepidoptera) of the World. C. Bridges, Urbana, Illinois, 1,113 pp. VOLUME 104, NUMBER 2 Callaghan, C. J. 1995. Les types des Riodinidae du Muséum national d’Histoire naturelle de Paris (Lepidoptera, Rhopalocera). Bulletin de la Société Entomologique de France 100(2): 153-155. Comstock, J. H. and J. G. Needham. 1918. The wings of Insects. American Naturalist 32: 231—257. d’Abrera, B. 1994. Butterflies of the Neotropical Re- gion, Part VI. Riodinidae. Hill House, Victoria, Australia. Pp. 880—1,096. DeVries, P. J. 1997. The Butterflies of Costa Rica and their Natural History. Volume II. Riodinidae. Princeton University Press, Princeton, 288 pp. DeVries, P. J., I. A. Chacon, and D. Murray. 1994. Toward a better understanding of host use and bio- diversity in riodinid butterflies (Lepidoptera). Journal of Research on the Lepidoptera 31(1): 103-126. Eliot, J. N. 1973. The higher classification of the Ly- caenidae (Lepidoptera): a tentative arrangement. Bulletin of the British Museum of Natural History (Entomology) 28: 373—506. Felder, C. and R. Felder. 1862. Specimen faunae lep- idopterologicae riparum fluminis Negro superioris in Brasilia septentrionali. Wiener Entomologische Monatschrift 6(3): 65—80. Hall, J. B W. 1999. A Revision of the Genus Theope: Its Systematics and Biology (Lepidoptera: Riodi- nidae). Scientific Publishers, Gainesville. 127 pp. . 2001. A revision of the new riodinid butterfly genus Dachetola (Lepidoptera: Riodinidae). Jour- nal of the New York Entomological Society 109(2):183-195. Hall, J. PR W. and D. J. Harvey. 2001. Phylogenetic revision of the Charis gynaea group (Lepidoptera: Riodinidae) with comments on historical relation- ships among Neotropical areas of endemism. An- nals of the Entomological Society of America 94(5):63 1-647. Hall, J. RP W. and K. R. Willmott. 1998. Three new 389 species of Riodinini from the cloud forests of Ec- uador (Lepidoptera: Riodinidae). Tropical Lepi- doptera 9(Suppl. 1): 22-26. . 2000. Patterns of feeding behaviour in adult male riodinid butterflies and their relationship to morphology and ecology. Biological Journal of the Linnean Society 69(1): 1—23. Harvey, D. J. 1987. The Higher Classification of the Riodinidae (Lepidoptera). Ph.D. Dissertation, Uni- versity of Texas, Austin, Texas, 216 pp. Hewitson, W. C. 1866. Illustrations of New Species of Exotic Butterflies, Selected Chiefly from the Col- lections of W. Wilson Saunders and William C. Hewitson, Vol. 3. J. Van Voorst, London. Kaye, W. J. 1921. A catalogue of the Trinidad Lepi- doptera Rhopalocera. Memoirs of the Department of Agriculture of Trinidad and Tobago 2: 1-163. Kirkpatrick, T. W. 1954. Notes on minor insect pests of Cacao in Trinidad. Part 2. Lepidoptera. Reports on Cacao Research 1953: 67-72. Klots, A. B. 1956. Lepidoptera, pp. 97-110. Jn Tuxen, S. L., ed. Taxonomists’s glossary of genitalia in insects. Munksgaard, Copenhagen, Denmark. Maza, R. G. de la and J. de la Maza. 1993. Mariposas de Chiapas. Gobierno del Estado de Chiapas, Mexico, 224 pp. Rebillard, P. 1958. Contribution a la connaissance des Riodinidae Sud-Américaines. Mémoires du Mu- séum d’ Histoire Naturelle (A) 15: 135-216. Stichel, H. E E. J. 1910a. Lepidoptera Rhopalocera. Fam. Riodinidae, pp. 1—238. Jn Wytsman, J., ed. Genera Insectorum 112A. J. Wytsman, Brussels. 1910b. Vorarbeiten zu einer revision der Riodinidae Grote (Erycinidae Swains.) (Lep. Rhop.). Berliner Entomologische Zeitschrift 55(1): 9-103. . 1930-31. Riodinidae, pp. 1—795. Jn Strand, E., ed. Lepidopterorum Catalogus 38—41. W. Junk, Berlin. PROC. ENTOMOL. SOC. WASH. 104(2), 2002, pp. 390—436 A REVISION OF THE ANASTREPHA SERPENTINA SPECIES GROUP (DIPTERA: TEPHRITIDAE) ALLEN L. NORRBOM Systematic Entomology Laboratory, PSI, Agriculture Research Service, U.S. Depart- ment of Agriculture, Bldg. 005, Rm. 137, BARC-West, 10300 Baltimore Ave., Beltsville, MD 20705-2350, U.S.A. (e-mail: anorrbom @sel.barc.usda.gov) Abstract.—The Anastrepha serpentina species group is revised, with a key, descrip- tions, illustrations, and phylogeny provided for the 11 species recognized: A. anomala Stone, anomoiae, n. sp., bistrigata Bezzi, normalis, n. sp., ocresia (Walker), ornata Aldrich, pseudanomala, n. sp., pulchella, n. sp., pulchra Stone, serpentina (Wiedemann), and striata Schiner. Lectotypes are designated for Anastrepha bistrigata Bezzi, Dacus serpentinus Wiedemann, Urophora vittithorax Macquart, and Dictya cancellaria Fabri- cius. Key Words: Anastrepha, serpentina, striata, ornata, bistrigata, Tephritidae, fruit flies, taxonomy, phylogeny, identification, distribution, host plants The serpentina group comprises 11 spe- cies, including two of the most significant pests within the genus Anastrepha and sey- eral others of lesser agricultural importance. Anastrepha serpentina (Wiedemann) is an especially important pest of sapotaceous fruits, such as star apple, sapotes, and nis- pero, and A. striata Schiner is among the worst pests of Myrtaceae, especially gua- vas, although both species also sometimes attack mango, mombins, citrus, and various other fruits (see Norrbom, in press). As de- limited here, the serpentina group compris- es four previously included species, three species formerly recognized as the striata group (see Norrbom et al. 1999b), and four newly described species. The serpentina group is most diverse in Central America and northern South Amer- ica, although A. serpentina and striata ex- tend north to Mexico or southern Texas and A. serpentina extends south to Argentina, and A. ocresia is known only from the An- tilles and southern Florida. Host plants are known for eight of the 11 species. Although the host range for the species group is very diverse, the native hosts are mainly in three families. Two spe- cies (A. anomala Stone and A. normalis, n. sp.) have only species of Apocynaceae as known hosts, whereas A. pulchra Stone is known only from Sapotaceae, which also includes the main hosts of the generalist species A. serpentina. The only native host known for A. ocresia (Walker) belongs to the Myrtaceae, but it has been reared from an exotic species of Sapotaceae. The only known native host plant of A. ornata A\- drich, and most of those of A. striata and A. bistrigata Bezzi, belong to the family Myrtaceae. MATERIALS AND METHODS Terminology follows the glossary of White et al. (1999). Acronyms for institu- tions where specimens are deposited follow Thompson (1999). Phylogenetic analysis was conducted using Hennig86 (© J. S. VOLUME 104, NUMBER 2 Farris) and the cladogram figures were pro- duced using Winclada (© K. C. Nixon). ile i) KEY TO THE SPECIES OF THE ANASTREPHA SERPENTINA GROUP Wing with C-band and S-band well separated (Fig. 21); middle section of S-band very nar- row, distal section broad; proximal arm of V-band connected along posterior wing mar- gin to basal extension of S-band along vein A,+Cu,. Thorax with darker areas mostly brown (Fig. 1D). Oviscape 3.40—3.95 mm long. Aculeus tip broadly triangular (Fig. 11A). Colombia, Ecuador C-band and S-band connected (Figs. 2A—H, 3A-—G) or at most very narrowly separated (Fig. 3H); middle section of S-band broad, or if narrow, distal section also narrow; proximal arm of V-band not connected to basal exten- sion of S-band along posterior wing margin except sometimes in A. pulchella. Other char- AGLEDS VallaDle~ oa) 5 fs cs Soe ees ee: 2 . Wing cell br with hyaline area posterior to pterostigma broadly extended to vein R,,; (Figs. 2D, 3F—H). Abdomen with all tergites mostly yellow to orange, without brown markings (Figs. 1A, H). Thoracic pleuron without brown areas. Epandrium with narrow, V-shaped, medial indentation in dorsal pos- terior margin; lateral surstylus with rounded subapical lateral lobe (Figs. 4K 7F). Aculeus tip broadly triangular, at least 0.17 mm wide (BigsamlVAI@)r cary. fs lees a ar ages Gees Cue Cell br entirely infuscated (Figs. 2C, 3A—C) or with hyaline area not extended to vein R,y,; (Figs. 2A—B, E-H, J, 3D-E). Abdomen with brown bands or spots on at least syntergite 1+2 and tergite 3, or more extensively brown (Figs. 1B—G). Thoracic pleuron usually with at least anepimeron partially brown (orange in A. ocresia). Epandrium with dorsal poste- rior margin evenly rounded or at most with slight indentation; lateral surstylus without rounded subapical lateral lobe. Aculeus tip variable in shape . Oviscape less than 2.8 mm long, less than 0.90 times mesonotum length (Fig. 1H). Acu- leus less than 2.5 mm long. Scutum with brown areas usually interrupted at transverse suture; setulae usually strongly contrasting white and dark brown, absent on narrow area lateral to postsutural nonmicrotrichose area. Distal section of S-band slightly narrowed to moderately broad, at apex of vein R,,, 0.44— 0.63 times width of cell r,,, (Figs. 3F—H). Widespread, Mexico to Bolivia and Brazil AR AE ie Ree ee oe A. striata Schiner ...A. ornata Aldrich — QOviscape more than 3.0 mm long, more than 0.90 times mesonotum length (Fig. 1A). Acu- leus more than 3.0 mm long. Scutum with brown area narrowed but uninterrupted at transverse suture; setulae yellow and brown, less dense but more or less continuous lateral to postsutural nonmicrotrichose area. Distal section of S-band slender, at apex of vein R,,; 0.36—0.44 times width of cell r,,, (Fig. 2D). Southern Brazil connected basally, cell br without hyaline area in apical half (Figs. 2C, 3A—C); C-band with yellow area posterior to pterostigma small, not extending beyond cell r, or level of basal third of pterostigma; ratio of length of section of vein M between crossveins R-M and DM- Cu: length of DM-Cu 0.48—0.69; distal sec- tion of S-band slender, its width at apex of vein R,,, less than 0.45 width of cell r,,; .. C-band and S-band less broadly connected, cell br with at least a small hyaline area in apical half (Figs. 2A—B, E-H, J, 3D-E); C-band with yellow area posterior to pteros- tigma large, extending into cells r, and r,,, at least to level of midlength of pterostigma; ra- tio of length of section of vein M between crossveins R-M and DM-Cu: length of DM- Cu 0.68—0.91; distal section of S-band vari- ablemimpwiGdthy ¢...o0. = «ac fees See = oe . Wing with proximal arm of V-band with an- terior end more proximal than posterior end and connected to S-band near middle of crossvein R-M (Fig. 2C); posterior end not extended basally along posterior wing margin. Aculeus 3.66 mm long; tip 0.86 mm long, serrate only on apex (Figs. 9B, 10D). Panama, Colombiale,.. .- A. anomoiae Norrbom, n. s Proximal arm of V-band with anterior end at least slightly more distal than posterior end, and separated from S-band or connected along vein R,,; (Figs. 3A—C); posterior end extended basally along posterior wing margin. Aculeus 4.00—5.70 mm long; tip less than 0.50 mm long, nonserrate or more than half serrate (acs: OIEI) 22.23.55 pee 9 ee . Wing cell r, with hyaline spot between S-band and vein R,,, (Fig. 3A); apex of proximal marginal hyaline spot in cell r, aligned prox- imal to crossvein R-M. Orbital plate yellow (Fig. 1E). Aculeus tip 0.24—0.26 mm long, more than half serrate (Fig. 10I). Panama . . Bree tt oo ens acre A. pulchella Norrbom, n. s Cell r, without hyaline spot between S-band and vein R,,,; (Figs. 3B—C); apex of marginal hyaline spot in cell r, aligned with or distal to crossvein R-M. Orbital plate with triangu- lar brown mark (Fig. 1F). Aculeus tip 0.33- eb ioe A. bistrigata Bezzi . Wing with C-band and S-band very broadly PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON es) Fig. 1. Dorsal habitus. A, Anastrepha bistrigata (USNMS53046, Brazil: ex. USP lab colony). B, A. normalis (USNM47450, Venezuela). C, A. ocresia (USNM51938, Dominican Republic: 2 km N Bayahibe). D, A. ornata (USNMS51868, Colombia: El Retiro). E, A. pulchella (USNM52982, Panama: Barro Colorado I.). EK A. pulchra (USNM00052975, Panama: La Campana). G, A. serpentina (USNM47625, Venezuela: Pueblo Nuevo). H, A. striata (USNM51843, Costa Rica: Jardin Las Cruces). VOLUME 104, NUMBER 2 0.46 mm long, nonserrate (Fig. 10J). Panama, Venezuela, n. Brazil A. pulchra Stone . Mesonotum with brown areas extensive, ex- tending to or nearly to anterior margin of scu- tum (Figs. 1B, E—G). Scutum microtrichose except presutural medial bare area. Wing with distal section of S-band slender to moderately broad, at apex of vein R,,; 0.32—0.64 times width of cell r,,,; distal arm of V-band usu- ally absent (except in sp. from Mexico) ... 8 Mesonotum with brown markings only near scuto-scutellar suture (Fig. 1C). Scutum with- out microtrichia except on extreme lateral margin. Wing with distal section of S-band very broad, at apex of vein R,,, at least 0.73 times width of cell r,,, (Fig. 2H); distal arm of V-band present from vein M to posterior wing margin. Greater Antilles, USA (southern Florida) A. ocresia (Walker) . Abdomen orange with brown bands on syn- tergite 1+2, tergite 3, and sometimes tergite 4 (Figs. 1B, E-F). Wing with distal section of S-band usually slightly narrowed to moder- ately broad, at apex of vein R,,, 0.47—0.64 times width of cell r,,, (Figs. 2A, E-G, J), rarely (Brazilian anomala) more slender (Fig. 2B). Oviscape 3.70—5.51 mm long, 1.02—1.56 times mesonotum length. Aculeus tip variable Abdomen mostly brown with T-shaped me- dial yellow mark (Fig. 1G). Distal section of S-band slender, at apex of vein R,, , 0.32—0.43 times width of cell r,,, (Figs. 3D—E). Ovis- cape 2.58-3.91 mm long, 0.79—1.02 times mesonotum length. Aculeus tip 0.37—0.46 mm long, 0.14—0.17 mm wide, finely serrate on more than distal half (Fig. 11B). Wide- spread, Mexico to Argentina Ree oe cons A. serpentina (Wiedemann) . Wing with distal arm of V-band complete (Fig. 2E). Southern Mexico . . Distal arm of V-band absent or at most con- sisting of small, faint spot in cell m (Figs. 2A-B, F-G, J) . . Anastrepha sp. . Wing with middle section of S-band entirely brown anterior and distal to crossvein R-M (Figs. 2A-B, J). Oviscape 3.70-5.51 mm long, 1.02—1.56 times mesonotum length. Aculeus 3.62—5.51 mm long; tip more than 2 times as long as wide, finely serrate on distal half or more (except some Brazilian anoma- la), 0.19-0.40 mm long (Figs. 10A—C, H). Lateral surstylus sometimes with strong ba- solateral lobe (Figs. 4A—B) Middle section of S-band mostly orange, in- cluding anterior and distal to crossvein R-M, brown only narrowly on margins (Figs. 2F— G). Oviscape 3.79—4.41 mm long, 1.07—1.21 times mesonotum length. Aculeus 3.62—4.14 mm long; tip less than 2 times as long as wide, nonserrate, 0.23—0.28 mm long (Fig. 1OF). Lateral surstylus without strong baso- lateral lobe (Figs. SA-B). Panama, Venezuela A. normalis Norrbom, n. sp. 11. Aculeus tip 0.09—0.12 mm wide, gradually ta- pered (Figs. 9A, 1OA—C). Oviscape 4.49-5.51 mm long, 1.44—1.56 times mesonotum length (except 3.70 mm long, 1.02 times mesonotum in Bahia, Brazil female). Aculeus usually 4.37-5.51 mm long (3.62 mm long in Bahia female). Lateral surstylus with strong baso- lateral lobe (Figs. 4A—B). Panama, Brazil A. anomala Stone — Aculeus tip 0.15 mm wide, with nonserrate basal part nearly parallel sided (Figs. 9D, 10H). Oviscape 4.11—4.28 mm long, 1.11— 1.18 times mesonotum length. Aculeus 4.16 mm long. Male unknown. Costa Rica, Pana- ma A. pseudanomala Norrbom, n. sp. Anastrepha anomala Stone (Figs. 2A—B, 4A-B, 8A, 9A, 10A-C) Anastrepha anomala Stone 1942: 29; Foote 1967: 7 [in catalog]; Steyskal 1977: 8 [in key]; Zucchi 1983: 280, 2000: 238 [Bra- zil, Bahia]; Norrbom et al. 1999a: 77 [in catalog]; Norrbom, in press [host data- base]. Recognition.—Anastrepha anomala is one of the species of the serpentina group with a hyaline area in cell br posterior to the pterostigma that is not extended to vein R,,;. Of the other species with this char- acter, A. anomala is most likely to be con- fused with A. normalis and A. pseudanom- ala. It differs from both species by its more slender, tapered aculeus tip (no more than 0.12 mm wide), and at least from A. nor- malis in having a strong basolateral lobe on the lateral surstylus (the male is unknown for A. pseudanomala). \t further differs from A. normalis in having the middle sec- tion of the S-band solidly dark brown an- terior and distal to crossvein R-M, and the aculeus tip finely serrate on the distal half or more (except in Maranhao, Brazil fe- male). Most females have slightly longer terminalia (Oviscape more than 1.4 times mesonotum length; aculeus more than 4.3 mm long) than in A. normalis and A. pseu- 394 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Wing. A—B, Anastrepha anomala (USNMS52954, Panama: La Campana. USNM53107, Brazil: Cruz das Almas). C, A. anomoiae (holotype, USNM50456, Colombia: Guaguaqui). D, A. bistrigata (USNM53050, Brazil: Belo Horizonte). E, A. sp. (USNM52942, Mexico: Tapachula). F—G, A. normalis (USNM52938-9, Ven- ezuela: Marcillal de la Costa). H, A. ocresia (USNMS51712, Cuba: Bolondron). I, A. ornata (holotype, USNM52985, Ecuador: Banos). J, A. pseudanomala (USNM50455, Costa Rica: 20 km. S Upala). danomala, but they are shorter in a single tae dark brown. Head: Yellow except ocel- female from Bahia, Brazil (see Comments). lar tubercle brown. Facial carina, in profile, Description.—Largely dark orange to concave. 3—5, usually 4, frontal setae; 1—2 dark red brown with yellow markings. Se- orbital setae, posterior seta well developed VOLUME 104, NUMBER 2 395 Bigs: Wing. A, Anastrepha pulchella (USNM52978, Panama: Arraijan). B—C, A. pulchra (Panama: El Cermeno), male (USNM51713), female (USNMS51714). D-E, A. serpentina (USNM53098, Peru: Malambo. and USNMS53097, Trinidad: nr. Teteron). F—H, A. striata (Colombia: Muhic. USNM53100, Trinidad: La Brea; and USNMS51718, Brazil: 62 km SE Ariquemes). if present. Ocellar seta weak, short to mi- nute. Antenna extended 0.75—0.90 distance to lower facial margin. Thorax: Mostly or- ange to dark red brown with following ar- eas yellow and distinctly contrasting: post- pronotal lobe; single medial and paired sub- lateral vittae on scutum, the slender medial vitta extended nearly full length of scutum, broadened posteriorly, but extended later- ally less than half distance from acrostichal seta to dorsocentral seta; sublateral vitta ex- tended from transverse suture to posterior margin, including intra-alar seta; scutellum except extreme base (brown area well sep- arated from basal seta); propleuron; dorsal margin and anteroventral corner of anepis- ternum; greater ampulla; dorsal margin of katepisternum; katepimeron; and most of anatergite and katatergite. Mesonotal pat- tern similar to A. normalis (see Fig. 1B); broad area bordering medial vitta, shorter and narrower area bordering mesal margin of sublateral vitta, and large sublateral pre- sutural area orange. Darker areas of anepis- ternum and katepisternum orange; those of anepimeron, meron, and katatergite mostly 396 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON dark brown. Subscutellum and mediotergite dark brown, narrowly dark orange medially. Mesonotum 3.06—3.66 mm long. Scutum microtrichose except for broad presutural bare area extended about %4 distance to transverse suture; setulae mostly yellow, brown on parts of darker areas. Katepister- nal seta weak, slightly to much shorter than postocellar seta, yellowish to red brown. Wing (Figs. 2A—B): Length 6.67—7.90 mm. Vein M strongly curved apically; section between BM-Cu and R-M 2.04—2.62 times as long as section between R-M and DM- Cu; section between R-M and DM-Cu 0.74—0.85 times as long as DM-Cu (0.69 times in Bahia female). Crossvein DM-Cu oblique, with anterior end more distal than posterior end. Pattern mostly dark brown. C-band and S-band broadly connected in cells r,,, and br, but separated basally by hyaline area in posterior half of br aligned with pterostigma, hyaline basal sixth of cell dm, and hyaline to yellowish area covering all of cell bm. C-band yellowish to subhya- line in cell be and posterior half to % of cell c; with large yellowish area in base of cell sc and cells r, and r,,, posterior to pteros- tigma, extending distally to or almost to level of apex of vein R,. S-band with large yellow area in cell dm usually extending into cell br and sometimes nearly touching crossvein R-M; rest of band dark brown, including areas distal to and anterior to R-M; distal section moderately broad, at apex of vein R,,,; 0.33—0.61 times width of cell r,,; (0.49-0.61 times in Panamania specimens; 0.45 times in Maranhao, Brazil female; in Bahia female, slender at apex of R,,3, 0.33—0.38 times width of cell r,,3, but slightly broader along vein R,,;); not ex- tended to apex of vein M. Hyaline spot in cell r, nearly triangular, extended well into cell r,,, and sometimes to vein R,,.;; its apex aligned with R-M or usually slightly basal to it. V-band with distal arm absent; proximal arm separated from S-band, ex- tended to vein R,,<,; extended basally along posterior wing margin almost to vein A,+Cu, but not connected to extension from base of S-band. Abdomen: Orange with yellow and brown markings. Synter- gite 1+2 and tergites 3—4 each with band, brown laterally, orange medially, on tergite 4 often entirely orange; posterior margin yellow, narrowing laterally and on succes- sive tergites. Tergite 5 and female tergite 6 orange. Male terminalia (Figs. 4A—B): Dor- sal posterior margin of epandrium evenly convex. Lateral surstylus moderately long; in lateral view slightly curved; in posterior view, with strong basolateral lobe, main part narrowly triangular, acute apically. Proctiger with lateral fold separating scler- otized areas. Phallus 6.2—7.1 mm _ long; 1.94—2.04 times as long as mesonotum. Glans 0.50—0.55 mm long; acrophallus rel- atively stout. Female terminalia: Oviscape 4.49-5.58 mm long, 1.44—1.56 times as long as mesonotum (3.70 mm, 1.02 times mesonotum length in Bahia female). Ever- sible membrane (Fig. 8A) with 35—40 large, hook-like dorsobasal scales in triangular pattern. Aculeus 3.62—5.51 mm long (4.37— 5.12 mm in Panamanian females; 5.51 mm in Maranhao female; 3.62 mm in Bahia fe- male); tip (Figs. 9A, 1OA—C) 0.19—0.32 mm long, 0.09—0.12 mm wide (0.27—0.32 mm long, 0.105—0.12 mm wide in Pana- manian females; 0.24 mm long, 0.10 mm wide in Bahia female; 0.19 mm long, 0.09 mm wide in Maranhao female), gradually tapered, very finely serrate on apical 0.60— 0.67 in Panamanian females, on apical 0.50 in Bahia female, nonserrate in Maranhao fe- male. Spermathecae ovoid. Egg: Similar in shape to A. serpentina, posterior end elon- gate and tapered. Anterior end without lobe (i.e., micropyle at apex). Length 1.75—1.82 mm, broadest width 0.20—0.22 mm. Distribution.—Anastrepha anomala is known with certainty only from Panama and Brazil, although the status of the Bra- zilian populations needs further study (see Comments). Records from Venezuela (Car- aballo 1981) were based on misidentifica- tions of A. normalis. The record from Gua- temala (Norrbom et al. 1999a) is uncon- firmed. It was based on a male (J. Lopez, VOLUME 104, NUMBER 2 37 Fig. 4. Epandrium and surstyli, lateral and posterior views. A—B, Anastrepha anomala (USNM52956, Pan- ama: Barro Colorado I.). C-D, A. anomoiae (USNM50457, Panama: El Valle). E-E A. bistrigata (USNM53047, Brazil: Univ. of Sao Paulo culture). 398 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON personal communication) that might be conspecific with the undetermined male from Mexico reported here. Biology.—The holotype and some of the paratypes were reared from seeds of Lac- mellea panamensis (Woodson) Markgr. (Apocynaceae) (Stone 1942, as Zschokkea panamensis). Comments.—The description of the egg is based on a sample of five dissected from the abdomen of the female from Arraijan, Panama (USNM52951). The aculeus tip is very finely serrate in the Panamanian spec- imens, but more extensively so than indi- cated by Stone (1942). The status of the populations represented by single females from Bahia and Maran- hao, Brazil (R. A. Zucchi, personal com- munication) needs further study. They may be distinct species. The Bahia female dif- fers from specimens from Panama in hav- ing shorter terminalia (oviscape 3.70 mm long and 1.02 times mesonotum length; aculeus 3.62 mm long), a slightly shorter, blunter, and less serrate aculeus tip, and a slightly narrower distal section of the S-band (less than 0.40 times width of cell T,3 at apex of vein R,,,). It also has a spur vein in cell r,,; anterior to crossvein DM- Cu, but this is probably an anomaly. The female from Maranhao has relatively long terminalia (oviscape 5.51 mm long and 1.56 times mesonotum length; aculeus 5.51 mm long) but the aculeus tip is short (0.19 mm) and nonserrate. The width of the distal sec- tion of its S-band is intermediate between the Panamanian and Bahian specimens. Type data.—Holotype 2 (USNM52966), PANAMA: Panama: Barro Colorado Island [9°9’17N 79°50'53W)], reared ex. seeds of Zschokkea panamensis [= Lacmellea pan- amensis|, 17 Mar 1937, J. Zetek 3814 [ex- amined]. Other specimen data.—BRAZIL: Ba- hia: Cruz das Almas, Fazenda Chapadin- ha, 14 Jun 1978, A. S. Nascimento, 1 2 (ESALQ, USNMS53107). Maranhéo: Ita- pecuru-Mirim, 2000, R. N. S. Lemos, 1 @ (ESALQ USNM31431). PANAMA: Pan- ama: Arraijan [8°57’N 79°39'W], 9 Jun 1949, J. Zetek 5401, 1 d (USNM52950); same, 21 «Sep. 19502". Zetek: S473 (USNM52951); same data as holotype ex- éept Janeil939. JinZetek 43829: 47 Gaia paratypes (USNM52954-8), | 2 paratype (TAMU USNM52959); same, 4—7 Jan 1937, J. Zetek 3777, 2 5 paratypes (TAMU USNM52952; BMNH USNM53115); same, 17-23 Mar 1937, J. Zetek 3814, 1 d par- atype (USNM52953), 1 6 paratype IOC USNMS53105). Anastrepha anomotae Norrbom, new species (Figs. 2C, 4C—D, 8B, 9B, 10D) Recognition.—Anastrepha anomoiae is one of three species of the serpentina group with the C- and S-bands broadly fused so that there is no hyaline area in the middle of cell br. It differs from all other known species of Anastrepha in having the proxi- mal arm of the V-band strongly oblique, with the anterior end more proximal than the posterior end, and connected to the S-band near the middle of crossvein R-M. Crossvein DM-Cu, which is covered by this band, is also oblique in this direction or per- pendicular to the long axis of the wing. The posterior end of the V-band does not extend basally along the posterior wing margin. The extremely long aculeus tip (0.86 mm long), which tapers slightly subbasally and is bluntly rounded distally, is also distinc- tive. The very slender distal section of the S-band, and the dark colors of the wing pat- tern and the darker areas of the body are additional useful diagnostic characters. Description.—Largely dark orange to dark red brown with yellow markings. Se- tae red brown to dark brown. Head: Mostly yellow. Frons, face and gena orange to red- dish brown in holotype (possibly discolor- ation). Ocellar tubercle brown. Occiput in holotype with paired small brown spot dor- sally and larger spot ventrally, extending to postgena. Facial carina, in profile, concave. 4—5 frontal setae; 2 orbital setae, posterior seta well developed. Ocellar seta weak. An- VOLUME 104, NUMBER 2 Fig. 5. Epandrium and surstyli, lateral and posterior views. A-B, Anastrepha normalis (USNM52938, Ven- ezuela: Marcillal de la Costa). C-D, A. ocresia (USNM52992, Cuba: Guantanamo airbase). tenna extended 0.75—0.85 distance to lower facial margin. Thorax: Mostly orange brown to red brown with following areas yellow and distinctly contrasting: postpron- otal lobe; single medial and paired subla- teral vittae on scutum, the slender medial vitta extended nearly full length of scutum, broadened posteriorly, but extended later- ally only slightly beyond level of acrosti- chal seta (at most half distance to level of dorsocentral seta); sublateral vitta extended from transverse suture to or almost to pos- terior margin, including intra-alar seta; scu- tellum except extreme base (brown area well separated from basal seta); dorsal mar- gin of anepisternum; most of greater am- pulla; large dorsal spot on katepisternum; katepimeron; and most of anatergite and ka- tatergite. Mesonotal darker areas mostly dark orange, but anterolateral corner, lateral margin of presutural area, margin along me- sal side of lateral vitta, somewhat U-shaped posterior area, postsutural lateral areas, and notopleuron dark brown; without orange vitta on dorsocentral line. Darker area of anepisternum about half to entirely brown. AOD PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Anepimeron mostly dark brown. Darker ar- eas of katepisternum, meron and katatergite dark orange or dark brown. Subscutellum and mediotergite dark orange to dark brown. Mesonotum 3.49—4.00 mm long. Scutum microtrichose except for broad me- dial anterior area extended %—% distance to transverse suture; setulae mostly brown, but yellow on medial stripe and in broad pre- sutural area. Katepisternal seta weak, no longer than postocellar seta, yellowish to pale brown. Wing (Fig. 2C): Length 7.5— 9.0 mm. Vein M moderately curved apical- ly; section between BM-Cu and R-M 2.4— 2.9 times as long as section between R-M and DM-Cu; section between R-M and DM-Cu 0.63—0.69 times as long as DM-Cu. Crossvein DM-Cu perpendicular to long axis of wing or oblique, with anterior end more proximal than posterior end. Pattern mostly dark brown. Basally with large in- fuscated area formed by fusion of C-band and base of S-band, which are separated only by hyaline to yellowish area covering much of cell bm; basal area mostly brown, but most of cells be and c and part of cell bcu yellowish to subhyaline, small basal spot in cell dm and small area in cells sc and r, posterior to bend in vein sc yellow; cell r,,,; basal to R-M and all of cell br brown. Remainder of S-band brown; distal section very slender, at apex of vein R,,, 0.27—0.33 times width of cell r,,,; closely following vein R,,, so that hyaline spot in cell r, is elongate along costa; not extended to apex of vein M. Hyaline spot in cell r, extended to vein R,,,;; its apex aligned with R-M. V-band with distal arm absent; prox- imal arm very slender, strongly oblique and connected to S-band in cell r,,; near middle of crossvein R-M; not extended basally along posterior wing margin. Abdomen: Orange with yellow and dark brown bands. Syntergite 1+2 with medial to subapical brown band, broader in male; posterior margin yellow, narrowing laterally. Tergites 3 and 4 with broad brown band, narrowed medially; posterior margin yellow, narrow- ing laterally. Male tergite 5 orange. Male terminalia (Figs. 4C—D): Dorsal posterior margin of epandrium convex with slight medial angle. Lateral surstylus moderately long; in lateral view very slightly curved; in posterior view with moderate basolateral lobe, main part strongly tapered, slender apically. Proctiger with lateral fold separat- ing sclerotized areas. Phallus 4.58 mm long; 1.31 times as long as mesonotum. Glans 0.65 mm long; acrophallus relatively stout. Female terminalia: Oviscape of ho- lotype damaged, but probably more than 3.7 mm long. Eversible membrane (Fig. 8B) with 60—70 large, hook-like dorsobasal scales in triangular pattern. Aculeus 3.66 mm long; base expanded; tip (Figs. 9B, 10D) 0.86 mm long, 0.15 mm wide, non- serrate except for very fine serrations on distal fifth, tapered rapidly subbasally to 0.12 mm wide, then parallel sided, extreme apex broad and bluntly rounded. Sperma- thecae not examined. Distribution.—Anastrepha anomoiae 1s known from Panama and Colombia. Comments.—The terminalia of the ho- lotype were previously dissected and slide mounted. Abdominal segments 4—6 were damaged and mostly lost and the oviscape was broken, but the other parts of the ter- minalia are in good condition. The sper- mathecae are not on the slide and may have been lost or they may be inside the remain- der of the abdomen. Biology.—The host plants of this species are unknown. Etymology.—The name of this species, a noun in apposition, is derived from the ge- nus Anomoia, which it resembles in wing pattern. Type data.—Holotype 2 (USNM50456), COLOMBIA: Boyaca: Guaguaqui [proba- bly Rio Guaguaqui, 5°46’N 74°29'W] [without date or collector]. Paratype and? ee= totype 2 Anastrepha bistrigata Bezzi by Norrbom 2001.’ Bezzi (1919a, b) stated that he described this species from i d and 2 2 syntypes from Brazil: Sao Paulo: Bau- ru, reared ex. fruit of “‘araxa” [Psidium gui- neense Sw.|, A. A. Barbiellini. The para- lectotypes are labeled similar to the lecto- type, except that the male has the number **59”’ and the female “57,” and the latter also has “‘Brasile’’ on its second label. There is also a damaged male of another Anastrepha species, possibly A. fraterculus, on the pin of the female. Other specimen data.—BRAZIL: Goias: Veadeiros, 30 Apr 1956, FE S. Truxal, 1 @ (LACM USNMS53051). Minas Gerais: Belo Horizonte, reared from goiabas [Psidium guajava L.], 3 Sep 1934, O. Monte, 1 @ (USNM53050). Sao Paulo: Sao Paulo, on guava, 11 Dec 1931, M. Kisliuk & C.E. Cooley, 1 2 (USNM53049); Univ. of Sao Paulo (Malavasi/Morgante) culture initiated from larvae reared from guava from Cam- pinas; Nov” 1986; «GJ.rSteck,ailinc, Mie (USNMS53047-8); same, 5 Jun 1991, 1 @ (USNM53046). VOLUME 104, NUMBER 2 Anastrepha normalis Norrbom, new species (Figs. 1B, 2F—G, 5A—B, 8C, 9C, 10F) Anastrepha anomala: Stone 1942: 29 [in part]; Caraballo 1981: 60 [Venezuelan specimens, host]. Anastrepha sp.: Boscan et al. [host]. Anastrepha n. sp. nr. anomala: McPheron et al. 1999: 346. 1980: 61 Recognition.—Anastrepha normalis is one of the species of the serpentina group with a hyaline area in cell br posterior to the pterostigma that is not extended to vein R,,5. It differs from the other species with this character in having a paler wing pat- tern. In particular, the middle section of the S-band anterior and distal to crossvein R-M is orange brown with only narrow dark brown margins. A. normalis further differs from all of these species, except A. ornata and one female of A. anomala (Maranhao, Brazil) in lacking fine serrations on the acu- leus tip (Fig. 1OF). It is most likely to be confused with A. anomala or A. pseudan- omala, from which it further differs in hav- ing the aculeus tip less than twice as long as wide. It differs further from A. anomala in lacking the basolateral lobe on the lateral surstylus and from A. pseudanomala in having the distal section of the S-band slightly broader and sometimes touching the apex of vein M. Description.—Largely dark orange to dark red brown with yellow markings. Se- tae dark brown. Head: Yellow except ocel- lar tubercle brown. Facial carina, in profile, concave. 3—5, usually 4, frontal setae; usu- ally 2 orbital setae, posterior seta absent on one side in 4 and on both sides in 3 of 29 specimens examined. Ocellar seta weak and small or minute. Antenna extended 0.70— 0.85 distance to lower facial margin. Tho- rax (Fig. 1B): Mostly dark orange to dark red brown with following areas yellow and distinctly contrasting: postpronotal lobe; single medial and paired sublateral vittae on scutum, the slender medial vitta extended 405 nearly full length of scutum, broadened posteriorly, but extended laterally only slightly beyond level of acrostichal seta (at most half distance to level of dorsocentral seta); sublateral vitta extended from trans- verse suture to or almost to posterior mar- gin, including intra-alar seta; scutellum ex- cept extreme base (brown area well sepa- rated from basal seta); propleuron; dorsal margin and anteroventral corner of anepis- ternum; greater ampulla and often border- ing area of anepimeron; dorsal margin of katepisternum; katepimeron; and most of anatergite and katatergite. Mesonotal pat- tern with broad orange area bordering me- dial vitta, large orange sublateral presutural area, usually with narrow orange area of varying length bordering mesal margin of sublateral vitta. Darker areas of anepister- num and anepimeron mostly dark brown. Those of meron and katatergite dark orange or dark brown. Darker area of katepister- num orange. Subscutellum and mediotergite dark brown, narrowly orange medially. Me- sonotum 3.41—3.74 mm long. Scutum mi- crotrichose except for broad presutural bare area extended about midway to transverse suture; setulae mostly yellow. Katepisternal seta weak, shorter than postocellar seta, yel- lowish. Wing (Figs. 2F—G): Length 7.24— 8.15 mm. Vein M strongly curved apically; section between BM-Cu and R-M 2.51-— 2.75 times as long as section between R-M and DM-Cu; section between R-M_ and DM-Cu 0.69—0.76 times as long as DM-Cu. Crossvein DM-Cu oblique, with anterior end more distal than posterior end. Pattern mostly brown and pale to dark orange brown. C-band and S-band broadly con- nected in cell br and often in r,,,;, but sep- arated basally by hyaline area in posterior *; of br aligned with pterostigma, usually a small basal hyaline area in cell dm, and hy- aline to yellowish area covering all of cell bm. C-band with cell be and posterior half to % of cell c yellowish to subhyaline, and large yellow area in base of cell sc and cells r, and r,,, posterior to pterostigma, gradu- ally merging with orange brown distal part 106 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Ven ° S00 33 » A an a'n 7% Fig. 8. Eversible membrane, dorsobasal scales. A, Anastrepha anomala (USNM52951, Panama: Arraijan). B, A. anomoiae (holotype). C, A. normalis (USNM52939, Venezuela: Marcillal de la Costa). D, A. pseudanomala (USNM50455, Costa Rica: 20 km. S Upala). VOLUME 104, NUMBER 2 of band; margins of band, including most of pterostigma moderate brown. S-band largely yellow to orange in cell dm, dark- ening to orange brown anteriorly but mid- dle section of band brown only narrowly on margins; distal section of band moderately broad, at apex of vein R,,, 0.51—0.64 times width of cell r,,3;; separated from or ex- tended to apex of vein M. Hyaline spot in cell r, nearly triangular, extended well into cell r,,,, Sometimes reaching vein R,,5; its apex aligned slightly basal to R-M. V-band with distal arm usually absent, rarely rep- resented by faint spot in cell M (1 dg, 1 9 from Venezuela); proximal arm extended to vein R,,;, usually separated from S-band but occasionally narrowly connected along R,.5; extended basally along posterior wing margin often almost to vein A,+Cu, but not connected to extension from base of S-band. Abdomen (Fig. 1B): Orange with yellow and brown markings. Syntergite 1+2 and tergite 3 each with brown band. Tergites 4—6 mostly orange. Posterior mar- gin of tergites yellow, this area narrower laterally and also narrower on successive tergites, nearly absent on male tergite 5 and female tergite 6. Male terminalia (Figs. 5A-B): Dorsal posterior margin of epan- drium evenly convex. Lateral surstylus moderately long; in lateral view very slight- ly curved; in posterior view without strong basolateral lobe, main part narrowly trian- gular, acute apically. Proctiger with lateral fold separating sclerotized areas. Phallus 5.11-5.42 mm long; 1.50—1.53 times as long as mesonotum. Glans 0.40 mm long; acrophallus relatively stout. Female termin- alia: Oviscape 3.79—4.41 mm long, 1.07— 1.21 times as long as mesonotum. Eversible membrane (Fig. 8C) with 40-50 large, hook-like dorsobasal scales in triangular pattern. Aculeus 3.62—4.14 mm long; tip (Figs. 9C, 10F) 0.23—0.28 mm long, 0.13-— 0.15 mm wide, gradually tapered and non- serrate. Spermathecae ovoid or teardrop shaped. Distribution.—Anastrepha normalis is known from Panama and Venezuela. 407 Etymology.—The name of this species is an adjective referring to the more normal color of the wing pattern (i.e., more com- mon in Anastrepha in general) compared to that of the closely related A. anomala. Biology.—Lacmellea panamensis (Wood- son) Markgr. (Apocynaceae) is probably the only known host. Boscan et al. (1980) reared an Anastrepha sp. from “‘chirica,”” which they reported as Jaquinia revoluta Jacq. (= J. armillaris Jacq.) (Theophrastaceae) and Caraballo (1981) identified these fly speci- mens as A. anomala without giving a sci- entific name for “chirica.”’ I reexamined several specimens of this series and found them to be A. normalis. Because chirica is a common name for L. panamensis (see Type data), and the related species A. anomala is also known from this host, it seems likely that all of the Venezuelan records pertain to this plant. Type data.—Holotype 2 (ZAM, USNM52943), VENEZUELA: Falcon: Mar- cillal [11°12’N 68°50’W], 125 m, 7 May 1993, K.P. Katiyar, reared ex. fruit Zschokkea panamensis |= Lacmellea panamensis| (“‘chirica’’?), MFAPK-00345. Paratypes: PANAMA: Panama: El Cermefio [8°44’N 79°51'W], 20 Jun 1939, J. Zetek 4465, 1 @ paratype A. anomala (USNM52949); same, J. Zetek 4467, 1 2 paratype A. anomala IOC USNM53106); same, J. Zetek 4475, 1 @ par- atype A. anomala (BMNH USNMS53114); El Cermefio, 27 Jun 1939, J. Zetek 4489, 1 3 paratype A. anomala (USNM52960); El Cer- mefio, 2 Apr 1940, J. Zetek 4643, 1 & (USNM52948). VENEZUELA: Falcoén: Mar- cillaly) (a 12 N68 50°W)p 125 im 7 May 1993, K.P. Katiyar, reared ex. fruit Zschokkea panamensis (“‘chirica”’), MFAPK-00345, 1 d (ZAM, UWSNM52944). 5 6, 8 2 (USNM47439—43, 47447-54), 1 6, 1 & (BMNH, USNM52946-47), 1 6, 1 2 (MEUP, USNM47445-6), 1 ¢, 1 2 (ES- ALQ, USNM47444, 52945); Maicillal [sic] de la Costa, 200-550 m, ex. larva en frutos de chirica, 29 May 1967, R. Mendez & R. Bandres, 2 6, 2 2 (USNM52938-—41), 42 d, 53 2 (ZAM). AOS PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON eens re ZA RF Ss Lie ——— —_ Bi el SS ~s 5 SSoss; Pa SUE YF Ser3> SS Fig. 9. Aculeus, apical part, ventral view. A, Anastrepha anomala (USNMS52951, Panama: Arraijan). B, A. anomoiae (holotype). C, A. normalis (USNM52939, Venezuela: Marcillal de la Costa). D, A. pseudanomala (USNMS50455, Costa Rica: 20 km. S Upala). Anastrepha ocresia (Walker) (Figs. 1C, 2H, 5C—D, 10G) Trypeta ocresia Walker 1849: 1016 [de- scription]; Loew 1873: 231 [taxonomy]. Trypeta (Acrotoxa) ocresia: Loew 1873: 337 [taxonomy]; Osten Sacken 1878: 189, 195 [in catalog]. Acrotoxa ocresia: Loew 1873: 231 [taxon- omy]. Anastrepha ocresia: Aldrich 1905: 602 [in catalog]; Bezzi 1909: 283 [in catalog and key]; Hendel 1914: 14 [in key]; Greene 1934: 158 [review]; Stone 1942: 24 [re- vision]; Aczél 1950: 221 [in catalog]; Foote 1964: 322 [type data], 1965: 673 [in catalog]; Weems 1968: 1 [review, habitus illustration, hosts]; Wasbauer 1972: 107 [host list]; Steyskal 1977: 8 [in key]; Norrbom and Kim 1988: 51 [host list]; White and Elson-Harris 1992: 148 [taxonomy, pest status]; Foote et al. 1993: 103 [taxonomy]; Fernandez et al. 1998: 34 [Cuba, host]; Norrbom et al. 1999a: 81 [in catalog]; Norrbom, in press [host database]. Anastrepha ochresia: Foote 1967: 14 [in catalog; misspelling]. Trypeta tricincta Loew 1873: 225 [descrip- tion]; Stone 1942: 24 [synonymy]. Trypeta (Acrotoxa) tricincta: Loew 1873: VOLUME 104, NUMBER 2 2 aa ys A \ y i Ma { NS ZA & S = aN | ral AA y N iZ (% {/\ a dita n aman % NN “A IS Gi ak v/\ RS | i aN * a\ ie OF GA LOZ ~ / WO mae \ ae) Seen Fig. 10. 409 i ¢ é a | | | .! r Ts i | al rR. S £ NG Wy it Aculeus tip, ventral view. A—C, Anastrepha anomala (USNM52951, Panama: Arraijan; USNMS53107, Brazil: Cruz das Almas; Brazil: Itapecuru-Mirim, USNM31431). D, A. anomoiae (holotype). E, A. bistrigata (copied from Stone 1942, fig. 3B). KR A. normalis (USNM52939, Venezuela: Marcillal de la Costa). G, A. ocresia (copied from Stone 1942, fig. IE). H, A. pseudanomala (USNMS50455, Costa Rica: 20 km. S Upala). I, A. pulchella (USNM51717, Panama: El Cermeno). J, A. pulchra (copied from Stone 1942, fig. 2B). 329 [classification]; Osten Sacken 1878: 189 [in catalog]. Acrotoxa tricincta: Loew 1873: 227 [clas- sification]. Anastrepha tricincta: Aldrich 1905: 602 [in catalog]; Bezzi 1909: 284 [in catalog and key]; Hendel 1914: 14 [in key]. Recognition.—Anastrepha ocresia is one of the species of the serpentina group with a hyaline area in cell br posterior to the pterostigma that is not extended to vein R,,;. It differs from all other species of the serpentina group in having the brown markings of the mesonotum restricted to the 410 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON areas near the scuto-scutellar suture, and the scutum without microtrichia except on ex- treme lateral margin. The broad distal sec- tion of the S-band (at apex of vein R,,, greater than 0.70 times width of cell r,,;) differentiates it from all other species ex- cept A. ornata. The distal arm of the V-band is always partially present, at least from vein M to posterior wing margin. A. ocresia is the only species of the serpentina group known from the Greater Antilles and Florida. Description.—Largely orange to red brown with yellow and dark brown mark- ings. Setae dark brown. Head: Yellow ex- cept ocellar tubercle brown. Facial carina, in profile, concave. 3—5, usually 4, frontal setae; 1—2 orbital setae, posterior seta ab- sent on at least one side in 9 of 25 speci- mens, but well developed if present. Ocellar seta weak, short to minute. Antenna ex- tended 0.70—0.85 distance to lower facial margin. Thorax (Fig. 1C): Mostly orange with following areas yellow and usually distinctly contrasting: postpronotal lobe; single medial and paired sublateral vittae on scutum, the medial vitta extended nearly full length of scutum, anterior %4 very slen- der, broadened posteriorly, but extended lat- erally only to or slightly beyond level of acrostichal seta; sublateral vitta extended from transverse suture to posterior margin, including intra-alar seta; scutellum except base (brown area well separated from basal seta or sometimes diffusely extended to it or slightly beyond); dorsal margin and an- teroventral corner of anepisternum; dorsal half to % of greater ampulla; katepimeron; and most of anatergite and katatergite. Me- sonotum mostly to entirely orange, usually with dark brown spot or band bordering scuto-scutellar suture, sometimes shaped like shallow U, the arms extended anteri- orly no more than to level of intra-alar seta. Darker areas of anepisternum, katepister- num, and anepimeron orange; those of mer- on and katatergite sometimes dark brown. Subscutellum and mediotergite usually dark brown, narrowly orange medially, but oc- casionally mostly orange with dark brown lateral margins. Mesonotum 2.50—3.45 mm long. Scutum without microtrichia except postsutural lateral margin (lateral to supra- alar seta); presutural setulae mostly yellow, brown on narrow area along dorsocentral line, postsutural setulae yellow medially, mostly brown lateral to level of acrostichal seta. Katepisternal seta usually slightly lon- ger than postocellar seta, usually moderate- ly developed but occasionally weak, red brown or occasionally yellowish. Wing (Fig. 2H): Length 5.27—-7.49 mm. Vein M strongly curved apically; section between BM-Cu and R-M 1.85—2.33 times as long as section between R-M and DM-Cu; sec- tion between R-M and DM-Cu 0.80—0.96 times as long as DM-Cu. Crossvein DM- Cu oblique, with anterior end more distal than posterior end. Pattern mostly moderate to dark brown. C-band and S-band narrow- ly to broadly connected in cell br and some- times r,,3, Separated basally by hyaline area in cell br aligned with pterostigma, hyaline basal 4—% of cell dm, and hyaline to yel- lowish area covering all of cell bm. C-band yellowish or subhyaline in cells be and cell c, except anterior margin; with large yel- lowish area in base of cell sc and cells r, and r,,; posterior to pterostigma, extending distally at least to apical *%4 of pterostigma, sometimes reaching level of apex of vein R,. S-band with narrow yellow area in cell dm; most of cell bcu yellowish to subhya- line; rest of band brown, including areas distal to and anterior to R-M; distal section broad, at apex of vein R,,, 0.73—1.00 times width of cell r,,,; extended to or occasion- ally almost to apex of vein M. Hyaline spot in cell r, usually nearly triangular, some- times rounded, usually extended into cell r,,; and often to vein R,,.;; its apex aligned slightly to distinctly basal to R-M. V-band with distal arm usually reduced, extended anteriorly beyond vein M, but only rarely reaching vein R,,; (one female from Do- minican Republic); separated from proxi- mal arm or sometimes connected along vein M (cf connected, never with hyaline space VOLUME 104, NUMBER 2 between band and vein M); proximal arm usually extended to vein R,,;, occasionally connected to S-band along vein R,,5; ex- tended basally along posterior wing margin often almost to vein A,+Cu, but not con- nected to extension from base of S-band. Abdomen (Fig. 1C): Orange with yellow and brown markings. Syntergite 1+2 and tergites 3—4 with narrow brown bands, nar- rowly medially interrupted occasionally on tergite 3, often on tergite 4; posterior mar- gin yellow. Tergite 5 and female tergite 6 orange. Male terminalia (Figs. 5}C—D): Dor- sal posterior margin of epandrium evenly convex. Lateral surstylus moderately long; in lateral view slightly curved; in posterior view with strong basolateral lobe, main part narrowly triangular, acute apically. Proctig- er with lateral fold separating sclerotized ar- eas. Phallus 3.54—4.51 mm long; 1.27—1.60 times as long as mesonotum. Glans 0.50 mm long; acrophallus relatively stout. Fe- male terminalia: Oviscape 2.95—4.00 mm long, 0.95—1.32 times as long as mesono- tum. Eversible membrane with 25—40 large, hook-like dorsobasal scales in triangular pattern. Aculeus 2.83—4.08 mm long; tip (Fig. 10G) 0.30—0.36 mm long, 0.11—0.13 mm wide, very gradually tapered basally, apical 0.53—0.67 finely serrate, triangular. Spermathecae ovoid. Distribution.—Anastrepha ocresia is known from the USA (Florida Keys), Cuba, Hispaniola, Puerto Rico, and Jamaica. The only records from the Florida Keys are from 1936, and A. ocresia may not be cur- rently established there. Biology.—The native host plants of A. ocresia are uncertain. It has been reared several times from Manilkara zapota (L.) P. Royen (Sapotaceae) (Foote 1964, as Achras zapota; Fernandez et al. 1998), but this plant is not native to the Greater Antilles. Weems (1968) listed Psidium guajava L. (Myrtaceae) as a host, but did not indicate the basis for this record, and the status of this plant as a host needs confirmation. Cit- rus X paradisi Macfad. (Rutaceae) was er- roneously listed as a host plant by White 411 and Elson-Harris (1992), citing Weems (1968), but the latter reported only an adult foliage record. Type data.—Trypeta ocresia: Lectotype 2 (BMNH), Jamaica, Mr. Gosse; designat- ed by inference of holotype by Greene (1934: 158) [examined; with following la- bels: [square, handwritten] ‘Jam. [another word, possibly “‘Gosse’’]”’; [long, hand- written] “‘Ocresia 1016’’; [handwritten, square] *“‘Anastrepha ocresia Walk. Type’”’; [circular, green-bordered] ““Type’’; [red bordered] “LECTOTYPE Trypeta ocresia Walker by Greene 1934: 158, by inference of holotype’’]. A female of Anastrepha sus- pensa (Loew) was in the BMNH collection with the lectotype, but is doubtfully a syn- type because Walker mentioned the incom- plete V-band in the original description (“‘a forked stripe, which is imperfect, nearly half of the tip fork being wanting’’) and it is complete in this specimen. However, be- cause Walker described 7. ocresia from an unstated number of female specimens it is best to regard Greene’s (1934) discussion of “the type female”’ as a lectotype designa- tion. According to Papavero (1973: 247) P. H. Gosse collected in Jamaica between 6 December 1844 and 11 July 1846. Trypeta tricincta: Lectotype 6 (MCZ), on shipboard, 60 mi. NW of St. Nicholas [probably M6le St.-Nicolas], Haiti, PR. Uhler; designated by inference of holotype by Greene (1934: 146) [examined; with fol- lowing labels: “‘60 miles N. W. of St. Ni- colas, Hayti, P. R. Uhler, on shipboard”’; “tricincta ILw.”; [red] “Type 132827; [red bordered] “LECTOTYPE Trypeta tricincta Loew by Greene 1934: 146, by inference of holotype” ]. Other specimen data.—CUBA: Baracoa de Banta, on mango, 19 Jun 1930, E. Kos- tal, 1 2 (MCZ USNMS50477). Guantanamo: Guantanamo Bay Naval Base, McPhail Tap) elec t975, 1 ¢ (USNMS29972). Isla de la Juventud [unspecified locality], jum 919245) Moznette,” 1 “29> (TAMU USNM52963). Matanzas: Central Conchita [= Puerto Rico Libre, 22°44’N 81°32'W], PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON reared from A. sapota [= Manilkara zapo- ta), 25. Jun) 1 946i Ca Scaramuzzay 2a? (USNMS50474—-5); same, reared 2-7 Jul 1946, from larvae collected 25 Jun 1946, 3 36, 6 2 (USNM52994—53002). Pinar del Rio: Péninsula de Guanahacabibes, Bolon- dron..[2.1°52/28N 1184-495. W is (ear 1924. S.C. Bruner,.no. 9041. 1G. (MGZ USNM50476); same, 11 Mar 1924, 1 @ (USNM51712). DOMINICAN REPUB- LIC: [unspecified locality], reared ex. Ach- ras sapota {|= Manilkara zapota|, Aug 1962, 2 6, 7 2 (USNM50464-72), 1 ¢ (MCZ USNM50463). Barahona: near Fili- pinas Larimar Mine, near La Travesia [18°07’N 71°07'W], at light, 20-26 Jun 1992, KR. E. Woodruthi & PE. Skelly, 1? (FSCA USNM51780). Distrito Nacional: Santo Domingo, 22 Jul 1919, H. Morrison, 1 2 (USNMS50473). La Altagracia: Baya- hibe: 2 kin Not, 138° 23"N 68°57 W, 10m. 5) Jul 992. ©. Young, (Ro “Davidson, s: Thompson, J. Rawlins, 1 2 (USNMS51938); Higuey, La Gima Colon [possibly La Jina (18°42'’N 68°37'W)], 730 m., McPhail trap in Citrus sinensis, 24 July 1990, C.W. For- ster, 2 d, 3 2 (USNM50458-—62). Peder- nales: Cabo Rojo, 26 km N of, 18°6’N 71°38'W, 730 m., wet deciduous forest, sweep sample, 13—25 Jul 1990, L. Masner, J Rawlinsy I 0) 0 G 0) DIDULOUD Oo @©@ € O © I Ona 0) I 0) 0) (0) C 0 @ I I 0 G (O) T 0) DISALIO i © © © I (0) 0 G 0) 0) ae) 0 Oo @ f © 0 0) C 0) S7]DULIOU Eé i @O ® © I I I © é C Ona GiO QO @ I I 0) I G G @ DIDUALO Gui 0 1 I 0) I 0) 0) 0) 0) 0 I Oo @® @® @ 0) I I 0 DIDS1A1S1qG Ce 30) eT I 0 I O c-0 0 0) 0) 0) oO @ @©O @ 0 I I 0 DIDIAIS Oo @ © @ @ 0) 0) 0 © @ @ 0) 0) ST/TWUISSIP SOR GH) ae 9 ¢ v € C I 0 6 SA FO aes v € C I soroedg CF I I I I I I I I I Joquinu 1aJOeIBY,) VOLUME 104, NUMBER 2 “OIBI I IY) S9}eIS dBSIPUL Sasoy IIe “ISI Poysi] St a}e}s JURUTWTOpaid ayy ‘sojovieys oqeiiea AT[eoyIoodse.nul 10,4 ‘staquunu 1aj9vIeYyS JO UOTeUL[dxa IOJ | a[qQuy_ sag ‘dno1is vuruadsas ay) JO satoads oy) 1OJ XLNeUT aIvIS JOIORIRYD “TZ aIqe], 130 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON dissimilis 2 559d [HOH _ I ornata 3 2021 2 Ne ie 6 1718 striata a : 9 2 bea fal bistrigata 26 4 ea 21 ‘laiet normalis 1 2 pseudanomala 39 19 3 ocresia 5 19 iH] = anomala 11 serpentina A ar ‘ == [+— pulchella el 67 Oo 19 2 4 21 { pulchra fo) 3 anomoiae dissimilis 21 normalis 1 9 11 Ey ornata 26 3 2021 22 eq] . ty 2 25 6 1718 SINE 2 [HI 9 Tg ial a" sey = bistrigata 1 2 3 9 F 1 L_] ocresia 22 anomala 19 pseudanomala serpentina 9 1119 11 FLA L_ pulchella B A A 67 19 2 4 21 = pulchra oO 3 anomoiae Fig. 12. Phylogenetic relationships among species of the Anastrepha serpentina group. Two of 95 most parsimonious trees resulting from cladistic analysis. All character state changes are shown, plotted using fast optimization. VOLUME 104, NUMBER 2 431 dissimilis 3 171820 striata bistrigata 1 normalis 3 1120 ornata ocresia anomala pseudanomala serpentina 11 pulchella 19 pulchra 21 anomoiae dissimilis 17182021 striata bistrigata 1 normalis ocresia anomala pseudanomala 3 112021 ornata serpentina 11 pulchella 19 pulchra anomoiae Fig. 13. Phylogenetic relationships among species of the Anastrepha serpentina group. Two of 95 most & g I g parsimonious trees resulting from cladistic analysis. All character state changes are shown, plotted using fast optimization. $32 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON cies known only from a male from Mexico also was excluded from the analysis. The sister taxon of the serpentina group is uncertain. Norrbom et al. (1999b) pro- posed that the serpentina and striata groups belong to a large clade also including the pseudoparallela, spatulata, ramosa, gran- dis, doryphoros, and fraterculus groups. Anastrepha dissimilis Stone, which belongs to the pseudoparallela group, was chosen as the outgroup taxon as a typical represen- tative of all of the possible sister taxa. Al- most all of the species of the other groups also would be coded O for the characters used in the analysis, except for aculeus tip shape, which is highly variable. The matrix was analyzed using Hennig86 option ie*. Ninety-five trees of 27 steps length resulted (ci = 0.74, ri = 0.76), four of which are shown in Figs. 12 and 13, which were pro- duced using WinClada. The serpentina group, as defined here, is supported as a monophyletic group mainly by the presence of brown markings on the mesonotum and subscutellum and medi- otergite (character 2), the only synapomor- phy present on all 95 trees. Except in A. ocresia, where the brown markings are re- stricted to the area bordering the scuto-scu- tellar suture, the brown area is extensive and either somewhat U-shaped or covering most or all of the darker areas of the me- sonotum. Similar extensive brown areas oc- cur in some species of the daciformis group, but other characters indicate that this is due to homoplasy (Norrbom et al. 1999b). Brown mesonotal markings also occur in a few other Anastrepha species (e.g., A. grandis, some species of the cryp- tostrepha and robusta groups), but the dif- ferent shapes of the markings and other characters also suggest that this is the result of homoplasy. The subscutellum and me- diotergite are usually mostly brown in spe- cies of the serpentina group, although in some specimens of some species it is brown only laterally, resembling the condition in most species of the fraterculus species group. Because the mostly to entirely brown condition is more common in these intraspecifically variable species, I regard this as a different character state than that in the fraterculus group, that further sup- ports the monophyly of the serpentina group. Outside of this group, it is uncom- mon in Anastrepha, occuring in some spe- cies of the grandis, daciformis, cryptostre- pha and robusta groups. Only the former species group appears to be relatively close- ly related to the serpentina group (Norrbom et al. 1999b, McPheron et al. 1999). McPheron et al. (1999) included four species of the serpentina group in their phy- logenetic analysis of Anastrepha based on mitochondrial DNA sequences. They found strong support for the monophyly of A. ser- pentina + normalis (as “‘n. sp. nr. anoma- la’) and also for A. striata + bistrigata. Both pairs of species were included in a large clade with species of the fraterculus, spatulata, pseudoparallela, and grandis species groups, but the relationships among these groups were poorly resolved. Studies of isozymes (Morgante et al. 1980, Matioli et al. 1992) have also shown strong simi- larity between A. striata and bistrigata, al- though A. serpentina was more similar to other species. Relationships among the species within the serpentina group were not well resolved by the present analysis (see Figs. 12-13), although two clades were consistent in all 95 trees: A. striata + bistrigata; and A. ser- pentina + (pulchella + (pulchra + ano- moiae)). The close relationship of A. striata + bistrigata is strongly supported by the shape of the posterior margin of the epan- drium and the ridge on the lateral surstylus (character states 17.1 and 18.1, respective- ly), which are unique to these two species. The clade of A. pulchella + (pulchra + an- omoiae) is also strongly supported, by the very broad connection of the C-band and S-band (character state 6.2), and the re- duced or absent yellow area of the C-band posterior to the pterostigma (character state 7.1). The relationship of A. serpentina to the latter clade is supported by one or more VOLUME 104, NUMBER 2 of the following character states: distal sec- tion of S-band slender (9.1); hyaline mar- ginal spot in cell r, elongated distally along costa (11.1; this can also be interpreted as arising independently in A. serpentina and in A. pulchra + anomoiae), and the lateral surstylus with a small basolateral lobe (19.1). None of these states is consistently hypothesized as a synapomorphy for these four species on every tree, however; char- acter state 9.1 is usually a synapomorphy, but on some trees it may be interpreted as a synapomorphy for these species plus A. ocresia or ornata, with subsequent change to state 2 in the latter species. The positions of the other species in the cladograms are highly variable. Anastrepha normalis is always relatively basal in the most parsimonious trees, arising as the sis- ter group to a clade including all other spe- cies of the serpentina group (Fig. 12B; sup- ported only if relatively darker wing pattern color (character state 5.1) is interpreted as a synapomorphy for the other species, with reversal in A. striata + bistrigata), as the sister group of a clade containing all of the other species except A. striata and _ bistri- gata (Figs. 13A—B), as the sister group of a clade with all species except A. ornata + (striata + bistrigata) (Fig. 12A), or as the sister group of the clade A. ornata + (stri- ata + bistrigata) (supported only if char- acter 21.1 is interpreted as a synapomorphy, with subsequent change to state 21.2 in the other three species). Anastrepha ocresia and anomala_ are usually somehow included in a clade also including the serpentina clade, A. pseudan- omala, and/or A. ornata, and they are not closely grouped with A. striata + bistrigata on any of the trees. Anastrepha pseudan- omala is highly variable in position, in part because characters 17—20 are unknown for it. It is usually within a clade including A. ocresia, anomala, the serpentina clade, and sometimes A. ornata. On some trees it is hypothesized as the sister group of A. or- nata + (striata + bistrigata), but this is supported only if A. pseudanomala is as- 433 sumed to have a subapical lateral lobe on the lateral surstylus (character state 20.1). The relationship of A. ornata among the other species of the serpentina group is also unresolved by the present analysis. It is highly variable in position on the 95 clad- ograms. In some trees it is the sister group of A. striata + bistrigata (Fig. 12), a hy- pothesis supported by scutal microtrichial pattern (character state 3.1), lateral sursty- lus shape (20.1) and aculeus tip shape @2i2)Pin’ other trees “(C1e-5 Fie, IS)rait is placed variably within clades including A. ocresia, anomala, pseudanomala, or the serpentina clade. The hypothesis that A. or- nata is more closely related to the species exclusive of A. striata and bistrigata is sup- ported by the more extensively brown bod- ies of these species (character state 2.2), their broad C-band, which extends into the distal half of cell br (6.1), and the relatively dark wing pattern of all of the latter species except A. normalis (character state 5.1). ACKNOWLEDGMENTS I am grateful to the curators of the insti- tutions who kindly loaned specimens for this study. Lucrecia Rodriquez prepared the wing illustrations, Taina Litwak, the habitus illustrations, and Elisabeth Roberts, most of the terminalia illustrations. I also thank Adrian Pont for interpretation of Bigot Col- lection labels, and EF C. Thompson for no- menclatural advice. Steve Gaimari, Steve Lingafelter, Gary Steck, and Roberto Zuc- chi kindly reviewed the manuscript. LITERATURE CITED Aczél, M. L. 1950. Catalogo de la familia “Trypetidae’ (Dipt. Acalypt.) de la region neotropical. Acta Zoologica Lilloana (1949) 7: 177-328. Aldrich, J. M. 1905. A catalog of North American Dip- tera (or two-winged flies). Smithsonian Miscella- neous Collections 46(2): 680 pp. . 1925. New Diptera or two-winged flies in the United States National Museum. Proceedings of the United States National Museum 66(18): 36 pp. [= No. 2,555]. Baker, A. ©, W. E. Stone, C. C. Plummer, and M. McPhail. 1944. A review of studies on the Mex- ican fruitfly and related Mexican species. 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Congresso Brasileiro de Entomologia Resumos 8: 280. . 2000. Chave para as espécies de Anastrepha conhecidas para o estado da Bahia, pp. 238-239. In Nascimento, A. S. and R. da Silva Carvalho, [Moscas-das-frutas nos estados brasileiros] Bahia, pp. 235-239. In Malavasi, A. and R. A. Zucchi, eds. Moscas-das-frutas de importancia econdmica no Brasil. Conhecimento basico e aplicado. Holos, Riberao Preto, 327 pp. PROC. ENTOMOL. SOC. WASH. 104(2), 2002, pp. 437-446 CONSTRUCTION OF A DEFENSIVE TRASH PACKET FROM SYCAMORE LEAF TRICHOMES BY A CHRYSOPID LARVA (NEUROPTERA: CHRYSOPIDAE) THOMAS EISNER, JAMES E. CARREL, EILEEN VAN TASSELL, E. RICHARD HOEBEKE, AND MARIA EISNER (TE, ME) Department of Neurobiology and Behavior, Cornell University, Ithaca, NY 14853, U.S.A. (e-mail: tel4@cornell.edu; mle3 @cornell.edu); (JEC) Division of Biolog- ical Sciences, University of Missouri, Columbia, MO 65211, U.S.A. (e-mail: carrelj@missouri.edu); (EVT) Department of Entomology, Michigan State University, East Lansing, MI 48824, U.S.A. (e-mail: vantasse @ pilot.msu.edu); (ERH) Department of Entomology, Cornell University, Ithaca, NY 14853, U.S.A. (e-mail: erh2 @cornell.edu) Abstract.—The behavior of a chrysopid larva from Arizona is described, identified as Ceraeochrysa lineaticornis (Fitch), which constructs its trash packet from the trichomes of sycamore leaves (Platanus wrightii S. Watson). The trichomes are particularly dense on the underside of the leaves. To fashion the packet, the larva uses trichomes stuck loosely to the leaf, and others that it plucks from the leaf surface. The packet on the mature larva weighs on average 2.2 mg, an equivalent of approximately two leaf under- sides worth of trichomes. Evidence is presented indicating that the packet provides the larva with protection against predation. It is argued that the trichomes are defensive in the sycamore tree itself, and that the chrysopid provides yet another example of an insect that benefits from utilization of a plant defense. Interestingly, the chrysopid benefits the sycamore tree itself. Although it usurps the tree’s defense, it aids the tree by preying on a specialist herbivore, the tingid, Corythucha confraterna Gibson, which feeds on the sycamore tree unbothered by the trichomes. Key Words: animal defense, Neuroptera, Chrysopidae, Ceraeochrysa lineaticornis, plant defense, trichome, insect-plant interaction Many chrysopid larvae have the habit of collecting exogenous materials and placing them on their backs, forming so-called trash packets that they retain throughout larval life (Smith 1922, Canard et al. 1984). The materials they use to fashion the packets are variable and may consist of vegetable mat- ter, arthropod remains, insect waxes, or general debris (Smith 1922, Slocum and Lawrey 1976, New 1969, Eisner et al. 1978, Canard et al. 1984). Existing evi- dence indicates the packets act as physical shields that provide the larvae with protec- tion against insectan predators (New 1969, Principi 1946, Eisner et al. 1978). Not all chrysopid larvae are trash carriers, but those that are, are obligatorily so, and the priority that larvae give to forming trash packets has a significant genetic component (Milbrath et al. 1993, Tauber et al. 1995). Here we describe the behavior of a chry- sopid larva that constructs its packet from trichomes that it takes from the leaves of sycamore trees (Platanus wrightii S. Wat- son) in Arizona. We present data on the feeding habits of the larva, as well as on PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON VOLUME 104, NUMBER 2 how the packet is constructed and used in defense. Our observations were made in the hamlet of Portal, Cochise County, Arizona, in 1967 (September) and 1982 (August). We refrained from publishing earlier be- cause we had failed to obtain a definitive identification of the chrysopid. This prob- lem has now been remedied, thanks to the courtesy of Catherine and Maurice Tauber, who have informed us that the chrysopid is Ceraeochrysa lineaticornis (Fitch) (voucher specimens have been deposited in the Cor- nell University Insect Collection). MATERIALS AND METHODS Field site——We first noted the larvae on a group of sycamore trees growing on the banks of Cave Creek, in Portal itself, on the grounds of the Cave Creek Ranch, where we were in residence (Fig. 1A). We subse- quently found them also on sycamore trees near Cave Creek, up to several miles up- stream from the original location. The lar- vae were readily spotted by their white trash packet, which rendered them extreme- ly conspicuous against the green color of the sycamore leaves on which they were found (Fig. 2A). As they scurried about, they resembled tiny ambulatory cotton wads. Careful scrutiny of the visually ac- cessible lower branches of the trees usually revealed presence of many larvae per tree. Maintenance of larvae.—Larvae were maintained on freshly clipped sycamore leaves in plastic containers of various sizes, including Petri dishes. In the field, larvae had been found on repeated occasions feed- ing on a tingid, Corythucha confraterna Gibson, whose colonies were of common occurrence on the sycamore leaves. Captive chrysopids were therefore always provided with some leaves that were tingid-infested. ee Fig. 1. 439 They took readily to the tingids, and ap- peared to feed on the nymphs only. Main- tained on this diet many of the larvae went on to pupate and develop into adults. As is typical for trash-carrying chrysopids, larvae retained the trash packet as an outer cov- ering of their cocoon when they pupated. The cocoons therefore have the same ap- pearance as the larvae, except that they are non-ambulatory. In the field we found sev- eral cocoons on the trunks of sycamore trees. Most larvae that we used in our ex- periments were probably in their last instar. Electronmicroscopy.—For examination with the scanning electronmicroscope, specimens (chrysopid larvae, pieces of syc- amore leaf) were preserved in the field in 70% ethanol, then critical point dried and gold coated in the laboratory. Predation tests.—Of the predators used, the reduviid, identified as Pselliopus latis- pina Hussey, is doubtless a natural enemy of the chrysopid. It occurred commonly on the leaves and branches of the sycamore trees themselves, and was once actually ob- served feeding on a C. lineaticornis in the field. The ant used in predation tests, Pogon- omyrmex barbatus (Smith), may not itself be a primary enemy of the chrysopid, but there can be little question that ants as such figure among the larva’s natural predators. We routinely observed ants foraging singly on the leaves, branches and trunks of the sycamore trees. Statistics —Numerical averages are giv- en as mean = S.E- RESULTS The trichomes.—Both surfaces of the sycamore leaves bear trichomes, but the structures are much more densely distrib- A, Stand of sycamore trees on which the chrysopid larva was discovered (Portal, AZ). B, Close-up of underside of sycamore leaf showing trichomes. C, Underside of sycamore leaf from which the trichomes have been removed; the detached trichomes form the wad seen in the ampule. D, Enlarged view of leaf underside (scanning electronmicrograph). E, Base of a trichome showing the hinge (arrow). F, Base of a trichome that has been torn off at the level of the hinge. Reference bars: D = 0.5 mm; E = 20 pm. 140 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON uted on the underside (Figs. 1B, D). Indi- vidually the trichomes are usually multiply branched, and they are flexibly hinged near their base, where there is a break in the core of the shaft and the wall is reduced to a thin-walled tube (Figs. 1E, F). If one pulls on individual trichomes with forceps, they tend to detach at the level of these hinges. They evidently detach spontaneously in large numbers from the leaf. surface, as ev- idenced by the fact that the underside of sycamore leaves usually has a loose coating of detached trichomes. Microscopic exam- ination verified clearly that these lose tri- chomes had broken off at the level of the hinges. Packet construction.—Removal of the trash packets from the larvae could be ef- fected easily by teasing away the trichomes with forceps. Eight larvae that were thus denuded and re-released into their leaf-con- taining enclosures, commenced reloading almost at once. In typical chrysopid fashion (New 1969, Principi 1940, Smith 1922), us- ing their jaws as a two-pronged fork, they scooped up load upon load of trichomes and placed these upon their backs (Figs. 2C—F). Most often they procured trichomes from the clusters of available loose ones, but they also pried many off with their jaws. It seemed clear, moreover, that they were programmed to utilize sycamore tri- chomes only. If released in a Petri dish with general particulate debris they tended to ig- nore such matter, but if then offered a syc- amore leaf, they usually returned promptly to the task of reloading. Like trash-carrying chrysopids generally (Canard et al. 1984), the larvae are equipped with a set of special hooked bristles that seem to serve specifi- cally for retention of the packet. The bris- tles project upward in rows from the back, and there are others that project outward in clusters from a series of lateral protuber- ances (Figs. 4A—-C). Packet construction appeared to proceed in accord to a fixed protocol. The first pluckings were always applied to the pos- terior third of the body. To gain access to the site, the larvae arched that region up- ward and forward every time that the head flexed backward to deliver a load. Later pluckings, delivered to the more accessible anterior regions of the back, were applied without simultaneous postural adjustments of the body. A visual demonstration of the sequence of trichome delivery during packet con- struction was obtained by giving denuded larvae access to trichomes of different col- ors. Packets were removed from a series of larvae and were then stained either in red (with acid fuchsin), in black (with chlorazol black), or kept unstained. Denuded larvae that were then given access to teased apart samples of these packets, in the sequence of black (for 30 min), to unstained (for 20 min), and to red (for 60 min), constructed packets in which the trichomes were laid out in three colored bands, in the sequence red, white, and black from fore to aft (Fig. 2B): Package repair.—Use of stained tri- chomes also yielded visual evidence of the precision with which the larvae are able to repair damage to their package. Four larvae, bearing natural unstained packets, were treated as follows: (1) Trichomes removed from center of packet, leaving the larva with a ring-shaped shield. (2) Posterior half of packet removed. (3) Left half of packet removed. (4) No trichomes removed (control). The larvae were then confined with a supply of black trichomes, and checked for condition of their packets after 24 hours. The results were as follows: (1) Center of ring filled with unstained trichomes. Periphery of packet loosely lad- en with black trichomes. Larva had evi- dently repaired the hole in the packet with trichomes from the periphery of the packet. New trichomes had been added to the pe- riphery to replace those used in the repair of the hole. (2) Posterior half of packet repaired, but VOLUME 104, NUMBER 2 44] Fig. 2. A, Full-grown larva. B, Larva that has formed a packet from pre-stained trichomes (see text). C, D, Larva, in lateral view, in the process of building its packet. In C it is scooping up trichomes with the mouthparts: in D it is adding the trichomes to the packet. E, K Same as preceding pair, but in frontal view. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1.0 pax 0.9 E =< 0.8 oO 6 oS 0.7 Qa. 5 06 © a = as i) GB 04 s : 0.3 0.2 0 2 4 6 8 Time (h) Fig. 3. Regeneration of packet by denuded larvae, expressed as mass of packet formed over time. Best fit equation is shown (y = 0.185 + 0.062; R* = 0.994, P < 0.001). with unstained trichomes. Front half of packet made up mostly of black trichomes. The larva evidently repaired the rear of packet with trichomes from the front, and replaced the trichomes removed from the front with ones newly acquired. (3) The missing left half of packet was restored with newly acquired black tri- chomes. Only few new trichomes had been added to the right half of the packet. (4) The entire packet had been aug- mented by a sparse uniform addition of new black trichomes. It is clear from these results that the lar- vae are able to repair their packet, and that in so doing they do not merely fill in dam- aged sites with newly acquired trichomes. They are able to effect repair by reposition- ing existing trichomes, and they correct for any resulting inequities in trichome distri- bution by bringing into place newly scooped up trichomes. Thus, the larvae have not only a sense of what is missing in a damaged packet but an elaborate mecha- nism for repairing the damage. Particularly remarkable is the larva’s ability to move tri- chomes from one area of the packet to an- other. Casual observation reveals that they effect such relocation both by use of the jaws and by a sort of wave-like peristaltic action of the abdomen, whereby trichomes are transferred from one set of bristles to the next. Such peristaltic action was also noted at times during the package rebuild- ing undertaken by denuded larvae. Trichome load of sycamore leaves.— Two procedures were adopted for determin- ing the mass of trichomes per unit surface area of sycamore leaf underside. One procedure involved taking a syca- more leaf, determining its surface area (by weighing a piece of paper in the shape of that leaf, and referencing this weight to that of a piece of known area of the paper), then removing the trichomes from the leaf un- derside (with forceps and by scraping with a scalpel) and weighing the trichomes (Fig. IC). Removal of trichomes was judged to be 90% complete. The leaf (90.3 cm? sur- face area) yielded 1.2 mg trichomes, an equivalent of 13.3 wg per cm? of leaf un- derside. The second procedure involved using de- nuded larvae to effect the trichome removal from a leaf. The leaf was exposed (under- side up, with the upper side inaccessible) to two sets of 3 larvae each, operating in se- quence for a combined period of 3 hours, by the end of which time the leaf underside was estimated to have lost 90% of its tri- chomes. The combined mass of the trash packets (which none of the larvae had com- pleted in the time allocated) was 1.0 mg. It follows from this value, and from the area of the leaf (74.2 cm?) that the leaf had yielded 13.5 wg trichomes per cm? of un- derside. Packet construction.—To obtain some measure of the rate at which the larvae con- struct their packets, 32 larvae were denuded and their packets weighed. They were then individually confined in plastic enclosures VOLUME 104, NUMBER 2 Table 1. Packet regeneration of denuded larvae. See also Fig. 3. N = 8 per category, except for 8-hour category, where N = 6. Time since New packet larvae were denuded Mass Proportion of (hours) (mg) original packet 1 0.263 + 0.050 0.149 + 0.047 2) 0.288 + 0.044 0.143 + 0.021 4 0.438 + 0.091 O22 es 101052 8 0.683 + 0.182 0.430 + 0.167 containing fresh sycamore leaves, and al- lowed to reload for 1, 2, 4, or 8 hours. Eight larvae were initially assigned to each time category, but the sample size of the 8-hour group was reduced to 6 because 2 of the larvae had affixed themselves, preparatory to pupation, during the experimental period. At the end of the assigned times, the larvae were again denuded and their packets weighed. From the data it was possible to calcu- late, first of all, the average mass of the initial trash packets of the larvae (in other words, the average mass of packets of near- ly full grown field-collected larvae). That mass was 2.16 + 0.13 mg (range = 0.70— 3.30 mg). In addition, a value could be ob- tained for each larva of the fraction of the initial packet that had been rebuilt in the time period assigned to that larva. The mean of such values, for each of the four larval categories, is given in Table 1, which also presents averages for the net mass of packet material gathered in the times allo- cated. A graphic representation of the latter values (Fig. 3), shows packet reconstruction to proceed at a steady rate, at least during the initial 8 hours of rebuilding. Predation tests: reduviid.—Five nymphs of P. latispina were set up in plastic con- tainers and individually offered equal num- bers (one of each, or four of each) of pack- et-bearing and denuded larvae. The test chambers were checked visually at inter- vals, and survivorship of the larvae was re- corded after 12—24 hr. Of the 12 denuded larvae that were offered, only three sur- 443 vived. By contrast, of the 12 shield-bearers, nine survived. Some of the packet-bearing larvae were used with more than one of the reduviids. Inspection of the cages during the experimental period had revealed some of the details of the encounters. Denuded larvae tended to be grasped the moment they were located, then promptly impaled on the reduviid’s proboscis and sucked out (Fig. 4G). Packet-bearing larvae, in con- trast, were held by the bug with the fore- legs, repeatedly probed with the proboscis (Fig. 4F), but then often released. We sus- pect that release was a consequence of the reduviid’s proboscis being too short to reach through the packet to the body of the larva. This interpretation is supported by an early observation in which a few larvae that were partly denuded and offered to P. la- tispina, were eaten. Predation tests: ants.—The tests were set up on barren ground, about 0.5 m from the nest entrance of a P. barbatus colony. Tap- ping the ground above the nest had thrown the ants into a state of agitation and they had emerged from the nest in numbers to mount guard around the entrance. Six of 8 denuded larvae released on the ground were encountered by individual ants and carried off. They were small relative to the ants and were not always noticed by the ants when first released. In fact, it was only upon the second or third presentation that the six lar- vae were taken. Of 8 packet-bearing larvae that were similarly released, only one was taken, even though they were all also re- tested individually two or three times. The larvae were encountered repeatedly by in- dividual ants, but they “‘froze’’ when con- tacted, and the ants, which in their palpa- tions appeared to touch the packet only (Fig. 4E), were quick to abandon the as- sault. DISCUSSION While our predation data are scant, and our predation tests not nearly as satisfactory in design as we would have liked, it seems reasonable to conclude that the trichome PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 4. A, Fully loaded larva in ventral view (scanning electronmicrograph); arrow points to one of the setose lateral protuberances that serves for retention of the packet. B, Enlarged view of a lateral protuberance. C, Detail of setae from a lateral protuberance; the barbs presumably help retain the packet. D, Nymph of the tingid Corythucha confraterna; note glistening droplets of secretion at tip of glandular hairs. E, An ant (Pogon- omyrmex barbatus) inspecting a chrysopid larva. K Reduviid nymph (Pselliopus latispina) probing a packet of a chrysopid larva. Most such inspections resulted in the larva being released. G, Reduviid feeding on a denuded larva. Reference bars: A = 1 mm; C = 20 pm; D = 0.5 mm. VOLUME 104, NUMBER 2 packet of the C. lineaticornis larvae is a protective structure. While ours may be the first demonstration that the trash packet of a larval chrysopid can shield against the at- tack of a reduviid, work with other chry- sopid larvae had shown the packets to pro- vide effective defense against other Hemip- tera (New 1969) and ants (Principi 1946, Eisner et al. 1978). The strategy of carrying overhead shields among insects is, of course, not restricted to chrysopid larvae. Classic examples are provided by chryso- melid beetle larvae, including notably tor- toise beetle larvae (Olmstead and Denno 1992, 1993; Gomez et al. 1999; Miiller and Hilker 1999; Eisner and Eisner 2000), in which the shields are usually fabricated in part with feces. By virtue of the fecal com- ponent, such shields have the capacity to deter by chemical in addition to physical action (G6mez et al. 1999), a property that sets them apart from chrysopid packets. To our knowledge, chemical noxiousness has never been demonstrated for a chrysopid packet. It was of some interest to find that the C. lineaticornis larva fed on a tingid, and spe- cifically on the nymphs thereof. Tingid nymphs, including those of C. confraterna (Fig. 4D), are endowed with glandular hairs (Livingston 1978), which in some species have been shown to produce a mixture of aromatic and alicyclic acetogenins (Oliver et al. 1990), potentially deterrent to preda- tors. Ceraeochrysa lineaticornis is evident- ly undeterred by this glandular material. Packet construction is clearly a matter of high priority for the larva. When denuded it always made prompt efforts to rebuild the packet, and when packets were partially de- stroyed it always proceeded to repair the structure. Packet construction has been shown to be a matter of priority also in an- other chrysopid, Chrysopa_ slossonae (Banks) (Eisner et al. 1978, Milbrath et al. 1993), and given that trash packets are doubtless defensive in chrysopids general- ly, they are probably maintained fastidious- ly by all larvae that possess them. In Cer- 445 aeochrysa cincta (Schneider), for instance, a species in which the larva cloaks itself with the wax of homopteran prey, the lar- vae engage in packet construction immedi- ately upon descending from the egg along the egg stalk (Mason et al. 1991). Packet construction from sycamore tri- chomes is a slow process. After a period of 8 hours, nearly full grown larvae that bore packets weighing on average about 2.2 mg before being denuded, had reconstituted an equivalent of only about 40% of the origi- nal shield. It can be inferred from this that the rebuilding of an entire shield should take such larvae over a day. In fact, if one assumes reloading to proceed indefinitely at the linear rate prevailing over the first 8 hours (Fig. 3), one can calculate from the best-fit equation pertinent to that rate that it would take larvae on average nearly 32 hours to reproduce the initial packet. The number of trichomes that go into the construction of the larval packet is substan- tial. A mature larva, bearing an average- sized packet of 2.2 mg, carries an amount of trichomes equivalent to what is obtain- able from two sycamore leaf undersides. Are the trichomes defensive for the syc- amore itself, and does C. lineaticornis ex- emplify yet another case of utilization by an insect of defenses that evolved in the first place for protection of a plant? Syca- more trichomes can be envisioned to serve in a multiplicity of capacities. They could, for example, act to trap a layer of “‘dead air’ directly adjacent to the leaf surface and by so doing provide for retardation of evap- orative water loss from the leaves. But this does not rule out the possibility that the tri- chomes serve also in defense. Tightly spaced and barbed, the trichomes could well be a hindrance to many an ambulatory arthropod. The chrysopid and its tingid prey could thus be viewed as specialists that have managed to access a niche not gen- erally open to colonization. The tingid has come to cope with the plant’s defense and has become the plant’s enemy. The chry- sopid in contrast, is essentially the plant’s 146 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ally. Although it usurps the plant’s defense, and uses the weaponry for it own purpose, it feeds on the tingid, and by so doing has become the plant’s “‘delousing”’ agent. Los- ing a fraction of its trichomes to the chry- sopid must thus be viewed as being bene- ficial to the sycamore. ACKNOWLEDGMENTS Supported in part by National Institutes of Health grant AI02908. We thank the late Robert Silberglied for identifying the re- duviid and the ant, and Catherine and Mau- rice Tauber for countless personal favors and advice on chrysopids, and for calling attention to citations we had missed. The late Ellis MacLeod deserves thanks as well for helpful comments on our findings. This is paper no, 177 imiethe series “Defense Mechanisms of Arthropods’”’; no. 176 is At- tygalle et al. 2001. Defense by foot adhe- sion in a chrysomelid beetle (Hemisphaer- ota cyanea): Characterization of the adhe- sive oil. Zoology 103: 1-6. LITERATURE CITED Canard, M., Y. Séméria, and T. R. New. 1984. Biology of Chrysopidae. W. Junk Publishers, The Hague. Eisner, T. and M. Eisner. 2000. Defensive use of a fecal thatch by a beetle larva (Hemisphaerota cyanea). Proceedings of the National Academy of Sciences USA 97: 2,632—2,636. Eisner, T., K. Hicks, M. Eisner, and D. S. Robson. 1978. ‘‘Wolf-in-sheep’s-clothing” strategy of a predaceous insect larva. Science 199: 790-794. Gomez, N. E., L. Witte, and T. Hartmann. 1999. Chem- ical defense in larval tortoise beetles: essential oil composition of fecal shields of Eurypedus nigro- signata and foliage of its host plant, Cordia cur- assavica. Journal of Chemical Ecology 25: 1,007— 1EO2TA Livingston, D. 1978. On the body outgrowths and the phenomenon of “‘sweating” in the nymphal in- stars of Tingidae (Hemiptera: Heteroptera). Jour- nal of Natural History 12: 377-394. Mason, R. T., H. M. Fales, M. Eisner, and T. Eisner. 1991. Wax of a whitefly and its utilization by a chrysopid larva. Naturwissenschaften 78: 28—30. Milbrath, L. R., M. J. Tauber, and C. A. Tauber. 1993. Prey specificity in Chrysopa: an interspecific com- parison of larval feeding and defensive behavior. Ecology 74: 1,384—1,393. Miiller, C. and M. Hilker. 1999. Unexpected reactions of a generalist predator towards defensive devices of cassidine larvae (Coleoptera, Chrysomelidae). Oecologia 118: 166-172. New, T. R. 1969. Notes on the debris-carrying habit in larvae of British Chrysopidae (Neuroptera). En- tomologist’s Gazette 20: 119-124. Oliver, J. E., W. R. Lusby, and J. W. Neal Jr. 1990. Exocrine secretions of the andromeda lace bug Stephanitis takeyai (Hemiptera: Tingidae). Journal of Chemical Ecology 16: 2,243—2,252. Olmstead, K. L. and R. EF Denno. 1992. Cost of shield defence for tortoise beetles (Coleoptera: Chryso- melidae). Ecological Entomology 17: 237-243. . 1993. Effectiveness of tortoise beetle larval shields against different predator species. Ecology 74: 1,394-1,405. Principi, M. M. 1940. Contributi allo studio dei neu- rotteri italiani. 1. Chrysopa septempunctata Wesm. e Chrysopa flavifrons Brauer. Bollettino di Entomologia della Universita di Bologna 12: 63— 144. . 1946. Contributi allo studio dei neurotteri it- aliani. 4. Notochrysa italica Rossi. Bollettino di Entomologia della Universita di Bologna 15: 85— 102. Slocum, R. D. and J. D. Lawrey. 1976. Viability of the epizoic lichen flora carried and dispersed by green lacewing (Nodita pavida) larvae. Canadian Journal of Botany 54: 1,827—1,831. Smith, R. C. 1922. The Biology of the Chrysopidae. Cornell Agricultural Experiment Station Memoir 58-121: 1,287—1,376. Tauber, C. A., M. J. Tauber, and L. R. Milbrath. 1995. Individual repeatability and geographical variation in the larval behavior of the generalist predator, Chrysopa quadripunctata. Animal Behaviour 50: 1,391-1,403. PROC. ENTOMOL. SOC. WASH. 104(2), 2002, pp. 447-457 TAXONOMIC REVISION OF THE GENUS MEGOURA BUCKTON (HEMIPTERA: APHIDIDAE) FROM THE KOREAN PENINSULA WITH THE DESCRIPTION OF A NEW SPECIES AND A KEY TO THE WORLD SPECIES SEUNGHWAN LEE, JAROSLAV HOLMAN, AND JAN HAVELKA (SL) Division of Entomology, National Institute of Agricultural Science and Technol- ogy, Suwon, 441-707 Korea (e-mail: seunglee@rda.go.kr); (JH, JH) Institute of Ento- mology, Czech Academy of Sciences, BraniSovska 31, Ceské Budéjovice, Czech Republic Abstract.—Three species of Megoura are recognized from the Korean Peninsula, among which Megoura nigra Lee, n. sp., and the fundatrix, hitherto unknown morph, of M. crassicauda Mordvilko 1919 are described. Megoura nigra can be distinguished easily from other species of Megoura by its dark brown body color in life. It also differs from the closely related species M. crassicauda and M. viciae Buckton 1876 by its relatively long ultimate rostral segment, 0.88—1.00 2HT (0.63—0.87X in the latter two species), mandibular laminae, antennal segment I, abdominal tergite II], and genital plate with 6— 8, 13-20, 16-21, and 24-33 hairs respectively (3—5, 8-15, 12-18, and 14—23 in the latter two species). After examining and measuring the specimens of all known Megoura spp.., a worldwide key to species is presented. Key Words: The Genus Megoura Buckton 1876 is a small genus of the tribe Macroshipini (He- miptera: Aphididae) with six valid species described from the Palearctic Region; three (M. crassicauda (Mordvilko 1919), M. les- pedezae (Essig and Kuwana 1918), and M. brevipilosa (Miyazaki 1971)) from East Asia, one (M. dooarsis (Ghosh and Ray- chaudhuri 1969)) from the Indian subre- gion, and two (M. viciae Buckton 1876 and M. litoralis Miller 1952 in Borner 1952) from Europe, Central Asia, and the Middle East (Remaudieére and Remaudiéere 1997, Blackman and Eastop 2000, Miyazaki 1971). This genus is characterized by hav- ing swollen siphunculi and, so far as is known, living only on limited genera of Le- guminosae: Vicia Tourn. ex Linn., Lathyrus Linn., Hedysarum Linn., Indigofera Linn., Cajanus DC., Desmodium Desv., and Les- pedeza Michx. All known species are green Hemiptera, Aphididae, Megoura, Korea, key to world species in life, and some species have antennae, legs, siphunculi, and cauda dark brown. In the Korean Peninsula, two species (M. crassicauda and M. lespedezae) have been recorded by Okamoto and Takahashi (1927) and Paik (1965, 1972). These two species are very common throughout the Korean Peninsula where their host plants occur. In 1999, we collected dark brown colo- nies of Megoura on Vicia venosa Maxim. Subsequently, we have collected and ex- amined many samples of Megoua through- out South Korea including Jeju Island. Moreover, many South Korean specimens stored in the National Institute of Agricul- tural Sciences and Technology, and North Korean specimens collected by Jan Havelka in 1985, 1987, and 1988, were also exam- ined. As a result, three species are recog- nized, among which the dark brown M. ni- gra is described as new to science and the 148 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON fundatrix, hitherto unknown morph, of ™. crassicauda is reported for the first time. In addition, after examining and measuring the specimens of all known Megoura spp., a worldwide key to the species of the genus Megoura is presented. Abbreviations used for descriptions in this paper are as follows: Ant.I, I, III, IV, V, VIb = antennal segment I, II, III, IV, V, and the base of Ant.VI, respectively; PT = processus terminalis; URS = ultimate ros- tral segment; ML = mandibular laminae; 2HT = second segment of hind tarsus; SIPH = siphunculus. Names of host plants were checked by “The Plant Names Project (1999). Interna- tional Plant Names Index. Published on the Internet; http://www.ipni.org [accessed 29 September 2001].” All specimens examined in this paper are housed in the National Institute of Agricul- tural Science and Technology (NIAST), Su- won, Korea, and the Institute of Entomol- ogy, Czech Academy of Sciences (IE CAS), Ceske Budejovice, Czech Republic. The holotype and paratypes of M. nigra, n. sp., are housed in NIAST. Megoura Buckton 1876 Megoura Buckton 1876: 64 (type species: Megoura viciae Buckton 1876). Drepaniella del Guercio 1913: 188 (type species: Aphis viciae Kaltenbach 1843, not Fabricius 1781 = Megoura viciae Buckton 1876). Neomegouropsis Ghosh, Basu and Ray- chaudhuri 1977: 584 (type species: Me- gouroparsus dooarsis Ghosh and Ray- chaudhuri 1969). Megoura nigra Lee, new species (Figs. 1-12, Table 1) Description.—Alate viviparous female. Color (in life): Entirely dark reddish brown, almost black. Color (in macerated speci- mens): Head including antennae and ros- trum dark brown. Prothorax pale brown; meso- and metathorax pale with spinal and marginal pigmented sclerites. Abdomen pale with marginal and occasionally dorsal small pigmented sclerites at base of hair; ante- and postsiphuncular sclerite well pig- mented; tergite VII and VIII with pale brown horizontal bands. Legs dark brown except bases of femora and tibiae from base to distal 1/5. SIPH and cauda dark brown. Morphology: Body 3.13—4.28 mm long. Head: Smooth with 4 pairs of acute dorsal hairs; antennal tubercle well developed, bearing 5—7 hairs, frons with | pair of ven- tral hairs. Antenna 3.43—4.63 mm long, as long as or longer (0.99—1.29X) than body length; Ant.[ smooth with 13—20 hairs; Ant.I] smooth with 6-8 hairs; Ant.III smooth with 28—64 secondary rhinaria; lon- gest hair on Ant.III 0.60—0.85 x the basal width of segment; Ant.[V and Ant.V im- bricated; primary rhinarium on Ant.V cili- ated, longest diameter distinctly shorter than middle width of Ant.V; Ant.VIb im- bricated with 3—4 hairs; PT imbricated, 3.42—-4.13X as long as base of Ant.VIb. Rostrum attaining frontal margin of hind coxae; clypeus with 4 hairs; mandibular laminae (ML) with 6—8 hairs; URS wedge- shaped, as long as or slightly shorter (0.88— 1.00) than 2HT with 2 pairs of secondary hairs. Thorax: Prothorax with 2—3 mesial and 3 marginal hairs anteriorly. Hind coxae spinulated with ca. 14 acute hairs; hind tro- chanter smooth with 3 hairs; hind femur smooth, more than 2 as long as SIPH; hind tibia smooth; first tarsal chaetotaxy 3: 3:3; 2HT imbricated with 2—4 dorsal, 6—7 ventral hairs. Abdomen: Abdominal dorsum membranous with marginal pigmented sclerites on abdominal tergites II-IV, and small pigmented sclerites at base of hairs; ante- and postsiphuncular sclerites large, well developed; 16—21 hairs on tergite III including marginal ones, 6—11 on tergite VI between SIPH, and 4-8 on tergite VIII; genital plate weakly pigmented, spinulated with 2—4 median long hairs and 22-30 short hairs on posterior margin. SIPH 1.09— 1.33 cauda, swollen in middle, middle di- ameter 2X as wide as the distal diameter. Cauda elongated, tapering to apex with 10— VOLUME 104, NUMBER 2 449 Figs. 1-12. Megoura nigra. 1-8, Apterous viviparous female. 1, Dorsal and ventral surface of head. 2, Antennal segment III (Ant.III). 3, Antennal segment IV (Ant.IV). 4, Antennal segment V (Ant.V). 5, Antennal segment VI (Ant.VIb + PT). 6, Tarsal segments. 7, Third and ultimate segment of rostrum (URS). 8, Siphunculus. 9, Cauda. 10-12, Alate viviparous female. 10, Antennal segment II (Ant.III). 11, Antennal segment IV (Ant.IV). 12, Antennal segment V (Ant.V). Scale bars equal 0.5 mm for Figs. 1-5, 8-12 and 0.25 mm for Figs. 6—7. +50 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON \5 hairs, denticulated at basal half, spinu- lated in a group of 2-3 spinules at distal half. Alate viviparous female. Color (in life): Thorax entirely dark brown. Abdomen with large marginal sclerite on each segment. Wings transparent with narrow dark pig- mentation along veins and stigma. Mor- phology: Ant. and Ant.IV with 62—89 and 19—37 secondary rhinaria respectively. Otherwise like apterous viviparous female. Apterous oviparous female. Hind tibia swollen with numerous pseudosensoria. Otherwise like apterous viviparous female. Alate male. About 24 and 18 secondary rhinaria on Ant.IV and Ant.V, respectively. Abdomen with additional dark horizontal pigmented sclerites on each segment. Oth- erwise like alate viviparous female. Measurements: See Table 1. Type material—Holotype. Apterous vi- viparous 2, South Korea: Gyounggi-do: Pocheon: Gwangreung National Arbore- tum, )i1-y.2001, Shde no 0105 11-sh- Ol(apt.5), on Vicia venosa Maxim. Paratypes. 32 apterous viviparous @, 39 alate viviparous 2, same collection data of holotype; 15 apterous viviparous 2, same locality of holotype, 21.v.1999, coll. #990521-16sh, on V. venosa; 2 alate 3, 3 Oviparae, same locality, 19.x.2000, coll. #001018-sh-33, on V. venosa. Biology and host plants.—So far, this species has been observed only on V. ven- osa in the Gwangreung National Arbore- tum, South Korea. It lives on young stems or undersides of young leaves. Males and Oviparae occur in the middle of October, and it is monoecious holocycly on V. ven- osa. Distribution.—South Korea. Etymology.—The species name nigra is derived from the dark reddish brown to al- most black body color in life, whereas all other species of Megoura are green in life. Notes.—Morphologically, this species is similar to M. crassicauda and M. viciae from which it can be distinguished by its entirely dark brown body color (green in the latter species), relatively long ultimate rostral segment, 0.88—1.00 2nd hind tar- sus (0.63—0.87X in the latter species), 6—8, 13-20, 16—21, and 24—33 hairs on mandib- ular laminae on each side, antennal segment I, abdominal tergite III, and genital plate respectively (3—5, 8—15, 12-18, and 14—23 in M. crassicauda and M. viciae), and rel- atively long siphunculi, 1.08—1.33 cauda (siphunculi shorter than cauda in M. cras- sicauda) (see Table 1 and Table 2). It also can be easily separated from other species of Megoura by dark brown body color. Megoura crassicauda Mordvilko 1919 (Figs. 13-21) Megoura viciae crassicauda Mordvilko ONO 327 Rhopalosiphum viciae var. japonicum Mat- sumura 1918: 10. (Invalid by Hiile Ris Lambers 1965.) Nectarosiphum moriokae Shinji 1923: 308. (Syn. by Moritsu 1948.) Megoura viciae japonica: Moritsu 1948: 84; Tao 1963: 183. Megoura japonica: Okamoto and Taka- hash 1927:2133: Megoura viciae coreana Moritsu 1948: 84; Paik 1965: 72. (Syn. by Hille Ris Lamb- ers 1965.) Nectarosiphum moriokae Shinji 1923: 308. (Syn. by Moritsu 1948.) Amphorophora lathyri Shinji 1924: 365. (Syn. by Moritsu 1948.) Megoura lathyri: Shinji 1941: 897. Amphorophora vicicola Shinji 1941: 773. Megoura crassicauda: Hille Ris Lambers 19652195; Description.—Fundatrix. Color (in life): Body green except head, antenna, legs, si- phunculi, and cauda black or dark brown. Color (in macerated specimens): Head in- cluding antenna and rostrum dark brown. Thorax pale with irregular dorsal and lateral dark sclerites; legs dark brown except ex- treme bases of femora pale brown. Abdo- men pale; antesiphuncular sclerite dark brown, postsiphuncular sclerite small; ter- VOLUME 104, NUMBER 2 Table 1. Biometric data of Megoura nigra. Length of (Gn mm) No. hairs on No. rhinaria on Part Body from antennal tubercle to cauda Whole antennae Antennal segment II (Ant. III) Antennal segment IV (Ant. IV) Antennal segment V (Ant. V) The base of antennal segment VI (Ant. VIb) Processus terminalis (PT) Ultimate rostral segment (URS) Hind tibia Hind femur Hind tarsus II (2HT) Siphunculus (SIPH) Cauda Antennal segment I (Ant. I) Ultimate rostral segment (URS) Mandibular laminae (ML) Tergite II] Tergite VI between SIPH Tergite VIII Genital plate Cauda Ant. III Ant. IV Apterous vivipara (n = 10) Min.—Max. (Avr.) 3.13—4.28 (3.75) 3.43-4.63 (4.13) 0.96—1.32 (1.17) 0.56—0.88 (0.77) 0.56—0.80 (0.71) 0.21—0.29 (0.26) 0.75—1.08 (0.98) 0.13—0.14 (0.139) 1.93—2.70 (2.39) 1.11—1.63 (1.44) 0.14—0.16 (0.150) 0.50—0.70 (0.60) 0.45—0.60 (0.51) 13—20 (15.75) 4—4 (4.00) 6-8 (6.90) 16-21 (19.00) 6-11 (7.50) 4-8 (5.70) 24—33 (28.10) 10-15 (12.20) 28-64 (53.30) 0-4 (0.20) Alate vivipara (n = 10) Min.—Max. (Avr.) 3.00—3.88 (3.37) 3.38-4.63 (4.21) 1.00—1.21 (1.10) 0.73—0.91 (0.82) 0.65—0.81 (0.73) 0.23—0.29 (0.25) 0.90-1.15 (1.06) 0.13—-0.15 (0.141) 2.20—2.95 (0.45) 1.19-1.63 (1.36) 0.12—0.15 (0.14) 0.45—0.58 (0.53) 0.38—0.50 (0.45) ISSO CS25) 44 (4.00) 6-8 (6.60) 20-24 (22.30) 5—9 (7.10) 4—6 (5.50) 23-31 (26.00) 12—14 (12.60) 62-89 (71.90) 19=37 (28-85) gite VII and VIII with transverse dark brown bands. SIPH, cauda, and genital plate dark brown. Morphology: Body oval or short spindle- shaped. Head: Smooth with 4 pairs of short hairs; antennal tubercle developed with 2— 5 hairs on each side; longest hair on dorsum shorter (0.7) than basal width of Ant.III. Antenna short, 0.56—0.70 body length; Ant.I and Ant.II smooth or slightly spinu- lated, bearing 6—10 and 3-6 hairs, respec- tively; Ant.III smooth with 1—4 secondary rhinaria on basal %; Ant.IV short, less than 0.5 Ant.III, weakly imbricated; Ant.V im- bricated, bearing small primary rhinarium, longest diameter less than 0.5 as long as middle width of Ant.V; Ant. VI strongly im- bricated: PT short 1.65=2:23X ‘base of Ant.VI; longest hair on Ant.III %X_ basal width of segment. Rostrum attaining me- socoxae; Clypeus with 4 hairs; ML with 2-4 hairs on each side; URS 0.92—1.08X and 0.71-1.00X as long as 2HT and Ant.VIb, bearing | pair of hairs. Thorax: Prothorax with | pair of spinal hairs and 2 pairs of marginal hairs anteriorly. Hind coxae spi- nulated with ca. 10 hairs; hind trochanter smooth with 2—3 hairs; hind femur spinu- lated, 1.50—1.89x SIPH, longest hair less than 0.5 basal width of segment; hind tib- ia smooth, longest hair shorter than middle width of segment; first tarsal chaetotaxy 3: 3:3; 2HT imbricated, bearing 3-5 dorsal hairs and 3—4 ventral hairs. Abdomen: Ab- dominal dorsum membranous with 9-11, 4—6, and 5-7 hairs on tergite III, tergite VI between SIPH, and tergite VIII respective- ly; longest hair on abdomninal dorsum less than 0.5X basal width of hind femur; an- tesiphuncular sclerite well developed; post- siphuncular sclerite small or undeveloped; tergites VII and VIII with transverse dark 452 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 14, 19 SPARE RYU AIF SUPsesPEOREACD 13, 15-18, 20-21 Figs. 13-21. Fundatrix of Megoura crassicauda. 13, Dorsal and ventral surface of head. 14, Third and ultimate segment of rostrum (URS). 15, Antennal segment II (Ant.III). 16, Antennal segment IV (Ant.IV). 17, Antennal segment V (Ant.V). 18, Antennal segment VI (Ant.VIb + PT). 19, Tarsal segments. 20, Siphunculus. 21, Cauda. Scale bars equal 0.5 mm for Figs. 13, 15-18, 20-21 and 0.25 mm for Figs. 14, 19. VOLUME 104, NUMBER 2 bands; genital plate well-pigmented, weakly spinulated with 2—6 median hairs and 12— 18 short hairs on posterior margin. SIPH 1.10—1.43X cauda, 0.62—0.78 Ant.III, and 0.53—0.67X hind femur, cylindrical, slightly swollen and widest in middle, smooth at basal half, weakly spinulated on distal half. Cauda elongated tongue-shaped, spinulated ventrally in a group of 1—4 spinules, bear- ing 8—13 hairs. Measurement (mininum-maximum (av- erage) in mm): Body, 2.87—3.43 (3.12). Antenna total, 1.80—2.13 (1.93); Ant.I, 0.15—0.17 (0.16); Ant.II, 0.11—0.12 (0.112); Ant.III, 0.61—0.73 (0.66); Ant.IV, 0.20—0.28 (0.25); Ant.V, 0.29-0.36 (0.32); Ant.VIb, On3—O) 17 (0.154); PT. 025-0 33%(0.29). Hind tibia, 1.25—-1.48 (1.32); hind femur 0.72-0.85 (0.77); 2HT, 0.12—-0.13 (0.124). SIPH, 0.45-—0.50 (0.47). Cauda, 0.34—0.41 (037) URS, 0:25—0.332(0.29). Specimens examined.—Five fundatrix, South Korea: Jeju-do: Bukjeju: Oo-do: Joil- ri, 19.iv.2000, Coll.# 00417-sh-39, on Vicia angustifolia L. Numerous apterous and alate viviparous 2, oviparous @ and alate 3d throughout the Korean Peninsula, on Pis- um sativum Linn., Vicia spp. (amurensis Oettingen, angustifolia L., unijuga A. Br.), and Lathyrus japonicus Willd. Biology and host plants.—This species lives on young stems or underside of leaves. It is holocyclic on Vicia spp. (amu- rensis, angustifolia, cracca, faba, segetalis Thuill., wnijuga), and Lathyrus spp. (davidii Hance, japonicus). Also collected on Pisum sativum in Korea. Distribution.—Korean Peninsula, Russia (Siberia, Primorskii), Japan, China, Taiwan. Note.—Hille Ris Lambers (1965) consid- ered M. viciae subsp. crassicauda Mordvil- ko 1919 as a separate species based on ‘“*having numerous protruding rhinaria over about Y,—7,, of antennal segment III along one side of the segment in apterae and also the antennal segment IV covered with a number of rhinaria in alate.”” According to 453 our examination and measurements for Eu- ropean samples of M. viciae and East Asian samples of M. crassicauda from Korea and Japan, it was found that some European samples from Slovakia have up to 26 sec- ondary rhinaria scattered on Ant. III in ap- terae. Conversely, some specimens of M. crassicauda from Korea have only 20 sec- ondary rhinaria, mostly located in a line as like M. viciae. The fundatrix of crassicauda described here is also closely related to that of M. viciae described by Heie (1995). In spite of these overlapping chracteristics and similarity, all alate samples of M. crassi- cauda from Korea and Japan could be se- perated by having more than 17 secondary rhinaria on antennal segment IV, whereas European M. viciae have no secondary rhi- naria or rarely 1—7 on basal % of Ant.IV Gee Mable 2): Megoura lespedezae (Essig and Kuwana 1918) Rhopalosiphum lespedezae Essig and Ku- wana 1918: 57. Myzus lespedezae: Shinji 1927: 59. Amphorophora_ lespedezae: Shinji 1941: 744; Tao 1963: 184; Paik 1965: 73. Megoura abnormis Ghosh 1970: 7. (Syn. by Ghosh 1973.) Megoura cajanae Ghosh, Ghosh and Ray- chaudhuri 1971: 385. Megoura lespedezae: Miyazaki, 1971: 49; Remaudiére and Remaudiere 1997: 118. Specimens examined.—Numerous apter- ous viviparous @, alate viviparous °, and oviparous 2 throughout the Korean Penin- sula on Lespedeza bicolor Turcz. Biology and host plants.—This species is holocycly on Lespedeza spp. (bicolor Turez., cyrtobotrya Miq.). It lives on the young stem or underside of leaves. Distribution.—Korean Peninsula, Japan, China, Taiwan, India, Switzerland (recently discovered by Giacalone and Lampel 1996). 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VOLUME 104, NUMBER 2 i) KEY TO WORLD SPECIES OF MEGOURA Apterous Viviparous Females . Cauda dark brown or black. On Vicia spp. or LATS HUSSSPPie s spore ces oo Roane ON Ee ke eo 2 Cauda pale yellow, at most fuscous. Not on Vicia spp. . Body totally dark brown or black in life. Tibia pale yellow except apical /; in macerated spec- imens. Ultimate rostral segment (URS) as long as or slightly shorter (0.88—1.00) than 2nd hind tarsus (2HT). Antennal segment I (Ant.I), mandibular laminae (ML), abdominal tergite Il, and genital plate with 13—20, 6-8, 16-21, and 24—33 hairs respectively. Ant.III with usu- ally more than 50(28—64) secondary rhinaria. On Vicia venosa Maxim. South Korea M. nigra Lee, n. sp. Body green except antenna, legs, siphunculus, and cauda black in life. Tibia black or dark brown in macerated specimens. URS distinctly shorter (0.63—0.87 *) than 2HT. Ant.I, mandib- ular laminae, abdominal tergite III and genital plate with fewer hairs, 8—15, 4—5, 15-18, and 14—23 respectively. Ant.III with usually less than 40 (5—42) secondary rhinaria . Antenna with more than 20 secondary rhinaria on Ant.III scattered irregularly over % or throughout the segment. SIPH as long as or frequently longer than cauda. On Vicia spp. (amurensis Oettingen, angustifolia L., cracca Linn, faba Linn., segetalis Thuill., unijuga A. Br.), Lathyrus spp. (davidii Hance, japonicus Willd), and Pisum sativum Linn. East Asia (Korea, China, Taiwan, Japan, Russia (Far East)), and India .... M. crassicauda Mordvilko Antenna usually with less than 20 secondary rhinaria on Ant.III, confined to basal half or % in a line. SIPH usually shorter than cauda, at most equal. On Vicia spp. (cracca, faba, sativa Linn.) and Lathyrus spp. (pratensis Linn, mon- tanus Bernh.,). Europe, Central Asia, Middle BastwethiOpials a6 on saa M. viciae Buckton . SIPH pale, shorter (0.67—0.95) than cauda. Ant.If with more than 10 secondry rhinaria. All legs pale except distal end of tibiae and tarsi pale brown. Processus terminalis (PT) 3.55—3.67X as long as base of Ant.VI. On Lathyrus maritimus Bigel. Northern Europe (Denmark, Sweden, Finland, Norway, Poland, northrn Germany) M. litoralis Miiller SIPH dark brown or black, distinctly longer than cauda. Ant.II] with fewer than 10 second- ary rhinaria. Legs dark brown, at least fuscous on distal half of femur and tibiae. PT more than 4.5 as long as base of Ant.VI . URS 0.89-1.17X as long as 2HT, 0.68—0.88 x Ant.VIb. Antennal tubercle weakly developed. i) Frons more than twice as wide as median depth. Antenna short, 0.83—0.94 as long as body length. SIPH 1.07—1.31X as long as Ant.IIl. On Lespedeza spp (bicolor Turez., cyr- tobotrya Miq.), Cajanus cajan Druce, Desmo- dium trifolium (L.) DC. East Asia (Korea, Ja- pan, China, Taiwan), India, Swizerland (re- cently discovered by Giacalone and Lampel 1996) M. lespedezae (Essig and Kuwana) URS 0.71-0.83X as long as 2HT, 0.47-0.54X Ant.VIb. Antennal tubercle well developed. Frons V-shaped, as wide as median depth. An- tenna at least 1.3% as long as body length. SIPH shorter than Ant.III . Cauda short, 0.5 as long as SIPH. SIPH with narrow base, basal diameter shorter than mid- dle diameter. Hairs on Ant.IIl 0.5 as long as basal width of Ant.III. On Indigofera (dosua Wall., gerardiana R. Grah., teysmanni Miq.), Hedysarum campanulatum. Indian Subregion (India, Pakistan, Afghanistan, Kashmir), and Thailand: 22 See a ee ee ee 3e8 ah pene M. dooarsis (Ghosh and Raychaudhuri) Cauda elongated, more than 0.7 as long as SIPH. SIPH widest at base. Hairs on Ant.III very short, 4X as long as basal width of Ant.lI. On Lespedeza bicolor. Alate vivipa- rous female unknown. Japan M. brevipilosa Miyazaki Alate Viviparous Females . Cauda dark brown or black. On Vicia spp. or EGU FUSES Pps vane ares pee eS 2 Cauda pale yellow, at most fuscous. Not on Vicia spp. . Ultimate rostral segment (URS) as long as (0.93-1.17) 2HT. Ant.I, mandibular laminae (ML), abdominal tergite III, and genital plate with 13-19, 6-8, 20—24, and 23-31 hairs re- spectively. Ant.III with 62—89 secondary rhi- naria. Body entirely dark brown or black in life. Tibia pale yellow except extreme base and apical '/; in macerated specimens b SUP Rey aos na ten a aha ae M. nigra Lee, n. sp. URS distinctly shorter (0.66—0.86) than 2HT. Ant.I, ML, abdominal tergite III, and genital plate with fewer hairs, 9-15, 3-5, 15—22, and 16-24, respectively. Ant.[II with relatively fewer secondary rhinaria (23—60). Body green except head and thorax including antenna, legs, siphunculi, and cauda black in life. Tibia black or dark brown in macerated specimens ..... 3 . Antenna with 46-64 secondary rhinaria on Ant.UI. Ant.1V with 17-31 secondary rhinaria scattered throughout the segment. SIPH as long as or frequently longer (1.00—1.11) than cau- Ga. Oe Peis ee M. crassicauda Mordvilko 456 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Antenna usually with less than 40 (23-38) sec- ondary rhinaria on Ant.UI. Ant.[V without or rarely 1—7 secondary rhinaria on basal % in a line. SIPH usually shorter (0.76—1.00X) than cauda M. viciae Buckton 4. SIPH pale, shorter (0.88—0.90) than cauda. Ant.U1 with more than 30 (37-38) secondary rhinaria scattered throughout the segment. Ant.I1V with 6—8 secondary rhinaria in a line. Processus terminalis (PT) 3.88—4.16 Ant.VIb Se Mn ene or ca tae Seem age oe M. litoralis Miller — SIPH dark brown or black, distinctly longer (1.30—1.81) than cauda. Ant.III with less than 15 secondary rhinaria in a line. Ant.1V without or rarely 1—2 secondary rhinaria. PT more than SOX Ant Vibes. Aas om cries tie ok 5 5. SIPH 0.93-1.05X as long as Ant.III. Ant. and Ant.IV with 10—15 and 1—2 secondary rhi- naria respectively. URS as long as (0.93- 1.09) 2HT and 0.63-0.78 Ant.VIb. Anten- na slightly longer (1.07—1.21x) than body length M. lespedezae (Essig and Kuwana) — SIPH distinctly shorter (0.64—0.66) than Ant.Iil. Ant.II] with 6—9 secondary rhinaria and Ant.IV without secondary rhinaria. URS distinctly shorter (0.67—0.84) than 2HT and 0.45—0.50* as long as Ant.VIb. Antenna dis- tinctly longer (1.50—1.70X) than body length M. dooarsis (Ghosh and Raychaudhuri) ACKNOWLEDGMENTS We thank Drs. R. L. Blackman, London, UK, M. Sorin, Mie-ken, Japan, and M. Mi- yazaki, Tsukuba, Japan for sending materi- als for comparison and the information on the fundatrix of Megoura spp. Many thanks are also due to Dr. Manya B. Stoetzel, Sys- tematic Entomology Laboratory, USDA, Beltsville, Maryland, and unknown review- er for reading the manuscript and giving many useful comments, and Dr. B. K. Byun, Kwangreung, Korea for permitting us to collect in the National Arboretum. This is a partial result of the Cooperative Research Project, ““Aphids, their parasit- oids, and the biological control of aphids on the Korean Peninsula” between the Na- tional Institute of Agricultural Sciences and Technology, Rural Development Adminis- tration of Korea and the Institute of Ento- mology, Czech Academy of Sciences. LITERATURE CITED Blackman, R. L. and V. E Eastop. 2000. Aphids on the world crops. An Identification and Information Guide. Second Edition. London, 466 pp. Borner, C. 1952. Europae Centralis Aphides. Die Blatt- lause Mitteleuropas. Namen, Synonyme, Wirts- pflauzen, Generationszyklen. Weimar 4: 1—488. Buckton, G. B. 1876. Monograph of the British Aphi- des. Vol. I, 190 pp. Essig, E. O. and S. I. Kuwana. 1918. Some Japanese Aphididae. Proceedings of the California Acade- my of Sciences, 4th Series 8: 35-112. Fabricius, J. C. 1781. Rhyngota. Species Insectorum. 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Trans- actions of the Sapporo Natural History Society 7(1): 1-22. VOLUME 104, NUMBER 2 Miyazaki, M. 1971. A revision of the tribe Macrosi- phini of Japan (Homoptera: Aphididae: Aphidi- nae). Insecta Matsumurana 34(1): 1-247. Mordvilko, A. 1919. Aphidodea. Faune de la Russie et de pays limitrophes, Insectes. Hémiptéres 1: 237-508. Moritsu, M. 1948. The genus Megoura Buckton in Ja- pan, with a note on the variation of the external characters in Megoura viciae japonica (Matsu- mura). Mushi 18: 83-88. Okamoto, H. and R. Takahashi. 1927. Some Aphididae from Corea. Insecta Matsumurana 1(3): 130-148. Paik, W. H. 1965. Aphids of Korea. Publishing Center, Seoul National University, Seoul, Korea, 160 pp. . 1972. Illustrated Encyclopedia of Fauna & Flora of Korea. Vol. 13. Insecta (V), 751 pp. Remaudiere, G. and M. Remaudiére. 1997. Catalogue 457 of the world’s Aphididae. Homoptera Aphidoidea. Institut National de la Recherche Agronomique, 473 pp. Shinji, O. 1923. New aphids from Saitama and Mori- oka. Zoological Magazine 35(417): 301-309. (In Japanese.) . 1924. New aphids from Morioka. Zoological Magazine 36(431): 343-372. (In Japanese.) . 1927. Studies on the germ cells of aphids with special reference to the evolutional significance of the chromosomes. Bulletin of Morioka Imperial College of Agriculture and Forestry 11: 1-121. . 1941. Monograph of Japanese Aphididae (in Japanese). Tokyo: 1,215 pp. Tao, C. C. 1963. Revision of Chinese Macrosiphinae (Aphididae, Homoptera). Plant Protection Bulle- tin, Taiwan 5(3): 162-205. PROC. ENTOMOL. SOC. WASH. 104(2), 2002, pp. 458-467 BIOLOGY, IMMATURE STAGES, AND REDESCRIPTIONS OF HYDRELLIA PERSONATA DEONIER (DIPTERA: EPHYDRIDAE), A LEMNA MINER J. B. Kemper, D. L. DEONIER, J. JIANNINO, M. SANFORD, AND W. E. WALTON (JBK) Department of Invertebrate Zoology, Cleveland Museum of Natural History, 1 Wade Oval Drive, Cleveland, OH 44106, U.S.A. (e-mail: jkeiper@cmnh.org); (DLD) Ad- junct Professor of Systematic Entomology, Department of Biology, Pittsburg State Uni- versity, Pittsburg, KS 66762, U.S.A.; (JJ, MS, WEW) Department of Entomology, Uni- versity of California, Riverside, CA 92521, U.S.A. Abstract.—Hydrellia personata Deonier (Diptera: Ephydridae) is a rarely collected shore fly found in the western and midwestern United States. We encountered a population in a constructed wetlands in southern California where the larvae were miners of duck- weed (Lemna minor L.). Eggs were inserted between the upper and lower layers of epi- thelial tissue. Newly hatched larvae either bored into a duckweed thallus or separated the upper and lower epithelial layers to gain access to the photosynthetic tissue. Each instar mined several plants prior to molting inside of a plant thallus, and pupariation occurred inside a hollowed-out thallus. Several pupae were parasitized by the braconid Cyrtogaster clavicornis Walker. Overall, the biology and morphology of the immature stages were very similar to that of the closely related genus Lemnaphila. We describe all immature stages and redescribe the adult (male and female) based upon the newly collected mate- rials. The unusual adult morphology of this species prevents its placement into any of the existing species groups of Hydrellia. Key Words: Lemna, wetlands Shore flies (Diptera: Ephydridae) repre- sent a species-rich family of acalyptrate Diptera with a world-wide distribution (Mathis and Zatwarnicki 1995). Most spe- cies are intimately linked to aquatic and semi-aquatic habitats, and this family dis- plays vast adaptive radiation in larval feed- ing habits. Shore flies exploit detritus, bac- teria, cyanobacteria, diatoms, green algae, plants, decaying animal carcasses, and prey on heterospecific invertebrates (Foote 1995). Although many shore flies are as- sociated with aquatic macrophytes as sec- ondary stem borers (e.g., Deonier 1999, Keiper et al. 2001) or use plant stands as refugia (Todd and Foote 1987, Keiper et al. shore flies, Hydrellia, aquatic insects, leaf miner, herbivory, duckweed, 1998, Keiper and Walton 2000), the genera Hydrellia Robineau-Desvoidy, Lemnaphila Cresson, and Cavatorella Deonier represent the only known ephydrid leaf miners from aquatic habitats (Deonier 1998). Duckweed (Lemna minor L.: Lemna- ceae) is a characteristically minute plant whose mature individuals are 1-3 mm long and 0.5—1 mm wide (Mason 1957). Despite its small size, certain species of Hydrellia (Deonier 1998) and all Lemnaphila_ spp. (Scotland 1934, 1939; Mathis and Edmiston 2000) exploit duckweed as a host plant. The best studied examples of ephydrids associ- ated with Lemna are Lemnaphila scotlan- dae Cresson and Hydrellia williamsi Cres- VOLUME 104, NUMBER 2 son (Williams 1938, Scotland 1939, Mathis and Edmiston 2000). Larvae mine the pho- tosynthetic tissues of multiple plants prior to pupariation, and puparia are formed within the last larval host plant. Mansor and Buckingham (1989) discussed the possible use of L. scotlandae in biocontrol efforts of large populations of duckweed and dem- onstrated its restricted host range using ovi- positional and larval development studies. During investigations of the distribution and abundance of Diptera in the Prado Con- structed Wetlands (CA, Riverside Co.), we encountered mined duckweed plants con- taining puparia. Lemnaphila scotlandae and H. griseola (Fallén) (Grigarick 1959) were the only two previously known Nearctic miners of duckweed. The distribution of L. scotlandae is restricted to areas east of the Mississippi River (Mathis and Zatwarnick1 1995, Mathis and Edmiston 2000) and H. griseola is highly polyphagous (Deonier 1998). Adults reared from the duckweed plants proved to be Hydrellia personata Deonier (1971), a relatively rare species for which few specimens are available. We pro- vide a redescription of the male and female based on the new material obtained, and de- scribe the immature stages and general bi- ology. MATERIALS AND METHODS The Prado Constructed Wetlands are a series of freshwater marshes interconnected by water control structures, encompass more than 125 ha, and are supplied with water from the Santa Ana River. The wet- lands support a mosaic of emergent and submerged vegetation, most notably Cali- fornia bulrush (Schoenoplectus californicus [Meyer] Sojak), cattails (Typha spp.), lesser duckweed (Lemna minor L.), emergent and submerged species of buttercups (Ranun- culus spp.), and pennywort (Hydrocotyle ranunculoides L.). Nearby aquatic habitats include periodically flooded duck club ponds and low gradient intermittent streams. Immature specimens were collected by 459 scooping up duckweed clusters with mos- quito dippers. Samples were scanned with a dissecting microscope at 6-12 to find plants with eggs, larvae, or puparia. Spec- imens were reared in petri dishes, and ac- tive larvae were given undamaged plants to observe feeding. Plants with puparia were placed in petri dishes, kept at laboratory temperatures (18—20° C), and a 16:8 light: dark photoperiod maintained with incandes- cent lights. Representatives of all immature stages were fixed in KAA solution and pre- served in 70% ethanol. Field-collected adults were placed in breeding cages with marsh water and duck- weed for observations of mating behavior, Oviposition, and adult feeding. Small cages manufactured from plastic cups inverted on petri dishes were ineffective, as adults spent most of their time resting at the screened openings. Larger, 2 liter plastic boxes with screened lids seemed to provide a better adult habitat as the flies spent considerable time among the floating duckweed placed within. Breeding cages were exposed to the same photoperiod and temperature regime as larvae. RESULTS AND DISCUSSION The following adult description is based upon the original type series and 31 speci- mens collected by JBK, MS, JJ, and WEW at the Prado Constructed Wetlands, River- side County, CA. For methods, indices, and other terminology see Deonier (1998). Hydrellia personata Deonier (Figs. 1-13) Hydrellia personata Deonier 1971: 86.— Mathis and Zatwarnicki 1995: 85.— Deonier 1998: 35, 42. Diagnosis.—Maxillary palp dark brown, smoothly angular and slightly spathulate with 3—4 apical setae about 0.3+ of palpal length; 5—8 (usually 5—6) dorsal aristal rays; antenna dark brown (velvety in dorsal view); antennomere 3 with sparse (occa- sionally dense) light golden-brown micro- 160 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON pubescence visible in dorsomedial view; frontal vitta and parafrontalia velvety dark- brown pruinose in dorsal view; antenno- mere 2 with usally 2 prominent spinoid ap- icodorsal setae; face, in profile with lower 0.5 slightly convex (not bulging) and with a slight median carina; face sericeous sil- very or light-gray pruinose, contrasting with dark brown, narrow, unilinear parafa- cialia; 3—4 primary facial setae, with usu- ally 1 minute upper secondary facial setula; ocular index 4.0—5.0; head width/head height 1.3—1.7; 1 postsutural (nearly sutur- al) dorsocentral macrochaeta present; pleu- ron densely olive- or reddish-brown prui- nose; legs, except yellowish orange tro- chanters, tibial apices, and tarsal venters, dark brown; tibiae not dilated or expanded; mesonotal disc and abdomen semiglossy or glossy brown in dorsolateral view. Male length 1.28-1.80 mm; female 1.45—2.20 mm. Male postabdomen as in Figs. 1—3 and 5; female postabdomen as in Figs. 4 and 6. Description.—Head: Face, in profile, with lower 0.5 slightly convex (not bulg- ing) and with a slight, but noticeable me- dian carina; face sericeous silvery or light- gray pruinose, contrasting with dark-brown, narrow, unilinear parafacialia; antennal fo- veae indistinct; epistoma squarely recessed (sometimes slightly concave with median indentation) and congruent with dark brown anteclypeus; 3—4 primary facial setae in | row, with 1—3 (usually |) minute, porrect or declinate secondary facial setulae; antenna dark brown (velvety in dorsal view); anten- nomere 2 with usually 2 prominent, spinoid apicodorsal setae; antennomere 3 with sparse (occasionally dense) light golden- brown micropubescence visible in dorso- medial view; 5—8 (usually 5—6) dorsal ar- istal rays; frontal vitta and parafrontale of- ten scarcely differentiated, both appearing velvety dark-brown pruinose in dorsal view (except ocellar triangle sometimes light- brown pruinose); fronto-orbital area con- colorous with parafrontale; anterior fronto- orbital seta 0.3—0.5 length of posterior seta; frons moderately sloping; 12—16 postocular setae in fairly regular row nearest posterior orbit; maxillary palpus dark brown, smoothly angular and slightly spathulate with 3—4 apical setae about 0.3+ of palpal length. Epistomal index 1.0—1.4; mesofacial index 1.5—2.0; vertex index 5.5—7.0; ocular index 4.0—5.0; subcranial index 1.3—2.0; head width/head height 1.3—1.7. Thorax: Postpronotum and notopleuron usually densely olive-brown pruinose, but sometimes dark-brown or light yellowish- brown pruinose; mesontal disc, in dorsolat- eral view, glossy dark brown with sparse to moderately dense olive-brown pruinosity; 3—4 antesutural (1—2 * macrochaetous) and | postsutural (macrochaetous) dorsocentral setae [Riverside Co. CA population with only postsutural (nearly sutural) dorsocen- tral seta macrochaetous]; minute, paired auxiliary apical scutellar setulae usually present between apical scutellar macrochae- tae; pleuron usually concolorous with no- topleuron, but sometimes moderate reddish- brown pruinose; | mesokatepisternal seta (macrochaetous); legs, except yellowish or- ange trochanters, apical 0.2 of tibiae, and tarsal venters, dark brown with sparse to moderately dense olive-brown pruinosity; meso- and metatibiae not dilated or ex- panded. Wing lenth 1.25—2.04 mm; veins dark brown; 6-8 setae on basal end of cos- ta; 3—6 dorsal and 6—9 anterior interfractur- al cotal setae; costal-section ratios: I: I 1.8— 2.2; Ill: IV 3.5—4.0; V: IV 3.0—3.6; M,,, index 1.2—1.5. Abdomen: Terga dark brown with sparse to moderately dense light-brown pruinosity in lateral view, but glossy or semiglossy in dorsolateral and posterodorsal views. Male postabdomen: median 0.3 of sternum 5 broadly concave; anterolateral margin of sternum 5 rounded through 95°—100° angle; copulobus truncate to diagonally truncate posteriorly and somewhat irregularly se- tose. Postgonite bifurcate and paralleling distiphallus for over 0.5 latter’s length; me- dian branch 2.0-—2.5x length of lateral branch of postgonite and with slightly curved postgonite uncus about 0.5 length of VOLUME 104, NUMBER 2 461 Figs. 1-4. Hydrellia personata. 1, Male genitalia, ventral view. 2, Male genitalia, left lateral view. 3, Male genitalia emphasizing phallapodeme, bifurcate postgonite, and distiphallus, left lateral view. 4, Female postab- domen, ventral view. more nearly straight uncus of lateral branch; both postgonite unci directed me- diad toward distiphallus; pregonite much smaller, straight, and covered by fused sur- styli; distiphallus long, digitiform, and often slightly expanded at midlength, upcurved and slightly tapering to nearly blunt apex in lateral view; basiphallus concealed in ven- tral view by fused surstyli (in cleared spec- imens appearing as slightly wider continu- 5 0.18MM Figs. 5—6. ation of distiphallus); phallapodeme, in lat- eral view, darkly sclerotized and forming right angle above basiphallus, but showing no distinct condylar scar or process. Fused surstyli with nonpubescent, acute, antero- medial papilliform projection and with sin- gle (paired) anteriorly projecting macro- chaeta inserted anterolaterally; fused sursty- li length:cercus length (ventral view) about 4.8:1.0. Epandrium (syntergum 9+10) Hydrellia personata. 5, Photomicrograph of male postabdomen with exserted genitalia, left lateral view. 6, Photomicrograph of female postabdomen (partly cleared), left lateral view. Abbreviations: CE, cercus; DP, distiphallus; E, egg; PAP, phallapodeme; PO, postgonite; SS, surstyli; S8, sternum 8; VR, ventral receptacle. 162 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON evenly rounded (semicircular) posteriorly. Female postabdomen: sternum 8 slightly narrower basally than 7, about 1.5 longer than wide, and tapering conically posteriad; tip of sternum 8 with 6—8 long, slightly in- curved setae arranged in a semirosette in ventral view; sterna 6 and 7 nearly qua- drangular, 5 noticeable wider distally than basally; cercus, in lateral view, diagonally to roundly truncate distally, 1.2-1.4x as long as wide, and directed straight posteri- ad. Ventral receptacle cupuliform, about 1.5 deeper than wide. Types.—Hydrellia personata Deonier 1971: 86 [USA. Washington. Grand: O’Sullivan Dam; HT d, Washington State University- G22); 19985 35,42 [revi- sion].—Mathis and Zatwarnicki 1995: 85 [world catalog]. Additional specimens examined.—Cali- fornia: Riverside County, Prado Wetlands, pan trap (VII-2-1999, coll: J. B. Keiper), 3 36,2 2; (VII-10-1999, collector: J. B. Kei- per), 1 6, 3 2; (V-4-2000, collector: J. B. Keiper) 4 6, 18 2 (1 ¢ in Deonier Collec- tion, remainder in collection of Cleveland Museum of Natural History). Distribution.—Found from Washington to southern California, east to Iowa and Texas. Remarks.—The identification of pregon- ite and postgonite was erroneously reversed in the original description. DLD has still not been able to place this species in a spe- cies-group. It differs from the H. prudens and H. tibialis species-groups noticeably by its wide head, and its normal, unexpanded male mesotibia. For the original type-series, the only habitats recorded were sedge meadow and margin of Mono Lake, Cali- fornia. Immature stages.—Egg (n = 5): Length, 0.36—-0.38 mm (x = 0.37). White, ends bluntly rounded, with longitudinal ridges along entire length; ridges interconnected with narrow cross ridges. Micropyle small and inconspicuous (Fig. 7). First instar (n = 2): Similar to third in- star except in the following: Length, 0.87— 0.89 mm (x = 0.88); maximum width, 0.14-0.15 mm (* = 0.15). Body white, somewhat transparent; little distinction among abdominal and cephalic segments. Cephalopharyngeal skeleton reduced, es- pecially dorsal cornu. Breathing tube lack- ing spines (Fig. 8). Second instar (n = 6): Similar to third instar except in the following: Length, 1.13-—1.65 mm (x = 1.40); maximum width, 0.20—0.32 mm (x = 0.26). Body nearly opaque yellowish; little distinction among abdominal and cephalic segments. Cephal- opharyngeal skeleton somewhat reduced. Breathing tube with minute brown spines only. Third instar (n = 7): Elongate, muscoid (Fig. 9). Maximum length, 2.24—2.77 mm (x = 2.47); maximum width, 0.43—0.53 mm (x = 0.47); widest at approximately poste- rior 2/3; yellow in life, fading to opaque white in preservative. Short antennae ante- riorly, dark brown; facial mask generally triangular, with median carina (Fig. 12). Pseudocephalic segment lined with rows of narrow, minute spines that increase in length posteriorly; anterior spines approxi- mately 0.2 length of longest posteior spines. Anterior spiracles absent. Abdomi- nal segments well-defined; anterior abdom- inal segments not strongly adorned, poste- rior segments adorned with 9-10 rows of short brown spines dorsally; breathing tube short, representing <1/10 of total body length, lacking spines; posterior spiracles tipped with brown conical spines ~0.04 mm long, each spine bordered by 4 palmate groups of hydrofuge hairs (Fig. 13); ab- dominal segments 2—7 with fleshy ventral creeping welts adorned with fine spinules; perianal pad oblate, tapering laterad, ~0.15 mm wide. Cephalopharyngeal skeleton typ- ical of Hydrellia; mouthhook strongly curved, no accessory teeth; dental sclerite small and triangular; dorsal cornu tapering posteriorly, posterior cornu truncate poste- riorly, both lacking windows (Fig. 10). Puparium (n = 17): Length, 1.52—2.16 mm (< = 1.91); maximum width, 0.63—0.94 VOLUME 104, NUMBER 2 463 Figs. 7-11. 10, Same, cephalopharyngeal skeleton. 11, Puparium, ventral view. mm (x = 0.74). Amber; segmentation con- spicuous, margin of puparium smooth; broadly rounded anteriorly, tapering poste- riorly, widest anteromedially. Posterior spi- racles dark, curved ventrad (Fig. 11). Third instar cephalopharangeal skeleton pressed flat, visible through puparium. Remarks.—The eggs of H. personata are easily distinguished from those of L. scot- landae (Mathis and Edmiston 2000). Those of L. scotlandae are flattened ventrally, have only four longitudinal ridges, and have a light brown chorion, whereas those of H. personata are not flattened, have more than 4 longitudinal ridges, and the chorion Hydrellia personata. 7, Egg, dorsal view. 8, First instar, lateral view. 9, Third instar, lateral view. is white. In contrast, the mature larvae are very similar in these two Lemna-consuming species, including the general shape, ventral creeping welts, conical posterior spiracles, and morphology of the mouth-hooks of the cephalopharyngeal skeleton. A slight dif- ference exists in the puparia, as L. scotlan- dae are widest posteromedially and H. per- sonata are widest anteromedially. Biology and larval feeding habits.— Adults of H. personata, some of the small- est in the genus, have been collected in five states from the Pacific coast to the Missis- sippi River. This distribution is entirely within the distributional range of the now 464 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 12-13. Hydrellia personata. 12, Scanning electron micrograph of third instar facial mask. 13, Same, posterior spiracles. VOLUME 104, NUMBER 2 known single larval host-plant species, Lemna minor L. Although this host-plant species has now been confirmed through rearings, much remains unknown about the natural history of this rare litthe Hydrellia. Despite records of populations in lowa and Texas, prolonged surveys with floating ad- hesive traps or floating detergent traps by DLD on the many year-round Lemna pools in southeastern Kansas have failed to dis- cover its presence there. Many L. scotlan- dae (over 100) and a few H. griseola and H. bilobifera Cresson along with four Se- tacera sp. and one Discocerina obscurella (Fallén) were the only ephydrids trapped. Adult populations of the Prado Con- structed Wetlands were located only in ar- eas of dense duckweed growth (at least three plants cm-*), where they were ob- served walking or resting on duckweed plants. Some areas of the wetlands with dense growths harbored no adults (or evi- dence of immatures), whereas other areas supported flies in abundance. Their small size and habit of walking on duckweed plants allowed us to observe adults or cap- ture them with vials easily. Although adults exhibited the capacity to skate on the water surface, they did so only rarely and ap- peared to prefer the solid substrate offered by the floating plants. Adults fed by spong- ing the dorsal surface of duckweed plants with their mouthparts and sometimes touched their probosci to the water surface. Unlike L. scotlandae adults (Scotland 1934, Mathis and Edmiston 2000), H. personata did not appear to damage the plants. Flies moved slowly over the surface of thick growths of duckweed, and flew short distances if disturbed. Intraspecific aggres- sion was apparent when conspecifics wan- dered closely (approximately 1—1.5 cm) to a stationary female. The female would rap- idly approach the intruder causing it to re- treat. Males did not display aggressive be- havior during our observations. Eggs were infrequently collected in na- ture, and only one egg was laid during lab- oratory observations. All eggs were insert- 465 ed about half way into the peripheral mar- gin of duckweed plants and positioned so the micropylar end protruded freely from the plant. Three to four eggs, situated ad- jacent to each other, were inserted into each of five field-collected plants; the single egg laid in the laboratory was placed similarly. The small size of the immatures made it impossible to locate them in nature. How- ever, population sizes were large enough that scooping duckweed from between stems of emergent plants produced ample specimens for study and rearing. Newly hatched larvae burrowed into the host plant or stretched their bodies and probed the sur- rounding area until a neighboring plant was located. Larvae latched onto the nearby plant with their mouthhooks and pulled themselves onto the new host. Older larvae used their mouthparts to separate the dorsal and ventral halves of plant fronds and fed on the exposed photosynthetic tissues. Lar- vae crawled into the hollowed area of the plant as tissue was removed. The posterior spiracles remained outside of the plant and in contact with the atmosphere. Two first instars did not separate the dorsal and ven- tral halves of the host frond, but moved to the underside of the plant where they bur- rowed through the epithelial tissue to gain access to the photosynthetic tissues. Young larvae did not hollow plants completely, but created U-shaped mines prior to exiting and moving to another host plant. Conversely, second and third instars always attacked plants by separating the two halves of the frond and consuming all the photosynthetic tissues or nearly so. Molting occurred inside duckweed plants and exuviae remained within the hosts. Pu- paria were formed within hollowed-out duckweed fronds, but the posterior spiracles did not extend to the atmosphere. Damaged duckweed plants remained floating on the water surface, and atmospheric air probably diffused into the damaged plant through the epithelial break formed by third instars. Adults exited plants through the epithelial break. Five adult Cyrtogaster clavicornis 466 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Walker (Hymenoptera: Braconidae) (2f, 7 Oct 1998; 3f, 9-15 Jan 2000) emerged from field-collected puparia situated within duck- weed plants. This parasitoid has also been reared from puparia of H. griseola (Fulmek 1962). Because of difficulty with laboratory rearings and the intimate trophic association of H. personata with duckweed in nature, the precopulation period, fecundity, and seasonal distribution remain unknown. The first specimens collected were puparia taken 22 September 1998, and further larvae and puparia were collected 26 September 1998 and 29 June 1999 for rearing. Adults were observed during January 2000, and puparia within plants were collected among a se- nescing population of duckweed at that time. No larvae or eggs were found during the winter months. Sporadic collections of adults and all immature stages indicate that H. personata is multivoltine and exhibits many overlapping generations in the lati- tude of southern California. The first, sec- ond, and third larval stadia were 2—3, 2-3, and 4—5 days, respectively, with a pupal du- ration of 8 days. The control certain of pestiferous aquatic plants using dipterans has been successful (Center et al. 1997), and has been tested for Lemna (Mansor and Buckingham 1989). Although biocontrol programs against large blooms of duckweed have not been imple- mented rigorously, Hydrellia personata represents such an opportunity. Biocontrol efforts with this shore fly in the western United States may allow for use of a local species rather than having to import an ex- otic one from a geographically distant area. ACKNOWLEDGMENTS Observations were made during a larger investigation of the Prado Constructed Wet- lands funded by the Orange County Water District and the Northwest Mosquito and Vector Control District (CA). We thank Bri- an Baharie of the Orange County Water District for access to the marsh. Apprecia- tion is expressed to J. Triplett, Chairman, Department of Biology, Pittsburg State Uni- versity (KS) for providing research space and equipment. We thank J. Cumming and the Canadian National Collection of In- sects, Arachnids, and Nematodes for the loan of specimens. M. Gates (USDA, Washington, DC) identified the parasitic wasp. J. EK Edmiston (Quincy University, Quincy, IL), B. A. Foote (Kent State Uni- versity, Kent, OH), and W. N. Mathis (Smithsonian Institution, Washington, DC) examined specimens and offered helpful suggestions. Kelli Payto and Wendy Don- kin (Cleveland Museum of Natural History, Cleveland, OH) kindly prepared the pho- tographs. B. A. Foote, J. T. Hannibal (Cleveland Museum of Natural History), and W. N. Mathis reviewed the manuscript. LITERATURE CITED Center, T. D., M. J. Grodowitz, A. E Cofrancesco, G. Jubinsky, E. Snoddy, and J. E. Freedman. 1997. Establishment of Hydrellia pakistanae (Diptera: Ephydridae) for the biological control of the sub- mersed aquatic plant Hydrilla verticillata (Hydro- charitaceae) in the southwestern United States. Bi- ological Control 8: 65—73. Deonier, D. L. 1971. 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Biology and immature stages of Brachydeutera sturtevanti (Diptera: Ephydridae), a hyponeustic generalist. Annals of the Entomological Society of America 93: 468-475. Keiper, J. B., P. L. Brutsche, and B. A. Foote. 1998. Acalyptrate Diptera associated with water willow, VOLUME 104, NUMBER 2 Justicia americana (Acanthaceae). Proceedings of the Entomological Society of Washington 100: 576-587. Keiper, J. B., J. Jiannino, M. Sanford, and W. E. Wal- ton. 2001. Biology and immature stages of Ty- popsilopa nigra (Williston) (Diptera: Ephydridae), a secondary consumer of damaged stems of wet- land monocots. Proceedings of the Entomological Society of Washington 103: 89-97. Mansor, M. and G. R. Buckingham. 1989. Laboratory host range studies with a leaf-mining duckweed shore fly. Journal of Aquatic Plant Management 27: 115-118. Mason, H. L. 1957. A Flora of the Marshes of Cali- fornia. University of California Press, Los Ange- les. 878 pp. Mathis, W. N. and J. K Edmiston. 2000. A revision of the shore-fly genus Lemnaphila Cresson (Diptera: 467 Ephydridae). Proceedings of the Entomological Society of Washington 102: 652—677. Mathis, W. N. and T. Zatwarnicki. 1995. World Catalog of Shore Flies (Diptera: Ephydridae). Memoirs on Entomology, International 4. Associated Publish- ers, Gainesville. 423 pp. Scotland, M. B. 1934. The animals of the Lemna as- sociation. Ecology 15: 290—294. . 1939. The lemna fly and some of its parasites. Annals of the Entomological Society of America 32: 713-718. Todd, J. L. and B. A. Foote. 1987. Spatial and temporal distribution of shore flies in a freshwater marsh (Diptera: Ephydridae). Proceedings of the Ento- mological Society of Washington 89: 448—457. Williams, FE X. 1938. Biological studies in Hawaiian water-loving insects, Part III, Diptera or flies. A. Ephydridae and Anthomylidae. Proceedings of the Hawaiian Entomological Society 10: 85-119. PROC. ENTOMOL. SOC. WASH. 104(2), 2002, pp. 468-478 HAMLET CLARK: PUBLICATIONS AND PROPOSED TAXA C. L. STAINES 3302 Decker Place, Edgewater, MD 21037, U.S.A. Abstract.—Hamlet Clark (1823-1867) was an English cleric who was interested in natural history. He became an authority on the Chrysomelidae, Dytiscidae, Gyrinidae, Haliplidae, and Noteridae (Coleoptera). From 1842 until 1867 he published 39 papers in which he described 71 genera and 709 species. This article presents a brief biographical sketch, a bibliography of his published works, and a list of proposed taxa. Key Words: BIOGRAPHICAL SKETCH The information summarized below was taken from the obituaries of Clark (Anon- ymous 1867; Newman 1867a, b, c; and Clark’s articles 14 and 35). Hamlet Clark was born 30 March 1823 in Navenby, Lincolnshire, England. He was the eldest son of Rev. Henry Clark, Vicar of Harmston, Lincoln. He attended Beverly Grammar School and then studied with the Rev. Mr. Scott where he first became inter- ested in natural history. Clark was a sick child and as a result he was unable to par- ticipate in many activities. Clark attended Corpus Christi College at Cambridge University and was a student with the the Coleopterist T. Vernon Wollas- ton (Jesus College). Cambridge at this time did not offer much formal training in nat- ural history, but Clark read much natural history and purchased many natural history specimens from residents in the Cambridge area. Clark’s early natural history interests were birds, spiders, and Lepidoptera. He devoted the last ten years of his life to the study of Coleoptera, especially water bee- tles and leaf beetles. Clark traveled and collected throughout Great Britain. In the spring of 1856 he took Clark, biography, bibliography, proposed taxa a two month cruise on John Edward Gray’s (Lepidopterist and explorer from the British Museum) yacht visiting many localities in Spain and Algeria. In December of the same year he traveled to Rio de Janeiro, Brazil also with Gray. While in Brazil he collected at Corcovardo Mountain, Pijica (just north of Rio), Constancia, Presidencia (in the Organ Mountains), and Paraihiba. Clark became a fellow of the Entomo- logical Society of London in 1850. He served on the council in 1864—5 and as Vice President in 1864. Clark attended and participated in many Society meetings when he was in London. He became a fel- low of the Linnean Society of London on 21 June 1860. Clark was described as tall and hand- some with a friendly outgoing personality. He served as curate of All Saints Parish in Northampton from 1848 until 1856, and he was conscientious in his ministerial duties. From 1856 to 1863 he lived in London. Due to poor health he left the ministry in 1863. Most of his private journals, field notes, and manuscript notes were destroyed ‘““due to a most untoward accident” prior to his death. At the time of his death he was engaged in working on a catalog of the Hy- dradephaga for the British Museum. He VOLUME 104, NUMBER 2 died 10 June 1867 at Rhyl, Wales, and is buried in the Rhyl Cemetery. Clark’s British Coleoptera and Lepidop- tera were sold at auction in 1865 by J. C. Stevens. His Elateridae, Hydradephaga, and Phytophaga were sold to a Mr. Higgins and were deposited in The Natural History Mu- seum, London, in 1867. Clark’s material was accessioned as number 67-56 and most specimens from the collection should bear a label with this number. ANALYSIS OF CLARK’S PROPOSED TAXA One measure of the value of a taxono- mist’s work is how proposed taxa have been accepted by other scientists. In 39 publica- tions Clark proposed 71 genus group names and 709 species group names. Of the 71 generic names, 17 (23.9%) are now syno- nyms and 6 (8.5%) are homonyms (Seeno and Wilcox 1982). The 23 names represent 32.4% of the names proposed. The species group names were analyzed using the most recent catalogs on each group (Blackwelder 1946; Clavareau 1914; Heikertinger and Csiki 1939, 1940; Jolivet POWs Mvronros 19585, 1959) Weise 1916; Wilcox 1971, 1973; Zimmermann 1920). Of the 709 names proposed, 74 (10.4%) are synonyms and 7 (1%) are homonyms. These 81 names represent 11.4% of the taxa proposed. Clark’s rate of unacceptable names is very high, especially at the generic level. This level of invalid names is fairly normal for a worker who described species rather than doing revisionary work. Many of the genera containing Clark species group names have not been revised since his death, so the percentage of unacceptable names may rise. BIBLIOGRAPHY The author is H. Clark unless otherwise specified. The numbering system was de- vised for this publication. I have attempted to present Clark’s articles in the order they were published. N 469 1842. Captures near Towcester. Entomologist 26: 409-410. 1845. Occurrence of Ceropacha octogesima near Cambridge. Zoologist (1)3: 1,006. 1845. Ctenicerus sanguinicollis. Zoologist (1)3: 1,016. . 1845. Omaseus aterrimus. Zoologist (1)3: 1,016. 1845. Mesosa nubia. Zoologist (1)3: 1,016. . 1845. Captures in the Cambridge fens. Zoologist DSF ONG: . 1845. Scarcity of certain Dytiscidae. Zoologist (1)3: 1,016. 1845. Occurrence of the pine grosbeak near Rochdale. Zoologist (1)3: 1,025. 1848. Notes on the Coleoptera of Northampton- shire. Zoologist (1)6: 2,045—2,046. 1852. Notes on the Coleoptera of Whittlesea Mere. Zoologist (1)10: 3,704—3,707. 1854. Notes on the capture of some of the rarer British Hydrocantharidae. Zoologist (1)12: 4,529—4,532. 1855. Notice and description of a new species of spider. Annals and Magazine of Natural History (2)16: 329-330. 1855. Notes on certain British Hydrocantharidae. Zoologist (1)13: 4,769—4,771. 1855. Synonymic list of the British carnivorous water beetles, together with critical remarks, and notices of foreign allied species. Zoologist (1)13: 4,846—4,869. 1856. A rearrangement of the nomenclature and synonymy of those species of British Coleoptera which are comprised under the sections Geoad- ephaga, Hydradephaga and part of the Philhydri- da, being the first portion of a general British catalogue. London. J. EK Dawson & H. Clark. 8 PPp- 1856. Synonymic list of the British species of Philhydridae, with notices and localities, &c. Zo- ologist (1)14: 5,005—5,008, 5,048—5,056. 1856. Notes on localities for Agabus brunneus and Hydroporus opatrinus. Zoologist (1)14: 5/508) 1857. Proceedings of natural-history collectors in foreign countries. Zoologist (1)15: 5,561—5,565, 5,649—5,652. 1860. Capture of Haltica atropae in Britain. Zo- ologist (1)18: 7,266. 1861 [1860]. Catalogue of Halticidae in the col- lection of the British Museum. Part I. Physapodes and Oedipodes. London. 301 pp. 1862. On the Mexican species of Hydropori. An- nals and Magazine of Natural History (3)10: 173-184. 1862. Description of a new species of Hydropo- rus, Clairv. Annals and Magazine of Natural His- tory (3)10: 326-327. 1863. Descriptions of new east-asiatic species of 20. 34. oS) Nn PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Haliplidae and Hydroporidae. Transactions of the Entomological Society of London (3)1: 417—428. 1864. Catalogue of the Chrysomelidae of South Africa. Annals and Magazine of Natural History (3)14: 115-124, 169-174. 1864 [1862]. Catalogue of the Dytiscidae and Gyrinidae of Australasia, with descriptions of new species. Journal of Entomology 1: 399—421. 1864. Descriptions of species of the genus Hy- droporus Clairy., new to the European or British Catalogues. Journal of Entomology 1: 468—474. 1866 [1864]. Notes on the genus Hydaticus (Leach), with descriptions of new species. Trans- actions of the Entomological Society of London (3)2: 209-222. 1865. An examination of the Dejeanian genus Coelomera (Coleoptera Phytophaga) and its af- finities. Annals and Magazine of Natural History (3)15: 256—268, 315-325. 1866 [1864]. Notes on the genus Schematiza (Phytophaga, Gallerucidae), with descriptions of new species. Transactions of the Entomological Society of London (3)2: 259-270. 1866 [1865]. Descriptions of species of Phyto- phaga received from Pulo Penang or its neigh- bourhood. Annals and Magazine of Natural His- tory (3)15: 139-148. 1866 [1865]. Descriptions of new Phytophaga from western Australia. Transactions of the En- tomological Society of London (3)2: 401—421. 1866. Catalogue of the Dytiscidae and Gyrinidae of Australasia, with descriptions of new species. Journal of Entomology 2: 14—23. 1866. Catalogue of Halticidae; being a continu- ation of the British Museum Catalogue, Part I. 1860. Journal of Entomology, 2: 163-174. 1866. Catalogue of the Dytiscidae and Gyrinidae of Australasia, with descriptions of new species. Journal of Entomology 2: 214—219. 1866. Descriptions of new Australian Phytopha- ga. Journal of Entomology 2: 247-263. 1866 [1864]. An examination of the Halticidae of South America. Journal of Entomology 2: 375-412. 1866. A catalogue of Phytophaga (Coleoptera, Pseudotetramera), Part 1. I. Sagridae, I. Dona- cidae, III. Crioceridae, IV. Megalopidae. With an appendix. Containing descriptions of new species by W. H. Bates and Rey. H. Clark. Williams and Norgate, London, 49 pp. 1866. Descriptions of new species of Phytophaga by W. H. Bates and Rev. H. Clark. Being the appendix to the catalogue of Phytophaga by the Rev. H. Clark. Williams and Norgate, London, 88 pp. 1867. Letters home from Spain, Algeria, and Brazil, during past entomological rambles. J. van Voorst, London, 178 pp. LETTERS AND MENTIONS IN SOCIETY MINUTES A. 1864. John Gray’s insects, collecting at Cape de Verdes. Transactions of the Entomological Society of London (3)2: 17. B. 1865. A letter to the Trustees of the British Mu- seum on the condition of the National Collection of Invertebrata. Robert Hardwicke, London. C. 1865. The flashing of fire-flies. Journal of the Asi- atic Society of Bengal 34: 190-192. D. 1865. The flashing of fire-flies. Transactions of the Entomological Society of London (3)2: 94—95. E. 1865. Note on the saltatorial power of Thaumas- tomerus viridis. Transactions of the Entomological Society of London (3)2: 94. FE 1865. New form of Parnides from Kaffaria. Trans- actions of the Entomological Society of London (3)2: 97-98. G. 1865. About wood-lice. Transactions of the Ento- mological Society of London (3)2: 95. [discussion with W. W. Saunders, J. O. Westwood, and J. J. Weir]. OBITUARIES AND ARTICLES ON HAMLET CLARK, HIS WorK, AND His COLLECTION Anonymous. 1867. Obituary. Proceedings of the Lin- nean Society of London 1867: c—ci. Blackwall, J. 1862—1863. Descriptions of newly dis- covered spiders captured in Rio Janeiro by John Gray, Esq., and the Rev. Hamlet Clark. Annals and Magazine of Natural History (3)10(1862): 348-360, 421—439, (3)11(1863): 29-45. Carpenter, M. M. 1945. Bibliography of biographies of entomologists. American Midland Naturalist 33: 1-116. Derksen, W. and U. Scheiding. 1963. Index Literaturae Entomologiae. Serie Il: Die Welt-Literatur tiber die gesamte Entomologie von 1864 bis 1900. Band I, A-E. Deutsche Akademi der Landwirt- schaftswissenschaften zu Berlin, 697 pp. Gilbert, P. 1977. A compendium of the biographical literature on deceased entomologists. British Mu- seum (Natural History), 455 pp. Horn, W. 1935-1937. Uber entomologische Sammlun- gen, Entomologen & Entomo-Museologie. Ento- mologische Beihefte. Berlin-Dahlem, 536 pp. Horn, W. and S. Schenkling. 1928. Index Literaturae Entomologiae. Serie I: Die Welt-Literatur tiber die gesamte Entomologie bis 1863. Band I. Berlin- Dahlem, 352 pp. Marshall, T. T. 1864. Descriptions of new genera and species of Eumolpinae from the collection of the Rev. Hamlet Clark. Annals and Magazine of Nat- ural History (3)13: 380-389. Musgrave, A. 1932. Bibliography of Australian Ento- mology 1775—1930 with biographical notes on au- VOLUME 104, NUMBER 2 thors and collectors. Royal Zoological Society of New South Wales, 380 pp. Neave, S. A. 1933. The history of the Entomological Society of London, 1833—1933. London, 224 pp. Newman, E. 1867. The late Rev. Hamlet Clark, M.A., EL.S. Entomologist’s Monthly Magazine 4: 43— 44. Newman, E. 1867. Death of Rev. Hamlet Clark. En- tomologist 3(43): 111. Newman, E. 1867. Death of Rev. Hamlet Clark. Zo- ologist (2)2: 840. Royal Society of London. 1867. Catalogue of scientific papers (1800-1863). Volume I. Eyre and Spottis- woode, London, 960 pp. Royal Society of London. 1877. Catalogue of scientific papers (1864-1873). Volume III. C. J. Clay and Sons, London, 1,047 pp. TAXA PROPOSED BY CLARK Taxa are listed alphabetically by genus under each higher taxonomic catagory; the numbers are the publication number fol- lowed by the page number. Class Arachinda Order Araneida Salticidae Salticus blackwalli 8: 329 Class Insecta Order Coleoptera Chrysomelidae Alticinae Aedmon 16: 129 Aedmon sericellum 16: 131 Allochroma 16: 181 Allochroma assimile 16: 191 Allochroma balii 16: 187 Allochroma coccineum 16: 184 Allochroma fasciatum 16: 186 Allochroma festivum 16: 192 Allochroma flavovittatum 16: 188 Allochroma generosum 16: 194 Allochroma humerale 16: 183 Allochroma lunatum 16: 188 Allochroma nigromarginatum 16: 190 Allochroma piceum 16: 184 Allochroma quatuorpustulatum 16: 191 Allochroma sexmaculatum 16: 185 Allochroma sexsignatum 16: 193 Allochroma venustum 16: 189 Apalotrius 16: 166 Apalotrius pubescens 16: 168 Argopus angulicollis 26: 148 Asphaera decipiens 32: 380 Asphaera fallax 32: 380 Asphaera marginata 32: 380 471 Asphaera subfasciata 32: 380 Aspicela 32: 378 Aspicela balyii 32: 381 Aspicela discoidalis 32: 381 Atyphus 16: 175 Atyphus carbonarius 16: 176 Atyphus comes 16: 177 Atyphus flaviventris 16: 177 Atyphus furcipes 16: 178 Cacoscelis bicolorata 32: 408 Cacoscelis clythraefromis 32: 407 Cacoscelis coeruleipennis 32: 407 [homonym, re- placed by coerulea Csiki 1939] Cacoscelis feldneri 32: 406 Cacoscelis fimbriata 32: 408 Cacoscelis flava 32: 407 Cacoscelis nigripennis 32: 408 Cacoscelis testacea 32: 407 Caeporis 32: 398 Caeporis marginata 32: 399 Caeporis subcostata 32: 399 Calipeges 16: 206 Calipeges crispus 16: 207 Caloscelis 32: 408 Caloscelis azuereipennis 32: 409 Cerichrestus 16: 194 Cerichrestus apicalis 16: 197 Cerichrestus balii 16: 196 Cerichrestus batesii 16: 199 Cerichrestus chevrolatii 16: 202 Cerichrestus deyrollii 16: 198 Cerichrestus exiguus 16: 201 Cerichrestus flavicans 16: 204 Cerichrestus humilis 16: 203 Cerichrestus marginicollis 16: 204 Cerichrestus tenuicornis 16: 200 Clamophora clypeata 32: 384 Clamophora generosa 32: 384 Clamophora sanguinicollis 32: 384 Coelocephalus 16: 144 Coelocephalus amoenus 16: 146 Coelocephalus fuscocostatus 16: 148 Coelocephalus pulchellus 16: 145 Coelocephalus pygmaeus 16: 147 Cryptosphaerus 32: 386 Cryptosphaerus ferrugineus 32: 386 Cyrton 16: 267 Cyrton anisotomoides 16: 269 Cyrton sanguineum 16: 269 Diphulaca apicalis 32: 388 Diphulaca costulata 32: 387 Diphulaca dimidiata 32: 388 Diphulaca erythrodera 32: 387 Diphulaca marginata 32: 388 Diphulaca nigroapicata 32: 388 Diphulaca rubens 32: 387 Diphulaca sulcata 32: 386 Diphulaca viridipennis 32: 387 172 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Disonycha adumbrata 32: 401 Disonycha trifasciata 32: 401 Disonycha viridipennis 32: 402 Eratosthenes 31: 261 Eratostehenes flavus 31: 261 Eupeges 16: 107 Eupeges nigrifrons 16: 109 Eupeges praeclara 16: 108 Eupeges scabrosa 16: 109 Euphenges 16: 25 Eupheneges lemoeides 16: 28 Eupheneges sericeus 16: 27 Eutornus 16: 64 Eutornus africanus 16: 65 Exartematopus 16: 169 Exartematopus nobilis 16: 170 Exartematopus scutellaris 16: 171 Febra 31: 261 Febra venusta 31: 262 Gethosynus 16: 179 Gethosynus sanguinicollis 16: 180 Glenidion 16: 93 Glenidion rubronotatum 16: 95 Homammatus 16: 116 Homammatus nitidus 16: 119 Homammatus turgidus 16: 118 Homotypus 16: 120 Homotypus asper 16: 125 Homotypus cionoides 16: 285 Homotypus fuliginosus 16; 124 Homotypus holosericus 16: 12 Homotypus lacunosus 16: 122 Homotypus maculicornis 16: 127 Homotypus squalidus 16: 126 Homotypus vellereus 16: 123 Homotypus wollastonii 16: 128 Hydmosyne 16: 172 Hydmosyne inclyta 16: 174 Hylodromus 16: 142 Hylodromus dilaticornis 16: 143 Hypantherus 16: 95 Hypantherus ambiguus 16: 98 Hypantherus assimilis 16: 99 Hypantherus batesii 16: 100 Hypantherus concolor 16: 97 Hypantherus deyrollii 16: 100 Hypantherus rufotestaceus 16: 101 Hypolampsis 16: 230 Hypolampsis aestivalis 16: 257 Hypolampsis alboguttata 16: 238 Hypolampsis anceps 16: 255 Hypolampsis atra 16: 246 Hypolampsis balii 16: 234 Hypolampsis campestris 16: 243 Hypolampsis costulata 16: 249 Hypolampsis dohrnii 16: 238 Hypolampsis elegantula 16: 234 Hypolampsis fallax 16: 244 Hypolampsis ferrugineonotata 16; 242 Hypolampsis flavonotata 16: 252 Hypolampsis fragilis 16: 247 Hypolampsis fryella 16: 254 Hypolampsis fusca 16: 254 Hypolampsis gibba 16: 259 Hypolampsis inaequalis 16: 236 Hypolampsis lacordairii 16: 248 Hypolampsis melanotus 16: 233 Hypolampsis meridionalis 16: 249 Hypolampsis miersii 16: 237 Hypolampsis minima 16: 245 Hypolampsis multicostata 16: 235 Hypolampsis murraii 16: 239 Hypolampsis nana 16: 241 Hypolampsis nigrina 16: 245 Hypolampsis parallela 16: 243 Hypolampsis porculus 16: 258 Hypolampsis pumilio 16: 242 Hypolampsis robusta 16: 256 Aypolampsis signaticornis 16: 238 Hypolampsis squamata 16: 250 Hypolampsis suborbicularis 16: 260 Hypolampsis sylvatica 16: 251 Hypolampsis vicina 16: 253 Imatium 16: 260 Imatium rotundatum 16: 263 Imatium tomentosum 16: 262 Imatium venutinum 16: 264 Lactica azureipennis 32: 396 Lactica basalis 32: 397 Lactica marginata 32: 396 Lactica pallida 32: 397 Lactica quadrimaculata 32: 395 Lactica seminigra 32: 397 Lactica sponsa 32: 396 Leptotrichus 16: 135 Leptotrichus castaneus 16: 136 Lithonoma africana 16: 272 Litosonycha 32: 377 Litosonycha adumbrata 32: 382 Litosonycha bifasciata 32: 382 Litosonycha nigripennis 32: 381 Litosonycha quadrimaculata 32: 382 Loxoprosopus coeruleus 16: 154 Loxoprosopus humeralis 16: 153 Loxoprosopus marginatus 16: 152 Metriotes 16: 226 Metriotes robinsonii 16: 228 Monoplatus 16: 2 Monoplatus angulatus 16: 20 Monoplatus apicatus 16: 10 Monoplatus bimaculatus 16: 11 Monoplatus croceus 16: 11 Monoplatus dimidiatipennis 16: 19 Monoplatus distinguendus 16: 7 Monoplatus ephippium 16: 280 Monoplatus fryii 16: 278 VOLUME 104, NUMBER 2 Monoplatus grayii 16: 8 Monoplatus impunctatus 16: 21 Monoplatus jucundus 16: 13 Monoplatus miersii 16: 9 Monoplatus nigricans 16: 17 Monoplatus nigrimanus 16: 18 Monoplatus nigripes 16: 5 Monoplatus presidenciae 16: 6 Monoplatus quatuornotatus 16: 14 Monoplatus semichalybeus 16: 15 Monoplatus semiviolaceus 16: 17 Monoplatus sexsignatus 16: 14 Notozona 32: 409 Notozona elegans 32: 411 Notozona humilis 32: 412 Notozona macularia 32: 410 Notozona marmorata 32: 410 Notozona novemmaculata 32: 411 Notozona quatuordecimaculata 32: 408 Notozona rufofusca 32: 411 Notozona sanguinea 32: 412 Notozona sparsa 32: 409 Notozona tenella 32: 412 Notozona transversenotata 32: 411 Octogonotes bicinctus 16: 165 Octogonotes binotatus 16: 163 Octogonotes brunneus 16: 160 Octogonotes sumptuosus 16: 164 Omototus 16: 208 Omototus artitus 16: 219 Omototus bimaculatus 16: 221 Omototus binotatus 16: 222 Omototus bituberculatus 16: 218 Omototus braccatus 16: 212 Omototus cayensis 16: 212 Omototus dohrnii 16: 214 Omototus fulvopubescens 16: 217 Omototus fuscatus 16: 220 Omototus humeronotatus 16: 223 Omototus morosus 16: 209 Omototus nodosus 16: 213 Omototus nubilus 16: 215 Omototus quadripes 16: 210 Omototus sericeopubescens 16: 225 Omototus sexmaculatus 16: 222 Omototus sexnotatus 16: 224 Omototus transversonotatus 16: 225 Omototus tuberculatus 16: 211 Oxygonus 32: 390 Oxygonus adumbratus 32: 392 Oxygonus exornatus 32: 391 Oxygonus fuscomaculatus 32: 394 Oxygonus interruptus 32: 391 Oxygonus luridulus 32: 393 Oxygonus nigripennis 32: 393 Oxygonus rubidus 32: 392 Oxygonus sexnotatus 32: 394 Oxygonus simplex 32: 393 473 Oxygonus succinctus 32: 391 Oxygonus violaceipennis 32: 391 Pachyonychis 16: 61 Pachyonychis paradoxus 16: 63 Panchrertus 16: 137 Panchrertus inconspicuus 16: 140 Panchrertus pulcher 16: 139 Panchrertus rubicundus 16: 139 Panchrertus rufescens 16: 141 Pedilia 32: 384 Pedilia rufa 32: 385 Pelonia 32: 399 |homonym, replaced by Heikertinger- ia Csiki 1939] Pelonia nigripennis 32: 400 Pelonia nigroviolacea 32: 400 Pelonia rubra 32: 400 Pelonia rufotestacea 32: 401 Pelonia vittata 32: 400 Peribleptus 16: 155 [|homonym, replaced by Zeteticus Harold 1875] Peribleptus laevigatus 16: 157 Phaedromus 16: 66 Phaedromus waterhousii 16: 68 Phylacticus 16: 110 Phylacticus amabilis 16: 116 Phylacticus modestus 16: 112 Phylacticus olivaceus 16: 113 Phylacticus pollinosus 16: 115 Phylacticus prasinus 16: 115 Phylacticus ustulatus 16: 113 Physimerus 16: 69 Physimerus adumbratus 16: 84 Physimerus agilis 16: 76 Physimerus allardi 16: 284 Physimerus ambiguus 16: 75 Physimerus angulofasciatus 16: 86 Physimerus batesii 16: 89 Physimerus bilineatus 16: 88 Physimerus bituberculatus 16: 87 Physimerus brevicollis 16: 85 Physimerus emphippium 16: 83 Physimerus fascicularis 16: 80 Physimerus griseostriatus 16: 92 Physimerus impressus 16: 71 Physimerus inornatus 16: 78 Physimerus irroratus 16: 89 Physimerus juvencus 16: 81 Physimerus labialis 16: 74 Physimerus luteicollis 16: 77 Physimerus minutus 16: 79 Physimerus nebulosus 16: 90 Physimerus obscurus 16: 79 Physimerus pruinosua 16: 91 Physimerus quatuorlineatus 16: 72 Physimerus revisus 16: 77 Physimerus rusticus 16: 283 Physimerus suboculatus 16: 83 Physimerus trivialis 16: 81 174 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Physimerus virgatus 16: 73 Physimerus vittatus 16: 72 Physimerus vulgaris 16: 282 Physonychis 16: 29 Physonychis smaragdina 16: 31 Pleurochroma 16: 131 Pleurochroma balteatum 16: 133 Pleurochroma nitidulum 16: 134 Pleurochroma pallidum 16: 134 Poebates 16: 229 Poebates nigripes 16: 230 Procalus 24: 260 Procalus mutans 24: 261 Psilapha 32: 389 Psilapha flava 32: 389 Rhinotmetus 16: 33 Rhinotmetus assimilis 16: 42 Rhinotmetus canescens 16: 52 Rhinotmetus cruciatus 16: 38 Rhinotmetus crucifer 16: 40 Rhinotmetus cyaneus 16: 41 Rhinotmetus cyanipennis 16: 46 Rhinotmetus depressus 16: 43 Rhinotmetus deyrollii 16: 49 Rhinotmetus elegantulus 16: 47 Rhinotmetus flavidus 16: 51 Rhinotmetus humilis 16: 50 Rhinotmetus inornatus 16: 48 Rhinotmetus marginatus 16: 37 Rhinotmetus neglectus 16: 280 Rhinotmetus nigricornis 16: 52 Rhinotmetus pallipes 16: 50 Rhinotmetus ruficollis 16: 45 Rhinotmetus spectabilis 16: 39 Rhinotmetus sulcicollis 16: 47 Rhinotmetus waiterhousti 16: 40 Rhopalotoma 32: 394 Rhopalotoma tuberculatum 32: 394 Rhopalotoma viridipenne 32: 395 Roicus 16: 22 Roicus sexmaculatus 16: 24 Sebaethe flava 26: 147 Sophraena fasciolata 32: 383 Sophraena simplex 32: 383 Sparnus 16: 265 Sparnus globosus 16: 266 Systena brunnipennis 32: 404 Systena discicollis 32: 403 Systena humeralis 32: 404 Systena lugubris 32: 404 Systena marginicollis 32: 403 Systena mustela 32: 405 Systena novemmaculata 32: 405 Systena pectoralis 32: 403 Systena plagiata 32: 403 Systena sinuatovittata 32: 403 Systena suturalis 32: 405 Systena testaceovittata 32: 404 Systena tincta 32: 405 Tenosis 32: 397 Tenosis purpureipennis 32: 398 Tetragonotes 16: 53 Tetragonotes angulicollis 16: 59 Tetragonotes atra 16: 56 Tetragonotes calceata 16: 57 Tetragonotes elegans 16: 55 Tetragonotes hexagona 16: 60 Tetragonotes subanchoralis 16: 58 Tetragonotes vittata 16: 59 Thrasygoeus 16: 102 Thrasygoeus exaratus 16: 104 Thrasygoeus eximius 16: 103 Thrasygoeus obscurus 16: 105 Thrasygoeus undatus 16: 106 Chrysomelinae Australica aeneonitens 27: 416 Australica paropsoides 31: 251 Australica pyrrhocephala 31: 251 Chalcomela pilula 31: 251 Chalcomela subpunctata 27: 417 Chalcolampra laticollis 27: 416 Chalcolampra undulatipennis 27: 415 Chalcolampra verrucosa 3\: 250 Paropsis amoena 27: 405 Paropsis apicata 27: 405 Paropsis captiosa 27: 406 Paropsis incurva 27: 412 Paropsis intertincta 27: 411 Paropsis maculicollis 27: 407 Paropsis mediovittata 27: 404 Paropsis nervosa 27: 413 Paropsis nigritula 27: 411 Paropsis nigroconspersa 27: 409 Paropsis nigropicta 27: 412 Paropsis perparvula 27: 413 Paropsis purpureoaurea 27: 407 Paropsis purpureoviridis 31: 250 Paropsis sanguineotincta 27: 409 Paropsis tessellata 27: 408 Paropsis transversomaculata 27: 410 Paropsis verrucipennis 27: 414 Polysticta clathrata 20: 173 Polysticta consimilis 20: 172 Polysticta eburnipennis 20: 118 Polysticta figurata 20: 172 Polysticta flavosparsa 20: 174 Polysticta hebe 20: 173 Polysticta laevigata 20: 120 Polysticta lineolata 20: 123 Polysticta macularis 20: 119 Polysticta marshalli 20: 121 Polysticta modesta 20: 170 Polysticta multifida 20: 171 Polysticta nigra 20: 121 Polysticta nigroaenea 20: 121 Polysticta nigrofasciata 20: 120 VOLUME 104, NUMBER 2 Polysticta nigrosignata 20: 116 Polysticta picturata 20: 122 Polysticta pulchella 20: 120 Polysticta soluta 20: 122 Polysticta subcruciata 20: 117 Polysticta vigintiguttata 20: 169 Polysticta vigintimaculata 20: 169 Criocerinae Crioceris aestivalis 34: 66 Crioceris callizona 34: 64 Crioceris consobrina 34: 67 Crioceris constricticollis 34: 69 Crioceris crassipennis 34: 65 Criocerus fuscomaculata 31: 249 Crioceris fuscopunctata 34: 67 Crioceris inconspicua 34: 68 Crioceris locuples 34: 64 Crioceris multipunctata 31: 249 Crioceris nigropicta 34: 70 [homonym, replaced by nigroornata 33: 40] Crioceris pusilla 34: 68 Crioceris rufosanguinea 34: 70 Crioceris sanguinea 34: 65 Lema aeneopicta 34: 56 Lema affinis 34: 33 Lema angulicollis 34: 36 Lema angustata 34: 28 Lema antennata 34: 42 Lema atrocaerulea 34: 32 Lema balyi 33: 24 [replacement name for Lema sellata Baly] Lema biannularis 34: 46 Lema bisbivittata 34: 49 Lema bituberculata 34: 35 Lema bizonata 34: 61 Lema bohemani 33: 24 [replacement name for Lema russula Boheman] Lema calabarica 34: 39 Lema chalybea 34: 33 Lema chalybeonotata 34: 37 Lema cingulata 34: 41 Lema circumvittata 34: 41 Lema clavipennis 34: 24 Lema constricta 34: 39 [|homonym, replaced by Lema stricta 33: 28] Lema corinthia 34: 27 Lema crucifera 34: 49 Lema divalis 34: 45 Lema exarata 34: 60 Lema flavomarginata 34: 40 Lema flavofasciata 34: 43 Lema fryii 34: 60 Lema histrio 34: 29 Lema immaculata 34: 38 [homonym, replaced by Lema pura 33: 27] Lema inconstans 34: 26 Lema indeterminata 34: 22 Lema infecta 34: 47 475 Lema innotata 34: 47 Lema latefasciata 34: 53 Lema latipennis 34: 25 Lema lecontei 33: 31 [replacement name for Lema tri- virgata LeConte] Lema maculifrons 34: 59 Lema maculipennis 34: 54 Lema mitis 34: 25 Lema nigroarcuata 34: 45 Lema nigroazurea 34: 27 Lema nigrocoerulea 34: 58 Lema nigrofrontalis 34: 40 Lema nigroornata 34: 46 Lema nigrosparsa 34: 48 Lema nigrosuturalis 34: 37 Lema pallida 34: 21 Lema pallidetestacea 34: 28 Lema piceipennis 34: 59 Lema picipes 34: 34 Lema praeclarior 33: 27 [replacement name for Lema praeclara 34: 38] Lema pubipes 34: 52 Lema punctatipennis 34: 52 S Lema purpureoaenea 34: 57 Lema quinquenotata 34: 50 Lema retucilosa 34: 43 Lema rufoadumbrata 34: 22 Lema rufofemorata 34: 32 Lema rufoornata 34: 30 Lema rufotestacea 34: 29 Lema rufotincta 34: 36 Lema rufozonata 34: 53 Lema semivittata 34: 31 Lema senegalensis 34: 24 [homonym, replaced by Lema bambotana Weise 1913] Lema simplicipennis 34: 34 Lema sinuatonotata 34: 54 Lema sinuatovittata 34: 31 Lema swartzii 33: 25 [replacement name for Lema quadripunctata Swartz] Lema tuberculosa 34: 61 Lema unicolor 34: 23 Lema vicina 34: 51 [homonym, replaced by Lema bra- ziliensis Heinze 1927] Lema violaceofasciata 34: 42 Donaciinae Donacia indica 34: | Eumolpinae Colasposoma aeneovirdie 26: 142 Colasposoma metallicue 26: 142 Colaspoides pulchella 26: 142 Corynoeides 26: 139 Corynoeides tuberculata 26: 140 Damelia 31: 255 Damelia marshalli 31: 256 Edusa aureovirdis 27: 419 Edusa hispidula 27: 420 A76 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Edusa nigroaenea 27: 420 Edusa setosa 27: 419 Geloptera duboulaii 27: 417 Geloptera nodosa 27: 418 Marsaeus 31: 252 Marsaeus flavus 31: 255 Marsaeus nigrocyaneus 31: 253 Marsaeus ruficollis 31: 254 Marsaeus rufoflavus 31: 255 Marsaeus rufus 31: 253 Marsaeus simplex 31: 254 Ocnus 27: 420 Ocnus viridis 27: 421 Psathyrocerus cyanipennis 34: 20 Rhyparada atripennis 26: 141 Rhyparada nitidia 31: 252 Rhyparada rufa 26: 141 Thaumastomerus 27: 418 [homonym, replaced by Aju- bus Aslam 1968] Thaumastomerus viridis 27: 419 Galerucinae Adorium dorsosignatum 31: 258 Adorium fryii 31: 258 Adorium laetabile 31: 259 Adorium seminigrum 31: 258 Alphidia 24: 258 Aulacophora simplicipennis 26: 145 Clitena cyanea 24: 259 Monocesta Monocesta Monocesta Monocesta Monocesta Monocesta Monocesta Monocesta . Monocesta | Monocesta Monocesta Monocesta Monocesta Monocesta Monocesta Monocesta Monocesta Monocesta Monocesta Monocesta Monocesta circumcincta 24: 316 consularis 24: 266 depressa 24: 267 ducalis 24: 265 elegantula 24: 267 equestris 24: 265 flavocincta 24: 316 frontalis 24: 317 fuscescens 24: 318 glauca 24: 317 hopfneri 24: 267 illustris 24: 266 imperalis 24: 264 klugit 24: 317 nigricornis 24: 318 nigriventris 24: 316 obliquenotata 24: 315 rubiginosa 24: 317 sanguinicolis 24: 268 spectanda 24: 315 splendida 24: 266 Nestinus 24: 324 Coelomera Coelomera Coelomera Coelomera Coelomera aerata 24: 322 binotata 24: 32 induta 24: 321 maculicollis 24: 321 modesta 24: 319 8) Coelomera parallela 24: 322 Coelomera Coelomera Coelomera Coelomera Coelomera rufofusca 24: 319 submetallica 24: 322 tenuicornis 24: 323 tibialis 24: 321 violaceopennis 24: 322 Coraia 24: 323 Coraia maculicollis 24: 324 Dercetis 26: 146 [homonym, replaced by Dercetina Gressitt & Kimoto 1963] Dercetis bifasciata 26: 147 Dercetis depressa 26: 147 Dircema 24: 262 Dircema cinctipenne 24: 263 Dircema ruficrus 24: 264 Gallercua semipullata 31: 257 Goniopleura viridipennis 26: 146 Hymenesia 24: 259 Menippus 31: 257 Menippus cynicus 31: 257 Monocesta 24: 264 Monocesta atricornis 24: 318 Monocesta balyi 24: 267 Monocesta carbonaria 24: 316 Monocesta cincta 24: 315 Nestinus bimaculatus 24: 325 Nestinus incertus 24: 325 Nestinus regalis 24: 325 Ochralea 26: 144 Ochralea nigricornis 26: 144 Oedecerus rufofuscus 26: 146 Orthoxia 24: 260 Orthoxia boisduvallii 24: 260 Pachytoma 24: 261 [homonyn, replaced by Eupachy- toma Laboissiére 1940] Pacyhtoma flava 24: 262 Pachytoma westermanni 24: 261 Pyesia 24: 260 Rhombopala 26: 143 Rhombopala duodecimpunctata 26: 143 Rhombopala pectoralis 26: 144 Rupilla 31: 260 Rupilla ruficollis 31: 260 Rupilla viridiaenea 31: 260 Schematiza aequinoctialis 25: 263 Schematiza Schematiza Schematiza Schematiza Schematiza Schematiza Schematiza Schematiza Schematiza Schematiza Schematiza Schematiza Sphenoraia Sphenoraia Sphenoraia amplicornis 25: 261 annulicornis 25: 267 antennalis 25: 267 apicalis 25: 268 emarginata 25: 269 hispiformis 25: 269 lineaticollis 25: 266 nigricollis 25: 266 praeusta 25: 265 scutellaris 25: 268 venusta 25: 263 vicina 25: 262 24: 262 flavicollis 24: 262 nigripennis 24: 262 VOLUME 104, NUMBER 2 Megalopodinae Agathomerus cyaneus 34: 84 Agathomerus egregius 33: 45 [Nomen Nudum] Agathomerus incomparabilis 34: 81 Agathomerus nigricollis 34: 82 Agathomerus notaticollis 34: 82 Agathomerus rubrinotatus 34: 83 Agathomerus viduus 34: 84 Mastostethus bizonatus 34: 77 Mastostethus duplocinctus 34: 71 Mastostethus frontalinotatus 34: 76 Mastostethus notaticollis 34: 75 Mastostethus robustus 34: 74 Mastostethus sexnotatus 34: 80 Mastostethus stramineus 34: 73 Megalopus flavofusciatus 34: 87 Megalopus melipona 34: 84 Poecilomorpha mutillaria 34: 87 Megascelinae Megascelis ambigua 34: 15 Megascelis brunnipennis 34: 18 Megascelis crucifera 34: 13 Megascelis dilecta 34: 8 Megascelis dorsalis 34: 12 Megascelis dryas 34: 17 Megascelis exclamationis 34: 14 Megascelis frontalis 34: 17 Megascelis grayii 34: 7 Megascelis humeronotata 34: 19 Megascelis larvata 34: 14 Megascelis postica 34: 5 [homonym, replaced by par- allela Harold 1870] Megascelis purpureipennis 34: 2 Megascelis purpureotincta 34: 13 Megascelis rufotestacca 34: 4 Megascelis sacerdotalis 34: 3 Megascelis semipurpurea 34: 16 Megascelis titan 34: 18 Megascelis viridipallens 34: 4 Megascelis viridisimplex 34: 8 Sagrinae Mecynodera balyi 31: 248 Zeugophorinae Pedrillia murrayii 34: 87 Dytiscidae Agabus bakewelli 28: 19 Agabus latissimus 28: 18 Agabus reticulosus 28: 19 Agabus tasmaniae 28: 18 Batrachomatus wingii 28: 15 Celina australis 28: 14 Colymbetes adumbratus 28: 17 Colymbetes lanceolatus 28: 16 Colymbetes pulcher 28: 16 477 Copelatus acuductus 28: 21 Copelatus australiae 28: 20 Copelatus simplex 28: 21 Copelatus victoriae 28: 22 Hydaticus adamsii 23: 211 Hydaticus aruspex 23: 212 Hydaticus bakewellii 23: 210 Hydaticus bowringii 23: 214 Hydaticus fulvonotatus 23: 221 Hydaticus histrio 23: 221 Hydaticus matruelis 23: 216 Hydaticus nigromarmoratus 23: 220 Hydaticus nigrovittatus 23: 222 Hydaticus paganus 23: 215 Hydaticus parallelus 23: 219 Hydaticus ussherii 23: 214 Hydaticus verecundus 23: 213 Hydroporus aberrans 19: 426 Hydroporus adumbratus 17: 183 Hydroporus aequinoctialis 17: 178 Hydroporus amabilis 21: 420 Hydroporus andalusiae 22: 469 Hydroporus apicatus 17: 184 Hydroporus badius 19: 424 Hydroporus bakewellii 21: 413 Hydroporus bistrigatus 21: 419 Hydroporus blakeii 21: 411 Hydroporus bryanstonii 17: 175 Hydroporus carbonarius 19: 423 Hydroporus celatus 22: 473 Hydroporus charlottii 17: 182 Hydroporus compactus 21: 421 Hydroporus decemsignatus 17: 176 Hydroporus derelictus 22: 471 Hydroporus emilianus 17: 183 Hydroporus fryii 17: 181 Hydroporus gardnerii 21: 408 Hydroporus gemellus 21: 421 Hydroporus gilbertii 21: 414 Hydroporus gravidus 21: 413 Hydroporus hamatus 21: 407 Hydroporus hansardii 21: 417 Hydroporus howittii 21: 406 Hydroporus infacetus 17: 180 Hydroporus infaustus 17: 179 Hydroporus insculptilis 21: 411 Hydroporus interrogationis 21: 408 Hydroporus kingii 17: 178 Hydroporus leconteti 17: 175 Hydroporus magensis 17: 182 Hydroporus malaccae 19: 425 Hydroporus meadfootii 21: 419 Hydroporus multimaculatus 21: 417 Hydroporus nigroadumbratus 21: 410 Hydroporus orientalis 19: 427 Hydroporus penicillatus 21: 415 Hydroporus portmanni 17: 174 Hydroporus pudicus 19: 426 A78 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Hydroporus roffii 17: 176 Hydroporus rubescens 19: 426 Hydroporus shuckardii 21: 420 Hydroporus sinuatocollis 21: 418 Hydroporus tenellus 19: 427 Hydroporus thoreti 21: 409 Hydroporus tinctus 18: 326 Hydroporus undecimmaculatus 21: 412 Hydroporus wardii 17: 177 Hydroporus wollastonii 21: 416 Hyphidrus australis 21: 405 Hyphidrus bisulcatus 19: 422 Hyphidrus blanchardii 21: 404 Hyphidrus caledoniae 21: 406 Hyphidrus eximius 19: 421 Hyphidrus humeralis 21: 403 Hyphidrus johnsonii 21: 405 Hyphidrus nigronotatus 19: 421 Hyphidrus orientalis 19: 419 Hyphidrus pulchellus 19: 420 Hyphidrus rufoniger 19: 423 Hyphidrus rufus 19: 423 Gyrinidae Enhydrus assimilis 30: 217 Enhydrus latior 30: 217 Enhydrus howittii 30: 215 Enhydrus rivularis 30: 218 Haliplidae Haliplus australis 21: 400 Haliplus eximius 19: 418 Haliplus fuscatus 21: 401 Haliplus gibbus 21: 402 Haliplus pulchellus 19: 418 Haliplus simplex 19: 419 Haliplus testudo 21: 400 Haliplus variabilis 19: 417 Hygrobiidae Pelobius australasiae 21: 402 Pelobius niger 21: 403 Scirtidae Ora 32: 385 Ora chevrolatii 32: 385 Ora grayii 32: 385 ACKNOWLEDGMENTS I thank my wife, S. L. Staines for edi- torial assistance. Funding for this research was entirely personal. LITERATURE CITED Aslam, N. A. 1968. Nomenclatorial notes on Chryso- meloidea (Coleoptera). Journal of Natural History 2: 127-129. Blackwelder, R. E. 1946. Checklist of the coleopterous insects of Mexico, Central America, the West In- dies, and South America, Part 4. Bulletin of the United States National Museum 185: 551-763. Clavareau, H. 1914. Coleopterorum Catalogus, Chry- somelidae: 11. Eumolpinae. pars 59: 1—215. Csiki, E. 1939. Jn Heikertinger, KF and E. Csiki, Co- leopterorum Catalogus, Chrysomelidae, Haltici- nae I. pars 166: 1-336. Gressitt, J. L. and S. Kimoto. 1963. The Chrysomeli- dae (Coleopt.) of China and Korea, part 2. Pacific Insects Monograph 1B: 301—1,026. von Harold, E. 1870. Geainderte Namen. Coleopterol- ogische Hefte 6: 119. 1875. Neue Coprophagen. Deutsche Ento- mologische Zeitschrift 19: 209-217. Heikertinger, E and E. Csiki. 1939. Coleopterorum Ca- talogus, Chrysomelidae, Halticinae I. pars 166: 1— 336. . 1940. Coleopterorum Catalogus, Chrysome- lidae, Halticinae I. pars 169: 337-635. Heinze, E. 1927. Beitrag zur Kenntnis Criocerinen (Col. Chrysomelid.). Entomologische Mitteilun- gen 16: 138-142. Jolivet, P. 1971. Coleopterorum Catalogus, Supple- menta. Chrysomelidae: Donaciinae. pars 51(2): 1-71. Laboissiére, V. 1940. Galerucinae (Coleoptera Phyto- phaga), Fam. Chrysomelidae. Exploration du Pare National de l’Albert Mission G. FE Witte (1933-— 1935). Fascicule 31: 1—93. Monroés, E 1958. Coleopterorum Catalogus, Supple- menta. Chrysomelidae: Sagrinae. pars 51(1): 1I— 19. . 1959. Los generos de Chrysomelidae (Cole- optera). Opera Lilloana 3: 1—337. Seeno, T. N. and J. A. Wilcox. 1982. Leaf beetle gen- era (Coleoptera Chrysomelidae). Entomography 1: 1-221. Weise, J. 1913. Synonymische Mitteilungen. Wiener Entomologische Zeitung 32: 218-220. . 1916. Coleopterorum Catalogus. Chrysome- lidae: 12. Chrysomelinae. pars 68: 1—255. Wilcox, J. A. 1971. Coleopterorum Catalogus, Supple- menta. Chrysomelidae: Galerucinae, Oidini, Gal- erucini, Metacyclini, Sermylini. pars 78(1): 1— 220. . 1973. Coleopterorum Catalogus, Supplemen- ta. Chrysomelidae: Galerucinae, Luperini: Luper- ina. pars 78(3): 433-664. Zimmermann, A. 1920. Coleopterorum Catalogus. Dy- tiscidae, Haliplidae, Hygrobiidae, Amphizoidae. pars 71: 1-326. PROC. ENTOMOL. SOC. WASH. 104(2), 2002, pp. 479-484 THE STEM-BORING SAWFLY GENUS CEPHUS LATREILLE (HYMENOPTERA: CEPHIDAE) IN JAPAN DAVID R. SMITH AND AKIHIKO SHINOHARA (DRS) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, % National Museum of Natural History, Smithsonian Insti- tution, Washington, DC 20560-0168, U.S.A. (e-mail: dsmith@sel.barc.usda.gov); (AS) Department of Zoology, National Science Museum (Natural History), 3-23-1 Hyakunin- cho, Shinjuku-ku, Tokyo 169-0073, Japan (e-mail: shinohar@kahaku.go.jp) Abstract.—Two species of Cephus occur in Japan, C. hyalinatus Konow from Hokkaido and C. brachycercus Thomson, newly recorded from Japan, from Honshu. Clarification of past records of Cephus in Japan are presented. A key, descriptions, and illustrations of the abdomen are given for both species. Key Words: Cephus is primarily a Eurasian genus of 20 to 25 species, all of which are associated with grasses. Gussakovski (1935) keyed and discussed 23 Palearctic species, Benson (1946) estimated 25 world species, and Mu- che (1981) keyed and discussed 31 world species. Two species occur in North Amer- ica, but one, Cephus pygmaeus (L.) is an introduction, and the other, Cephus cinctus Norton, is still open to question as to whether it is native or an early introduction from eastern Asia. Larvae bore and feed in the stems of grasses and can be especially destructive in cultivated grain crops such as wheat. Takeuchi (1938) recorded four species of Cephus from northeastern Asia, but two of them, C. nigripennis Takeuchi and C. sach- alinensis Takeuchi, are now placed in Cal- ameuta Konow (Benson 1946). Of the other two, one, Cephus hyalinatus Konow, is known from eastern Asia including Japan (Hokkaido), and the other, C. camtschat- calis Enslin, from Kamtchatka and Sakha- lin. Takeuchi (1938) corrected his previous record of Cephus camtschatcalis Enslin stem-borers, Cephidae, Cephus, Japan from Hokkaido (Takeuchi 1936) to Cephus hyalinatus. Takeuchi later recorded C. cam- tschatcalis from Kunashiri Island, but this is corrected to C. hyalinatus in this paper. Togashi (1997, 1998) gave collection re- cords of Cephus hyalinatus from Akan, Hokkaido, and Niigata Prefecture, Honshu, respectively, but the latter was corrected to Stenocephus oncogaster Shinohara by Shi- nohara (1999). Specimens of this genus are not com- monly collected in Japan, and only a few from Hokkaido and Honshu are represented in collections. Based on about 35 specimens available, we record two species from Ja- pan, Cephus hyalinatus from Hokkaido and Cephus brachycercus Thomson from Hon- shu. The former is widespread in eastern Asia, and the latter is widespread in Eurasia and is reported here for the first time from Japan. Based on the specimens available, there are rather constant color differences between those collected in Honshu and those from Hokkaido. Thus, at present we prefer to treat them as separate species, and we attempt to clarify past records from Ja- pan. 0 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Cephus belongs to the tribe Cephini, all of which are associated with grasses. It is separated from the other two genera of the tribe, Calameuta and Trachelus, by the fol- lowing: Distance between antennae sube- qual to distance between antenna and ten- torium; female sheath in dorsal view not broadened toward apex; male eighth ster- nite without concavity, with only a patch and apical fringe of modified flattened se- tae. In Calameuta, the distance between the antennae is much shorter than the distance between the antenna and tentorium (as 1.0: 1.5 to 2.0), and in Trachelus the female sheath is broadened at its apex and the male eighth sternite has a deep concavity with modified setae. KEY TO JAPANESE SPECIES OF CEPHUS Ihelemialets eter eis Pete a ieee 2 Male Sea et. Aono RS Sie O oe SS aaa 3 2. Abdomen (Fig. 1) with lateral yellow spots on segments 3—7, appearing almost as a lateral yellow longitudinal stripe, yellow transverse bands on posterior margins of abdominal seg- ments 4 and 6, apex of 9th tergite broadly yel- low: wings commonly moderately infuscated ata LOR ae ees frat EG on hyalinatus Konow — Abdomen (Fig. 2) with lateral yellow spots on segments 3 or 4—7, spots small and separated, not appearing as a lateral longitudinal stripe, without yellow transverse bands, though 6th segment sometimes with narrow incomplete band on posterior margin, 9th tergite narrowly yellow at apex; wings hyaline ......... brachycercus Thomson 3. Abdomen (Fig. 3) with transverse yellow bands on posterior margins of segments 3, 4, 6, and 7, narrow or incomplete on 3 and 7, segments 3 or 4—7 with lateral yellow spots, sometimes minute spots on 2 and 8; wings commonly moderately infuscated: venter of cervical sclerite usually with yellow spot; me- sepisternum with yellow spot on extreme upper corner and yellow spot usually on lower ante- rior area hyalinatus Konow — Abdomen (Fig. 4) with transverse yellow bands on posterior margins of segments 4 and 6, lateral yellow spots on segments 3 or 4—7; wings hyaline; cervical sclerites and mesepis- ternum usually black except sometimes inner ventral surface of cervical sclerite with yellow spot and possibly a small yellow spot on dorsal COLNEH OF MESEPIStTe nnn eee Cephus hyalinatus Konow (Pigs 15-3) Cephus hyalinatus Konow 1897: 173.— Takeuchi 1938: (Hokkaido).—Togashi 1997: (Hokkaido). Cephus camtschatcalis: Takeuchi 1936: 59.—Takeuchi 1955: 9 (misidentifica- tions). This species was described from ‘‘Sibiria or. (Irkutsk).”” It has been recorded from various localities in eastern Russia (Gussa- kovskij 1935, Verzhutski1 1966), Kunashiri Island (Takeuchi 1955 [as C. camtschatcal- is|), and Hokkaido, Japan (Takeuchi 1936 [as C. camtschatcalis|), Takeuchi 1938). Female.—Length, 9.0—11.0 mm. Anten- na and head black; mandible yellow with apex reddish brown; maxillary palpal seg- ments 3—6 yellow. Thorax black, some- times extreme upper corner of mesepister- num with small yellow spot. Fore- and mid- legs black with tibiae and tarsi yellow. Hind- leg black with tibia yellow with about apical fourth black. Abdomen black with segments 3—7 yellow laterally, yellow ex- tending width of segments, thus appearing as a lateral longitudinal stripe; transverse yellow bands on posterior margins of seg- ments 4 and 6; apex of 9th tergite broadly yellow. Oblong plate (valvifer 2) yellow ventrally. Wings moderately, uniformly in- fuscated; costa and anterior margin of stig- ma yellowish, rest of stigma and veins black. Antenna with 18 or 19 segments; preapical segments slightly longer than broad. Hindtibia with 2 preapical spines. Tarsal claw with a minute inner tooth, much shorter than outer tooth. Male.—Length, 8.0—9.5 mm. Outer sur- face of antenna from segment 5 to apex yel- lowish. Head black, usually with yellow spot on lower inner orbit near malar area and spot on supraclypeal area bordering clypeus; these spots sometimes small or ab- sent. Thorax black with yellow spot on ex- treme upper corner of mesepisternum; per- apterum yellow; yellow spot on lower an- terior mesepisternum, and yellow spot on VOLUME 104, NUMBER 2 481 Figs. 1-4. Abdomen, lateral view. 1, Cephus hyalinatus, female. 2, C. brachycercus, female. 3, C. hyalinatus, male. 4, C. brachycercus, male. venter of cervical sclerite. Fore- and mid- legs with coxae yellow anteriorly, black posteriorly; trochanters black; outside mar- gin of femora black, inside margin of fem- ora and tibiae and tarsi yellow; hindleg sim- ilar but black mainly at base of coxa with apical half or more yellow. Abdomen black with lateral yellow spots on segments 3 or 4—7, sometimes minute spots on 2 and 8, spots separated; yellow transverse bands on posterior margins of segments 3, 4, 6, and 7, usually narrower on 3 and sometimes in- complete on 3 and 7. Apex of hypandrium and harpes of male genitalia yellow. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Japan records.—Hokkaido: Wakkanai, 7-VII-1977, col. Ko Baba (279. 3.6); same except 5-VII-1977 (1 ¢); Rishiri Is., 2-VII- 1968, T. Naito leg. (2 2, 1 3); Kawayu, 25-VI-1969, T. Naito (1 3); Teshio, 2-VII- 1930, Takeuchi (identified as C. hyalinatus by Takeuchi) (1 ¢). Other specimens examined.—Russia, Sa- khalin: Konuma, 7-VII-1930 (identified as C. hyalinatus by Takeuchi) (2 2); Karafuto (= Sakhalin), 1922, coll. M: Yano (““753”’ K. Sato Collection 1975) (2 2); Sakhalin, Yuzhno-Sakhalinst City, meadows near for- est stream, 15-VI-1971, V. Ermolenko (3 3); Sakhalin, Shenbunino Vil., meadow along Japanese Sea, 19-VI-1971, V. Ermo- lenko (1 2, 1 ¢). Russia, Primorski Kray, Primorskoe Vil., Reserve Kedzovaya “Padj”*, 11=VI-1966; Gs Anutriev,, Cy )): Kuriles, Kunashiri, Yambetsu, 22—25-VII- 11935" Uchida (l 2.5’ ¢); Etorofu; Porosu 14—15-VII-1936 (2 6), Seseki 16—18-VII- 1936, Y. Sugihara (1 2) (identified as C. camtschatcalis by Takeuchi). Kunishir Is- land, Alekhino Vil., wet meadow along for- est stream, 27-VI-1971, V. Ermolenko (2 S). Shikotan Island, Krabozavodsk Vil., meadows along Pacific Ocean, on flower of Ranunculus sp., 6-VII-1071, V. Ermolenko (7 2,4 2). Shikotan Island, Malo-Kuril’sk Vil., meadows along Pacific Ocean, 9-VII- 1971, V. Ermolenko (1 ¢). Remarks.—Ivie and Zinovjev (1996) synonymized C. hyalinatus under Cephus cinctus Norton 1872 described from North America; thus, concluding that the same species occurs in Asia and North America and that the North American populations may have been the result of an early intro- duction from Asia. This synonymy is still open to question, and we prefer to call the Japanese specimens C. hyalinatus pending additional proof for the synonymy. This species was recorded from Hokkai- do by Takeuchi (1936, 1938), the latter ref- erence correcting his 1936 identification of C. camtschatcalis to C. hyalinatus. We have seen the specimen on which Takeuchi based this record and confirm his identification as C. hyalinatus. Takeuchi (1955) recorded C. camtschatcalis from Kunishiri Island. We have examined those specimens, deposited in Hokkaido University, Sapporo, and con- firm that they are actually C. hyalinatus. Togashi’s (1998) record of C. hyalinatus from Honshu was erroneous and is actually Stenocephus oncogaster (Shinohara 1999). Cephus brachycercus Thomson (Figs. 2, 4) Cephus brachycercus Thomson 1871: 322. This species was described from Europe, but it occurs across the Palearctic (Gussa- kovskij 1935, Verzhutskii 1966) and is here newly recorded from Honshu, Japan. Female.—Length, 6.5—8.3 mm. Antenna and head black; mandible yellow with apex reddish brown. Thorax black. Fore- and midlegs black with tibiae and tarsi yellow; hindleg black with tibiae yellow except for black on about apical fourth. Abdomen black with small lateral spots on segments 3 or 4—7, spots small and separated; some- times a narrow, incomplete yellow trans- verse band on posterior margin of segment 6; apex of 9th tergite narrowly yellow. Ob- long plate (valvifer 2) narrowly yellow on margin. Wings hyaline; costa and anterior margin of stigma yellowish, rest of stigma and veins black. Antenna with 18 to 22 seg- ments; preapical segments about as broad as long. Hindtibia with 2 preapical spines. Tarsal claw with small inner tooth, much smaller than outer tooth. Male.—Length, 6.0—8.0 mm. Head and thorax black; inner ventral surface of cer- vical sclerite sometimes with yellow spot, and sometimes small yellow spot on dorsal corner of mesepisternum; perapteron yel- low. Fore- and midlegs with coxae yellow anteriorly, black posteriorly; trochanters yellow anteriorly; outside margin of femora black, inside margin of femora and tibiae and tarsi yellow; hindleg similar but black mainly at base of coxa with apical half or more yellow. Abdomen with lateral yellow spots on segments 3 or 4 to 7, sometimes VOLUME 104, NUMBER 2 small on 3; transverse yellow bands on pos- terior margins of segments 4 and 6. Japan records.—Honshu: Mt. Hakkoda, Aomori Pref., 20-VH-1965, T. Naito leg. (2 6); Sukayu, Aomori Pref., 20-VII-1965, col. T. Naito (2 2,5 6); Mt. Hakkoda (To- wadako), Aomori Pref., 7-VIII-1988, M. Yamada (1 6); Mt. Iwaki, Aomori Pref., 7-VII-1995, M. Yamada (1 2); Sasayama, Hyogo Pref., 27-V-1954, S. Momoi (1 @, 3 3). Remarks.—Cephus brachycercus occurs in Europe and has been recorded from Si- beria, at least to the Irkutsk Region (Gus- sakovskij 1935, Verzhutskii 1966, Zhelo- chovtsev and Zinovjev 1996). We have not seen records from farther east, but it could occur in more eastern regions and into Ja- pan, or it could be an introduction into Ja- pan. The specimens agree with C. brachy- cercus specimens from Europe and key to C. brachycercus in existing keys (e.g., Gus- sakovskij 1935). It is not a species treated by Takeuchi (1938). Many species of Cephus appear to be based on color differences, but coloration varies, and it is difficult to evaluate the de- scribed species. The Japanese specimens are close to C. camtschatcalis, but C. cam- tschatcalis lacks yellow on the mandibles and the hindleg is black with only the base of the tibia and tarsus dark brown (Enslin 1926). Also, Zhelochovtsev and Zinovjev (1996) treated C. camtschatcalis as a ques- tionable synonym of C. hyalinatus, and these Japanese specimens are distinct from C. hyalinatus. Benson (1951) mentioned that Cephus nigrinus Thomson, known from Europe to Siberia, is close to C. brachycercus. Cephus nigrinus usually has the abdomen entirely black, but Benson noted that the abdomen sometimes may have small yellow spots. He separated C. brachycercus by its slightly broader flagellum, with the preapical seg- ments broader than long, its more infuscat- ed wings, and by the sawsheath (valvula 3) not being set in a direct line with the oblong plate (valvula 2). In the Japanese speci- 483 mens, the preapical antennal segments are about as broad as long, and the sawsheath is in a direct line with the oblong plate. Because of the favorable similarity of the Japanese specimens with C. brachycercus, we believe it is best to refer these speci- mens to that species. Since we are not sure of the significance of the characters used by Benson, and since the entire genus needs revisionary study, we prefer not to describe a separate species at least until taxonomic problems within the genus are resolved. ACKNOWLEDGMENTS We thank H. Hara, Hokkaido Forestry Research Institute, Bibai; M. Ohara, Hok- kaido University, Sapporo; T. Hirowatari, University of Osaka Prefecture, Sakai; T. Naito, Kobe University, Kobe; and M. Ya- mada, Hirosaki, for the loan or gift of ma- terial. LITERATURE CITED Benson, R. B. 1946. Classification of the Cephidae (Hymenoptera Symphyta). Transactions of the Royal Entomological Society of London 96: 89— 108. . 1951. Hymenoptera 2, Symphyta, section (a). In Royal Entomological Society of London, Handbooks for the Identification of British In- sects, Vol. VI, Part 2(a), 49 pp. Enslin, E. 1926. Die Tenthrediniden (Hymenoptera) der Kamtschatka-Expedition 1908-1909. Annu- aire du Musée Zoologique de 1’ Académie des Sci- ences de l’URSS 27: 363-381. Gussakovskij, V. V. 1935. Faune de 1’URSS, Insectes Hymeénopteres, T. I, vol. 1, Chalastogastra (P. 1). Institut Zoologique de 1] Académie des Sciences de 'URSS, Nouvelle Série no. 1, Edition de Académie des Sciences de lURSS, Moscou, Leningrad, 452 pp. Ivie, M. A. and A. G. Zinovjev. 1996. Discovery of the wheat stem sawfly (Cephus cinctus Norton) (Hymenoptera: Cephidae) in Asia, with the pro- posal of a new synonymy. The Canadian Ento- mologist 128: 347-348. Konow, FE 1897. Neue palaearctische Tenthrediniden. Wiener Entomologische Zeitung 16: 173-187. Muche, H. 1981. Die Cephidae der Erde (Hym., Ce- phidae). Deutsche Entomologische Zeitschrift, N.E 28: 239-295. Norton, E. 1872. Notes on North American Tenthre- dinidae, with descriptions of new species. Trans- 484 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON actions of the American Entomological Society 4: 77-86. Shinohara, A. 1999. A study on stem boring sawflies (Hymenoptera, Cephidae) of the tribe Hartigiini from Japan and her adjacent regions. Japanese Journal of Systematic Entomology 5: 61—77. Takeuchi, T. 1936. Tenthredinoidea of Saghalien (Hy- menoptera). Tenthredo 1: 53-108. 1938. A systematic study on the suborder Symphyta (Hymenoptera) of the Japanese Empire (1). Tenthredo 2: 173-229. . 1955. Sawflies of the Kurile Islands (1). In- secta Matsumurana 19: 9-22. _ Thomson, C. G. 1871. Hymenoptera Scandinaviae. Tom. 1. (Tenthredo et Sirex Lin.). Lundae, 342 pp. Togashi, I. 1997. Symphyta (Hymenoptera) collected by Dr. Y. Nishijima in Hokkaido, Japan. Bulletin of the Biogeographical Society of Japan 52(1): 1-6. 1998. Sawfly from Niigata Prefecture (3). Sawfly fauna [of] Hokuetsu District. Transactions of the Essa Entomological Society, Niigata (77): 6—10. (In Japanese.) Verzhutskii, B. N. 1966. Sawflies of Baikal Region. Izdatel’stvo ““Nauka’”’, Moskva. 162 pp. Zhelochovtsev, A. N. and A. G. Zinovjev. 1996. A list of the sawflies and horntails (Hymenoptera, Sym- phyta) of the fauna of Russia and adjacent terri- tories. 2. Entomologicheskoe Obozrenie 75: 357— By/9} PROC. ENTOMOL. SOC. WASH. 104(2), 2002, pp. 485-504 REVIEW OF THE NORTH AMERICAN GRAY PYLA GROTE (LEPIDOPTERA: PYRALIDAE: PHYCITINAE) WITH DESCRIPTION OF A NEW SPECIES FROM WESTERN UNITED STATES JOHN H. WILTERDING III AND GEORGE J. BALOGH (JHW) Department of Entomology, A. J. Cook Arthropod Research Collection, Mich- igan State University, East Lansing, MI 48824-1115, U.S.A.; Olivet College, Olivet, MI 49076, U.S.A. (e-mail: wilterdi@pilot.msu.edu); (GJB) 6275 Liteolier, Portage, MI 49024, U.S.A: Abstract.—Eleven North American species of the “‘gray”’ section of the genus Pyla Grote are diagnosed by keys based on male and female genitalia. Pyla westerlandi Wil- terding and Balogh, new species, is described from the Lake Tahoe region of California and central Oregon. The female of Pyla criddlella Dyar, previously known from only the male holotype, is described and Phycitia bilineata Curtis is placed as a new synonym of Pyla fusca (Haworth). The distribution, habitat, and flight period information for the gray Pyla species are summarized. Key Words: life history, habitat, alvar, Aweme Heinrich (1956) grouped nine species (four described as new) of Pyla Grote based on the color of the forewings in what he termed the “gray” section of the genus. Shortly thereafter, McDunnough (1958) de- scribed Pyla gaspeensis from the Gaspé re- gion of Québec. Balogh and Wilterding (1998) described Pyla arenaeola from sandy habitats, particularly dunes, of central North America, and determined that the fe- male of that new species was previously misassociated as the female of Pyla aequti- voca Heinrich. In addition, P. gaspeensis was recognized as a synonym of P. aequi- voca. During the course of that study, Julian Donahue (LACM) brought to our attention another new gray Pyla species based on material collected by the late Nils Wester- land in the Lake Tahoe region of California and allowed us the opportunity to describe this distinctive moth. Examination of nearly 1,000 specimens disjunction, genitalia, Criddle, Westerland, of North American gray Pyla affords us the opportunity to review the group, add con- siderable new distributional and habitat in- formation, and refine the identification of the species. The female of Pyla criddlella Dyar (1907), known to previous authors only from the male holotype is described here for the first time. Many of the gray Pyla species are similar in wing color and maculation and cannot be reliably deter- mined without examination of the genitalia. The eleven North American gray Pyla are readily recognized based on distinctive gen- ital features found in both sexes as illus- trated in Heinrich (1956), Balogh and Wil- terding (1998), and this paper (Figs. 1—6). All Pyla species, including both the ‘“‘oray’’? and ‘“‘bronzy brown’’ species groups of Heinrich (1956), and with the ex- ception of the Holarctic gray species Pyla fusca Haworth are limited to North Amer- ica. The bronzy brown Pyla species are re- 486 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON stricted to montane and alpine habitats of western North America. In contrast, many gray species have western (typically north of a line from central California to Colo- rado) and/or transcontinental distributions in the northern states and boreal Canada (Figs. 8-10; see also P. aequivoca, Balogh and Wilterding 1998). Life history information for most Pyla species is lacking; only the life histories of the holarctic P. fusca and the dune species, P. arenaeola, are known (Balogh and Wil- terding 1998). In common with many phy- citine larval strategies, the widespread P. fusca webs together the shoots and leaves of its hosts, primarily Ericaceae. The larvae of P. arenaeola, a species much more re- stricted in distribution occurring largely on sand dunes of the Great Lakes (Balogh and Wilterding 1998), constructs silk-lined sand tubes and feeds concealed beneath its host, bearberry, Arctostaphylos uva-ursi (L.) Sprengel (Ericaceae). If the life history strategy of P. arenaeola is representative for other Pyla species, larval dependence on particular substrates for shelter construc- tion, specific ericaceous hosts, or both, might account for restricted and disjunct distribution patterns. For example, Pyla in- sinuatrix Heinrich, a species associated with fens, alvar, and coastal bogs in Nova Scotia, is distributed in a narrow band from the Atlantic coast to western Alberta (Fig. 8). Pyla impostor Heinrich, known from many localities in the west, has an appar- ently disjunct population in eastern Canada (Fig. 9). The new species described in this paper is primarily known from one locality on the northwest side of Lake Tahoe, Cal- ifornia, and a single specimen from Crook County in central Oregon. Specimens examined originated from the following institutions and private collec- tions: AMNH American Museum of Natural History, New York, NY (EF H. Rindge). BGS Brian G. Scholtens, Charleston, SC: CNEe Canadian National Collection, Ottawa, Ontario, Canada (J. E Landry, B. Landry). CUI Cornell University Collection, Ithaca, NY (James Leibherr). EGV E. G. Voss, Mackinaw City, MI. GJB George J. Balogh, Portage, MI. JAB James A. Bess, Wanatah, IN. JHW John Wilterding, East Lansing, MI. JKM John K. Morton, Waterloo, On- tario, Canada. KS Ken Stead, Brantford, Ontario, Canada. LACM Los Angeles County Museum, Los Angeles, CA (J. P. Donahue). LAF Leslie A. Ferge, Middleton, WI. MCZ Museum of Comparative Zoolo- gy, Harvard University, Cam- bridge, MA (P.D. Perkins). MSU Michigan State University, East Lansing, MI (FE W. Stehr). NMNH_ National Museum of Natural His- tory, Smithsonian Institution, Washington DC (M. A. Solis). UCB Essig Museum, University of California, Berkeley, CA (J. A. Powell). UM University of Minnesota, St. Paul, MN (P. J. Clausen). UMBS _ University of Michigan Biologi- cal Station, Pellston, MI (B. G. Scholtens). UMMZ_ University of Michigan Museum of Zoology, Ann Arbor, MI (M. EF O’Brien). UZMH Zoologiska Muset, Universitets Helsinki, Finland (K. Mikkola). KEYS TO THE GRAY PYLA Males In the following key the term clasper (sensu Heinrich) refers to a process from the mesial aspect of the valva (including valva base) and should be distinguished from the costal process, a projection from VOLUME 104, NUMBER 2 the sclerotized costal edge of the valva. All Pyla species included in this key have the aedeagus divided distally into two variably sclerotized lateral elements, with one or both divisions developed posteriorly into sclerotized spines or processes. In some species one division is reduced, sometimes greatly so as in Pyla fusca where the right division is straight, very thin, and normally appressed against the much larger left di- vision and thus inapparent in many prepa- rations. i) . Aedeagus with one or both lateral divisions terminating in a single, outcurved, pointed process, or the divisions symmetric and each with a single, subapical, laterally directed SINC Bucs s cen kee ees At, Soe ys Pee, Vas 2, Aedeagus with each lateral division terminat- ing as a nearly straight, or slightly incurved, blunt or pointed process, or one or both di- visions with multiple distal spines (these spines may be laterally outcurved) . Aedeagus asymmetric, only one division ter- minating as an outcurved pointed process .. 3 Aedeagus symmetric, each division terminat- ing as an outcurved pointed process or each division with a single laterally directed sub- apical spine . Aedeagus slender, the left division outcurved and pointed, the right division very small and inapparent; vesica with a fine hairlike cornu- tus; valva with clasper at about % length, clasper knoblike and tipped with spines (emnich M956 is-9369)) 5 5-5: fusca (Haworth) Aedeagus wider and both divisions apparent, the right division outcurved to a sharp point, the left division shorter and straight; vesica without cornutus; valva with clasper near base, clasper toothlike or triangular (Heinrich 1956, fig. 368) criddlella Dyar . Each aedeagus division terminating distally as aAMZOULCURVeEd pointed }processy.. 4-56). 4: 5S) Each aedeagus division with a single subapi- calmlateralllysdinectedispine yess ane 6 . Valva with costal process at %4 length, and clasper beyond middle of valva; juxta without spine clusters (Balogh and Wilterding 1998, figs. 1-5) ... arenaeola Balogh and Wilterding Valva without costal process and clasper near base of valva; juxta with comb-like clusters of spines (Heinrich 1956, fig. 370) hypochalciella (Ragonot) . Tegumen with two pairs of projecting spined lobes from ventrolateral margin; valva with clasper at % valva length, clasper semicircular i) 487 and much shorter than valva width (Heinrich 1956, fig. 367) aenigmatica Heinrich Tegumen without spined lobes; valva with clasper near base, clasper a thick, pointed curved process and longer than valva width (Figs. 1-3) westerlandi Wilterding and Balogh, n. sp. . Aedeagus with multiple distinct distal spines on one or both divisions Aedeagus with each division terminating in a blunt or pointed process and at most a single subapical spine (P. impostor may have a few widely scattered spinules along length of dis- tally pointed processes) . Aedeagus divisions with one or more stout posterolaterally directed spines, more numer- ous on left division; juxta with paired, long, slender and pointed posterior processes; valva with clasper short and tooth-like or triangular (Heinrich 1956, fig. 366; Balogh and Wilter- ding 1998, figs. 6-10) .... aequivoca Heinrich Aedeagus divisions with multiple small apical spines; juxta without pointed elongate pro- cesses; valva with clasper an elongate spine (Heinrich 1956, fig. 371) .... hanhamella Dyar . Aedeagus with right division longer than left, the left with a subapical spine, each division with a helix-like twist at middle third (Hein- rich 1956, figs. 362, 363) .. . fasciolalis (Hulst) Aedeagus divisions nearly equal in length and without a helix-like twist .......-....-: 10 . Aedeagus divisions pointed distally; valva with clasper a long, thin, pointed process near base of valva, and valva without costal pro- cesses (Heinrich 1956, fig. 364) impostor Heinrich Aedeagus divisions blunt distally; valva with clasper a short slightly irregular ridge at % valva length as well as a thick and heavily spined costal process at base (Heinrich 1956, fig. 365) insinuatrix Heinrich Females . Ostium bursae with dense ventrolateral scler- otization, sclerotized projections, or scobinate lobeston plates: tas. cho tee cue ee aa 2D Ostium bursae membranous, or only lightly sclerotized with fine granulations or folds .. 9 . Corpus bursae with extensive convolute scler- otized folds on posterior half (Heinrich 1956, fig. 850) Corpus bursae membranous or with at most weakly developed small patches of scleroti- zation impostor Heinrich . Ductus bursae sclerotized posteriorly and with lateral margins subparallel, the scleroti- zation extending onto ostium bursae and var- 188 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON fear ater Sete 2 Figs. 1-3 uesiree'= ufeeee= 3 Pyla westerlandi, male genitalia. 1, Genital capsule, valves spread, aedeagus removed, ventral view. 2, Eighth sternite, ventral view, without scale tufts. 3, Aedeagus, ventral view. iably developed into a ventrolateral sclero- tized rim Ductus bursae without dense sclerotization, outer margins bulge laterally, or if lateral mar- gins subparallel, ductus bursae semimembran- ous; ostium bursae with paired ventrolateral sclerotized plates or scobinate lobes ...... Ostium bursae ventrolateral sclerotization a sinuate, liplike band; ductus seminalis from right-sided lobe of the corpus bursae (Hein- meh 9565 fit SOO) ieee eee hanhamella Dyar Ostium bursae otherwise modified; ductus seminalis from the dorsal aspect of the corpus bursae . Ostium bursae sclerotization paired convolute sclerotized lobes with deep central notch (Heinrich 1956, fig. 853). . . aenigmatica Heinrich Ostium bursae sclerotization funnel shaped VOLUME 104, NUMBER 2 10. and without deep central notch (Heinrich 1956, fig. 854) hypochalciella (Ragonot) . Ostium bursae with paired, pad-like, rounded, scobinate ventrolateral lobes; ductus bursae strongly dorsoventrally flattened, lateral mar- gins subparallel and distinctly narrower than the width of posterior aspect of corpus bursae (Heinrich 1956, fig. 856) . . Ostium bursae with paired, densely sclero- tized, ventrolateral convolute and/or scobinate plates; ductus bursae not stongly dorsoven- trally flattened, lateral margins bulging and subequal in width to posterior aspect of cor- pus bursae (when the bursae is well inflated) insinuatrix Heinrich . Ostium bursae with elongate, triangular, scler- otized, ventrolateral plates (Heinrich 1956, fig. 855 [as aequivoca]; Balogh and Wilter- ding 1998, figs. 11—13) arenaeola Balogh and Wilterding Ostium bursae with convolute, sclerotized, ventrolateral plates that are not distinctly tri- angular in shape . Ostium bursae paired sclerotized plates as long or longer than wide; ostium bursae a large rectangular chamber; ductus bursae membranous, with lateral walls bulging some- what asymmetrically to the left (Fig. 4) westerlandi Wilterding and Balogh, n. sp. Ostium bursae paired sclerotized plates short- er than wide; ostium bursae not rectangular in shape; ductus bursae partially sclerotized lat- erally with walls symmetrically bulging lat- erally (Balogh and Wilterding 1998, figs. 14— 16) aequivoca Heinrich . Ostium bursae membranous and weakly con- stricted at junction with ductus bursae; ductus and corpus bursae with extensively devel- oped, convolute, sclerotized bands that extend onto anterior half of corpus bursae (Heinrich IPED, ile 4B ara eo 5 a Ble ao ene fasciolalis (Hulst) Ostium bursae with fine granulations or par- tially sclerotized folds and obviously con- stricted at junction with ductus bursae; ductus and corpus bursae sclerotization smooth or convoluted but not extending onto anterior halizoisxcorpus) bursa eee ee a 10 Ductus and corpus bursae elongate with smooth elongate bands of sclerotization and lacking diverticula (other than the lobe at in- sertion of ductus seminalis) (Heinrich 1956, HU OAES SD) me. War) Saeco ee fusca (Haworth) Ductus and corpus bursae globular in shape; an oblique, convolute band of sclerotization near ductus-corpus bursae junction and mul- tiple adjacent diverticula (in addition to the lobe at insertion of ductus seminalis) (Figs. 5, (5) Aestest dete iat edie sient a criddlella Dyar 489 Pyla fasciolalis (Hulst) (Fig: 7; Heinrich 1956; figs. 362, 363, 849) Pinipestis fasciolalis Hulst 1886: 162. Ho- lotype 6, “Spencers Bridge’ (Rindge 1955) [= Spence’s Bridge, NW of Nicola (J. D. Lafontaine, personal communica- tion 1999)], British Columbia, Canada (AMNH). [In the original description the type locality is in error as Nevada, see Heinrich (1956) and Rindge (1955).] [Type examined. ] Pyla fasciolalis is known from scattered localities in western North America (Fig. 7), with most localities represented by sin- gle specimens. Recorded elevations in the northern portion of the distribution range from 2,100 feet (3 mi W Nelson, British Columbia) to 5,500 feet (Manning Park, Valley View, British Columbia) and the moth has been collected as high as 10,150 feet farther south (Crooked Creek Lab, Mono Co., California). In northeastern Utah (Sheep Creek and Highway 44) a small se- ries was collected in mixed sage-juniper and riparian habitat (GJB). Flight dates range from 7 June to 13 August. The life history is unknown. Heinrich (1956) noted variation in genital details perhaps worthy of additional study as additonal material becomes available. However, the twisted asymmetric lateral processes of the male aedeagus (Heinrich 1956, fig. 362a) and configuration of the female corpus bursae (Heinrich 1956, fig. 849) distinguish P. fasciolalis from all other Pyla species. Material examined: 13 36, 7 2 [AMNH, CNC, GJB, LACM, NMNH, UCB, UM]. Pyla impostor Heinrich (Fig. 9; Heinrich 1956, figs. 364, 850) Pyla impostor Heinrich 1956: 143. Holo- type 6d, Slate Peak, 6,500-—7,000 ft, Whatcom County, Washington, United States (NMNH). [Type examined. ] Pyla impostor is a widely distributed western species with most documented lo- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 490 Oe 8 ae ene ee a8 wert PRED eee re VOLUME 104, NUMBER 2 491 aco , © westerlandi * hypochalciella @ fasciolalis Fig. 7. Known distribution of Pyla westerlandi, P. hypochalciella, and P. fasciolalis in western North Amer- ica. Hemrich 1956, fig. 368) Pyla criddlella Dyar 1907: 110. Holotype 3d, Aweme, Manitoba, Canada (NMNH). [Type examined. | (Figs. 5, Diagnosis.—Pyla criddlella is the small- est of the gray Pyla with brownish gray wings and indistinct maculation (Figs. 11, 12). The genitalia are diagnostic in both 194 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON @ impostor * aenigmatica yeh jis, ©), sexes. The male aedeagus is asymmetric with the lateral divisions separated by a membranous gap, and the right division de- veloped distally into a posterolaterally di- rected pointed spine (Heinrich 1956, fig. 368a). The female ostium bursae is semi- membranous without heavily sclerotized plates or processes; the posterior portion of the corpus bursae has a short convolute band of sclerotization with three adjacent diverticula (Figs. 5, 6). Redescription.—Alar expanse: 18.1 to 22.4 mm (mean = 20.8 mm, n = 7) Ves- titure: A mixture of semilustrous light and dark brownish gray scales, many scales white tipped, darker and browner than most other gray Pyla species. Head: Frons con- vex, scaled brownish gray; male vertex shallowly concave with appressed central lustrous brownish scaling, female vertex flat with loose central pale-tipped scaling, ocelli and chaetosemata well developed; la- Known distribution of Pyla impostor and P. aenigmatica in North America. bial palpus thick, porrect, length 2%—3x width of eye, 2nd segment longest and broadest, scaled with brownish gray and pale-tipped scales, base and mesial aspect paler; maxillary palpus with tufts of narrow scales extending to % length labial palpus; tongue well developed, covered proximally with brownish gray scales; antenna lami- nate, basal segments of male flagellum modified forming a mesial sinus, each mod- ified segment with a distal spine, spines progressively larger distally, sinus bordered medially and laterally by rows of gray scales, inner row of scales forming medial border of sinus modified, thick, dark, each with curved central spine-like tooth, base of female flagellum unmodified, unmodified segments of flagellum in both sexes scaled pale brown, scape nearly uniform brownish gray. Thorax: Dorsum, tegula, and collar and ventral aspect of thorax gray or brown- ish gray, scales sometimes faintly pale- VOLUME 104, NUMBER 2 tipped; legs similar in color, mesial aspect paler, tarsal segments white-tipped distally, protibia with spine-tipped epiphysis from middle extending just distal to first tarsal joint, mesotibia with one and metatibia with two pair of spurs. Wings (Figs. 11—12): Forewing dark semilustrous brownish gray sprinkled with white-tipped scales; mark- ings faintly contrasting, antemedial and postmedial lines indicated by slightly great- er concentration of white-tipped scales, bor- dered with slightly darker shade, anteme- dial oblique, postmedial indented opposite cell and in lower third, discal mark and ter- minal line absent; fringe paler brownish gray white-tipped scales; underside brown- ish gray, paler along inner margin, vague indication of postmedial line, fringe nearly unicolorous, paler at base. Hindwing brownish gray, lighter than forewing, veins slightly darker; fringe paler, a thin pale line at base; underside concolorous with upper- side of hindwing and underside of fore- wing, fringe paler. Abdomen: Brownish gray, slightly paler than thorax. Male genitalia (Heinrich 1956: fig. 368): Uncus subtriangular and hood-like; gnathos with a stout central hook; valva narrow, ta- pered in distal third, slightly greater degree of sclerotization along proximal %4 of costa and proximal % of sacculus; clasper (sensu Heinrich) at costal aspect of valva base, a slightly irregular sclerotized ridge devel- oped into a short medial triangular or tooth- like process; tegumen unmodified; juxta de- veloped into a rounded pocket anteriorly, weakly sclerotized lateral to the aedeagus, without posterior processes; aedeagus (Heinrich 1956, fig. 368a) divided distally, divisions asymmetric, sclerotized, the right longer, thicker, semi-cylindrical, developed into a lateral sclerotized spine distally, ve- sica with fine scobinate patch, cornuti ab- sent; anellar membrane with small pouches dorsal to basal process of valva; eighth ab- dominal sternite with a pair of hair pencils. Female genitalia (Figs. 5—6): Ostium bursae finely scobinate, semimembranous, funnel-shaped, lamella antevaginalis with 495 weakly sclerotized folds radiating to junc- tion of ostium bursae and ductus bursae; junction of ostium bursae and ductus bursae weakly convoluted, membranous and slightly fleshy; ductus bursae short, curved, arising ventrally from ostium bursae; cor- pus bursae largely membranous, posterior half coarsely wrinkled with an oblique con- voluted band of sclerotization, three small membranous diverticula from posterior half of corpus bursae, two dorsal, one ventral; ductus seminalis arising from a sclerotized dorsal outpouching on the left; ovipositor telescoped, less than half length of genital capsule, ovipositor lobes slightly concave laterally. Holotype-—d ‘“‘Criddle, Aweme, Man., 1O.V1.04"’ = “WVype* 2 * (Genitalia, Slide, By 101,103, USNM’’ [green label]; ““Type, No. 10344, U.S.N.M.” [red label]; “‘[male sym- bol] genitalia on, slide 30 Jan 1942, C.H. 2780” (NMNH). Material examined.—(6 ¢, 12 2) CAN- ADA. Alberta: 1 2, 28 Jun 1923, Waterton Lakes (CN@): 1G, 1 July 1923, Waterton Lakess(@NG)3a)g,. 172, 2k 1922, Bantt (CNC). British Columbia: 1 2, 11 Jul 1937, Jesmond (CNC); 2 2, 11 Jul 1937, Jes- mond, 1,500’ (CNC); 1 2, 9 Jul 1923, Jes- mond,,.3;500;, (ENG); tie ?3.13 Jul 1937 Jes- mond (CNC); 1 6, 22 July 1937, Jesmond (CNC); 1 2, 22 Jul 1937, Jesmond, 7,500’ (ENCG)> 1), 6.923: Sun, 1938-4 Canim, Wake (CNC); 1 ¢, 28 Jun 1937, Kamloops (CNG) 2yit-Gse30e une 1955s Atlins 92.200) (CNC). Manitoba: 1 6, 1 2, 16 Jun 1904, Aweme (CNC); 1 @, 10 Jun 1926, Aweme (CNC). Saskatchewan: 1 2, 3 June 1926, Cypress Hills, nr Maple Creek (CNC). Discussion.—Pyla criddlella was previ- ously described from a single male (Dyar 1907, Heinrich 1956). The female is de- scribed here for the first time. Unlike other gray Pyla, P. criddlella, P. hanhamella Dyar, and P. hypochalciella (Ragonot) are brown or brownish gray moths but not dis- tinctly lustrous bronzy-brown. Pyla crid- dlella is the smallest of the gray Pyla and its distribution is poorly known (Fig. 8). 196 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON The documented distribution includes southwestern Manitoba (Aweme), southern Saskatchewan (Cypress Hills), southwest- ern Alberta (Waterton Lakes, Banff) and south central British Columbia (Jesmond, Kamloops, Canim Lake); specimens ex- amined from these localities were collected between 1904 and 1938. A single male was collected at Atlin in northwestern British Columbia in 1955. Flight dates range from 3 June to 22 July and the life history and habits are unknown. Perhaps the small number of records and lack of recent col- lections reflects narrow habitat require- ments. History and etymology.—Pyla criddlella is named for the Canadian entomologist Norman Criddle (1875-1933), a well known illustrator and agricultural entomol- ogist who avidly studied and collected in- sects in southwestern Manitoba. The Crid- dle homestead and farm, southeast of Bran- don, Manitoba, is in a region of mixed sandy prairie, forest (primarily white spruce, Picea glauca (Moench) A. Voss; and poplar, Populus tremuloides Michaux), and wetland supporting tamarack (Larix laricina (DuRoi) K. Koch) and black spruce (Picea mariana (Miller) BSP). A fossil sand dune system, the Carberry Desert or Bald Head Hills, is evident as actively moy- ing open sand deposits to the east of the homestead. Norman Criddle frequently made day collecting trips into these sand hills. Despite the local diversity of habitats, all Criddle material is labeled Aweme, a name given to his entomological lab and the local post office (Criddle 1973). Bird (1927) and Criddle (1915) described the habitats in the vicinity of Aweme. Norman’s father Percy Criddle (1844— 1918) and other members of the family had diverse interests in natural history. Percy was unquestionably proud of Norman’s ac- complishments and his ego bolstered, if he was not somewhat amused, as insects took on the Criddle namesake: “T expect to find myself elected correspond- ing member of at least six learned societies in Ontario and elsewhere immediately, on ac- my butterfly collection—buffalo bones—and chunks of old stones. While my new friends anticipate a brilliant future and immortality owing to my possible discovery of some new insect or other in this unex- plored district which will be of course called *‘Criddle-de-diddle-ensis’ or some other fancy family name.” [Entry from Percy Criddle’s diary, one year after immigrating to Manitoba from England, following his visit with Ernest Thompson Seton and ‘Mr. Christie’ in 1883 (Criddle 1973: 94).] count of Pyla fusca (Haworth) (Fig. 10; Heinrich 1956, figs. 369, 852) Phycis fusca Haworth 1811: 493. England. [Location of type unknown. ]} Phycis spadicella Zincken 1818: 168. 3 and 2 syntypes indicated in original de- scription, Augsburg, Germany. [Types not examined. | Phycis janthinella carbonariella Duponchel 1836: 292; plate 281, fig. 2. Syntypes not itemized in original description, Domo d’Ossola, Italy. [Types not examined. ] Phycis posticella Zetterstedt [1839]: 996. Syntypes 1 6, 1 @, Lapland (Riksmu- seum, Stockholm, Sweden). [Physis pos- ticella Zetterstedt is treated as a synonym of Salebria fusca Haworth by Benander (1940).] [Types not examined. ] Phycis annulatella Zetterstedt [1839]: 997. Syntypes 1 6, 2 2, Lapland (Riksmu- seum, Stockholm, Sweden). [Phycis an- nulatella Zetterstedt is treated as a syn- onym of Salebria fusca (Haworth) by Benander (1940), and is noted to be a ju- nior synonym of Pyla fusca (Haworth) by Speidel (1996). Speidel (1996) further notes that P. annulatella sensu Ragonot is the misidentified type species of Po- lopeustis Ragonot.] [Types not exam- ined. | Phycitia bilineata Curtis 1850: 114. 1 ¢ is indicated in the original description, Nor- folk, England (syntypes in Melbourne Museum, Australia). [This was an unpub- lished new synonymy, M. Schaffer per- VOLUME 104, NUMBER 2 497 Fig. 10. Known distribution of Pyla fusca in North America, including Greenland. sonal communication 1999.] [Types not United States, five from St. Martin’s examined. | Falls, and one from Nova Scotia.] [Types Nephopteryx moestella Walker 1863: 53. not examined. | Extant syntypes include | 2 from Nova Paedisca procellariana Walker 1863: 379. Scotia, 4 2 from St. Martin’s Falls, On- Holotype 2, Arctic America (BMNH). tario, and possibly | 2 with no data that [Type not examined. ] carries Walker’s name label; no lectotype Eudorea ‘*?” frigidella Packard 1867: 53. designated (BMNH). [Note: Walker’s de- Syntype d [abdomen missing], Caribou scription clearly indicates that both sexes Island [= Great Caribou Island], Labra- were represented in the type series and dor, Canada (MCZ). [Note: Packard did he itemized three specimens from the not indicate the number or sex of the 498 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON specimens in his type series. The date of publication follows Miller and Hodges (1990). The MCZ syntype is labeled ‘“*Labrador”’ and likely is one of a series of specimens from which Packard de- scribed frigidella. Subsequently Packard (1873) recognized that his frigidella was a synonym of Pempelia fusca (Haworth) and the syntype is so labeled by Packard. This specimen also bears a label “*Eu- dorea borealis n. sp.,”’ a combination not published by Packard but likely a misla- beling as the name Pyrausta borealis Packard was published on the same page as Eudorea ? frigidella. Pyrausta boreal- is Packard is now treated as a subspecies of Pyrausta subsequalis (Gn.) (Hodges et al. 1983)]. [Type examined. | Pinipestis cacabella Hulst 1887: 133. Ho- lotype 3d, New York, United States (AMNH). [Note: New York is given as the type locality in the original descrip- tion but the type lacks a locality label as noted by Heinrich (1956) and Rindge (1955).] [Type examined. ] Salebria triplagiella Dyar 1904: 109. Ho- lotype 6, Winnipeg, Manitoba, Canada (NMNH). [Type examined. ] Pyla fusca is the most commonly en- countered and widely distributed Pyla in North America and ranges from the arctic south to New Jersey, North Carolina, and the Great Lakes region in the east and to Colorado and northern California in the west (Fig. 10). As the only holarctic Pyla, it is also widespread in northern and central Europe and Asia, and recorded from Green- land and Iceland (Speidel 1996, Sinev 1997, Wolff 1971). Based on European re- cords the larvae primarily feed on the shoots and leaves of Ericaceae (Goater 1986, Palm 1986). Darlington (1952) cites a rearing from “blueberry” at Anglesea, New Jersey. Flight dates range from 25 May to 23 September. Although P. fusca deviates from other Pyla species based on the presence of a cor- nutus on the male vesica, the aedeagus is divided as in other Pyla species but with one division very fine and closely ap- pressed to the opposite, distally curved, and more heavily sclerotized division. The pres- ence of two divisions is not apparent in published figures of Heinrich (1956, fig. 369a). We are indebted to Michael Shaffer (The Natural History Museum, London) for per- mitting us to publish the above synonomy for Pyla fusca. Material examined: 107 6, and 128 2 [AMNH, CNC, -CULE, (GJB, JKMikS LACM, MCZ, MSU, NMNH, UM, UMMZ, UZMH]. Pyla hypochalciella (Ragonot) (Fig. 7; Heinrich 1956, figs. 370, 854) Nephopteryx ovalis hypochalciella Ragonot 1887: 7. Type a single specimen of in- determinate sex and in poor condition [E. Cudel, personal communication 1998], Washington Territory (Paris Museum). [Type not examined. ] Pyla blackmorella Dyar 1921: 68. Holotype d, Mt. Tzouhalem, Duncan, British Co- lumbia, Canada (NMNH). [Type exam- ined. | Pyla hypochalciella is an infrequently collected species from the Pacific North- west (Fig. 7), most specimens are known from central and southern Vancouver Is- land, British Columbia, with single speci- mens examined from Tenino, Washington and Baker, Oregon, all collected between 1913 and 1947. Flight dates range from 1 June to 4 July. Life history and habits are unknown. Specimens of P. hypochalciella are found mixed in collections with a superfi- cially similar phycitine, a brown species of Catastia Hiibner that occurs to the east in the Cascade Mountains. Py/a and Catastia are readily distinguished by genitalia in both sexes (Heinrich 1956). Males can also be separated by examination of the modi- fied basal segments of the antenna, which are developed into a sinus in both genera. VOLUME 104, NUMBER 2 In P. hypochalciella the spines within the sinus are short and exceeded and partially covered by a tuft of long curved scales along the mesial aspect of the sinus; in the Catastia species the spines are long, the distal spines exceeding the shorter mesial scale tuft. The row of scales along the inner border of the sinus are modified distally into spine-like processes in P. hypochal- ciella (as in other gray Pyla) whereas these scales are rounded distally with multiple fine teeth in the Catastia species. Material examined: 7 d and 5 @ [CNC, NMNH]. Pyla hanhamella Dyar (Fig. 8; Heinrich 1956, figs. 371, 860) Pyla hanhamella Dyar 1904: 109. Holotype 6, Winnipeg, Manitoba, Canada (NMNH). [Type examined. ] Pyla hanhamella is recorded from rela- tively few localities in west central North America (Fig. 8) including central and southern Manitoba (Aweme; 2 miles west of Stockton; Onah; Winnipeg; “‘C[entral] Manitoba’), western North Dakota (Co- lumnar Juniper Area, Slope county; not mapped), western Saskatchewan (Harlan), and southwestern Alberta (Hillcrest; Water- ton Lakes). A single male was flushed from short-grass prairie in the badlands of Slope Co., North Dakota near midday, June 12, 1991 (GJB) (see photograph of habitat in McCabe and Post 1977, fig. 24). Flight dates range from 20 May to 25 July. The life history is unknown. Material examined: 13 ¢, 12 2 [AMNH, CNC, GJB, NMNH]. Pyla arenaeola Balogh and Wilterding (Balogh and Wilterding 1998, figs. 1—5, [1—13, 17, 19-22) Pyla arenaeola Balogh and Wilterding 1998: 705. Holotype 6, Saugatuck Dunes State Park, Allegan County, Michigan, USA (42.39'18’N, 86.12'07”"W) (NMNH). Pyla arenaeola is known only from the dunes of the Great Lakes and southwestern 499 Manitoba (Balogh and Wilterding 1998, fig. 17); the larvae live hidden in sand tubes and feed on Bearberry (Arctostaphylos uva- ursi). Details of the life history, habits, and biogeography of P. arenaeola can be found in Balogh and Wilterding (1998). Addition- al specimens collected on the southern shore of Lake Huron were examined since publication of the original description (CANADA: Ontario, Lambton County, Port Franks, | d, 30 Jul 1997 and 1 2, 7 Aug 1998 [KS]). Material examined: 162 36, 95 @ [BGS, ENC, (GIB; JAB; JHW:. IKM, KS. jBAR MSU, NMNH, UMMZ]. Pyla westerlandi Wilterding and Balogh, new species (Figs. 1-4, 7, 13, 14) Diagnosis.—This species is on average the largest gray Pyla species, but like other species of the group, is best recognized by features of the male and female genitalia. Males have an unusually long and stout clasper (sensu Heinrich) near the base of the valva and the aedeagus has symmetric laterally directed thorn like spines distally (Fig. 3). Females have an unusually large rectangular ostium bursae with a pair of rounded heavily sclerotized ventrolateral plates (Fig. 4). Description.—Alar expanse: 29.7 to 36.2 mm (mean = 33.2 mm, n = 22). Ves- titure: A mixture of gray, pale gray, whit- ish, and pale tipped gray scales. Head: Frons convex, a broad tuft of pale-tipped gray scales superiorly, paler scaling ven- trally; male vertex concave with appressed central scaling and an anterior row of thick- er and narrower brownish scales, female vertex nearly flat with loose central scaling, in both sexes most scales gray, often pale tipped, a tuft of elongate pale-tipped gray scales posteriorly; ocelli and chaetosemata well developed; labial palpus ascending to level of vertex, 2nd segment longest and broadest and in male with medial groove for maxillary palpi, 3rd segment thin, 4% length of 2nd, scaled with gray and whitish PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 11-14. scales, pale scales predominate on base and mesial aspect, 3rd segment darker and often pale at tip; male maxillary palpus extends nearly to % length 2nd segment labial pal- pus, Ist segment thin and elongate, 2nd bul- bous distally, pale tufts of elongate thin scales from 2nd and 3rd segments extend to distal aspect 2nd segment labial palpus, female maxillary palpus short, 3rd segment reduced, scale tufts shorter; tongue well de- veloped, covered proximally with gray and pale scales; antenna laminate, basal 5—6 segments of male flagellum modified form- ing a mesial sinus, first several segments of Pyla criddlella and P. westerlandi adults. 11-12, P. criddlella. 11, Alberta. 12, female, Cypress Hills, Saskatchewan. 13-14, P. westerlandi. 13, Holotype male, Ward Creek, 2 mi S. Tahoe City, California. 14, Paratype female, same locality. male, Waterton Lakes, sinus may be fused or partially fused, each modified segment with a distal spine, spines progressively larger distally, sinus bordered medially and laterally by rows of gray and pale-tipped gray scales, inner row of scales forming medial border of sinus modified, thick, dark, each with curved central spine- like tooth, base female flagellum unmodi- fied, unmodified segments of flagellum in both sexes scaled with fine bands of gray and pale scales, scape with broad posterior band of pale scales. Thorax: Dorsum, te- gula, and collar scaled with pale-tipped gray scales, whitish scaling predominates VOLUME 104, NUMBER 2 on ventral thorax; legs scaled with mixture of gray and whitish scales, white predomi- nates on mesial aspect, mesotibia with dark band at *%rds, metatibia sometimes with vague darker banding, tarsi contrastingly dark, each segment white-tipped distally, protibia with spine-tipped epiphysis from middle extending just distal to first tarsal joint, mesotibia with one and metatibia with two pair of spurs. Wings (Figs. 13-14): Forewing pale gray with little variation, pale scales nearly uniformly distributed in basal, medial, and subterminal spaces, often a vague darker patch at anal margin of me- dian space; antemedial line slightly paler than rest of wing, oblique, variably dentate, indented and most contrasting in lower third, margined proximally by dark gray patch along lower third, and distally by a darker line which is widest at costal half; postmedial line pale, dentate, indented op- posite cell and in lower third, margined proximally and distally by dark line, the proximal dark line usually better defined; terminal line a thin row of dark gray spots; discal spot dark gray, bilobed; fringe pale gray white-tipped scales; underside pale gray, paler along inner margin, weakly de- veloped dark subterminal mark at costa, ter- minal line faint. Hindwing pale brownish gray, slightly translucent, darker at outer margin, veins slightly darkened; fringe con- trasting, pale, a brownish gray band proxi- mal third; underside light gray, slightly darker at costal and outer margins. Abdo- men: Scaled with gray and whitish scales forming lighter bands at posterior margin of segments. Male genitalia (Figs. 1—3): Uncus sub- triangular, hood-like; gnathos with a stout central hook; valva with base broad and concave mesially, flattened and tapering distally, proximal *%4 costa slightly thickened and with small smooth bulge at base, sac- cular margin thickened in proximal 2 and forming a bulbous rounded basal protuber- ance; clasper (sensu Heinrich) large, arising from base of valve as a thick heavily scler- otized hollow spine, midportion slightly in- 501 curved, pointed distally, %—% length of valve, basal portion finely spiculate; tegu- men unmodified; juxta U-shaped, forming a pocket anteriorly, posterolateral paired pro- cesses slightly incurved, often with a short spine at tip of each process; aedeagus (Fig. 3) slightly keeled ventrally, divided distally, divisions symmetric, sclerotized, each di- vision with a single short ventrolateral rose thorn-like process proximal to posterior end, cornuti absent; anellar membrane de- veloped into wrinkled membranous folds mesial to valve base; eighth abdominal ster- nite sclerotized as in Fig. 2 with prominent hair pencils. Female genitalia (Fig. 4): Ostium bursae large, rectangular, nearly equal in length to corpus bursae, lamella postvaginalis largely membranous with weakly developed bands of sclerotization, lamella antevaginalis de- veloped into paired, convoluted, heavily sclerotized plates with broadly rounded posteromedial margins, anterior aspect of ostium bursae produced into a squared-off pocket which protrudes dorsal to ductus bursae; junction of ostium bursae and duc- tus bursae membranous, flanked internally by fleshy folds; ductus bursae short, mem- branous, about *%4 width of corpus bursae, a slightly asymmetric bulge on left; corpus bursae membranous, broadest in anterior half, nearly symmetric; ductus seminalis dorsal, from posterior margin of corpus bur- sae; Ovipositor telescoped, less than half length of genital capsule, ovipositor lobes slightly rounded laterally. Holotype.—