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Ty £ wa) ion ay 4) n tel Yon" I c “u a ' 4s o ral DY eS ee ‘TT Lat vy ty 4 / . + i ¥ i } ' ' uy y hf ai eee | er i ' 4 ae m } ie z f 7 , if fi i ee ; = iy wd a? i ‘ ; 4 ! ran i ih \ (, en ul iV i 1 i | i we ee a re ee } { i i hy Fj i A i LM } , i f 4s } T) | - any ‘alt ’ AM a oe 1 ie ap sit! in ii ve hn wh Fi TR OY Ak TR 1 ' } tt i ( Whe) La | i ay, inh) ihe RU ket ie MME Wels mar me ny ed AON HAT FU en i A Yea 8 Ye, ii i SNE tay Wy 4 Wey ir iy Carel | YN: de } Ait Wi A i \ As, Y) ‘ ee! i Ly ae iL ety hie Wf NA Me a ae at Loe ; f i ne 4 vy Wao ! ae Rea Av Pas Pe ee me at We ey ial ea ures nah ’ i i enh id f y f von ant an cee ry te ayy ay ey i wy ‘ } ial ee ak Mate (|e yi ’ 1A ry j yh ih ; te Bh ve i Van a AK I Al iss i" q cas na —— it es ! Ne yi iy) ae wi) Se eet eM Ns a | hiv i) ae) ; le a f mo la , : ' i : i he , Pe pe , a Nie y ion ori iM Lin it Fae | VOR; 99 JANUARY 1997 NO. 1 lo | 29K (ISSN 0013-8797) Pai eh PROCEEDINGS of the ENTOMOLOGICAL SOCIETY of WASHINGTON PUBLISHED CONTENTS BATRA, SUZANNE W. T.—Bionomics of Lasioglossum (Evylaeus) matianense (Blithgen) (Hymenoptera: Halictidae), the predominant pollinating bee in orchards at high altitude in ther GreatiHimalayajohGarhwall (UCP = india ace serrate oir tora Aare srattiegstet eee cles tee oteieten eee terse 162 COSCARON, MARIA DEL C. and JUAN J. MORRONE—Cladistics and biogeography of the assassin bug genus Melanolestes Stal (Heteroptera: Reduviidae) ...................00..eeee eee Sys) EVANS, HOWARD E. and STUART M. FULLERTON—Report on a collection of Bethylidae @aymenoptera)iiroml central Hloriday 4 nrst oh wecloicrn vale ole ier erent satay enter 174 FITZGERALD, SCOTT J. and BORIS C. KONDRATIEFF—A new species of Pseudonomo- neura Bequaert (Diptera. Viydidae) from Mexico a o.n acceso soueiece = erect ei en ee 171 GAGNE, RAYMOND J., CORNELIA OTT, and SUSANNE S. RENNER—A new species of gall midge (Diptera: Cecidomyiidae) from Ecuador associated with flowers of Clavija (Theo- (od a) EIST Yoho) | Rote aR ane A eRe Re RSME AG ATS GONE ceiind sot sao gem pHooUpoSad an nace orate oAdeHe 9: ‘i 110 GOEDEN, RICHARD D. and JEFFREY A. TEERINK—Notes on life histories and descriptions of adults and immature stages of Procecidochares kristineae and P. lisae new species (Dip- tera: Tephritidae) on Ambrosia spp. in southern California ..................0.ee ee eeeee eee ee ees 67 GOEDEN, RICHARD D. and JEFFREY A. TEERINK—Life history and description of imma- ture stages of Procecidochares anthracina (Doane) (Diptera: Tephritidae) on Solidago cal- iornrcauNuttall sin, Souther) Califormiaw y. 1. cerca. : seem soon eee eee ea etree te aie 180 KROMBEIN, KARL V. and BETH B. NORDEN—Nesting behavior of Krombeinictus nordenae Leclercq, a sphecid wasp with vegetarian larvae (Hymenoptera: Sphecidae: Crabroni- ip feet) Meta 0S a ol Sen as Aan RE Serene Oe Ati ear aaa Me tac AamA AK Gh tas nh acrid dan ion cide Sanam Adee 42 LOYA, LANE J. and J. E. MCPHERSON—Life history and laboratory rearing of Oedancala dorsalis (Say) (Heteroptera: Lygaeidae), with descriptions of immature stages ............... 89 MACKAY, WILLIAM P.—A revision of the Neotropical ants of the genus Camponotus, sub- eenus Myrmostenius (Hymenoptera: HOLMICIGAC)!Lovinsa as same nes islelee te need gelonsteeet = ace ress 194 MATILE, LOIC—Fenderomyia Shaw, a valid North American taxon in Macrocerinae (Diptera: Mivcetophilordea mero attdae) i -.cietms cts aareetatsrs cle slas nlettatetclofeteinke eyetclelety eee tertiaae aii -efone aieios se) siete es 50 (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ORGANIZED MARCH 12, 1884 OFFICERS FOR 1997 M. ALMA SOLIS, President MICHAEL G. POGUE, Treasurer WARREN E. STEINER, JR., President-Elect Davip G. FurtTH, Program Chair HAROLD HARLAN, Recording Secretary Davip ADAMSKI, Membership Chair HOLuis B. WILLIAMS, Corresponding Secretary RALPH P. ECKERLIN, Past President Gary L. MILLER, Custodian DAVID R. SmiTH, Editor Publications Committee THOMAS J. HENRY WAYNE N. MATHIS Gary L. MILLER, Book Review Editor Honorary President CurRTIS W. SABROSKY Honorary Members LoulIsE M. RUSSELL ALAN STONE KARL V. KROMBEIN All correspondence concerning Society business should be mailed to the appropriate officer at the following address: Entomological Society of Washington, % Department of Entomology, MRC-168, Smithsonian Insti- tution, Washington, D.C. 20560. 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 8 P.M. Minutes of meetings are published regularly in the Proceedings. MEMBERSHIP.—Members shall be persons who have demonstrated interest in the science of entomology. Annual dues for members are $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, MRC-168, Smith- sonian Institution, Washington, D.C. 20560. Members in good standing receive the Proceedings of the Ento- mological 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. 615-616 OF THE JULY 1996 ISSUE FOR INFORMATION REGARDING PREPARATION OF MANUSCRIPTS. STATEMENT OF OWNERSHIP Title of Publication: Proceedings of the Entomological Society of Washington. Frequency of Issue: Quarterly (January, April, July, October). Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash- ington, D.C. 20560. Editor: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560. Books for Review: Gary L. Miller, Systematic Entomology Laboratory, ARS, USDA, Building 046, BARC- West, Beltsville, MD 20705. Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 25 February 1997 Second Class Postage Paid at Washington, D.C. and additional mailing office. PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PROC. ENTOMOL. SOC. WASH. 99(1), 1997, pp. 1—27 ERETMOCERUS HALDEMAN (HYMENOPTERA: APHELINIDAE) IN THE UNITED STATES, WITH DESCRIPTIONS OF NEW SPECIES ATTACKING BEMISIA (TABACI COMPLEX) (HOMOPTERA: ALEYRODIDAE) MIKE ROSE AND GREGORY ZOLNEROWICH Department of Entomology, Texas A&M University, College Station, TX 77843, U.S.A. Abstract.—A key to species of Eretmocerus that occur in the continental United States is provided. Eretmocerus eremicus, n. sp., E. joeballi, n. sp., E. staufferi, n. sp., and E. tejanus, n. sp., all reared from Bemisia (tabaci complex) collected in the United States, are described. Redescriptions of Eretmocerus californicus Howard, E. corni Haldeman, E. haldemani Howard, and E. portoricensis Dozier are provided. Eretmocerus debachi Rose and Rosen, FE. furuhashii Rose and Zolnerowich, and E. illinoisensis Dozier are discussed. Lectotypes are designated for Eretmocerus californicus Howard, E. haldemani Howard, and E. portoricensis Dozier. Key Words: Species of Eretmocerus Haldeman, 1850 (Hymenoptera: Chalcidoidea: Aphelinidae) are attracting widespread interest because of population explosions of Bemisia (tabaci complex) (Homoptera: Aleyrodidae) around the world (Rose et al. 1996). All known species of Eretmocerus are primary para- sites of whitefly, and species of Eretmoce- rus have been purposefully utilized in ef- fective biological control programs (Rose and DeBach 1991-1992; Rose and Rosen 1991-92; Rose 1988). This paper, which is part of a larger study of Eretmocerus spe- cies of the world, is designed to clarify characterizations of named species in the U.S., and to describe new species that at- tack Bemisia (tabaci complex) in the U.S. Because of confusion regarding the use of the names Bemisia tabaci (Gennadius) A-strain, B. tabaci B-strain, and B. argen- tifolii Bellows and Perring on specimen la- bels and in the literature (Brown et al. 1995), we refer to hosts from this group as Bemisia (tabaci complex). Species of Be- misia (tabaci complex) have emerged as a Eretmocerus, Aphelinidae, Bemisia, Aleyrodidae, biological control major agricultural pest attacking a variety of food, cash, and ornamental crops in the United States and abroad. Bemisia (tabaci complex) attacks over 500 plant species in 74 families (Mound and Halsey 1978; Brown and Bird 1992) and new host plants are continually being added (Gill 1992; Costa et al. 1993). Crop damage occurs due to feeding, honeydew production and resul- tant sooty mold, and disease transmission. Species in the Bemisia (tabaci complex) transmit more than 15 viruses (Byrne et al. 1990) that cause more than 40 plant dis- eases (Brown and Bird 1992). Reflecting the current U.S. situation, Bemisia (tabaci complex) is a major pest of crops in Mex- ico, the Caribbean, Central and South America, the Middle East, India, and parts of Africa. Biological control research to attain pop- ulation regulation of Bemisia (tabaci com- plex) in the U.S. is emphasizing evaluation of naturally occurring species of parasitic Hymenoptera and importation of exotic populations and species of parasites. The 2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON majority of native and imported species are found in Eretmocerus and Encarsia Foers- ter (Hymenoptera: Chalcidoidea: Aphelini- dae). Polaszek et al. (1992) provided a key and discussion of Encarsia species that at- tack Bemisia (tabaci complex). Schauff et al. (1996) furnished a pictorial key and dis- cussion of species of Encarsia that attack whitefly in North America. Species of Ami- tus Haldeman (Hymenoptera: Platygastri- dae) also have been reared from Bemisia (tabaci complex) (Viggiani and Evans 1992): Surveys to obtain samples of Bemisia (tabaci complex) and its parasites from var- ious host plants in the U.S. have been un- dertaken primarily in Arizona, California, Florida, and Texas. These states have suf- fered the greatest damage to field crops and subsequent losses to Bemisia (tabaci com- plex), so biological control research efforts have been emphasized in these areas. It is vitally important that researchers in these and other areas are able to identify the bi- ological entities they are discovering, im- porting, colonizing and evaluating. There are currently 8 described species of Eretmocerus from the New World, and 30 species are known from the Old World. Given that there are nearly 1200 described species of whitefly, and that species of Er- etmocerus are known from all continents where whitefly occur, there are undoubtedly many more undescribed species of Eret- mocerus. Because most species of Eretmo- cerus have been reared from agricultural pests, little is known of the actual range of these species. Likewise, little is known about species found outside of agricultural settings. The literature encompassing the taxono- my of Eretmocerus is disparate in its com- prehensiveness, and researchers have relied on historical taxonomy to provide names for numerous species of Eretmocerus that are currently being studied and transferred nationally and internationally. These histor- ical names often find their way into new literature and are then generally adopted, which may obscure the actual species being studied. For example, two of the species de- scribed in this paper were called E. califor- nicus or E. sp. nr. californicus for many years. Previous studies (Rose and Rosen 199]— 1992, Rose and Zolnerowich 1994, Rose et al. 1996, Hunter et al. 1996) have empha- sized developing criteria for the character- ization of species based on morphology, pigment patterns of males, reciprocal mat- ing trials, and electrophoretic analysis. These studies all indicate that minor mor- phological differences in the size, shape, or chaetotaxy of the habitus, antennae, and forewings, and pigment patterns of males, are significant species characters. Males of different species of Eretmoce- rus are very similar unless they have fea- tures such as distinct pigment patterns. The poor condition of male specimens available for Eretmocerus haldemani and E. illinois- ensis preclude their separation in the key to species. Males are unknown for Eretmoce- rus joeballi and E. portoricensis. METHODS High quality microslide mounts are nec- essary to correctly examine and _ identify species of Eretmocerus. Because clearing and mounting specimens in balsam removes all or most of their color, most specimens used in this study were mounted in Hoyer’s medium as described by Rosen and DeBach (1979), and the cover slips were sealed with two coats of red GLPT, a nonconducting insulating varnish used in electronics (GC Electronics, Rockford, Illinois). Other spec- imens were critical point-dried and mount- ed on cards prior to mounting in balsam. Most primary type material was mounted in balsam. Position of the body and antenna on mi- croslides is critical for proper identification. The antenna should be examined from the lateral aspect. Viewing the antenna from any other angle obscures the true length and shape of the segments. In many instances, the antennae were removed from the head VOLUME 99, NUMBER 1 fll —_ ro fl 1 2 3 Figs. 1-3. 1, 2 antenna. 2, Forewing. 3, Venation of forewing. c = length of club, fl = length of first funicular segment, fll = length of second funicular segment, 1 = length of forewing, m = length of marginal vein, p = length of pedicel, r = length of radicle, s = length of scape, sm = length of submarginal vein, st = length of length of stigmal vein, w = greatest width of disc, wl = width I. and mounted separately on the same slide. Wings should be positioned away from the body to reveal their shape and allow accu- rate measurements to be made. Length of the body was measured from critical point-dried specimens using an eye- piece micrometer when possible. Other measurements of specimens mounted in balsam or Hoyer’s were taken using an eye- piece micrometer or a digitizing tablet. The maximum length of each antennal article in lateral view was measured (Fig. 1). The an- tennal ratios given in descriptions are the ratios of the lengths of the radicle : scape : pedicel : club. Unless expressed as a range, the length : width ratios represent average values. Length and width of the forewing were measured as shown in Fig. 2. The maxi- mum length (1) and width of the wing across the disc (w) were measured. Width I (wl) is the distance between the distal end of the frenal fold and the anterior margin of the forewing above the distal end of the stigmal vein. Points of measurement for the lengths of the submarginal, marginal, and stigmal veins are shown in Fig. 3. The ra- tios of the lengths of the longest anterior and posterior alary fringes (Fig. 5) to the width of the wing are given, as is the num- ber of tubercles (Fig. 11), which are present on the ventral side of the wing. In many instances, the distal boundary of the marginal vein and the advent of the stig- mal vein were not clearly delimited. In those cases, the end of the marginal vein and the beginning of the stigmal vein were taken at the base of the distal large seta on the marginal vein (Fig. 3, st). Descriptions of color for critical point- dried and specimens mounted in Hoyer’s may differ because of differences in speci- £ PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Species of Eretmocerus in the U.S. and original collection data. Species Author Year Original Host Insect corni types lost Haldeman 1850 Tetraleurodes corni (Haldeman) [= Aleurodes corni Haldeman] corni neotype Dozier 1932 Trialeurodes packardi (Morrill) [= Trialeurodes morrilli (Britton)] californicus Howard 1895 unknown ‘‘Aleyrodes’”’ haldemani Howard 1908 Aleuroplatus coronata (Quaintance) [= Aleyrodes coronata Quaintance] illinoisensis Dozier 1932 unknown portoricensis Dozier 1932 Aleurothrixus floccosus (Maskell) debachi Rose & Rosen 1991-92 Parabemisia myricae (Kuwana) furuhashii Rose & 1994 Parabemisia myricae (Kuwana) Zolnerowich eremicus Rose & 1997 Bemisia (tabaci complex), Zolnerowich possibly Trialeurodes Joeballi Rose & 1997 Bemisia (tabaci complex) Zolnerowich staufferi Rose & 1997 Bemisia (tabaci complex), Zolnerowich Trialeurodes abutiloneus (Haldeman) tejanus Rose & 1997 Bemisia (tabaci complex) Zolnerowich ' Host plant probably Quercus (R. Gill, personal communication). men preparation. Specimens mounted in Hoyer’s do not show colors as well as crit- ical point-dried material, but do show fus- cous pigment patterns well. Descriptions of color or pigment are given for both critical point-dried and Hoyer’s-mounted speci- mens where appropriate. For new and described species, label data for primary types is presented exactly as recorded on the specimen labels, with the data for each individual label enclosed by quotation marks and each line of the label separated by a slash. Label data for material other than primary types are standardized and multiple records from the same locality may be combined. Table | provides sum- mary information such as original locality, host, and host plant for the species of Er- etmocerus discussed here. The following acronyms represent insti- tutions or individuals who loaned material for study, are repositories for type material, or are otherwise mentioned in the text: BMNH, The Natural History Museum, London, United Kingdom; CDFA, Califor- nia Department of Food and Agriculture, Sacramento; HU, The Hebrew University, Rehovat, Israel; INHS, Illinois Natural His- tory Survey, Champaign; MJR, personal collection of M. J. Rose; NSM, National Science Museum, Tokyo, Japan; TAMU, Texas A&M University, College Station; UCR, University of California, Riverside; USNM, National Museum of Natural His- tory, Smithsonian Institution, Washington, Dic KEY TO SPECIES OF ERETMOCERUS IN THE UNITED STATES 1 Female, antenna with 2 funicular seg- moe (Yt EMTS, I) ae ac occa dcaon DZ _ Male, antenna without funicular seg- THENUS SS. hoe oe sence eee esse ate ces 1 2(1) Mesosoma brown to brown orange, con- trasting strongly with head and metasoma (Fase 25) i. x teartupceettays E. staufferi, n. sp. - Body yellow, mesosoma not contrasting with head and metasoma ............ 3 3(2) Mesoscutum with 4 setae (Fig. 15) ..... 4 VOLUME 99, NUMBER I iablewle Continued. Nn Original Host Plant Original Locality, Date Cornaceae: Cornus sericea, dogwood Balsaminaceae: /mpatiens biflora, jewel weed Fagaceae: Quercus agrifolia, California live oak unknown! unknown Burseraceae: Bursera simaruba Zygophyllaceae: Guaiacum officinale Rutaceae: Citrus spp. Moraceae: Morus sp., mulberry Malvaceae: Gossypium hirsutum, cotton Moraceae: Morus sp. Solanaceae: Solanum Solanaceae: Lycopersicon, tomato Cruciferae: Brassica oleracea, cabbage Pennsylvania, 1847 Pennsylvania, White Clay Creek, 1929 California, Los Angeles, 1887 California, Berkeley, 1908 Illinois, Elizabethtown, 1932 Puerto Rico: Bayamon & Central Aguirre, 1925 California, Orange County, 1982 Japan: Honshu, 1979 Arizona, Phoenix, 1991 California, Bakersfield, 1993 Texas, College Station, 1993 Texas, Mission, 1994 5(3) 6(5) 7(6) Mesoscutum with 6 setae Parapsis with 2 setae; club 4.1—4.5 times as long as its greatest width (Fig. 8) BAe ARI Ree Se es, Le La E. debachi Parapsis with 3 setae (Fig. 15); club 4.3- 5.6 times as long as its greatest width (Fig. 13) E. furuhashii Club 3.2-3.8 times as long as wide, with dorsal surface convex and contrasting with straight ventral surface (Fig. 20) Betts Pitas ton ses LEIS, SePEEITSE E. portoricensis Club 4.5—8.3 times as long as wide, and with dorsal and ventral surfaces more or less parallel (e.g., Figs. 4, 26) Dorsal surface of club apically tapered so that apex forms a broad point (Fig. 4); rad- icle 0.6 times or more as long as scape E. californicus Apex of club truncate; radicle 0.35—0.50 times as long as scape (e.g., Figs. 16, 26) Pedicel short, 2—2.3 times as long as wide; club 4.5—5.1 times as long as wide (Fig. 18); ovipositor 1.5 times as long as club and 1.3 times as long as midtibia E. joeballi, n. sp. Pedicel longer, 2.5—4 times as long as wide; club 5.9-8.3 times as long as wide (e.g., Figs. 6, 16); ovipositor shorter than or equal to length of club or midtibia ... 8 9(8) 10(9) 11(1) 12(11) Scape at least 6.5 times as long as wide (Fig. 16); midbasitarsus 8.5 times as long as wide E. haldemani Scape 3.5—5.5 times as long as wide (e.g., Fig. 10); midbasitarsus 5.5—8.3 times as long as wide Pedicel 2.4—3.1 times as long as wide, and 0.25—0.3 times as long as club (Fig. 6); reared from Trialeurodes ........ E. corni Pedicel 3—4 times as long as wide, and 0.29—0.39 times as long as club (Fig. 10); reared from Bemisia (tabaci complex) and possibly Trialeurodes Gastral tergite II usually with | pair, but occasionally 2 pairs, of lateral setae (Fig. 12); gastral sternites anterior to base of Ovipositor usually with a group of 4—5 se- tae, occasionally with 3—6 setae 3 sat gd SRE sciyiyl ne cme umes aaa as E. eremicus, 0. sp. Gastral tergite II usually with 2 pairs, but occasionally 1 pair, of lateral setae (Fig. 28); gastral sternites anterior to base of Ovipositor usually with a group of 7-8 se- tae, occasionally with 5—6 setae BO, Ora Ere Ps cna roe ae E. tejanus, n. sp. Mesoscutum with 4 setae (Fig. 15) .... 12 Mesoscutum with 6 setae Parapsis with 2 setae; mesoscutum with dark fuscous “‘T’’ shape; scutellum com- pletelyatuscouss(Biga29) pa -eaee- E. debachi 6 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON = Parapsis with 3 setae (Fig. 15); mesoscu- tum with light fuscous *““T’’ shape; scutel- lum medially fuscous, lateral areas unpig- isacsynieerol (Vente SN) 55555505000 E. furuhashii 13(11) Entire mesosoma brown to brown orange, contrasting strongly with head and meta- GYopmnkey (IFW, BY) ooo ou aod E. staufferi, n. sp. - Mesosoma not brown to brown orange, al- though fuscous pigment patterns on pro- notum, mesoscutum, or scutellum may be present or absent (e.g., Figs. 30,33) ... 14 14(13) Mesoscutum with longitudinal fuscous bandsi(Hic=33) ie oe. o E. tejanus, n. sp. - Mesoscutum without longitudinal fuscous bands cc BWR ne Goac. Gas seems IS) 15(14) Mesoscutum fuscous along anterior margin (Fig. 30), or with a light fuscous “‘T” shape (as in Fig. 31), or evenly fus- COWS! i 2e5 as ee eee een oar 16 - Mesoscutum not fuscous............ 197; 16(15) Hind tibia fuscous ...... E. eremicus, n. sp. - Hind tibia not fiscous) 2-25-24. - E. corni 17(15) Scape 2.5—2.8 times as long as pedicel .. . MN Ds ate ee ar eRe Lc, hepa ls E. californicus - Scape 3.2—4 times as long as pedicel .. . ENC ele ice ours E. haldemani, E. illinoisensis Eretmocerus californicus Howard (Figs. 4—5) Eretmocerus californicus Howard, 1895: 16 [USNM, examined]. Type material. Lectotype here designat- ed, 2 mounted in balsam on a slide la- belled, ‘“‘Name Eretmocerus/californicus/ Howard/REMOUNT/Det HOWARD 19 1895/Coll D. W. Coquillett/No. USNM 2699/ Com" LECTOTYPE? "Ci" “toc. Tos Angeles/California/Date VI-9 19 1887/Host Aleyrodes/Det 19/On Quercus/agrifolia’’. There are an additional five 2 and 16 d paralectotypes mounted on 15 slides [USNM]. Diagnosis. Females of E. californicus can be distinguished by the length of the radi- cle, which is 0.6 or more than the length of the scape, the extremely reduced first fu- nicular segment, and the tapered shape of the club (Fig. 4). Eretmocerus californicus is most similar to E. eremicus and E. teja- nus, which differ in having the radicle only 0.4—0.5 the length of the scape, the first funicular segment is not as reduced, and the club is not tapered (Figs. 10, 26). Males of E. californicus lack pigment patterns on the mesoscutum, and the scape is 2.5-2.8 X as long as the pedicel. Eret- mocerus eremicus and E. tejanus differ by having distinct fuscous pigment patterns on the mesoscutum (Figs. 30, 33). Female.—Length and body color could not be accurately determined from available slide-mounted specimens. Wings hyaline. Face and occiput with transverse substri- gulate sculpture, interscrobal area vertically substrigulate. Antenna (Fig. 4) with radicle 5.4X as long as wide; scape 3.9X as long as wide, 1.5—1.6X length of radicle, 1.6X length of pedicel, 0.5 length of club; ped- icel 2.6X as long as wide, slightly shorter than radicle, 0.6X length of scape, 0.3 length of club. Funicle I triangular, 1.2 as wide as long and much smaller than funicle II; funicle II 1.8 as wide as long. Club with apex apically tapered, 5.2—6.6X as long as wide, 2X length of scape, 3.2 length of pedicel. Antennal ratio 1:1.5:1.0: Sole Mesoscutum trapezoidal with 6 setae and with reticulate sculpture in the anterior %, remainder with elongate reticulate sculpture laterally and substrigulate sculpture medi- ally. Parapsis with 2 setae and faint reticu- late to substrigulate sculpture; axilla with 1 seta and sculpture similar to parapsis. Scu- tellum with 4 setae and 2 placoid sensilla lateral to and close to posterior setae, lat- erally with elongate reticulate sculpture, re- mainder with elongate to substrigulate sculpture. Propodeum smooth. Endophrag- ma extending to posterior half of gastral ter- gite II. Forewing (Fig. 5) 2.7X as long as wide at width I, 2.4 as long as maximum width of disc. Longest anterior alary fringe 0.2 width of disc, longest posterior alary fringe 0.4X< width of disc. Single seta at base of wing present or absent; distal portion of costal cell with 2—5 setae. Marginal vein with 3 long setae, 10—12 setae usually be- tween marginal vein and linea calva. Linea VOLUME 99, NUMBER 1 4 / af SS oo " ~ N= mets ae tS ERR a aay ee 2 —~ ~ a ~ SOE BRS SN es & ~\ mae ane ~ Sim) ~ = in ee SS $a 4 Sa ~ \ ~~ > TES; = a ae ea YN NY ~ Va SS es ee aa eng eit CN We \ = ie e ed ae a ~\s\ ply Vw Sens See SE Sats \, EN E ee SAIN ee Teal aa! j — aN Ne Sade Sey J a? me oe , Ss Re ees Bae Say Nan ANG OD Coe ~*~ eet e\ VN S e ad Sis Z Nees Ry aoa SS by SS Proc | Sy ay pr 7 = eye aS eee —— x ie SEEN Set iy, ee db ee GA? mn _ ry, es CF rN a AEE Gi Si AE EB re ee — mer = - Fe eerie — ; Cee ee sz - = ——— ae DSI IE (OTN mn 1 a Vas eee, Gp am Nw ee oz vr 3 me 7 cd (if 4 EDN RE = ; { a a - dy Tne Kacey pe ee He / Ly hd “¢ Fi i Te aol = = eS ghia ihbee8t eS oe NS ee Mr 55 x Ly rr Sj eas ‘ Sear aS we ca Pfr sare = ee 72 re or eo 4 : — rm : OE as Sal a _ [eg ae rm ap %& A 4 LV wn oo Wo i —- pe te ean a, ee y, a a ~~~ 2S mea le ee \ yy S a = — 3 i, - apa UF ? r=) art Hi, oo erie PN Sa, Bente hwy, SOS GN erst a er MA v fr i¢ = — _ ~ 2 “7 Umeete Eyes ERS eS a2 ° = Fs $e ~ CN eta LU TMG oS ei oe - t — oy Sc SS 2 Figs. 4-9. 4-5, Eretmocerus californicus. 4, 2 antenna. 5, 2 forewing. 6—7, E. corni. 6, 2 antenna. 7, 2 forewing. 8-9, E. debachi. 8, 2 antenna. 9, 2 forewing. af = anterior alary fringe, pf = posterior alary fringe. calva closed posteriorly by setae, with 12— 15 tubercles on ventral surface of wing near posterior end of linea calva; a group of 31— 43 setae including those forming distal edge of linea calva point toward anterior margin of wing, remaining 171—175 setae in disc point to distal apex of wing. Ratio of sub- marginal:marginal:stigmal veins 2.8:1.1:1. Hind wing 6.9 as long as wide, with 7— 9 setae in center. Gastral tergite I covered with strong re- ticulate sculpture; lateral margins of gastral tergites imbricate with dense stippling; gas- tral tergites I-VI with paired setae as fol- lows: 1, 1, 2, 2-3, 2, 1. Syntergum with 4 setae. 8 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Ovipositor slightly exserted and equal to length of club, 3X length of pedicel, 1.9 length of scape, and 1.1 length of midtib- ia. Male.—Body color could not be accu- rately determined from available slide- mounted specimens. Mid- and hind tarsi I— III fuscous. Marginal and stigmal veins fus- cous, base of forewing and costal cell slightly fuscous. Host.—Reared from an unknown species of Aleyrodes. Discussion.—Howard’s original descrip- tion and figures of E. californicus were based on a series of female and male spec- imens reared from an undetermined Aley- rodes collected by D. W. Coquillett in Los Angeles, California, in 1887. Although Howard (1895) deposited type material in the USNM, and Dozier examined the ma- terial in 1932, Compere (1936) was unable to examine E. californicus and placed it in his key based on the original description. The type material was lost from about 1932 until 1992. In 1992, G. Evans found the missing slides of E. californicus at the USNM. The specimens on one slide (five female, eight male) were crumpled in dark orange balsam and not suitable for study. All the speci- mens on this slide have been individually remounted in balsam. G. Evans also re- mounted broken specimens, including one female, from two other slides labelled as USNM Type No. 2699. The label data for these two new slides, which list “‘Astero- diaspidis”’ as the host, do not agree with the label data from the other type slides. We consider this to be an error. Our figures and redescription are derived from the six remounted females and eight remounted males, and seven male speci- mens from a single original Howard slide. There are now a total of 16 slides bearing E. californicus specimens labelled with the original collection data and identified as USNM Type No. 2699. It is interesting to note that FE. californi- cus was reared from an unknown species of Aleyrodes on oak. Aleuroplatus, Tetraleu- rodes, and Trialeurodes all occur on Quer- cus in southern California. M. Rose and J. B. Woolley (TAMU) were unable to obtain Eretmocerus from numerous collections of whitefly, particularly Tetraleurodes stanfor- di (Bemis), from oaks in Los Angeles County over the past 15 years. Lack of accurate characterization of E. californicus has resulted in misapplication of this name, particularly to species of Er- etmocerus reared from Bemisia (tabaci complex) in the U.S. We have not examined any specimens of E. californicus reared from Bemisia (tabaci complex). Eretmocerus corni Haldeman (Figs. 6-7) 1850: Eretmocerus corni Haldeman, 110 [USNM, examined]. Type Material—Neotype ¢ mounted in balsam on a slide labelled. ““Eretmocerus corni Hald./Reared from white-/fly, Aster- ochiton/sp. on Impatients/biflora./White Clay Creek/Pa. Sept. 16, 1929/H.L Dozier” ‘“‘Eretmocerus/corni Hald./Designated as/this long-lost/species./2 + ¢ Neotype/U.S.N.M.” Diagnosis.—Females of Eretmocerus corni can be distinguished by the combi- nation of the elongate club that is 5.9-7X as long as wide and the relatively short ped- icel, which is only 2.4—3.1X as long as wide, and 0.25—0.29X, rarely 0.30X, as long as the club (Fig. 6). Eretmocerus corni is similar to E. eremicus and E. tejanus, which differ in having the pedicel 3—4X as Jong as wide and 0.28—0.39% as long as the club (Figs. 10, 26). Eretmocerus haldemani also has an elongate club, but it is 7.5-8.3X as long as wide (Fig. 16). Males of Eretmocerus corni have the me- soscutum and scutellum fuscous, and the hind tibia is not fuscous. They are most similar to males of E. eremicus and E. te- janus. Males of E. eremicus differ in having the hind tibia fuscous, and males of E. te- janus can be distinguished by the longitu- VOLUME 99, NUMBER 1 dinal fuscous bands on the mesoscutum (Fig. 33). Female.—Length of specimens in Hoy- er’s 0.47—0.55 mm. Specimens in Hoyer’s with body and antennae pale yellow. Head amber; eyes amber with inner red pigment, ocelli pale. Wings hyaline. Face and occiput with transverse substri- gulate sculpture, interscrobal area vertically substrigulate. Antenna (Fig. 6) with radicle 4x as long as wide; scape 4.9 as long as wide and 2.1X length of radicle, 2.2X length of pedicel, 0.4 length of club; ped- icel 2.4—3.1X as long as wide, 0.4 length of scape, 0.25—0.29X, and rarely 0.30, length of club. Funicle I triangular, 1.25 as wide as long; funicle II subquadrate. Club with apex truncate, 5.9—7X as long as wide, 2.5 length of scape, 4.9 length of pedicel. Antennal ratio 1:2.1:1.6:4.7. Mesoscutum trapezoidal with 6 setae and reticulate sculpture anteriorly, remainder with elongate reticulate sculpture. Parapsis with 2 setae and reticulate to substrigulate sculpture; axilla with | seta and reticulate to substrigulate sculpture. Scutellum with 4 setae and 2 placoid sensilla lateral to and slightly closer to posterior setae than to an- terior setae, with fine substrigulate sculpture medially, remainder with reticulate sculp- ture. Propodeum medially reticulate, faintly reticulate on lateral margins. Endophragma extending nearly to posterior margin of gas- tral tergite IV. Forewing (Fig. 7) 2.9X as long as wide at width I, 2.7 as long as maximum width of disc. Longest anterior alary fringe 0.16 width of disc; longest posterior alary fringe 0.3X width of disc. Single seta at base of wing present or absent; distal portion of costal cell usually with 3, occasionally 2— 5, setae. Marginal vein with 3 long setae; 10-11 setae, occasionally 10—15, between marginal vein and linea calva. Linea calva closed posteriorly by setae, with 12—16 tu- bercles on ventral surface of wing near pos- terior end of linea calva; a group of 32—45 setae including those forming distal edge of linea calva point toward anterior margin of wing, remaining 183-238 setae in disc point to distal apex of wing. Ratio of sub marginal:marginal:stigmal veins 3:1.3:1. Hind wing 7.5X as long as wide, with 7— 15 setae in center. Gastral tergite I with faint transverse sub- strigulate sculpture medially and reticulate sculpture on anterior half of lateral margins; lateral margins of tergites II-VI imbricate with stippling; tergites I-VI usually with paired setae as follows: 1, 1, 2, 2, 2, 1, but occasionally 1, 1—3, 2—3, 2—3, 2, 1. Synter- gum with 4 setae. Ovipositor slightly exserted, 0.8 length of club, 3.4 length of pedicel, 1.8 length of scape, equal in length to midtibia. Male.—Specimens mounted in balsam with head orange. Apex of scape, pedicel, and multiporous plate sensilla fuscous. Pro- notum fuscous. Mesoscutum fuscous, darkest along anterior margin and becoming paler posteriorly; tegula slightly fuscous. Scutellum fuscous, darkest along anterior and posterior margins. Metanotum and pro- podeum slightly fuscous. Aedeagus slightly fuscous. Fore- and hind tarsi slightly fus- cous; distal portion of midtibia and midtarsi I-III fuscous, midtarsus IV less fuscous. Submarginal vein, costal cell, and edges of marginal and stigmal veins fuscous; base of forewing and venation of hind wing slightly fuscous. Host.—Reared from Trialeurodes pack- ardi (Morrill), the strawberry whitefly. Discussion.—Haldeman’s (1850) original description and figure of the type species, E. corni, were based on “‘Two mutilated specimens”’ reared from Tetraleurodes cor- ni (Haldeman) [= Aleurodes corni Halde- man]. Unfortunately, Haldeman did not designate a depository for his specimens. Dozier (1932), who was unable to locate the original material, provided a redescrip- tion of the species and designated a neotype female based on a series of specimens (15 2, 3 3) reared from Trialeurodes packardi (Morrill) [as 7. morrilli (Britton)] and an additional field-collected female. The slide labelled as the neotype lists 10 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ‘‘Asterochiton”’ as the host, but Dozier only mentions Trialeurodes morrilli as a host in the redescription. Mound and Halsey (1978) do not list any species of Asterochi- ton in the U.S. Dozier’s original identifica- tion of the host is probably an error. Al- though Dozier stated the slide bearing the neotype female also contained one addition- al female and one male specimen, the slide with the red USNM type label only has a single female and single male. Despite these discrepancies, we accept this slide with the red USNM type label as the one with the neotype. Eretmocerus corni did not successfully reproduce on Bemisia (tabaci complex) in limited trials at TAMU. It may prove to be important in the biological control of spe- cies of Trialeurodes. Other material examined.—USA: Mary- land: Blackwater Refuge, reared from 7. packardi on wild strawberry, 19.x, 24, 26.x.1942, H. L. Dozier (1 2, 2 6, USNM); Cambridge, 24, 26, 30.vili., 9.1x.1943, Ex. Trialeurodes on Euphorbia hirsuta, H. L. Dozier (1 2, 6 6, USNM); Church Creek, Ex. Trialeurodes on Lizard’s Tail, 2, Axe 94). sit OE tad SINM): = Church Greek:) Ex, Lizard?s: Vail}, 101942) (io USNM); New York: Ithaca, Cornell Plan- tations, 29.ix.1992, Trialeurodes packardi on Impatients pallida, T92047, G. W. Fer- rentino (8 2, 4 ¢, MJR); Ithaca, S. of Bee- be Lake, 10.viii.1988, ex. Trialeurodes packardi, M. Hunter (1 6, MJR). Eretmocerus debachi Rose and Rosen (Figs. 8-9, 29) Eretmocerus debachi Rose and Rosen, 1991-1992: 200 [USNM, examined]. Type material.—Holotype 2 mounted in balsam on a slide labelled, “‘Name Eret- mocerus &/debachi/Holotype sp.nov./Bal- sam/Det Rose 1982/Coll M. Rose/No. B2.76/Corr. USNM/Rose&Rosen 91-92” ‘‘Loc Rancho Mission/Viejo, Orange Co., Cal./Date VII.15 1982/Host Parabemisia/ myricae/Det Rose 1982/On Valencia or- ange/and lemons’”’. There are five 2 and one d paratypes with the same label data as the holotype. These are deposited with the BMNH, HU, MJR, TAMU, and USNM. Diagnosis.—Females of E. debachi can be distinguished by the deflexed apex of the club (Fig. 8), mesoscutum with 4 setae, and each parapsis with 2 setae. It is most similar to E. furuhashii, which differs in having a longer club (Fig. 13) and 3 setae on each parapsis (Fig. 15). Eretmocerus joeballi can have the apex of the club slightly deflexed (Fig. 18) but differs in having 6 setae on the mesoscutum. Males of E. debachi have each parapsis with 2 setae, a fuscous ““T’’ shape on the mesoscutum, and the scutellum is com- pletely fuscous (Fig. 29). Males of E. fu- ruhashii differ in having each parapsis with 3 setae, a very light fuscous ““T”’ shape on the mesoscutum, and the lateral areas of the scutellum are unpigmented (Fig. 31). Host.—Reared from Parabemisia myri- cae (Kuwana), the bayberry whitefly. Discussion.—The description of E. de- bachi was based on series of specimens reared from Parabemisia myricae collected on citrus in southern California. The tax- onomy, biology and history of E. debachi in California is well documented (Rose and DeBach 1991-1992, Rose and Rosen 1991— 92, Rose and Zolnerowich 1994). This spe- cies has been introduced and established in Israel and Turkey, where it is responsible for control of P. myricae. Introductions are planned for Spain and France (J. C. Onil- lon, personal communication) and Morocco (D. Rosen, personal communication). Eretmocerus eremicus Rose and Zolnerowich, new species (Figs. 10—12, 30) Diagnosis.—Females of E. eremicus can be distinguished by the club that is 6.5— 7.3X as long as wide (Fig. 10), pedicel that is 3—4X as long as wide and 0.3—0.39X as VOLUME 99, NUMBER 1 Figs. 10-12. long as the club, mesoscutum with 6 setae, and gastral tergite II usually with 1 pair, but occasionally 2 pairs, of lateral setae (Fig. 12): Females of E. eremicus are very similar to E. corni and E. tejanus. Females of E. corni have the pedicel 2.4—3.1 as long as wide and 0.25—0.30 as long as the club, and have not been reliably recorded from Bemisia (tabaci complex). Eretmocerus te- janus has the anterior alary fringe of the forewing slightly shorter (Fig. 27), and gas- tral tergite II usually has 2 pairs, but oc- casionally 1 pair, of lateral setae (Fig. 28). Males of E. eremicus have the meso- scutum fuscous along the anterior margin (Fig. 30), or with a light fuscous ““T”’ shape (as in Fig. 31), or evenly fuscous, and the hind tibia is fuscous. Males of E. corni do not have the hind tibia fuscous, and males of E. tejanus can be distinguished by the SS SS SSN SS Sey > ~~ Si “N EN oe ‘N ~ SSS Sore os SSE SR SSSR VX ERR RS RES TSS NR 4, — Soa TR Were \ 8S Se St eS es? Wheeler (lc: UTA); Saddlebunch Keys, 29 Dec 1953, H. V. Weems (1 ¢, 1 2; USNM). North Car- olina. Onalow Co., Ashe Island, 11 Aug 1975, J Cy Dukes “(1"2>-USNM)- Diphuia nitida. 1, Sth tergum and sternum, lateral view. 2, 5th tergum, dorsal view. 3, Male Distribution.—Nearctic: USA (FL, NC, NY). Neotropical: Bahamas, Belize, Ber- muda, West Indies (Dominican Republic, Grand Cayman, Jamaica). Natural history—The vast majority of specimens from the Belizean cays were col- lected by sweeping just above mangrove peat that is well shaded most of the day. A VOLUME 99, NUMBER | few specimens, apparently feeding, were collected on flowers. The association with mangrove peat must be opportunistic, as the species occurs in areas where mangrove does not now exist. Diagnosis.—This species is distinguished from congeners, especially D. anomala and D. zatwarnickii, by the sparsely microto- mentose mesofrons; the subshiny mesono- tum that is very sparsely invested with fine, golden brown microtomentum; the sparsely, whitish gray microtomentose anepisternum; the shiny anterior half of the katepisternum; and several characters of the male termi- nalia (see description and figures). Remarks.—Since 1989, when the re- search from my earlier revision (Mathis 1990) was completed, I have studied longer series of specimens from additional locali- ties. These studies indicate that D. nasalis and D. nitida are conspecific, with the latter being the senior synonym. Although no more specimens from the type locality (New York: Long Island, Douglaston) of D. nitida have become available, I now strong- ly suspect that the difference noted earlier between the nominate species has mostly to do with the poor condition of the holotype female of D. nitida. This specimen, which is the entirety of the type series, appears to have been “‘rubbed”’ (antennae and several setae are missing), which could also ac- count for the shinier frons, the only distin- guishing character. Diphuia flinti Mathis, new species (Figs. 9-16) Description.—Small to moderately small shore flies, length 1.60 to 2.10 mm. Head: Frons sparsely and uniformly in- vested with fine brownish to grayish micro- tomentum. Ventral portion of face with me- dian, vertical, densely microtomentose, sil- very white stripe extended dorsad from and connected to microtomentose ventral mar- gin. Thorax: Mesonotum, including postpro- notum and notopleuron, sparsely invested 33 with fine brownish to golden brown micro- tomentum, mostly subshiny; anepisternum, anepimeron, and posterior half of katepi- sternum sparsely invested with fine, whitish gray microtomentum; anterior half of kat- episternum with oval spot bare of micro- tomentum, shiny. Wing with costal vein ra- tio 0.58; M vein ratio 0.40. Abdomen: 5th tergum with anterior mar- gin emarginate medially (Fig. 16); Sth ster- num a narrow arch that is connected ante- riorly with 5th tergum (Fig. 11), apex with 2 flanges (Fig. 12). Male terminalia (Figs. 9—10, 13-15) as follows: epandrium higher than wide in lateral view, ventral margin pointed (Fig. 10); surstylus longer than cer- cus, obtusely angulate in lateral view, apex truncate and pointed medially and anteri- orly (Figs. 9-10); gonite in lateral view with basal half subrectangular, with narrow, sinuous, pointed process apically (Figs. 13— 14); aedeagal apodeme in lateral view ir- regularly triangular (Fig. 13); aedeagus rounded apically in lateral view (Fig. 13), pointed (not bifurcate) in dorsal view (Fig. 14); hypandrium in ventral view wider than long, anterior margin deeply arched with medial margins recurved, posterior margin evenly arched except for a short, median process (Fig. 15). Type material——The holotype ¢ is la- beled ““DOMINICAN REPUB. El Seibo: near Rincon[,] 18°45.3'N, 68°55.7’W{[,] 12 May 1995, WNMathis/USNM ENT 00136995 [bar code label].”’ The allotype 2 and 14 paratypes (10 6, 4 2) bear the same locality label as the holotype. Other paratypes are as follows: DOMINICAN RE- PUBLIC. “EV Setbo> EV Seibo"G@* kim E; 18°44.73'N, 68°59.2'W; 120 m), 12 May 1995, W.N. Mathis (1 ¢,5 2; USNM). The holotype is double mounted (minuten in sil- icon plastic block), is in excellent condi- tion, and is deposited in the USNM. Distribution.—Neotropical. Dominican Republic (El Seibo). Diagnosis.—I can distinguish this species only on the basis of structures of the male terminalia, especially the shape of the 5th 34 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON SORES CEES S37 rhe S vA 9 Figs. 9-16. Diphuia flinti. 9, Epandrium, cerci, and surstyli, posterior view. 10, Same, lateral view. 11, 5th tergum and sternum, lateral view. 12, Apex of Sth sternum, ventral view. 13. Gonite, aedeagus, and aedeagal apodeme, lateral view. 14, Left gonite and aedeagus, dorsal view. 15, Hypandrium, ventral view. 16, 5th tergum, dorsal view. Scale = 0.1 mm. tergum, surstylus, gonite, aedeagus, and hy- pandrium as described and illustrated. Natural history.—The type series was collected along the banks of the Rio Qui- sibani, which is a freshwater stream. Etymology.—It is a pleasure to name this species after Dr. Oliver S. Flint, Jr, who accompanied me on the field work that re- sulted in the collection of this species. PHYLOGENETIC CONSIDERATIONS Diphuia is in the tribe Hecamedini (Mathis 1991a, 1991b, 1993) and belongs to the clade including Hecamede Haliday and Allotrichoma Becker, sensu lato, in- cluding Pseudohecamede Hendel (Mathis 1991la). This relationship is evidenced by the retracted fifth tergum of the male, which is moderately elongate, almost tubular. Di- phuia may be the sister group to the rest of the Allotrichoma clade, following Cresson’s (1944) original assessment, although the evidence supporting this relationship is not compelling, and Diphuia could be mostly closely related to Hecamede Haliday. The monophyly of Diphuia within He- camedini is well established (see generic di- agnosis for a partial list of synapomor- phies), and the remainder of this section will be devoted to the phylogenetic rela- tionships among the species. First the char- acter evidence is presented, followed by a brief discussion that is intended to comple- ment the cladogram. Character evidence.—The characters used in the cladistic analysis are listed be- VOLUME 99, NUMBER 1 Table 1. analysis of Diphuia. Matrix of characters used in the cladistic Characters Taxa 1 2 Sy 4.5 6 7 8 9 Outgroug ORO OO. tO) 105 OR SOR 0) D. anomala 1 0 1 2 | | 1 i] 1 D. zatwarnickii 1 (0) 1 2 1 1 1 1 1 D. nitida 0 1 0 0 10) 10) 0 0 0 D. flinti Oe 05. 10 1 1 1 1 1 0 low and the distribution of their states is shown in the matrix (Table 1) and discussed here. For each character, the plesiomorphic condition is indicated by a O, and the rela- tive apomorphic condition(s) is indicated by al or 2. — . Vestiture of mesonotum: (0) sparsely mi- crotomentose, subshiny; (1) moderately densely microtomentose. . Vestiture of anepisternum: (0) sparsely, uniformly microtomentose; (1) antero- ventral % to % bare, shiny. . Length of 5th tergum of male: (0) longer than wide (Figs. 1—2, 11, 16); (1) wider than long (Figs. 1—2, 16—17 in Mathis 1990). . Shape of anterior margin of 5th tergum of male: (O) straight, even; (1) shallowly emarginate; (2) deeply emarginate, V-shaped (Figs. 2, 17 in Mathis 1990). Shape of ventral margin of epandrium: (O) broadly formed, somewhat truncate; (1) tapered to point. Shape of surstylus: (0) rounded; (1) truncate ventrally, pointed. Shape of gonite: (0) trapezoidal with sides more or less equal (Fig. 4); (1) somewhat rectangular, longer than high (Bias 13): . Shape of posterior margin of hypan- drium: (0) smooth and even; (1) with a small, median projection. Nw eS) & fo) SN =a (oe) 9. Shape of aedeagus: (0) slender, 3—4> longer than high in lateral view; ( more robust, length in lateral view about twice height (Figs. 7, 21 in Mathis 1990). 1) Discussion.—Analysis of the character evidence (Table 2), all morphological and primarily from the male terminalia, resulted in a single cladogram (Fig. 17). The single tree resulted from the implicit enumeration (ie*) option of Hennig86 and has a length of 10 steps and consistency and retention indices of 1.00 and 1.00, respectively. The analysis of characters is given in Table 2. The cladogram (Fig. 17) for the four spe- cies consists of three bifurcations, with D. nitida at the base and the remaining species as the sister group. The monophyly of the latter assemblage is established by charac- ters four (1) and five through eight. The next bifurcation has D. flinti, which 1s known thus far only from the Dominican Republic, branching off and as the sister group to D. anomala and D. zatwarnickii. The monophyly of the D. anomala and D. zatwarnickii group is indicated by four characters (1, 3;:-4(2), 9): ACKNOWLEDGMENTS I am grateful for the assistance in the field from Oliver S. Flint, Jr. (USNM) and Kelvin A. Guerrero (Museo Nacional de Historia Natural, Santo Domingo). For the opportunity to examine specimens housed in their collections, I thank David Grimaldi and Julian Stark (AMNH) and Donald Azu- ma (ANSP). For critically reviewing a draft of this paper, I thank Allen L. Norrbom. The illustrations were skillfully inked by Elaine R.S. Hodges. Funding for this research project, es- Table 2. Analysis of characters used in the cladistic analysis. Character: 1 2 3 Steps: 1 2 Consistency Index: 100 100 100. Retention Index: 100 100 100 4 5 6 if 8 9 | 1 1 1 1 | 100 100. 100 100 100 100 100 100 100 100 100 100 36 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON outgroup (2) Fig. 17. consistency index 1.00, retention index 1.00. pecially the field work in Belize and the Dominican Republic, was provided by the Caribbean Coral Reef Ecosystems (CCRE) and Biodiversity Program (Biological Sur- veys and Inventories, BSI), National Mu- seum of Natural History, Smithsonian In- stitution. This is contribution number 501 of the CCRE project, which is partially sup- ported by a grant from the Exxon Corpo- ration. LITERATURE CITED Cresson, E. T., Jr. 1944. Descriptions of new genera and species of the dipterous family Ephydridae. Paper XIV. Notulae Naturae. The Academy of Natural Sciences of Philadelphia 135: 1-9. . 1946. A systematic annotated arrangement of the genera and species of the Neotropical Ephyd- ridae (Diptera). 1. The subfamily Psilopinae. Transactions of the American Entomological So- ciety 71: 129-163. Griffiths, G. C. D. 1972. The Phylogenetic Classifi- cation of Diptera Cyclorrhapha with Special Ref- erence to the Structure of the Male Postabdomen. Series Entomologica 8: 1-340. W. Junk, Hague. Mathis, W. N. 1986. Studies of Psilopinae (Diptera: Ephydridae), I: A revision of the shore fly genus Placopsidella Kertész. Smithsonian Contributions to Zoology 430: 1-30. . 1990. A revision of the shore-fly genus Di- phuia Cresson (Diptera: Ephydridae). Proceedings of the Entomological Society of Washington 92(4): 746-756, 22 figures. 1991a. Studies of Gymnomyzinae (Diptera: Ephydridae), Il: A revision of the shore-fly sub- asaya Cladogram depicting hypothetical relationships among species of Diphuia. Tree length 10 steps, D. nitida D. flinti D. anomala D. zatwarnickiil genus Pseudohecamede Hendel of the genus AI- lotrichoma Becker. Smithsonian Contributions to Zoology 522: 28+iii pp. . 1991b. Classification of the shore flies (Dip- tera: Ephydridae), past, present, and future, pp. 209-227. In Weismann, L., et al., eds., Proceed- ings of the Second International Congress of Dip- terology. VEDA, Bratislava, SPB Academic Pub- lishing, 368 pp. The Hague. 1993. Studies of Gymnomyzinae (Diptera: Ephydridae), IV: A revision of the shore-fly genus Hecamede Haliday. Smithsonian Contributions to Zoology 541: 46+iu1 pp. Mathis, W. N. and T. Zatwarnicki. 1995. A world cat- alog of the shore flies (Diptera: Ephydridae). Memoirs on Entomology, International 4: vi+423 Pp. McAlpine, J. E 1981. Morphology and terminology- adults, pp. 9-63. In McAlpine, J. F, et al., eds., Manual of Nearctic Diptera. Ottawa. [Volume | is Monograph 27 of Research Branch Agriculture Canada. | Sturtevant, A. H. and M. R. Wheeler. 1954. Synopses of Nearctic Ephydridae (Diptera). Transactions of the American Entomological Society 79: 151— PSM fe Wirth, W. W. 1956. The Ephydridae (Diptera) of the Bahama Islands. American Museum Novitates 1817: 1-20. 1965. Family Ephydridae, pp. 734-759. In Stone, A., et al., eds., A catalog of the Diptera of America north of Mexico. U.S. Department of Ag- riculture, Agriculture Handbook 276. 1968. 77. Family Ephydridae, pp. 1—43. In Papavero, N., ed., A catalogue of the Diptera of the Americas south of the United States. Depar- tamento de Zoologia, Secretaria de Agricultura, Sao Paulo. PROC. ENTOMOL. SOC. WASH. 99(1), 1997, pp. 37—41 A NEW SPECIES OF DRYMUS FIEBER FROM MEXICO, WITH A KEY TO SPECIES AND A CHECKLIST OF WESTERN HEMISPHERE DRYMINI! (HEMIPTERA: LYGAEIDAE) JAMES A. SLATER AND HARRY BRAILOVSKY (JAS) Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs, CT 06269, U.S.A; (HB) Departamento de Zoologia, Instituto de Biologia, UNAM, Apartado Postal No. 70-153, Mexico 04510, D. FE, Mexico. Abstract.—The first Mexican species of Drymus Fieber, is described as Drymus mex- icanus, n. sp., from the mountains of the state of Veracruz. A key to the Western Hemi- sphere species is included together with a list of the Western Hemisphere species of the tribe Drymini. A discussion of the distribution of the tribe is given and the affinities of the Nearctic fauna analyzed. Key Words: The zoogeography of the Rhyparochrom- inae was reviewed in detail by Slater (1986). In that paper he noted the interest- ing distribution of the members of the tribe Drymini and how closely the overall distri- bution paralleled that of the tribe Rhypar- ochromini. Both of these tribes are abun- dant, diverse and speciose in the Palearctic, Ethiopian-Oriental (= Paleotropical) and Australian regions. However, both make up only a very small proportion of the Nearctic rhyparochromine fauna and both are absent from the the true Neotropical Region (in- cluding the West Indies). Of the Nearctic fauna most species belong to genera that are also found in the Palearctic. The Drymini tend to be associated with woodland margins and to live in moderately mesic habitats. The Rhyparochromini occur for the most part in relatively dry savannah- like habitats. The absence from the Neotropics of Drymini, despite their abundance in Afri- ca, suggests a radiation subsequent to 90 million years b.p. or subsequent to the Drymus, Mexico, biogeography, distribution, checklist breakup of West Gondwanaland. It also suggests that despite the abundance of the Drymini in Australia they are relatively re- cent elements in that fauna since there was apparently no movement of such a fauna across Antarctica (and because there are no native species in New Zealand). The most plausible scenario for the occurrence of Drymini in the Western Hemisphere ap- pears to be as a Holarctic element that was present during, or before, the onset of the Pleistocene. If this is true, it means that two taxa have been able to reach generic status during this period (both of which are confined to areas of California that escaped glaciation). The Mexican drymine fauna is found in montane areas and all species are members of genera also found in the United States and Canada as well as in the Palearctic. Drymus is the most unusual of these in that the two previously known species are known from only a few localities in the western United States. All measurements are in millimeters. 38 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Drymus mexicanus, Slater and Brailovsky, new species (Figs. 1, 2) Robust, subelliptical. Head, anterior pronotal lobe, scutellum and antennae dark chocolate brown, almost black. Explanate pronotal margins, posterior pronotal lobe, entire hemelytron, legs and labium dull red- dish-brown. Fourth antennal segment with apical two-thirds white. Coarsely punctate Over entire surface except wing membrane, OR Re ~~ Sai Sey iy, Drymus mexicanus. Dorsal view, head and pronotum. the punctures on head and anterior pronotal lobe relatively small. Head acuminate, non-declivent; tylus at- taining distal one third of first antennal seg- ment; vertex convex. Eyes set well away from anterior margin of pronotum. Length head 0.84, width 1.00, interocular space 0.82. Pronotal calli confluent or nearly so, mesally, elevated above surface of posterior lobe. Lateral pronotal margins broadly ex- planate (Fig. 1); posterior margin of pro- notum shallowly concave. Length prono- VOLUME 99, NUMBER I Fig. 2. Drymus mexicanus. Abdomen, lateral view. tum 1.20, width 2.00. Scutellum with a me- dian elevation distally, mesally concave on basal half. Length scutellum 1.12, width 1.20. Hemelytra with lateral margins broad- ly convex. Clavus with four rows of punc- tures. Length claval commissure 0.60. Mid- line distance apex clavus-apex corium 1.24. Midline distance apex corium-apex mem- brane 0.64. Metathoracic scent gland auri- cle curved posteriorly. Evaporative area only slightly exceeding end of auricle, dis- tally truncate, covering only mesal one- third to one-fourth of metapleuron. Fore fe- mur only moderately incrassate, armed be- low on distal one fourth with a short sharp spine with three minute spines distad. La- bium extending to metacoxae, first segment not attaining base of head. Length labial segments I 0.62, II 0.70, III 0.54, [IV 0.44 (approx.). Length antennal segments I 0.50, II 0.80, TT 0.62, [V 0.68. Total body length 5:65. All abdominal spiracles located below sternal shelf. Posterior pair of trichobothria on sternum four located dorsoventrally, the ventral trichobothrium reduced and obso- lete (Fig: 2): Holotype 2 MEXICO: Veracruz: Na- 39 olinco, 1.VIHI.1977, 2100 m. In Universi- dad Nacional Autonoma de Mexico collec- tion. This is a very distinctive species at once separable from the other Western Hemi- sphere species by the white distal portion of the fourth antennal segment, the much more strongly explanate lateral pronotal margins, and the shorter antennae. Actually this is not the only species of Drymus with a pale distal end to the fourth antennal seg- ment; several Palearctic species have a somewhat paler apex on segment four, al- though none that we have examined have a white terminal end to the antennae. KEY TO WESTERN HEMISPHERE SPECIES OF DRYMUS 1. Distal half of fourth antennal segment white, strongly contrasting with dark coloration of ba- sal half of antennae; explanate margins of pro- notum broad, subequal to width of second an- tennal segment; second antennal segment sub- equal in length to length of head Ee Cenc ee Ia eer et oot ae en mexicanus N. sp. — Fourth antennal segment unicolorous dark brown to black; explanate lateral pronotal mar- gins relatively narrow, much narrower than width of second antennal segment; second an- 40 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tennal segment considerably longer than length Ofte ad! Oe spe Fee os hee) aon die eee 72 . Large (6.5—7 mm); very dark almost black; an- terior and posterior pronotal lobes nearly even- ly punctate; expanded lateral margins of pro- notum concolorous with doraal surface of pro- NOUN seh Se EG oes eee crassus Van Duzee — Smaller (rarely over 5 mm); dull brown; ante- rior pronotal lobe more finely punctate than posterior lobe; expanded lateral margins of pronotum usually slightly paler than surface of PLONOtUM: «5 0% Ges. ges eenate siete unus (Say) i) CHECKLIST OF WESTERN HEMISPHERE DRYMINI See Slater 1964 and Slater and O’Don- nell 1995 for complete references. Drymus Fieber crassus Van Duzee 1910. New England south to Florida, west to Texas and South Dakota. mexicanus, n. sp. Mexico. unus (Say) 1831. Eastern Canada, south to North Carolina, west to Colorado. Eremocoris Fieber arnaudi Brailovsky 1982. Mexico. borealis (Dallas) 1852. Western U.S.A. canadensis Walley 1929. British Columbia; Idaho. chalmaensis Brailovsky and Barrera 1981. Mexico. cupressicola Ashlock 1979. California. depressus Barber 1928. New England south to Florida and Louisiana coastal areas. dimidiatus Van Duzee 1921. Colorado. extremus Brailovsky and Cervantes 1989. Mexico. ferus (Say) 1832. Widespread over much of U.S.A. and southern Canada. garciai Brailovsky and Barrera 1981. Mex- ico. guerrerensis Brailovsky and Barrera 1981. Mexico. inquilinus Van Duzee 1914. California. legionarius Brailovsky and Barrera 1981. Mexico. lopezformenti Brailovsky and Barrera 1981. Mexico. melanotus Walley 1929. British Columbia; Idaho. mimbresianus Brailovsky and Cervantes 1989. Mexico. obscurus Van Duzee 1906. California; Ida- ho; British Columbia. opacus Van Duzee 1921. California. semicinctus Van Duzee 1921. California; Idaho. setosus Blatchley 1926. Eastern U.S.A. squalidus Brailovsky and Barrera 1981. Mexico. Gastrodes Westwood arizonensis Usinger 1938. Arizona. conicolus Usinger 1933. California. intermedius Usinger 1938. British Colum- bia. pacificus (Provancher) 1886. Nebraska and Colorado, west to Calfornia and British Columbia. walleyi Usinger 1938. Ontario, British Co- lumbia. Scolopostethus Fieber atlanticus Horvath 1893. Northern U.S.A. and southern Canada; Newfoundland to British Columbia south to New Jersey and New Mexico. diffidens Horvath 1893. Northern U.S.A. and southern Canada: Nova Scotia to British Columbia south to New Jersey. pacificus Barber 1918. California, Idaho, British Columbia. thomsoni Reuter 1874. Almost throughout Holarctic. tropicus (Distant) 1882. California to Gua- temala. Thylochromus Barber nitidulus Barber 1928. California. Togodolentus Barber wrighti (Van Duzee) 1914. California. ACKNOWLEDGMENTS We express our appreciation to Mr. Er- nesto Barrera (UNAM) Mexico for prepa- ration of the illustrations and to Dr. Jane VOLUME 99, NUMBER 1 O’Donnell, University of Connecticut, for assistance with the manuscript. LITERATURE CITED Slater, J. A. 1964. A Catalogue of the Lygaeidae of the World. 2 Volumes. University of Connecticut. Storrs. 1688 pp. 4] . 1986. A synopsis of the Zoogeography of the Rhyparochrominae (Heteropteraa: L) Journal of the New York Entomological Society 94: 262-280. Slater, J. A. and J. E. O'Donnell. 1995. A Catalogue of the Lygaeidae of the World (1960-1994). New York Entomological Society, 410 pp. gaeidae). PROC. ENTOMOL. SOC. WASH. 99(1), 1997, pp. 42—49 NESTING BEHAVIOR OF KROMBEINICTUS NORDENAE LECLERCQ, A SPHECID WASP WITH VEGETARIAN LARVAE (HYMENOPTERA: SPHECIDAE: CRABRONINAE) KARL V. KROMBEIN AND BETH B. NORDEN Department of Entomology, National Museum of Natural History, Smithsonian Insti- tution, MRC 165, Washington, DC 20560, U.S.A. Abstract.—Nesting behavior of the recently described Sri Lankan wasp, Krombeinictus nordenae Leclercq, is discussed. Females nest in the hollow internodes of the leguminous myrmecophyte, Humboldtia laurifolia Vah\. The biology of this stem-nesting crabronine iS unique among Sphecidae in several aspects. An adult female exhibits remarkable ma- ternal care, rearing one larva at a time, and feeding it progressively. Progressive provi- sioning has not been noted previously for any Crabroninae. Nests lack cell partitions and mature larvae are transported to the basal regions of their stems for cocoon spinning. The cocoon is also unlike that of any other known crabronine species, exhibiting adaptations to internode morphology and allowing movement of adults within the nest cavity. Finally, K. nordenae is remarkably different from all other known Sphecidae in feeding pollen rather than paralyzed arthropod prey to its larvae. Key Words: cocoon, pollen Sri Lanka has been called the land of ser- endipity. A recent example is the amazing nesting behavior of a Ceylonese wasp, a newly described genus and species, Krom- beinictus nordenae Leclercq (1996). This pretty little wasp (Fig. 1), 5—6 mm long, has creamy to pale yellow markings on its black head and thorax, and a mostly light red ab- domen bearing narrow, transverse, brown to black stripes on some of the dorsal seg- ments. Krombeinictus belongs to the Sphe- cidae, normally a family of predaceous, mostly solitary wasps. MATERIALS AND METHODS Our first encounter with this species was when we received a single male of K. nor- denae from a colleague, Prof. Fred R. Rick- son. It was among a few wasps that Sphecidae, Crabroninae, Humboldtia, Sri Lanka, internode, myrmecophyte, emerged from a dozen internodes of the myrmecophyte, Humboldtia laurifolia Vahl, that he collected in the Sinharaja Forest Re- serve in Sri Lanka in 1992. We visited Sri Lanka in 1993, and spent five days (18-20 Jul and 2—3 Aug) in the rainforest near Gilimale, Ratnapura District, 06°46'N, 80°26’E. We hoped to make be- havioral observations on Krombeinictus during this brief period, but intermittent rains of the delayed monsoon season pre- cluded nesting activity by the wasps. How- ever, we censused about a thousand inter- nodes from H. laurifolia, and placed several hundred unopened stems directly into al- cohol for subsequent study. These inter- nodes were split open carefully in the lab- oratory to avoid damaging associated or- ganisms, and notes were made on their con- tents. VOLUME 99, NUMBER | 43 Figs. 1-2. Humboldtia laurifolia Vahl (Figs. 3-6, 12—15) This botanical section is condensed from a detailed account of the plant’s morphol- ogy in Krombein et al. (in prep.). Hum- boldtia laurifolia (Fabaceae) (Fig. 3) occurs only in Sri Lanka; three other species of the genus and one variety occur in southern In- dia. Humboldtia laurifolia is a small under- story tree, growing to a height of about 10 m, found in the lowland rainforest of the southwestern quadrant of Sri Lanka. It usu- ally grows along streams or in seeps, is highly gregarious, and occurs in groups of ten to several hundred trees. Humboldtia laurifolia and two of its In- dian congeners, H. brunonis Wallich and H. decurrens Beddome ex Oliver, are of par- ticular interest to naturalists because they are myrmecophytes that have coevolved with several species of ants. The trees pro- vide swollen internodes, each with a self- opening entrance, that serve as domatia for the ants. The trees also provide an abun- dance of extrafloral nectaries on leaves, stipules and inflorescences whose secre- tions are attractive to the ants. In turn, the ants protect the foliage, especially the tender, young leaves, from herbivory. Humboldtia laurifolia is typical of leg- umes in having pinnately compound leaves, each leaf having four to six pairs of oppo- Krombeinictus nordenae, female. 1, Lateral view. 2, Frontal view of head. site leaflets (Fig. 4). As in many legumes there is a developmental period during which a flush of four or five new inter- nodes, each with an associated leaf, is pro- duced over a period of two months. As the internode develops, the apical part becomes inflated, and is filled with pith. When the internode matures, the pith collapses against the inner wall, forming a hollow cavity, and a slit-like opening develops near the apex (Fig. 5). The opening widens grad- ually until ants or other small organisms can access the hollow domatium. Some oc- cupants apparently gnaw at the more or less elliptical opening, transforming it to an oval or circular opening that is gradually rimmed by a callus (Fig. 6), the plant’s response to the injury. Aculeate Hymenoptera (ants, wasps, bees) remove the pith to make the cavity larger, except for one species of sol- itary crabronine wasp, Crossocerus muka- lanae Leclercq which uses the pith in con- structing its nest in the cavity. We found aculeates nesting in internodes ranging in length from 4.3 to 11.2 cm. The basal end of the internode is woody to a variable extent, so the length of the cavity varies from 3.2—6.3 cm. The woody outer wall of the internode is 0.3—0.7 mm thick. The internode entrances are elliptical to cir- cular in shape with width to length mea- surements ranging from a minimum of 1.1 x 1.1 mm to a maximum of 1.2 X 1.8 mm. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON hes ‘ os dd Figs. 3-6. Humboldtia laurifolia. 3, Habitat; collecting on plant. 4, Leaf; note swollen internode at lower right. 5, Internode opening early in development. 6, Internode opening to Krombeinictus nordenae nest; note callus around hole. VOLUME 99, NUMBER I Krombeinictus nordenae Leclercq (Figs. 1, 2, 7-11) Apparently an uncommon wasp, we found only eight specimens (6 @, 2 d) in Humboldtia internodes at Gilimale as com- pared to several hundred of its fellow cra- bronine, Crossocerus mukalanae. It has un- usual morphology in that it is the only ge- nus of Crabroninae, other than the Oriental Vechtia Pate, in which both sexes possess a triangular lamella overhanging the deep scapal basin (Fig. 2). The female is unusual morphologically in the Crabroninae in lacking a pygidium de- limited by carinae on the last abdominal ter- gum. Instead there is a median brush of sev- eral rows of close-set, erect setae (Fig. 7). The last abdominal segment of females usu- ally bears the residue of a secretion from abdominal glands (Fig. 8) that we believe may function as an ant guard. We suspect that a nesting female uses its brush of setae to smear this secretion around the nest en- trance to deter predators and parasites from entering while the adult wasp is foraging. The Oriental Piyumoides Leclercq, con- sidered by him (1996) to be the genus most closely related to Krombeinictus, also lacks a pygidium. Females, however, lack the me- dian brush of setae on the last abdominal tergum. There is no secretion from abdom- inal glands on this segment in females of three of the four known species in our col- lection. Nest.—We found only ten nests of K. nordenae compared with about 75 of C. mukalanae, and adult females were present in only four of them. We believe that the missing females took flight during the pe- riod between gathering Humboldtia branch- es, and placing the internodes in alcohol. Several females were found sheltering in empty internodes. One typical nest exhibited the sequence of nesting activity (Fig. 9). The internode was 6.3 cm long, and the maximum width was 4.0 mm. The interior cavity was 5.0 cm long and had a maximum width of 3.4 mm. 45 When we split the internode, we found a small wasp larva, 2 mm long, on the wall of the cavity just below the entrance hole The female was lower in the cavity, prob- ably a reaction to being immersed in alco- hol when the internode was _ preserved. There was a wasp pupa with well developed adult coloration in its cocoon at the bottom of the cavity. Ten mm above this cocoon was a second cocoon that contained a post- defecated larva just prior to pupation. From these data, and observations in oth- er internodes with nesting K. nordenae, we deduce the following behavioral sequence. The foundress lays the first egg on the inner cavity wall just below the entrance. When the larva hatches, she feeds it progressively until mature. The wasp then transports it to the bottom of the cavity where it subse- quently spins its cocoon, pupates, and slow- ly begins to develop adult coloration. After the wasp takes the mature larva below, she lays a second egg just below the entrance. That larva is fed progressively, and, when mature, it is transported lower in the cavity for cocoon spinning. The same cycle is re- peated again with the wasp depositing an- other egg below the entrance. Typically, many sphecid wasps that nest in pithy stems or borings in wood construct nests containing a linear series of cells. The cells are sealed by partitions that separate siblings, thus preventing cannibalism. Fur- ther, nests are normally closed before eggs hatch so that females have no contact with their progeny. Thus, the progressive feed- ing, maternal attention, and lack of parti- tions is noteworthy in K. nordenae. Cocoon.—The cocoon also is unique among the Crabroninae. The typical crab- ronine cocoon is more or less ovoid, cir- cular in cross section, the posterior end ta- pers rather narrowly, and there is a pore at the anterior end, as in Ectemnius paucima- culatus (Packard) (Krombein, 1964). The cocoon of Krombeinictus (Fig. 10) is broad- ly ovoid, tapers very slightly posterad, and lacks a pore at the anterior end. The upper surface is only slightly convex so that in PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 7-11. Krombeinictus nordenae. 7, Female, apex of abdomen lateral oblique; note median brush of setae on last tergum. 8, Female, apex of abdomen, lateral oblique from rear; note dried secretion on last segment, and on fifth sternum. 9, Nest diagram. 10, Cocoon in section of internode from which adult emerged. 11, Cocoon in section of internode, oblique; note ample space for passage of adults. VOLUME 99, NUMBER 1 cross section the cocoon is more curved on the side that is appressed against the round- ed inner wall of the internode. Cocoon di- mensions are 6—9 mm long, 2.6—3.4 mm wide, and 2.0—2.1 mm high. There is a space at least 2 mm high be- tween its upper surface and the opposite in- ner wall of the cavity (Fig. 11). A cocoon of this shape permits the mother to crawl over a cocoon to carry a mature larva to- ward the lower end of the cavity, or for a newly emerged adult from lower in the in- ternode to crawl over a higher cocoon to reach the nest entrance. Larva.—The data above support our con- clusion that the female feeds her larva pro- gressively. We found larvae in various stages of development, but never with any prey or inedible prey fragments such as wings and legs that one would expect to find in the nest of a predaceous wasp. The problem of larval food identity was finally solved when we examined the exuviae of post-defecated larvae, and adjacent fecal wastes. Unmistakably, small grains of pol- len had been excreted with the meconium (Fig. 12) which were identical in size and appearance with grains of Humboldtia pol- len (Fig. 13). Also, we noted that freshly preserved larvae had a distinct yellow color reminiscent of that seen in bee larvae that have fed on pollen. Variations in larval bee color are attributed to pollen color (Norden, 1984). Later in our investigation, Rickson sent us a female of K. nordenae that he collected from an internode in Gilimale. We found clumps of Humboldtia pollen grains on the mandibles and hypostomal setae beneath the head (Figs. 14, 15). Due to the oily pol- lenkitt that envelops the pollen grains, they tend to adhere to each other and to insects that come in contact with the stamens. We infer from these data that the female probably gathers a quantity of the clumped Humboldtia pollen on the hypostomal area, returns to the internode, and deposits the pollen on the inner wall next to the head of the larva. Again, this feeding behavior is 47 remarkable and in contrast to all known sphecids whose larval food consists of par- alyzed arthropods. DISCUSSION Behaviorally, K. nordenae is unusual or unique among Sphecidae in several char- acteristics of its life history. The female manifests extraordinary maternal care, rear- ing one larva at a time, and feeding it pro- gressively. Progressive provisioning has not been noted previously for any Crabroninae. Evans (1966) reported that it developed in- dependently at least four times in the Nys- soninae. Evans also noted that eggs are pro- duced more slowly in progressive provi- sioners than in mass provisioners. However, the slow development that characterized our nests may actually be related to scarcity of pollen and thus a slower rate of feeding rather than to the rate of egg production. Humboldtia flowering occurs throughout the year, but the major production of inflo- rescences is usually February to June. The foundress with several brood in varying stages of development forms a sub- social group. Additional field observations are needed to ascertain whether this sub- social group reaches the level of social be- havior by having a daughter join the foun- dress in caring for later brood. There is also the possibility that K. nor- denae is so coadapted to Humboldtia that the wasp nests only in the plant’s inter- nodes, where nearby there is a plenitude of extrafloral nectaries for adult feeding, and pollen when the plant is in bloom. It is noteworthy that the wasp was never col- lected during the dozen years of the Smith- sonian’s “Ceylon Insect Project,” 1968— 1980. None of the project specialists col- lected on Humboldtia, but several of us, in- cluding the first author on a number of trips, collected at emergence holes in dead wood, and in tunnels in wood, and failed to find K. nordenae in these other plants. If K. nor- denae is in fact associated only with Hum- boldtia, we wonder if it or other species of Krombeinictus may have a similar relation- 48 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Seo) bee Figs. 12-15. Humboldtia laurifolia pollen. 12, Pollen mass in meconium within cocoon of Krombeinictus nordenae. 13, Grain from flower bud. 14, Pollen grains clumped on hypostomal setae behind mandibular apex of female K. nordenae. 15, Pollen grains massed on setae beneath head of female K. nordenae. ship in southern India with H. decurrens and H. brunonis. The cocoon also is unlike that of any oth- er known crabronine. Clearly it is an ad- aptation to the morphology of the internode cavity and nesting behavior of the foun- dress which permits movement of adults throughout the cavity. The cocoon of the normal twig-nesting crabronine is slightly less in diameter than that of the cavity. The larva spins a loose network of silken guy- lines against the entire wall of the cavity and then constructs the cocoon, suspending it within the network (Krombein, 1964, fig 7b, shows cocoons not in contact with cell wall). The cocoon of K. nordenae is spun directly in contact with the rounded inner wall of the internode, and there are only a few silken guy-lines along the cocoon mar- gins. Finally, and perhaps most significant, the species is extraordinarily different from all other known Sphecidae in feeding its larvae pollen rather than paralyzed arthropod prey. VOLUME 99, NUMBER I ACKNOWLEDGMENTS We are grateful to our late colleague and friend, PB. Karunaratne, for his many years of help to scientists conducting field studies in Sri Lanka. The Smithsonian Institution’s “Ceylon Insect Project’? (1968-1980) was highly successful in large part because of Karu’s dedication and knowledge. During our trip in 1993 he introduced the authors to Humboldtia habitat, and helped collect many of the internodes we examined. We dedicate this work to him. We also thank Arnold W. Norden who was with us during part of our 1993 visit, provided valuable support in field studies, and reviewed an earlier draft of the manu- script. We gratefully acknowledge Prof. Fred R. Rickson, Department of Botany and Plant Pathology, Oregon State University, Cor- vallis, for bringing H. laurifolia to our at- tention, explaining the morphology and phenology of the myrmecophyte, and for insect specimens from his collecting in Sri Lanka during 1992. We are further indebted to Prof. Jean Le- clercq, Liége, Belgium, for describing Krombeinictus nordenae. It is an honor to 49 have our names bestowed on this bizarre wasp with such unusual behavior Within the Smithsonian we thank Susann G. Braden who prepared most of the scan- ning electron micrographs, and George L. Venable who prepared the stem drawing and photographic plates. Finally, we thank Drs. Howard E. Evans, Colorado State University, Fort Collins, and Frank E. Kurczewski, State University of New York, College of Environmental Sci- ences and Forestry, Syracuse, for their re- views and suggestions. LITERATURE CITED Evans, H. E. 1966. The comparative ethology and evolution of the sand wasps. Harvard University Press, Cambridge, Massachusetts. 526 pp. Krombein, K. V. 1964. Natural history of Plummers Island, Maryland, XVIII. The hibiscus wasp, an abundant rarity, and its associates (Hymenoptera: Sphecidae). Proceedings of the Biological Society of Washington 77: 73-112. Leclercq, J. 1996. A new genus and species of Cra- bronini (Hymenoptera: Sphecidae) from Sri Lan- ka. Memoirs of the Entomological Society of Washington 17: 95-102. Norden, B. B. 1984. Nesting biology of Anthophora abrupta (Hymenoptera: Anthophoridae). Journal of the Kansas Entomological Society 57: 243- 262. PROC. ENTOMOL. SOC. WASH. 99(1), 1997, pp. 50-54 FENDEROMYIA SHAW, A VALID NORTH AMERICAN TAXON IN MACROCERINAE (DIPTERA: MYCETOPHILOIDEA: KEROPLATIDAE) Loic MATILE Muséum national d’ Histoire naturelle, Laboratoire d’Entomologie et EP 90 du CNRS, 45, rue Buffon, F-75005 Paris, France. Abstract.—The North American genus Fenderomyia Shaw, 1948, was long believed to be a junior synonym of Macrocera Meigen, based on an artifact in thorax pleura and several plesiomorphic states of characters. Identification of a specimen of the type species, F. smithi Shaw, shows that the thoracic structure described by Shaw was not an artifact, and makes it possible to point out several peculiar morphological characters of the taxon. A phylogenetic analysis shows that these characters are autapomorphic and justifies the resurrection of Fenderomyia. The genus is also present in the Neotropical Region. Key Words: analysis The genus Fenderomyia was erected by Shaw (1948) for a new North American species, F. smithi, allied to Macrocera Mei- gen, but which differed from this genus by the following characters: base of median vein distinct and reaching to the ““m-cu”’ crossvein, costa produced far beyond apex of wing, and mesepimeron not reaching the metapleuron. Coher (1963) rightly pointed out that the venation characters cited by Shaw for his new genus occured also in several species of Macrocera. He added that Shaw’s inter- pretation of the pleural sclerites was erro- neous, based on “‘a variable condition shown by a series of M. brunnea Brunetti, 1912, to be a result of distortion during dry- ing rather than of actual structure’. Ac- cordingly, Coher considered Fenderomyia a junior synonym of Macrocera. Laffoon (1965) recognized Fenderomyia as a valid genus, but it is doubtful that he knew of Coher’s paper before the comple- tion of his manuscript for the North Amer- ican Catalog, which includes only ‘“‘some Macrocerinae, Fenderomyia, valid generic status, morphology, character 1963 names’’. Thompson (1975), discuss- ing the pleural morphology of Lygistorrhi- na, mentions the synonymy established by Coher, and Vockeroth (1981) does not in- clude Fenderomyia in his key to the genera of Nearctic Mycetophilidae, which means that he also accepts Coher’ synonymy. Matile (1990), revising the generic clas- sification of Macrocerinae, discussed the status of Fenderomyia and generally agreed with Coher, noted that all three venation characters were plesiomorphic for the group, and accepted the thoracic distortion explanation. In some unsorted material of Macroceri- nae at the National Museum of Natural His- tory, Smithsonian Institution (USNM), I identified a specimen belonging to Shaw’s species. It was collected in Oregon (Coos Co., Charleston, VI. 1963, NLH. Krauss), as was Shaw’s type-series. As a result of an examination of this specimen and compar- ison with the extensive collection of Kero- platidae available to me, it is now possible to resurrect Fenderomyia from synonymy VOLUME 99, NUMBER I on the basis of several morphological char- acters. It is significant that the attribution to an artifact of an exceptional feature, the abridged mesepimeron, and the obviously plesiomorphic venation characters cited by Shaw for his new genus, have diverted the attention of all subsequent authors, the pres- ent one included, from the other very sig- nificant characters cited in the original de- scription. MATERIAL AND METHODS The USNM specimen is a male (accord- ing to habitus) in rather poor condition: The antennae, most of the legs, and the tip of abdomen are broken (only the first five seg- ments are present). It was originally glued laterally to a piece of cardboard. The integ- ument of the thorax was slightly transpar- ent, thus showing the thoracic muscles; the chitin of the pleura was smooth, without trace of infolding or outfolding, and the mesepimeron was definitely shortened. The specimen was detached from the cardboard and gently heated in distilled water, which demonstrated that the ventral margin of the mesepimeron corresponded to a true suture, with an internal phragma. One of Shaw’s diagnostic characters of Fenderomyia was thus confirmed. The head was further boiled in KOH to check the absence of a membra- nous area between cerebral phragma and front or occiput, thus confirming its posi- tion in the Macrocerini rather than the Rob- sonomyiini. The specimen was then dried by the method of Sabrosky (1966), modi- fied by the replacement of cellosolve by monoethylene glycol (Matile 1994), and then glued back to its cardboard. The head was preserved in glycerine in a microvial. According to the original description, the type series of F. smithi should be in Shaw’s collection, presently at the University of Massachussetts. Dr. T.M. Peters was unable to locate it either under Macrocera or Fen- deromyia. Dr. R. J. Gagné was kind enough to make inquiries about the possible loca- tion of these specimens, but without suc- cess, and writes “we will just hav sume that the types were never distri to the intended institutions and are probably lost”’ (in litt. Apr. 1996). ~ -, CHARACTER ANALYSIS Four characters, presumably apomorphic, can be added to the character of the short- ened mesepimeron of Fenderomyia: the long stem of the anterior fork; the diagonal strip of dense, erect black setae of the met- episternite, briefly mentioned in the original description, which runs under the posterior spiracle and above the dorsal margin of the pleurite; the angular, short mediotergite, strongly projecting backwards behind the scutellum; and the presence of only one spur on all tibiae. I have examined these five features on about 100 species of Ma- crocera, described or undescribed, from all biogeographical regions, as well as in an extensive collection of other Keroplatidae and related families. The five characters are discussed below. 1. Mesepimeron.—The evolution of the mesepimeron has been discussed in Matile (1990: 378, 411), and outgroup comparison has shown that the most primitive condition in most Nematocera was a vertical plate reaching the lower margin of the pleura at the level of the metepisternite. In the My- cetophiloidea, the plate narrows ventrally, and the disappearance of its ventral part is undoubtedly apomorphic. The phenomenon has occured several times in the Keroplati- dae—at least twice in the Macrocerinae, e.g. in Vockerothia Matile (Macrocerini) and Micrepimera Matile and an unde- scribed genus in Robsonomyiini—and once in the Keroplatinae, namely in Nauarchia Matile (Keroplatini). It is also a trait of all Lygistorrhinidae and most Ditomyiidae. 2. Stem of anterior fork.—As regards the stem of the media in the studied species of Macrocera, the anterior fork always opens far basad of the base of R;, except in two Neotropical species, M. guayanasi Lane and M. unidens Edwards. Among the other Macrocerinae, a long stem as in Fendero- 52 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON myia is present only in a group of Austral- asian Paramacrocera Edwards, and in the Oriental genus Micrepimera Matile. In my character analysis of the anterior fork of the Mycetophiloidea (Matile 1990: 438), I not- ed that fossil evidence was in favor of the plesiomorphy of a long petiole, and inferred that its shortness in the Keroplatidae was apomorphic, and part of their groundplan. The condition of Fenderomyia, Micrepi- mera, the two South American Macrocera and the Australasian Paramacrocera should thus be a reversal, and its appearance in at least four groups of Macrocerinae the result of parallelism. 3. Postspiracular setae.—There is often in Keroplatidae a sparse patch of short setae at the posterior or ventral margin of the me- tepisternite, but the dorsal rows of dense setae in F. smithi do not really stand on the metepisternite, but above it, on the peris- piracular membrane, a few of the ventral setae extending to the extreme dorsal mar- gin of the sclerite (Figs la, b). These setae should in fact properly be named infraspi- racular setae. I have been unable to find this feature in any Macrocerinae, in Arachno- campinae and Keroplatinae (Keroplatini). I have checked also representatives of almost all the described genera of Orfeliini and have found this character absent also. It does not exist in the more primitive family Ditomyiidae, nor in the presumed sister- group of the Keroplatidae, the Diadocidi- idae (for a provisional phylogenetic analy- sis of the relationships of the families of Mycetophiloidea, see Matile 1990: 383— 386). The apomorphic state of the character seems therefore well founded. 4. Mediotergite.—In most Macrocera known to me, the mediotergite is high, sub- vertical in its upper half, then curves gently downward and forward to meet the meta- notum, thus not projecting behind the scu- tellum, or very slightly so. In a few species, the sclerite is evenly rounded and projects somewhat behind the scutellum, the condi- tion illustrated by Shaw for M. formosa Loew, but never in such proportions as in Fenderomyia (Fig. 1a; compare also Shaw’s Figs. 1 and 2). The evolution of the mediotergite has been studied (Matile 1990: 405), and out- group comparison has shown that its prim- itive condition in Mycetophiloidea was a vertical, high and slightly rounded sclerite. In fact, this character is one of the nine used to separate Macrocerini and Robsonomyi- ini, the two tribes of Macrocerinae. Within the subfamily, the projecting mediotergite was considered autapomorphic for the Rob- sonomyiini. The condition of Fenderomyia thus must have appeared independently once in Macrocerini. 5. Tibial spurs.—Only one tibia and three tarsomeres, as well as some isolated tarso- meres, remained on the piece of cardboard of the USNM specimen and these became detached as the glue dissolved; the absence of a sensorial crypt indicates that the re- maining tibia belongs to leg II or III; it has only one very short spur. Shaw states “‘tibia with one spur’’, and he had three specimens available, therefore I assume that his obser- vation refers also to both the median and posterior legs. The loss of the outer spurs II-III is an evolutionary trend of the Keroplatidae (Ma- tile 1990: 418) which occured indepen- dently once in the Keroplatinae Keroplatini (Xenokeroplatus Matile), and several times in the Orfeliini, but the state of this char- acter was not known up to now in the Mac- rocerinae, although the reduction in size is common. Most Macrocera known to me have two posterior spurs, the longest about twice as long as the width of the tibia at its apex. Occasionally, the spurs are reduced to one tibial width, or a little less. Only in M. guayanasi and M. unidens are there two very minute spurs as observed on the Ar- gentinian Fenderomyia mentioned below. DISCUSSION The validity of Fenderomyia is supported by five strong apomorphies. The short me- dian fork is shared by two species of Mac- rocera (both with a rather shortened medi- VOLUME 99, NUMBER 1 AG) lens, Al and infraspiracular setae. otergite), and does not exist elsewhere in the subfamily. The thoracic structure has appeared several times in the subfamily, but never in Macrocera. The projecting, short- ened mediotergite is unique for the Macro- cerini, while it is characteristic of the Rob- sonomyiini. The loss of the external tibial spurs II-III is not yet known to happen in the Macrocerinae. Lastly, the infraspiracu- lar rows of setae are unique in the family Keroplatidae and its closest relatives. This amply justifies the acceptance of Fendero- myia as a valid taxon, if only to emphasise this peculiar set of apomorphies. In the Duret Collection of Neotropical Mycetophiloidea, now preserved in the Mu- séum national d’ Histoire naturelle, Paris, I found an unidentified female of “‘Macro- cera” (Argentina, Salta, Campo Quijuano) which shares with F. smithi the disappear- ance of the ventral part of the mesepimeron, the long stem of the median fork and the projecting mediotergite, but lacks the dorsal row of infraspiracular setae. Sc, is definitely absent, but the basal fold of the media is Nn Las) a b Fenderomyia smithi (USNM specimen). a, Lateral view of thorax and coxae. b, Posterior spiracle very faint. The two anterior legs and one posterior leg are left; there is a short spur on tibia I, and two very minute spurs on the II, smaller than the basal diameter of the protarsus. I am reluctant to describe for- mally the species from a unique female, but the specimen certainly represents the clos- est known relative of F. smithi. Therefore, the infraspiracular setae would be autapo- morphic for F. smithi. Provisionally, pending a revision of Ma- crocera, | propose to follow Shaw and to recognize Fenderomyia as a valid genus close to Macrocera, with the following di- agnosis: Fenderomyia Shaw, 1948: 94. Type spe- cies: Fenderomyia smithi Shaw, 1948: 94 (original designation). A genus close to Macrocera, sharing with this genus the globular antennal scape and the long flagellum, but differing in the incomplete mesepimeron, interrupted at the level of lower third of the laterotergite, the 54 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON projecting mediotergite, the stem of the me- dial fork long, ending at level of the base of R;. In the type species, posterior spiracle with several rows of posterior setae and only one tibial spur II-III. In the Neotrop- ical species, spiracular setae absent, and hind tibia with two minute spurs. Additional characters of taxonomic or phylogenetic significance in Macrocera or Macrocerini are: Cerebral sclerite large, meeting the eyes at their anterior corner, bearing long anterior setae. Thorax and coxae as in Fig. la, acrostichal bristles ab- sent. Scutellum with short, fine marginal bristles. Mesanepisternite with long, erect, dorsal setae. Subscutellar membranous area narrow. Wing: C strongly produced after R,, Sc, visible as a trace (USNM specimen) or absent, basal fold of the media very strong (F. smithi) or faint (Neotropical spe- cies). Anal vein long, reaching to the mar- gin. Wing membrane without macrotri- chiae. Ciliation of veins, dorsal surface: C, R,, branches of the anterior fork, M,, Cu, and tip of anal vein. Male genitalia simple, of the Macrocera type (Shaw, 1948: fig. 4). Distribution: Nearctic and Neotropical. ACKNOWLEDGMENTS My thanks are due to Dr. Raymond J. Gagné for the loan of an important material of Keroplatidae from USNM, including the specimen discussed above, and for his search for the type series of Fenderomyia smithi. Drs. R. J. Gagné and D. Williams were kind enough to revise the language of the manuscript. I thank also Gilbert Hode- bert for the drawings. LITERATURE CITED Coher, E. I. 1963. Asian Macrocera Meigen, 1803, (Diptera: Mycetophilidae), with some remarks on the status of the genus and related genera. Bulletin of the Brooklyn Entomological Society 58(1): 23— 36. Laffoon, J. L. 1965. Family Mycetophilidae (Fungi- voridae), pp. 196-229. In Stone, A. et al., eds., A Catalog of the Diptera of America north of Mex- ico. U.S. Department of Agriculture, Agricultural Handbook 276, iv + 1—1696. Matile, L. 1990. Recherches sur la systématique et Vévolution des Keroplatidae (Diptera, Myceto- philidae). Mémoires du Muséum national d’ His- toire naturelle, sér. A, Zoologie 148: 1—682. . 1994. Les Diptéres d’Europe occidentale. I. Introduction, Techniques d’ étude et Morphologie. Nématoceéres, Brachycéres Orthorrhaphes et As- chizes. Paris, Boubée, 439 pp. Sabrosky, C. W. 1966. Mounting Insects from Alco- hol. Bulletin of the Entomological Society of America 12(3): 349. Shaw, E R. 1948. A new genus and species of fungus- gnats (Mycetophilidae). Bulletin of the Brooklyn Entomological Society 43(3): 94-96. Thompson, FE C. 1975. Notes on the genus Lygistor- rhina Skuse with the description of the first Ne- arctic species (Diptera: Mycetophiloidea). Pro- ceedings of the Entomological Society of Wash- ington 77(4): 434-445. Vockeroth, J. R. 1981. 14. Mycetophilidae, pp., 223— 246. In McAlpine, J. E, ed., Manual of Nearctic Diptera. Volume 1.Biosystematics Researc Centre, Research Branch. Agriculture Canada. Mono- graph No. 28. Ottawa, vi + 674 pp. PROC. ENTOMOL. SOC. WASH. 99(1), 1997, pp. 55-59 CLADISTICS AND BIOGEOGRAPHY OF THE ASSASSIN BUG GENUS MELANOLESTES STAL (HETEROPTERA: REDUVITIDAE) Maria del C. Coscar6n and Juan J. Morrone (MCC) Departamento Cientifico de Entomologia, Museo de La Plata, Paseo del Bosque, 1900 La Plata, Argentina; (JJM) Laboratorio de Sistematica y Biologia Evolutiva (LAS- BE), Museo de La Plata, Paseo del Bosque, 1900 La Plata, Argentina. Abstract.—Melanolestes Stal is a monophyletic genus of nine species, two Nearctic and seven Neotropical. A cladistic analysis of the genus was carried out using 28 char- acters. The cladograms were rooted with the genus Peirates Serville. The analysis yielded 20 equally parsimonious cladograms, with 34 steps, CI = 0.52, and RI = 0.44. A suc- cessive weighting procedure resulted in one cladogram with 77 steps, CI = 0.84, and RI = (0.87. The distribution of Melanolestes coincides in part with a previous study on Peiratinae, that showed that the former continuous Amazonian forest was separated into two parts by a diagonal line of open areas. In addition, the two Nearctic species M. picicornis and M. picipes are sister-taxa, so a single dispersal event accounts for the presence of Melanolestes in the Nearctic. Key Words: The New World assassin bug genus Me- lanolestes Stal (Heteroptera: Reduviidae: Peiratinae) is known from southeastern Canada to northern Argentina. The nine species belonging to this genus have been recently revised (Coscar6n and Carpintero 1994). Two of these species, M. picicornis Stal and M. picipes (Herrich-Schaeffer), are restricted to the Nearctic Region, whereas the remainder are Neotropical. Within the Neotropics, M. goiasensis Coscar6n and Carpintero, M. lugens Coscar6n and Car- pintero, M. minutus Coscar6n and Carpin- tero, and M. picinus Stal have very small distributional areas within the Amazonian and Chacoan domains. In a previous bio- geographic study (Morrone and Coscar6n 1996), we analyzed distributional patterns of the Neotropical Peiratinae, concluding that these patterns have been basically caused by the gradual development of a di- agonal line of open areas (Chaco-Cerrado- Peiratinae, Reduviidae, cladistics, biogeography Caatinga), which separated the former con- tinuous tropical forest into two parts. In this paper we provide a cladistic anal- ysis of Melanolestes, and discuss its bio- geographic patterns. MATERIAL AND METHODS This study is based on the revision of Melanolestes by Coscar6n and Carpintero (1994), and the examination of specimens borrowed from the following collections: American Museum of Natural History, New York, USA; The Natural History Museum, London, United Kingdom; Instituto Na- cional de Pesquisas Amazonicas, Manaus, Brazil; Museo Argentino de Ciencias Na- turales Bernardino Rivadavia, Buenos Ai- res, Argentina; Museo de La Plata, La Plata, Argentina; Museu de Zoologia de Sao Pau- lo, Sao Paulo, Brazil; Naturhistoriska Riks- museet, Stockholm, Sweden; Zoologisches Museum der Humboldt Universitat zu Ber- 56 character states; 1, 2 = apomorphic character states; ? PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Data matrix and characters of the species of Melanolestes used in the analysis. 0 = plesiomorphic missing data. No ONAN ff WwW \O 10. lite 1 13). 14. 15). 16. Ws 18. 19. 20. 2ile D2r 23): 24. 25: 26. PaTte 28. Peirates OVDD0D0D0DD0D0D0D00D0D0000000000000000 M. argentinus CLONE OOM SOLO LO LOR OLO Ts eal ete OOKOLORIsO sia M. degener AAO OS WO OOO POI eh OO) ea SL M. goiasensis ORB ECOR ORR OLO MS eile sie Oy Oise rales? M. lugens OZ MEO MEE ON 2 OMS Oss OM OFOTO OO Om eiiaG M. minutus OZMFOLO MOA TSO ORO ME OLO;OLOl ON 25 cela? M. morio ORS OLORE TOMO ce OR aaa ele OLOLO ORO RIEO M. picicornis COMTOO NT a Ose ae OLOTOLO ORIG M. picinus LPL OLOTO RL ORO ROLO RLS OF Seat is OL esr is tate M. picipes OFZ OORT 1 e ee OM OLO FI tials ee OrO OO RO maa . Body shape. [0] slender; [1] robust. . Eyes. [0] not attaining superior edge of head; [1] attaining superior edge of head; [2] surpassing superior edge of head. Treated as additive. . Ocelli. [0] not placed on a tubercle; [1] placed on a tubercle. . Lateral tubercle on neck. [0] present; [1] absent. . Metallic shine in pronotum. [0] absent; [1] present. . Pronotal granulations on anterior lobe. [0] absent; [1] present. . Pronotal granulations on posterior lobe. [0] absent; [1] present. . Pronotal sulci. [0] distinct; [1] not distinct. . Pronotal lateral internal sulci. [0] distally united; [1] medially united; [2] not united. Treated as non-additive. Lateral margin of pronotum. [0] carina absent; [1] carina present at entire length. Scutellum coloration. [0] unicolorous; [1] bicolored. Spongy fossa. [0] not occupying distal third of fore and hind tibiae; [1] occupying distal third of fore and hind tibiae. Female hemelytra. [0] macropterous; [1] brachypterous. Hemelytra. [0] surpassing apex of abdomen; [1] not surpassing apex of abdomen. Body color, [0] not uniform; [1] uniform. Hemelytral pale stripe on corium and clavus. [0] present; [1] absent. Color of fore femora. [0] bicolored; [1] unicolorous. Color of hind femora. [0] unicolorous; [1] bicolored. Color of fore tibiae. [0] unicolorous; [1] bicolored. Color of mid tibiae. [0] unicolorous; [1] bicolored. Connexivum. [0] dorsally visible; [1] dorsally not visible. Connexivum color. [0] unicolorous; [1] bicolored. Parameres shape. [0] subrectangular; [1] subtriangular. Gonocoxite IX internal edge hairs. [0] thin; [1] thin and thick. Gonocoxite IX unsclerotized area close to inner margin. [0] absent; [1] present. Shape of IX and X tergites. [0] subquadrangular; [1] subrounded. Intersegmental line of tergites IX and X. [0] not entire; [1] entire. Intersegmental line. [0] not strongly sclerotized; [1] strongly sclerotized. lin, Berlin, Germany; Zoological Museum, University of Helsinki, Helsinki, Finland; and the private collections of D. Carpintero, Argentina; L. Jir6n, Costa Rica; and the late J. Maldonado-Capriles, Puerto Rico. Melanolestes constitutes a monophyletic group that is distinguished from other Peira- tinae by the spongy fossa occupying distal third of fore and hind tibiae, body of uni- formly dark color, ocelli placed on a tubercle, pronotal granulations on anterior lobe, he- melytra lacking a pale stripe on the corium and clavus, hind femora unicolorous, and in- tersegmental line of tergites [IX and X entire. The nine species currently assigned to the genus are considered as terminal taxa: M. argentinus Berg, M. degener (Walker), M. goiasensis Coscar6n and Carpintero, M. lugens Coscar6n and Carpintero, M. minu- tus Coscaron and Carpintero, M. morio (Er- VOLUME 99, NUMBER 1 Peirates Figs = dotted rectangles. ichson), M. picicornis (Stal), M. picinus Stal, and M. picipes (Herrich-Schaeffer). The data matrix and the 28 characters used in this study are detailed in Table 1. The data were analyzed with Hennig86 ver- sion 1.5 (Farris 1988), applying the implicit enumeration (ie*) option for calculating the shortest trees. Consistency (CI) and reten- tion (RI) indices were calculated excluding uninformative characters (autapomorphies and synapomorphies of the genus). We used the successive weighting procedure in Hen- nig86, that calculates weights from the best fits of the characters on the most parsimo- nious cladograms using rescaled consisten- cies (products of the character consistency and the character retention index). These products are scaled in the range 0-10, and the weighting procedure is repeated succes- sively until the cladograms no longer change (Farris 1989). CLADOS version 1.1 (Nixon 1992) was used for examination of character distributions. RESULTS AND DISCUSSION The analysis using equal weights yielded 20 equally parsimonious cladograms, each Cladogram of the species of Melanolestes. M. picinus M. degener Synapomorphies = solid black rectangles; homoplasies with 34 steps, CI = 0.52, and RI = 0.44. When the successive weighting procedure was applied, one minimum-length clado- gram was selected from the original ones after the second round of weighting, with 77 steps, CI = 0.84, and RI = 0.87 (Fig. 1). Values for the number of steps, consis- tency index (ci), retention index (ri), and weight (ri X ci X 100) in the weighted trees are listed in Table 2. The phylogenetic se- quence from the basal to the most distal species is as follows: M. lugens, M. minu- tus, M. picicornis plus M. picipes, M. mo- rio, M. picinus plus M. goiasensis, and M. argentinus plus M. degener. Several conclusions can be deduced by comparing the cladogram obtained with the areas inhabited by the species of Melano- lestes (Fig. 2): (1) the more basal species (M. lugens) is restricted to the Paranaense province of the Amazonian domain; (2) M. minutus is restricted to the Cha- coan domain; (3) the more widespread species M. mo- rio and M. argentinus are among the most distal species of the cladogram; 58 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1000 2000 Km Fig. 2. Geographical distribution of the species of Melanolestes, with the cladogram superimposed. a, M. lugens; b, M. minutus; c, M. picicornis; d, M. picipes; e, M. morio, f, M. picinus; g. M. goiasensis; h, M. argentinus; i, M. degener. VOLUME 99, NUMBER 1 Table 2. Character consistencies and retention in- dices are the best fits of the 20 most parsimonious cladograms obtained applying ie. Final weights were obtained after the second round of the successive weighting procedure. Number of Consistency Retention Weight Final Character Steps Index (ci) Index (ri) (ri X ci X 100) Weight 1 1 1.0 1.0 100 100 2 4 0.5 0.6 30 0 3 1 1.0 1.0 100 0) 4 i 1.0 1.0 100 100 5 2 0.5 0.5 2) 25 6 DD 0.5 0.5 DS 25 7 2 1.0 1.0 100 0 8 2 0.5 0) 0 10) 9 2 0.5 0) 0 0) 10 1 1.0 1.0 100 100 11 i 1.0 1.0 100 100 12 | 1.0 1.0 100 100 13 I 1.0 1.0 100 100 14 1 1.0 1.0 100 100 15 1 1.0 1.0 100 100 16 1 1.0 1.0 100 100 7 1 1.0 1.0 100 100 18 ] 1.0 1.0 100 100 19 1 1.0 1.0 100 100 20 1 1.0 1.0 100 100 21 1 1.0 1.0 100 100 22 1 1.0 1.0 100 100 28 2 0.5 10) 0) 0) 24 2 0.5 0.5 25 0) 25 1 1.0 1.0 100 100 26 1 1.0 1.0 100 33 2 1 1.0 1.0 100 100 28 2 0.5 0.5 25 DS (4) the two Nearctic species (M. picicor- nis and M. picipes) are sister-taxa. These results corroborate, in part, our previous study (Morrone and Coscar6n 59 1996), because the Chacoan species M. minutus 1S one of the most basal species, whereas the Amazonian species are among the most distal species. Because M. pici- cornis and M. picipes are sister-taxa, a sin- gle dispersal event accounts for the pres- ence of Melanolestes in the Nearctic. ACKNOWLEDGMENTS We thank Thomas J. Henry and one anonymous reviewer for the critical reading of the manuscript. Our study was supported by grants from the University of Helsinki to the senior author and grant 4662-91 of the National Geographic Society to the ju- nior author. Support from the Consejo Na- cional de Investigaciones Cientificas y Téc- nicas (CONICET), Argentina, to which the authors belong, is gratefully acknowledged. LITERATURE CITED Cabrera, A. L. and A. Willink. 1973. Biogeografia de América Latina. Monografia 13, Serie de Biolo- gia, OEA, Washington D.C. Coscaron, M. del C. and D. L. Carpintero. 1993. Re- vision of the genus Melanolestes Stal (Heterop- tera: Reduviidae, Peiratinae). Entomologica Scan- dinavica 24(4): 361-381. Farris, J. S. 1988. Hennig86 reference. Version 1.5. Published by the author, New York. —. 1989. The retention index and the rescaled con- sistency index. Cladistics 5: 417—419. Morrone, J. J. and M. del C. Coscar6n. 1996. Distri- butional patterns of the American Peiratinae (Het- eroptera: Reduviidae). Zoologische Mededelingen Leiden. Nixon, K. C. 1992. CLADOS ver. 1.1. IBM PC-com- patible character analysis program. Published by the author, Port Jefferson, New York. PROC. ENTOMOL. SOC. WASH. 99(1), 1997, pp. 60-66 AULACIDAE (HYMENOPTERA) OF SRI LANKA DAVID R. SMITH Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart- ment of Agriculture, % National Museum of Natural History, MRC-168, Washington, DC 20560, U.S.A. Abstract.—Three species of Aulacidae from Sri Lanka are keyed, described, and illus- trated: Pristaulacus flavipennis (Cameron), P. signatus (Shuckard), and P. krombeini, n. sp. Lectotypes for Aulacus flavipennis Cameron and Aulacus magnificus Schletterer are designated. Key Words: Three species of the family Aulacidae are known from Sri Lanka, all of which were among collections of the Smithsonian Cey- lonese Insect Project of the 1970’s. Two are species previously described from Sri Lan- ka, and one was undescribed. All are in the genus Pristaulacus Kieffer. About 20 spe- cies of Aulacidae have been described from the Oriental Region, and six of them are from northern India, but the species from Sri Lanka are distinct. Species of Aulacidae are parasitoids of wood-boring Coleoptera (especially Cer- ambycidae and Buprestidae) and Hymenop- tera (Xiphydriidae). Hosts are not known for the Sri Lankan species, and there are only two host records from southeastern Asia. Pristaulacus beesoni Turner, de- scribed from **Thano, Siwalik Hills, United Provinces, India’’ was “‘taken emerging from a Sal log” (Turner 1922), and a form described as Pristaulacus nigripes var. du- porti Kieffer from Viet Nam was ‘‘Obtenu en abondance des larves de Xylotrechus quadripes |Cerambycidae]”’ (Kieffer 1921). KEY TO AULACIDAE OF SRI LANKA 1. Head, mesosoma, and legs mostly yellow; wings yellowish, forewing sometimes with Pe Aulacidae, Pristaulacus, Sri Lanka, parasitoids spot below stigma and apical margin black (Fig. 4), or entirely blackish apical to stigma; mesonotum reticulate (Fig. 2); head in dorsal view elongate behind eyes (Fig. 1) BS Can ee eae CAboe P. flavipennis (Cameron) Head and body nearly all black, at most with only antennal scape and pedicel and fore- and midlegs reddish; forewing hyaline except for black substigmatal spot and black spot at apex or with anterior half black; mesonotum with transverse ridges (Figs. 6, 10); head in dorsal view short and narrowing behind eyes (Figs. 5, Oy yea meet ete chad aye 2.4 ae ee Forewing hyaline with infuscated spot below stigma (Fig. 8); antennal scape and pedicel and fore- and midlegs reddish; mesosoma with rather dense silvery hairs obscuring sculpture (Fig. 7); anterolateral lobes of mesonotum pro- jecting forward in dorsal and lateral views (Figs. 6, 7) Forewing infuscated on anterior half, posterior part nearly hyaline (Fig. 12); head, body, and legs black, at most with mandible, tegula, and first metasomal segment reddish to brownish; mesosoma with relatively sparse hairs which do not obscure sculpture (Fig. 11); anterolateral lobes of mesonotum scarcely projecting for- ward in dorsal view (Fig. 10), projecting up- ward in lateral view (Fig. 11) PAA Pee Tee P. krombeini, new species Pristaulacus flavipennis (Cameron) (Figs. 1—4) Aulacus flavipennis Cameron 1887: 134 a eg dr P. signatus (Shuckard) VOLUME 99, NUMBER 1 61 Figs. 1-4. Pristaulacus flavipennis. 1, Head, dorsal. 2, Mesosoma, dorsal. 3, Mesosoma, lateral. 4, Wings. 1-3, Specimen from Kegalla District. 4, Lectotype of P. magnificus. (So): —Kietier 19253715372 (in key, ¢ fe- described). Pristaulacus flavipennis: Turner 1919: 386 (syn.: magnificus Schletterer).—Hedi- exe 1959278: Aulacus magnificus Schletterer 1890: AO SSOO Pt 2201 SIC Ls "s:). Pristaulacus magnificus: Kieffer 1900: 338.—Kieffer 1902: 13.—Kieffer 1912: 380, 388 (in key; d and @ redescribed). Female.—Length excluding ovipositor, 16-17 mm; forewing length, 14 mm; ovi- positor length, 16-17 mm. Antenna black. Head yellow; apex of mandible black. Me- sosoma yellow. Legs yellow with hindleg black except base and inner surface of coxa (lectotype of magnificus) or first segment of trochanter, apical third of outer surface of tibia, and tarsus black (specimen from Ke- galla District). Metasoma, except base and dorsal surface of first segment black (lec- totype of magnificus) or orange with apical third of metasomal terga 2—6 blackish (specimen from Kegalla District); sheath black. Wings yellow; forewing with stigma black and large spot below stigma and ap- icoventral margin black (lectotype of mag- nificus; Fig. 4) or yellow with basal portion of stigma blackish and very faint darker spot below stigma (specimen from Kegalla District). Head from above elongated be- hind eyes (Fig. 1), shining and impunctate; 62 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON occipital carina very short. Propleuron shin- ing and impunctate. Front margin of pro- notum with a forward projecting triangular tooth just above its midlength; mesonotum with reticulate sculpture; front margin of mesonotum rounded, slightly lobate, not overhanging pronotum in lateral view (me- sosoma as in Figs. 2, 3). Hindcoxa with dis- tinct, coarse cross wrinkles laterally and posteriorly. Tarsal claws with 5 teeth (in- cluding apical tooth) and small basal lobe. Ovipositor length about 1.2 forewing length. Male.—Length, 15 mm. Color similar to lectotype female (magnificus), except smaller substigmatal spot of forewing, which is about half width of stigma and does not enter cubital cells (paralectotypes of flavipennis and magnificus), or with apex of forewing from base of stigma black (lec- totype of flavipennis). Structure as for fe- male. Records.—SRI LANKA: Kandy District, Dikoya, 06°52'N, 80°30’E (types of A. fla- vipennis, spelled ““‘Dekaya’’ by Cameron, 1887). Kegalla District, Kitulgala, Makan- de Mukalana, 3-4-11-1979, K.V. Krombein, PB. Karunaratne, T. Wijesinhe, S. Siriwar- dane, T. Gunawardane (1 2). Types.—Aulacus flavipennis Cameron was described from two males from “De- kaya, Ceylon (Mr. George Lewis)’’ The specimens are in The Natural History Mu- seum, London. The colors of the two spec- imens are slightly different, and Cameron called them “‘two forms of this species.” One specimen is labeled “B.M. TYPE HYM 3.a.139’’, and this is hereby desig- nated lectotype. The specimen lacks anten- nae, except for the basal three segments of the left antenna, hindlegs, and the metaso- ma, except for the first segment. The other specimen, the second “‘form”’ described by Cameron, lacks a type label and is consid- ered a paralectotype. Schletterer described Aulacus magnificus from a female and a male. He stated “‘Type im kaiserl. naturhistorischen Hofmuseum zu Wien (2) und im kGnigl. naturhistorisch- en Museum zu Berlin (d).”” I have exam- ined both specimens. The lectotype here designated is the female at the Naturhisto- risches Museum Wien, Austria, labeled ‘Felder [spelling ?], Ceylon, 1861,” ““mag- nificus det. Schlett.”’, and a red label ““TY- PUS.”’ The specimen is in fair condition with the following parts missing: right an- tenna beyond basal 4 segments, left antenna beyond basal 6 segments, right forewing, right foreleg, and left midleg. The male, a paralectotype, is at the Zoologisches Mu- seum an der Humboldt-Universitat zu Ber- lin, Germany, labeled “‘Ceylon, Nietner.” “11548,” “‘type’’ [red label], “‘magnificus Schlett.,’’ and “‘Zool. Mus. Berlin.” It is in good condition. Discussion.—Even though there are col- or differences in the wings, metasoma, and hindlegs among the specimen from Kegalla District (female), the two forms of A. fla- vipennis Cameron (males), and the two specimens of A. magnificus Schletterer (fe- male and male), I regard these as probable variation or sexual differences and treat them as the same species. The structure, es- pecially sculpture, of all is similar. Much more study material will be necessary to determine if this is a species complex with perhaps more than one species involved. The specimen collected in Kegalla District is the palest: yellow with antenna, apex of mandible, apical portion of terga 2—6, sheath, ovipositor, first segment of hindtro- chanter, apical third of outer surface of hindtibia, and all hindtarsus black, and the forewing is yellow with only a faint black area beneath the stigma. The lectotype of A. flavipennis has most of the metasoma and hindtibia black and the forewing with the apex beyond the basal end of the stigma mostly black. The paralectotype of A. fla- vipennis is similar to the lectotype except the forewing has a large spot below the stigma and apical margin black, and the lec- totype female and paralectotype male of A. magnificus Schletterer are similar to the paralectotype of A. flavipennis. VOLUME 99, NUMBER I Figs. 5-8. Pristaulacus signatus (Shuckard) (Figs. 5-8) Aulacus signatus Shuckard 1841: 124.— Westwood 1844: 268.—Schletterer 1890: 509 (= ? stigmaticus Westwood).—Kieffer 1902: 12 (? Aulacus).—Kieffer 1912: 373 (repeats Shuckard’s description).—Hedicke 193919: Pristaulacus signatus: Kieffer 1904: 455 (? Pristaulacus). Female.—Length, excluding ovipositor, 9-11 mm, forewing length, 8-10 mm; ovi- positor length, 8-10 mm. Black except for following: mandible dark orange with apex reddish brown; anterior half or less of clyp- eus sometime brownish; antennal scape or Pristaulacus signatus. 5, Head, dorsal. 6, Mesosoma, dorsal. 7, Mesosoma, lateral. 8, Wings. scape and pedicel and fore- and midlegs be- yond trochanters dark reddish; hindleg be- yond coxae sometimes brownish; tegula brownish to reddish brown; first gastric seg- ment sometimes partly pale reddish to or- ange. Wings hyaline; forewing with small infuscated spot below stigma (Fig. 8). Head from above short and narrowing behind eyes (Fig. 5); shining with widely scattered punctures and silvery hairs, these both more dense between ocelli and antennae; occipi- tal carina short. Propleuron shining with scattered punctures, similar to top of head. Pronotum with a forward projecting trian- gular tooth just above its midlength; antero- lateral angles of mesonotum protruding an- teriorly in dorsal and lateral views and 64 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON mesonotum with coarse transverse ridges; dense silvery hairs somewhat obscuring sculpture, especially laterally (mesosoma as in Figs. 6, 7). Hindcoxa with distinct, coarse transverse wrinkles laterally and posteriorly. Tarsal claws with 4 teeth, in- cluding apical tooth, and small basal lobe. Ovipositor length about as long as forewing length. Male.—Length, 8-10 mm. Color and structure as for female. Records.—SRI LANKA: Amparai Dis- trict, Ekgal Aru, 12-VI-1976, K.V. Krom- bein, P.B. Karunaratne, S. Karunaratne (1 2); Ekgal Aru, 100 mtrs, Reservoir Jungle, 19-22-11-1977, K.V. Krombein, P.B. Karu- naratne, P. Fernando, D.W. Balasooryia (1 2), 11-12-VI-1976, K.V.Krombein, PB. Karunaratne, S. Karunaratne (1 2); Lahu- gala Tank, 14-15-VI-1976, K.V.Krombein, PB. Karunaratne, S. Karunaratne (1 °). An- uradhapura District, Padaviya Tank, 180 ft., 20-21-V-1976, K.V.Krombein, P.B. Ka- runaratne, S. Karunaratne, D.W. Balasoor- yia (1 2, 1 d); Padaviya, 180 ft., Irrigation Bungalow, 18-V-1976, K.V. Krombein, P.B. Karunaratne, S. Karunaratne, D.W. Bala- sooryia (1 2). Colombo District, Labuga- ma, 23-24-VI-1975, S.L. Wood & J.L. Petty (2 2). Kandy District, Hasalaka, 140-170 m, 15-17-IX-1977, K.V. Krombein, P.B. Karunaratne, T. Wijesinhe, M. Jayaweera (1 2). Kegalla District, Kitulgala, Makande Mukalana, 3-4-II-1979, K.V. Krombein, P.B. Karunaratne, T. Wijesinhe, S. Siriwar- dane, T. Gunawardana (1 d). Polonnaruwa District, 25 mi NW Polonnaruwa, 11-VI- 1975; Sil» Woodrc& Tk PettyaC +2) aN: Central Province, Ritigala Nat. Reserve, 8 mi NW Habarana 8-II-1962. Loc. 546:], swept on grass in forest, Lund University Ceylon Expedition 1962, Brinck-Anders- son-Cederholm (1 2). Puttalam District, 17 mi SE Puttalam, 18-VI-1975, S.L. Wood & J.L Petty (2 2). Ratnapura District, Uggal- kaltota, 350’, 20-VI-1976, K.V. Krombein, PB. Karunaratne, S. Karunaratne (1 Q); Gilimale, Induruwa Jungle, 5-7-11-1977, K.V. Krombein, P. Fernando, D.W. Bala- sooryia, V. Gunawardane (1 2). Trincom- alee District, China Bay Ridge Bungalow, 0-50 feet, 24-25-VIH-1978, K.V. Krombein, T. Wijesinhe, V. Kulasekare, L. Jayawick- rema (1 2, 2 d); Trincomalee, China Bay Ridge Bungalow, 25-50 ft., 26-II-1979, K.V. Krombein, T. Wijesinhe, S. Siriwar- dane, L. Jayawickrema, T. Gunawardane (2 2), 0-100’, 13-17-V-1976, K.V. Krombein, PB. Karunaratne, S. Karunaratne, D.W. Balasooryia (1 2), 16-17-V-1976, same collectors (1 2); Trincomalee, China Bay, 1-30 m, 8-11-X-1977, collected near ridge bungalow, K.V. Krombein, P.B. Karunarat- ne, P. Fernando, T. Wijesinhe, M. Jayaweera (1 ¢), in Malaise trap, same date and col- lectors (1 @). Type.—The type is presumably lost. It was not found in The Natural History Mu- seum, London (A. Shinohara, correspon- dence). Discussion.—Shuckard’s type was not found. His description is very brief: “*Ni- ger: scapo antennarum pedibusque 4 anticis rufo-testaceis: alis hyalinis, macula ad stig- ma brunnea. Long. 5% lin. Exp. alar. 9% lin.’ The sex is not given and cannot be determined since both sexes are colored similarly. The specimens from Sri Lanka agree with Shuckard’s description, and I re- gard them as A. signatus. This species ap- pears to be the most widely distributed spe- cies in Sri Lanka and is easily separated from the other two species by the black col- or with the reddish scape, pedicel, and fore- and midlegs and structural characters as given in the key and as illustrated. Pristaulacus krombeini Smith, new species (Figs. 9-12) Female.—Length, excluding ovipositor, 16 mm; forewing length, 13 mm; ovipositor length, 15 mm. Black; tegula reddish brown; anterior margin of clypeus, mandi- ble except apex, tarsi, and lateral and ven- tral portions of first metasomal segment slightly brownish. Forewing with anterior VOLUME 99, NUMBER 1 oO Nn Figs. 9-12. half black, posterior half of forewing and hindwing more hyaline to slightly blackish; veins and stigma black (Fig. 12). Head from above short and narrowing behind eyes (Fig. 9); with widely scattered fine punc- tures and silvery hairs, each more dense be- tween ocelli and antennae; occipital carina short. Propleuron shining with scattered punctures, similar to top of head. Pronotum with a forward projecting triangular tooth just above its midlength; anterolateral an- gles of mesonotum scarcely protruding and rounded in dorsal view, directed upward in lateral view, mesonotum with coarse trans- verse ridges; silvery hairs on mesosoma not obscuring sculpture (mesosoma as in Figs. 10, 11). Hindcoxa with coarse transverse Pristaulacus krombeini. 9, Head, dorsal. 10, Mesosoma, dorsal. 11, Mesosoma, lateral. 12, Wings. wrinkles laterally, posterior surface almost devoid of wrinkles. Tarsal claws with 4 teeth, including apical tooth, and a small acute basal lobe. Ovipositor length about 1.2 X forewing length. Male.—Unknown. Holotype.—Female, labeled “Sri Lanka: Tri. [Trincomalee] Dist., Tennamaravadi, 18-V-1976, K.V. Krombein, P.B. Karunar- atne, S. Karunaratne, D.W. Balasooryia.”’ In the National Museum of Natural History, Washington, D.C. Etymology.—Named for my colleague, Dr. Karl V. Krombein, Department of En- tomology, Smithsonian Institution, Wash- ington, D.C. Discussion.—The forewing with the an- 66 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON terior half infuscated and the posterior half lighter black to hyaline, entirely black col- oration, and structure of the mesonotum as shown in Figs. 10 and 11 separate this spe- cies from other Aulacidae described from southeastern Asia. A review of the descrip- tions of other species described from south- eastern Asia, especially those from northern India and Indochina, revealed various dis- tinguishing features from P. krombeini, es- pecially combinations of coloration, wing maculation, and mesonotum structure. These taxa include P. beesoni Turner, P. ir- idipennis (Cameron), P. nigripes Kieffer, P. rufobalteatus Cameron, P. rufobalteatus leviceps Kieffer, and P. bituberculatus (Cameron) from the Himalayas of northern India, and P. emarginaticeps Turner, P. ex- cisus Turner, P. nigripes var. duporti Kief- fer, P. tuburculiceps Turner, and P. tonki- nensis Turner from Indochina. ACKNOWLEDGMENTS I thank K.V. Krombein, Smithsonian I[n- stitution, for allowing study of material from the Ceylonese Insect Project, and the following for loaning types and other ma- terial: R. Danielsson, Museum of Zoology, Lund University, Lund, Sweden; S. Schédl, Naturhistorisches Museum Wien, Austria; and FE Koch, Zoologisches Museum an der Humboldt-Universitét zu Berlin, Germany. I am especially grateful to A. Shinohara, National Science Museum, Tokyo, Japan, who kindly compared photos with types and provided notes on the types of A. fla- vipennis during his visit to The Natural His- tory Museum, London, in 1995. LITERATURE CITED Cameron, P. 1887. Descriptions of one new genus and some new species of parasitic Hymenoptera. Pro- ceedings of the Manchester Literary and Philo- sophical Society 26: 117—136. Hedicke, H. 1939. Aulacidae. /n Hedicke, H., ed., Hy- menopterorum Catalogus, Pars 10. Dr. W. Junk, Gravenhage. 28 pp. Kieffer, J.-J. 1900. Note sur le genre Pristaulacus Kieff. [Hymén.]. Bulletin de la Société Entomo- logique de France 1900: 338-339. 1902. Hymenoptera, Fam. Evaniidae. In Wytsman, P., Genera Insectorum, Fascicule 2, 13 pp., | plate. Bruxelles. 1904. Les Evaniides. Jn André, E., Species des Hyménopteéres d’ Europe et d’ Algérie, Volume 7, part 2, pp. 347—482. Paris. 1912. Hymenoptera, Ichneumonidae, Evani- idae. Das Tierreich, Berlin, Volume 30, I-XIX + 431 pp. . 1921. Sur divers Hyménopteres destructeurs des Cérambycides nuisibles au Caféier et au Bam- bou. Bulletin Agricole de I’ Institut Scientifique de Saigon 3: 129-140. Schletterer, A. 1890. Die Hymenopteren-Gruppe der Evaniiden. Annalen des K. K. Naturhistorischen Hofmuseums (Separatabdruck aus Band IV), pp. 373-546, plates XIX-XXII. Shuckard, W. E. 1841. Art. XXII.—On the Aulacidae, a family of Hymenoptera pupivora; and that 77i- gonalys is one of its components: with the de- scription of a British species of this genus, and incidental remarks upon their collateral affinities. Entomologist 1: 115-125. Turner, R. E. 1919. On Indo-Chinese Hymenoptera collected by R. Vitalis de Salvaza.—lII. Annals and Magazine of Natural History (9) 4: 385-395. . 1922. XXXIV.—New Evaniidae and Bracon- idae in the British Museum. Annals and Magazine of Natural History (9) 10: 270-281. Westwood, J. O. 1844 (1843). XXXVIII. On Evania and some allied genera of Hymenopterous insects. Transactions of the Entomological Society of Lon- don 3: 237-278, plates XITV—XV. PROC. ENTOMOL. SOC. WASH. 99(1), 1997, pp. 67-88 NOTES ON LIFE HISTORIES AND DESCRIPTIONS OF ADULTS AND IMMATURE STAGES OF PROCECIDOCHARES KRISTINEAE AND P. LISAE NEW SPECIES (DIPTERA: TEPHRITIDAE) ON AMBROSIA SPP. IN SOUTHERN CALIFORNIA RICHARD D. GOEDEN AND JEFFREY A. TEERINK Department of Entomology, University of CA, Riverside, CA 92521, U.S.A. Abstract.—Procecidochares kristineae Goeden, n. sp., and P. lisae Goeden, n. sp., are described and figured as adults and distinguished from each other by their wing patterns, chaetotaxy, and genal color and by these and additional characters from all other North American congeners. Sexually dimorphic wing patterns in P. lisae are reported for the first time from this genus. First through third-instar larvae and puparia also are de- scribed and figured for both species, but few morphological differences were found. Pro- cecidochares kristineae and P. lisae are strictly monophagous tephritid flies forming ax- illary bud galls on branches of the native, shrubby, xerophytic ragweeds, Ambrosia du- mosa (Gray) Payne and A. eriocentra (Gray) Payne (Asteraceae), respectively. Both are circumnatal species that overwinter as first instars within incipient galls in southern Cal- ifornia. Further gall and larval development and host-plant regrowth and reproduction are stimulated concurrently by winter rainfall. Reproduction by P. kristineae is rarely biennial, usually univoltine or bivoltine, and rarely trivoltine, depending on local yearly rainfall patterns; whereas, P. /isae usually is univoltine or bivoltine. Key Words: Insecta, Procecidochares, Ambrosia, ragweed, biology, galls, taxonomy of adults and immature stages, sexual dimorphism, parasitoid Two undescribed species of Procecido- chares (Diptera: Tephritidae) were detected during faunistic surveys of native rag- weeds, Ambrosia spp. (Asteraceae: Ambro- Siinae), in southern California by Goeden and Ricker (1976a, b). The life history of one species forming galls on A. dumosa (Gray) Payne was studied by Silverman and Goeden (1980), but it has remained un- named until now. In this paper, both te- phritid species are named, their adult and immature stages are described and illustrat- ed, and new life-history information on each is reported. MATERIALS AND METHODS Our field studies on the tephritid infest- ing A. dumosa were conducted near sea lev- el in the low-elevation Colorado Desert (= northern Sonoran Desert) in southern Cali- fornia (Munz 1974) at various locations in Imperial, Riverside, and San Diego counties listed by Silverman and Goeden (1980) or mapped by Goeden and Ricker (1976a). Our principal study site for the fly on A. eriocentra (Gray) Payne was Mountain Pass at 1430-m elevation, San Bernardino Co., in the high-elevation, Mojave Desert; although this tephritid also was reared from galls obtained from surrounding locations named below in northeastern San Bernar- dino Co. during 1970—71 faunistic surveys (Goeden and Ricker 1976b). Galls contain- ing eggs, larvae and puparia were sampled most recently from A. dumosa near Snow 68 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Creek at 370-m elevation in the dry San Gorgonio River bed, Riverside Co., during February—April, 1993 and 1994, and from A. eriocentra at Mountain Pass during Feb- ruary—May, 1991-1995. Excised galls were transported in cold-chests in an air-condi- tioned vehicle to the laboratory and stored under refrigeration for subsequent dissec- tion, photography, description, and mea- surement. Twenty-one and 19 larvae and 4 and 3 puparia dissected from galls on A. dumosa and A. eriocentra, respectively, were preserved in 70% EtOH for scanning electron microscopy (SEM). All other fully grown larvae and puparia from A. eriocen- tra were placed in separate, glass shell vials stoppered with absorbant cotton and held in humidity chambers at room temperature for adult and parasitoid emergence. Specimens for SEM later were hydrated to distilled wa- ter in a decreasing series of acidulated EtOH. They were osmicated for 24 h, de- hydrated through an increasing series of acidulated EtOH, critically point dried, mounted on stubs, sputter-coated with a gold-palladium alloy, and studied with a JEOL JSM C-35 SEM in the Department of Nematology, University of California, Riv- erside. Most adults reared from isolated puparia obtained from galls on A. eriocentra were individually caged in 850-ml, clear-plastic, screened-top cages with a cotton wick and basal water reservoir and provisioned with a strip of paper toweling impregnated with yeast hydrolyzate and sucrose. These cag- ings were used for longevity studies and Oviposition tests in the insectary of the De- partment of Entomology, University of Cal- ifornia, Riverside, at 25 + 1°C, and 14/10 (L/D) photoperiod. Virgin male and female flies obtained from emergence vials as well as field-swept adults were paired in clear- plastic petri dishes provisioned with a flat- tened, water-moistened pad of absorbant cotton spotted with honey (Headrick and Goeden 1991) for direct observations, vi- deorecording, and still-photography of their courtship and copulation behavior. Plant names used in this paper follow Munz (1974); tephritid nomenclature and adult terminology follow Foote et al. (1993). Format used to describe the adults follows Blanc and Foote (1961). Morpho- logical terminology and telegraphic format used to describe the immature stages follow Goeden and Headrick (1992), Headrick and Goeden (1990, 1993), Goeden et al. (1994a, b, 1995a, b), Goeden and Teerink (1996a, b, c), Headrick et al. (1995), and our other works cited therein. Means + SE are used throughout this paper. The holotypes, allo- types, and five individually reared para- types of each sex of both new species have been deposited in the National Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM). Six additional, individually reared paratypes of each sex of both species also were deposited in the col- lection of the California Academy of Sci- ences (CAS). The holotype, allotype, and 22 paratypes used for measurements to de- scribe P. kristineae originated from one col- lection in 1980 at the Palm Springs study site used by Silverman and Goeden (1980); the same numbers of types were used to describe P. lisae from one collection in 1981 at Mountain Pass. All remaining para- types and voucher specimens not designat- ed as paratypes and reared parasitoids of both tephritids reside in the research collec- tions of RDG; preserved specimens of lar- vae and puparia are stored in a separate col- lection of immature Tephritidae maintained by JAT. RESULTS AND DISCUSSION TAXONOMY The new species from A. dumosa de- scribed below as P. kristineae is most sim- ilar to P. stonei Blanc and Foote, which it was first misidentified as (Silverman and Goeden 1980). However, adults of these two species are distinguished below mainly on the basis of their wing characters, in- stead of those often lacking in swept or pre- served specimens, e.g., distribution patterns VOLUME 99, NUMBER 1 of setae on the scutum, consistent with the philosophy embodied in the key to U.S. and Canadian species of Procecidochares in Foote et al. (1993). The new species from A. eriocentra described as P. lisae also is distinguished below primarily by wing characters. Accordingly, the following key couplets replacing couplets 1 and 5 in the key of Foote et al. (1993) enable one to distinguish these two new species: 1. Pterostigma lighter brown in basal fifth to half than in apical part; sometimes a second orbi- talbrisilo mee eer eat cents. cheer ese la — Pterostigma evenly brown; | pair orbital bris- WES A ards ot 884 youn doen cede taker Ok eran 2. la. Two orbital bristles, the second pair very small and delicate; basal and discal bands SCPalale wt ems ies alsa flavipes Aldrich — One pair of orbital bristles; basal and discal bands usually connected 1b. Gena with prominent dark brown spot at ven- tral margin of eye; discal band not extended posterior to vein A, + CuA, in male, usually crossing it in female, but fading towards pos- terior wing margin; basal and discal bands broadly connected in cells c, sc, br, and bm + Bug abt OfdLc sei aA OTE ENS ee lisae Goeden, n. sp. — Gena with ocherous to pale brown spot at ventral margin of eye; discal band extending distinctly to posterior wing margin in both sexes; basal and discal bands narrowly con- nected in cell sc or br . . kristinae Goeden, n. sp. Procecidochares kristineae Goeden, new species (Figs. 1, 3-7) Procecidochares stonei Blanc and Foote: Wasbauer 1972: 7 (in part, Ambrosia du- mosa host record). Procecidochares n. sp.: Goeden and Ricker 1976a: 49 (host record).—Silverman and Goeden 1980: 283-288 (host, gall de- scription, California distribution, biology, behavior, seasonal history, parasitoids, predators, gall inquiline).—Foote et al. 1993: 318 (taxonomic status) Female (Holotype).—Head: In profile 0.6 to 0.7 times as long as high, face and frons meeting at an angle of about 120°; parafacial 0.75 times as wide as third an- tennal segment; gena about 0.2 times as high as eye, which is 0.5 to 0.6 as wide as 69 high; frons ocherous brown to yellow or white, at vertex 1.4 to 1.6 times as wide as eye in lateral view, 1.2 to 1.3 times as wide as length from vertex to lunule; lunule half as high as its width between the antennae; face yellow to white, pollinose, concave, but raised medially and protruding slightly at middle of oral margin; antenna yellow, pollinose, third segment sometimes ocher- ous brown along anterior margin and apex, arista ocherous brown to black, lightest ba- sally. Usually 3-4 frontal bristles (rarely 5 or 6), all black; one pair of black orbital bristles; black genal bristle slender, situated immediately below lower curvature of eye; all postoculars white. Thorax: Pleuron mostly shining dark brown to black, especially the katepister- num, but with a very sparse pollinosity on anepisternum; katepisternal bristle black, most other pleural setae rather long and white; wing base and anepimeron densely silver pollinose over a dark brown to black ground-color; lateral third of mesonotum, including postpronotal lobe, shining dark brown to black; a wide, silvery pollinose stripe on a shiny black ground-color occu- pying median third of mesonotum from an- terior margin nearly to scutoscutellar suture, slightly widened at transverse suture and in- vested with short, white, blunt setae (some- times appearing pale yellow) as follows: scattered + uniformly over median polli- nose strip, except for anterior half of pre- sutural part of scutum, where confined to center and margins of strip, also encircling the lateral third of presutural part of scutum including along the transverse suture, and trailing posteriolaterad and posteriomediad in separate rows 3 to 4 setae-wide, the outer row narrowing and crossing the scutoscu- tellar suture to end in a cluster of 5 to 12 setae at base of anterior scutellar bristle. One dorsocentral bristle situated about half- way between transverse suture and level of the postsutural supra-alar bristle, and locat- ed on margin of median pollinose area; cluster of 5 to 8 short, white setae anterior- ad of base of postsutural supra-alar bristle. 70 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Right wings of Procecidochares kristineae: Fig. 1. (A) female; (B) male. Scutellum shining dark brown to black, bul- bous, two pairs scutellar bristles; postscu- tellum dark brown to black, pollinose. Hal- tere shaft ocherous yellow, slightly darker than knob. Femora of fore, mid, and hind legs dark brown but with yellow extremi- ties, the remainder of legs yellow to ocher- ous yellow. Wing pattern as in Fig. 1A, B; pterostigma about 2.0 times as long as wide, lighter in basal fifth to third; basal and discal bands narrowly joined in cell br and/or sc, subapical and apical bands usu- ally joined in cell(s) r, and/or r,,, (Fig. 1A) or sometimes narrowly separated (Fig 1B) and only joined by darkened portions of veins R,,, and R,,;; distance between crossveins r-m and dm-cu measured along vein M about equal to length of r-m. Abdomen: Dark brown to black, each ter- gum covered with white, flattened setae progressively increasing in length posteri- orly. Oviscape shining dark brown to black, basal half slightly swollen, about half as long as all terga, except the first, taken to- gether. Male (allotype)——Head: Like &, but parafacial 0.67 times as wide as third an- tennal segment; frons at vertex 1.1 to 1.3 times as wide as one eye, 1.1 to 1.4 times as wide as distance from vertex to lunule; lunule 0.5 to 0.7 times as high as its width at antennal base. Thorax: Like 2°. Abdo- men: Like 2, only external genitalia dark brown to black. Variation.—Examination of P. kristineae specimens with mostly intact setation con- firmed variation in numbers of frontal setae, noted for the genus to vary from two to five by Foote et al. (1993). The holotype and 19 ? paratypes (68%) of a total of 28 2 types examined had three pairs of frontal setae; one pair of these setae was white in two &, and one pair was reduced in length and thickness in another 2. One @ had four pairs of frontal setae; another 2 had five pairs. The frontal setae were not strictly paired in the remaining six @, as four @ had three and four frontal setae on different sides of their heads; and each of two &, four and five frontal setae located asym- metrically. The allotype and 32 d paratypes (64%) of a total of 50 d types examined had three pairs of frontal setae. Three d (6%) had four pairs of frontal setae. Among the remaining 15 6d, nine (18%) had three and four frontal setae on different sides of their heads, four d (16%) each had three and five frontal setae, and one d each had four and five or four and six frontal setae located asymmetrically. Regarding the wings, the subapical and apical bands also showed variation. The ho- lotype and 14 @ paratypes (48%) of a total of 31 2 types with intact wings had the subapical and apical bands broadly joined (Fig. 1A); 7 2 paratypes (23%) had the subapical and apical bands narrowly joined in at least one wing (in the other wing in 5 of these 2, these bands were narrowly sep- arated, Fig. 1B); and in the remaining 9 2 (29%), these bands were narrowly separat- ed in both wings. The allotype and 25 d VOLUME 99, NUMBER 1 types (49%) of a total of 51 d paratypes with intact, fully formed wings had the sub- apical and apical bands broadly joined (Fig. 1A); 8 3d paratypes (16%) had these bands narrowly joined in at least one wing (in the other wing in 1 d, these bands were nar- rowly separated); and in the remaining 18 3d paratypes (35%), these bands were nar- rowly separated in both wings. Diagnosis.—The two main morphologi- cal characters distinguishing the adults of P. kristineae are the pterostigma being lighter brown in the basal fifth to third than in the apical part (Fig. 1), in combination with one pair of orbital setae. All types of P. kristi- neae possessed these two characters. The former character is shared only with P. fla- vipes and P. lisae (see below); the latter character distinguishes P. flavipes which has two pairs of orbital setae (Foote et al. 1993). Procecidochares kristineae and P. lisae are distinguished below. In the partial key provided above, if the pterostigmal character is missed P. kristineae will run to P. stonei, the only other species with basal and discal bands connected. Most P. kris- tineae differ from P. stonei by having the subapical and apical bands broadly to nar- rowly joined. Of 179 reared voucher specimens of P. stonei with fully formed wings in the re- search collection of RDG (Green et al. 1993), 14 (8%) had a pterostigma in at least one wing that was light basally, but usually this basal area was small and only partially extended posteriorad across the cell, nor were the subapical and apical bands in the wings of these 14 flies joined. Similarly, an- other 12 voucher specimens of P. stonei had at least one wing with the subapical and apical bands joined, but all of these flies had pterostigmas that were evenly brown. Additional biological and ecological char- acteristics that distinguish P. kristineae and P. stonei are discussed below. Types.—Holotype, 2; 8 km NE of Palm Springs, 250-m elevation, Riverside Co., Galiformiay 2641198 1:,.R..;D...Goeden, coll.(hereafter, RDG, coll.); reared from ax- val illary bud gall on A. dumosa [at study site of Silverman and Goeden (1980)] (USNM); allotype, ¢6, same data as holotype (USNM). Paratypes: CALIFORNIA: 12 ¢ and 24 2; same data as holotype (5 d and 5 2 to USNM, 6 ¢ and 6 2 to CAS). 4 6 and 3 2; same location data as holotype; 10.11.1971; RDG and D.W. Ricker, coll. (hereafter, RDG & DWR, coll.). 1 2; Twen- tynine Palms, San Bernardino Co.; 25.iii.1970; RDG & DWR, coll. 4 6 and 1 2; Llano, San Bernardino Co.; 30.iv.1970; RDG & DWR, coll. 6 d and 1 2; Yucca Valley, San Bernardino Co.; 4.111.1971; RDG & DWR, coll. | 2; Borrego Springs, San Diego Co., 1.11.1973; RDG & DWR, coll. 5 ¢ and 7 2; Valliceto Valley, S end of Smugglers Canyon at 442 m, San Diego Co.; 11.11.1993; RDG and J.A. Teerink, coll. (hereafter, RDG & JAT, coll.).5 ¢ and 4 2; Snow Creek at 370 m, Riverside Co.; 7.iv.1994; RDG & JAT, coll. 3 6 and 3 &; Ocotillo, Imperial Co., 2.11.1995; RDG & JAT, coll. (Remaining 39 d and 20 & para- types along with numerous swept and/or damaged-reared, voucher and nonvoucher specimens examined and identified are held in research collection of RDG). Etymology.—Procecidochares kristineae is named for my younger daughter, Kristine Louise Gilbert (nee Goeden), mother of my two grandsons, Samuel Vanderpoel Gilbert V and Nikolaus Richard Gilbert. Procecidochares lisae Goeden, new species (Figs. 2, 8-12) Procecidochares n. sp.: Goeden and Ricker (1976b): 927 (host record). Female (holotype).—Head: In profile 0.5 to 0.6 times as long as high, face and frons meeting at an angle of about 120°; parafa- cial as wide as third antennal segment; gena with prominent, shiny, dark-brown spot from lower margin of eye to genal groove, gena 0.1 to 0.2 times as high as eye, which is 0.4 to 0.5 as wide as high; frons ocherous yellow to white, at vertex 1.3 to 1.7 times M2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON as wide as eye in lateral view, 1.3 to 1.6 times as wide as length from vertex to lu- nule; lunule 0.4 to 0.6 times as high as its width between the antennae; face pale yel- low to white, pollinose, concave, but raised medially; antenna mostly brown, pollinose, apex of second segment and sometimes posterior margin of third segment yellow, arista ocherous yellow to brown, lightest basally. Usually, 3—4 frontal bristles (rarely 2 or 5), all black; one pair of black orbital bristles; genal bristle black, arising behind brown part of genal margin, which is ex- tension of brown spot (noted above) below genal groove; all postoculars white. Thorax: Pleuron mostly shining dark brown to black, especially the katepister- num, but with a very sparse pollinosity on anepisternum; katepisternal bristle black, most other pleural setae rather long and white; wing base and anepimeron densely silver pollinose over a dark brown to black ground-color; lateral third of mesonotum, including postpronotal lobe, shining dark brown to black; a wide, silvery pollinose stripe on a shiny black ground-color occu- pying median third of mesonotum from an- terior margin nearly to scutoscutellar suture, slightly widened at transverse suture and in- vested with short, white, blunt setae (some- times appearing pale yellow) as follows: scattered + uniformally over median polli- nose strip, except for anterior half of pre- sutural part of scutum, where confined to center and margins of strip, also encircling the lateral third of the presutural part of scu- tum including along the transverse suture, encircling the anterolateral two-thirds of the scutum, and trailing posteriolaterad and crossing the scutoscutellar suture to end in a cluster of 8 to 18 setae at base of anterior scutellar bristle. One dorsocentral bristle situated just anterior to a line between the postsutural supra-alars and slightly laterad of the margin of median pollinose area. Scutellum shining dark brown to black, bul- bous, two pairs scutellar bristles; postscu- tellum dark brown to black, pollinose. Hal- tere shaft ocherous yellow, slightly darker B Right wings of Procecidochares lisae: (A) Fig. 2. female; (B) male. than knob. Femora of fore, mid, and hind legs dark brown but with ocherous yellow extremities, the remainder of legs yellow to ocherous yellow. Wing pattern as in Fig. 2A pterostigma about 2.0 times as long as wide, lighter in basal fifth to half; basal and discal bands broadly joined in cells c, sc, and br; discal band usually shortened, crossing cell cua, along and distal to vein CuA,, and nearly or just touching vein A, + CuA, growing faint if extending into anal lobe; subapical and apical bands usually separated, or sometimes narrowly joined in cell(s) 1, and/or 1,5; distance’ between crossveins r-m and dm-cu measured along vein M about equal to length of r-m. Abdomen: Dark brown to black, each ter- gum covered with white, flattened setae progressively increasing in length poste- riorad. Oviscape shining dark brown to black, basal half slightly swollen, about half as long as all terga, except the first, taken together. VOLUME 99, NUMBER I Male (allotype)—Head: Like @, but about 0.6 times as long as high, face and frons meeting at an angle of about 110°; parafacial 0.67 times as wide as third an- tennal segment; eye 0.5 to 0.6 times as wide as high; frons at vertex 1.1 to 1.5 times as wide as eye, 1.3 to 1.7 times as wide as length from vertex to lunule; lunule 0.5 to 0.7 times as high as its width at antennal base. Thorax: Like 2, but discal band of wing rarely extending into anal lobe and usually ending before and not touching vein A, + CuA,; subapical and apical bands usu- ally separated, but sometimes narrowly or widely joined in cell(s) r, and/or r,,, (Fig. 2B). Abdomen: Like @, only external gen- italia dark brown to black. Variation.—Examination of P. lisae specimens with mostly intact setation fur- ther confirmed variation in numbers of frontal setae for the genus (Foote et al. 1993). The holotype and 30 @ paratypes (50%) of a total of 62 2 types examined had three pairs of frontal setae. Six @ para- types (13%) each had four pairs of frontal setae; another @ had five pairs. The frontal bristles were not paired in the remaining 22 2, as 19 2 had three and four frontal setae on different sides of their heads; two 9 had two and three frontal setae located asym- metrically; and one @ had three and six frontal setae so located. The allotype and 25 6d paratypes (53%) of a total of 47 ¢ types examined had three pairs of frontal setae. Eight d paratypes (13%) each had four pairs of frontal setae. Among the re- maining 13 d paratypes, nine (19%) had three and four frontal setae on different sides of their heads; two d¢ (4%) each had two and three frontal setae, and one d each had three and five or four and five frontal setae located asymmetrically. Regarding the wings, the relationship of the subapical and apical bands showed vari- ation. The holotype and 48 @ paratypes (69%) of a total of 71 @ types with intact wings had the subapical and apical bands fully separated (Fig. 2A); 13 2 paratypes (18%) had the subapical and apical bands Us: narrowly separated; and in the remaining 9 2 (13%), these bands were narrowly or widely joined in cell r, in eight 2 and one 2, respectively. The allotype and 21 ¢d paratypes (45%) of a total of 49 d types with intact, fully formed wings had the sub- apical and apical bands fully separated (Fig. 2B); 19 ¢ paratypes (39%) had these bands narrowly separated in at least one wing (in the other wing in four d, these bands were narrowly joined in cell r, or r,,,); and in the remaining eight d paratypes (16%), these bands were broadly joined in four 6 or nar- rowly joined in four ¢ in cell r, or r5,3. In the holotype and 35 @& paratypes (51%) of the same 71 @ types with fully developed wings, the discal band extended across vein A, + CuA, into the anal lobe in both wings (Fig. 2A); in 27 2 paratypes (37%), this band ended at this vein in both wings; in two 2 paratypes (3%) each wing showed a different one of these two char- acters. In only three (4%) of the remaining ? paratypes, the discal band ended before and did not reach vein A, + CuA,. In the allotype and 39 ¢d paratypes (80%) of the same 49 ¢ types with intact, fully formed wings, the discal band ended before and did not touch vem Ay -- CuwA> (Fics 2B): or reached this vein in nine (18%) other 3 paratypes, but crossed this vein into the anal lobe in only one d paratype. These data, therefore, provide the first document- ed incidence of sexual dimorphism in the wing pattern of a species of Procecido- chares (Foote et al. 1993); although, this dimorphism pales in comparison with wing pattern sexual dimorphism recently report- ed by us for certain species of Aciurina (Goeden and Teerink 1996a, b, c), or as long known for certain Trupanea spp. (Foote, ctjala 1993): Diagnosis.—The incomplete discal band distinguishes P. lisae from all previously described species of Procecidochares. Two other characters distinguishing the adults of P. kristineae and P. lisae from other Pro- cecidochares spp. are the pterostigma being lighter brown in the basal fifth to half than 74 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON in the apical part, in combination with one pair of orbital setae. These species can then be readily separated on the basis of their wing patterns and the color of the genal spot as described in the partial key provided above. The pterostigmal character is shared only by P. flavipes, P. kristineae, and P. lisae, with P. flavipes separated by its pos- session of two pairs of orbital setae. The basal and discal bands are joined only in the wings of P. kristineae, P. lisae, and P. stonei, with P. lisae readily distinguished by the broad anterior juncture of these bands and by the shortened discal bands. Types.—Holotype, 2; Mountain Pass, 1430-m elevation, NE San Bernardino Co., California; 30.iv.1981; RDG, coll.; reared from axillary bud gall on A. eriocentra (USNM); allotype, 6, same data as holo- type (USNM). Paratypes: CALIFORNIA: 17 3 and 17 92; same data as holotype (5 6 and5 ¢ to USNM, 6 6 and 6 2 to CAS). 5 6 and 1 2,2 6 and 2 2; same location data as holotype; 19.iv.1971, 26.vi.1971, re- spectively, RDG & DWR, coll. 5 3 and 3 2,2 2,106 and 11 2; same location data as ‘holotype; 9:-v- 1991. 9:v.1992. 9.v. 1993; respectively; RDG & JAT, coll. 8 ¢ and 15 ?; Halloran Springs, NE San Bernardino Go 191iv.1971; RDG PWR colly 372: Halloran Springs, NE San Bernardino Co.; 291v. 1981) RDGcoll” 12'S and ie es Hackberry Mountain, NE San Bernardino Co-; 220% 1970.4 111-197 1 respectively: RDG & DWR, coll. 3 3d and 4 @; Granite Cove, NW San Bernardino Co., 25.v.1971; RDG & DWR, coll. (Remaining 36 d and 50 @ paratypes along with numerous swept and/or damage-reared, voucher and non- voucher specimens examined and identified are held in research collection of RDG). Etymology.—Procecidochares lisae is named for my older daughter, Lisa Marie Goeden, a unique woman of many accom- plishments. IMMATURE STAGES Procecidochares kristineae.—Egg: Eggs of P. kristineae (Fig. 3A, B), were de- Fig. 3. itus; (B) anterior end, aeropyles. Egg of Procecidochares kristineae (A) hab- scribed by Silverman and Goeden (1980). Chorion smooth (Fig. 3A); pedicel with few aeropyles (Fig. 3B). Third instar: Superficially smooth, elon- gate cylindrical, tapering anteriorly, lacking minute acanthae (Fig. 4A); gnathocephalon conical, broad dorsally, smooth with few rugose pads (Fig. 4B); paired dorsal sen- sory organs dorsomediad of anterior sen- sory lobes, consisting of dome-shaped pa- pilla (Fig. 4B-1, 4C-1); anterior sensory lobes bear terminal sensory organ (Fig. 4C-2), pit sensory organ (Fig. 4C-3), lateral sensory organ (Fig. 4C-4), and supralateral sensory organ (Fig. 4C-5); stomal sense or- gans ventrad of anterior sensory lobe (Fig. 4B-2); lateral sensillum ventrolaterad of stomal sense organs (Fig. 4B-3); mouth hooks tridentate, teeth conical, stout (Fig. 4B-4); median oral lobe fleshy, tapering an- teriorly, attached to labial lobe (Fig. 4B-5); labial lobe with two pore sensilla; ventro- VOLUME 99, NUMBER 1 wo the ; J {sku xie0@ oo1e “f8.00 Fig. 4. 15KUY K2666 6616. 1a. au ISKU 41166 8626 Third instar of Procecidochares kristineae: (A) habitus, anterior to left; (B) gnathocephalon, anterior view, 1—dorsal sensory organ, 2—stomal sense organ, 3—lateral sensillum, 4—mouth hooks, 5—median oral lobe, 6—ventrolateral sensillum; (C) anterior sensory lobe, 1—dorsal sensory organ, 2—terminal sensory organ, 3—pit sensory organ, 4—lateral sensory organ, 5—supralateral sensory organ; (D) anterior thoracic spiracles; (E) second abdominal lateral spiracular complex, 1—spiracle, 2—verruciform sensillum; (F) posterior spiracular plate, 1—rima, 2—interspiracular process. lateral sensillum ventrolaterad of mouth lu- men (Fig. 4B-6); prothorax smooth, verruciform sensilla circumscribe dorsal half of anterior margin, anterior thoracic spiracles on posterior margin consist of three ovoid papillae (Fig. 4D); meso- and metathoracic lateral spiracular complexes consist of an open spiracle; abdominal lat- eral spiracular complex consist of an open spiracle (Fig. 4E-1) and a single verruci- form sensillum (Fig. 4E-2); caudal segment bears posterior spiracular plates (Fig. 4F); PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON BSKY K2680 15KU x2688 @41i3 Fig. 5. 15KU ¥3660 6815 Second instar of Procecidochares kristineae: (A) gnathocephalon, anterior view, 1—dorsal sensory organ, 2—stomal sense organ, 3—mouth hooks, 4—ventrolateral sensillum; (B) anterior sensory lobe, 1—dorsal sensory organ, 2—terminal sensory organ, 3—lateral sensory organ, 4—supralateral sensory organ; (C) gnath- ocephalon, anteriolateral view, 1—mouth hooks, 2—median oral lobe; (D) anterior thoracic spiracles; (E) pos- terior spiracular plates, 1—rima, 2—interspiracular process; (F) compound sensillum, 1—verruciform sensillum, 2—-stelex sensillum. posterior spiracular plates with three ovoid rimae, ca. 0.024 mm in length (Fig. 4F-1), and four spiniform interspiracular process- es, longest measuring 0.01 mm in length (Fig. 4F-2); compound sensilla ventrad of posterior spiracular plates consist of a ver- ruciform sensillum and a stelex sensillum. Second instar: Superficially smooth, bar- rel-shaped, rounded anteriorly and posteri- orly; gnathocephalon conical, smooth with few rugose pads (Fig 5A); paired dorsal sensory organs consist of a dome-shaped papilla (Fig. 5A-1, B-1); anterior sensory lobe bears terminal sensory organ (Fig. VOLUME 99, NUMBER | 15KY x8 6@ae LT Fig. 6. First instar of Procecidochares kristineae: (A) gnathocephalon, anteriolateral view, 1—dorsal sensory organ, 2—mouth hooks; (B) anterior sensory lobe, 1—dorsal sensory organ, 2—terminal sensory organ, 3—pit sensory organ, 4—lateral sensory organ, 5—supralateral sensory organ; (C) gnathocephalon, anteriolateral view, 1—mouth hooks, 2—median oral lobe; (D) caudal segment, posterior spiracular plates, 1—rima, 2—interspi- racular process, 3—stelex sensillum. 5B-2), pit sensory organ, lateral sensory or- gan (Fig. 5B-3), and supralateral sensory organ (Fig. 5B-4); stomal sense organ ven- trad of anterior sensory lobe (Fig. 5A-2); mouth hooks tridentate, teeth conical, apical tooth longest (Fig. SA-3, C-1); median oral lobe laterally flattened, rounded apically (Fig. SC-2); ventrolateral sensillum ventro- laterad of mouth lumen (Fig. 5A-4); ante- rior thoracic spiracles on posterior margin of prothorax, consist of three ovoid papillae (Fig. 5D); lateral spiracular complex not observed; caudal segment bears posterior spiracular plates (Fig. 5E); posterior spirac- ular plates with three ovoid rimae, ca. 0.009 mm in length (Fig. SE-1) and four spini- form interspiracular processes, longest mea- suring 0.005 mm (Fig. 5E-2); compound sensilla ventrad of posterior spiracular plates, consist of a verruciform sensillum (Fig. 5F-1), and a stelex sensillum (Fig. 5F-2). First instar: Superficially smooth, barrel- shaped, rounded anteriorly and posteriorly, minute acanthae circumscribe segmental lines; gnathocephalon conical, smooth, lacking rugose pads (Fig. 6A); dorsal sen- sory organ consists of a dome-shaped pa- pilla (Fig. 6A-1, B-1); anterior sensory lobes bear terminal sensory organ (Fig. 6B-2), pit sensory organ (Fig. 6B-3), lateral sensory organ (Fig. 6B-4), and supralateral sensory organ (Fig. 6B-5); stomal sense or- gans indistinct; mouth hooks bidentate, api- cal tooth with v-shaped frontal groove along entire length (Fig. 6A-2, 6C-1); me- dian oral lobe rounded apically (Fig. 6C-2); anterior thoracic spiracles absent; lateral 78 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON spiracular complexes not observed; caudal segment bears posterior spiracular plates (Fig. 6D); posterior spiracular plates with two ovoid rimae, ca. 0.002 mm (Fig. 6D-1), and four ovoid interspiracular processes, longest measuring 0.001 mm (Fig. 6D-2); stelex sensillum ventrad of posterior spirac- ular plates (Fig. 6D-3). Puparium: Superficially smooth, elon- gate-ellipsoidal (Fig. 7A); anterior end bears invagination scar (Fig. 7B-1), and an- terior thoracic spiracles (Fig. 7B-2); caudal segment bears posterior spiracular plates (Fig. 7C); posterior spiracular plates with three ovoid rimae, ca. 0.03 mm in length (Fig. 7C-1), and four spiniform interspirac- ular processes, longest measuring 0.01 mm (Fig. 7C-2). Measurements of P. kristineae puparia were provided by Silverman and Goeden (1980). Procecidochares lisae.—Egg: Fifty ova of P. lisae were white, opaque, smooth; with an elongate-ellipsoidal body, 0.57 + 0.005 (range, 0.52—0.64) mm long, 0.17 + 0.002 (range, 0.16—0.20) mm wide, smooth- ly rounded at tapered posterior end; and with a short, peg-like anterior pedicel, 0.02 mm long by 0.03 wide. Third instar: Superficially smooth, elon- gate cylindrical, tapering anteriorly, lacking minute acanthae (Fig. 8A); gnathocephalon conical, broad dorsally, smooth with few rugose pads (Fig. 8B); paired dorsal sen- sory organs dorsomediad of anterior sen- sory lobes, consisting of dome-shaped pa- pilla (Fig. 8B-1); anterior sensory lobes bear terminal sensory organ (Fig. 8B-2), pit sensory organ (Fig. 8B-3), lateral sensory organ (Fig. 8B-4), and supralateral sensory organ (Fig. 8B-5); stomal sense organs ven- trad of anterior sensory lobe; lateral sensil- lum ventrolaterad of stomal sense organs (Fig. 8B-6); mouth hooks tridentate, apical tooth longest (Fig. 8C-1); median oral lobe fleshy, laterally flattened, tapering anterior- ly, (Fig. 8C-2); ventrolateral sensillum ven- trolaterad of mouth lumen (Fig. 8C-3); pro- thorax smooth, verruciform sensilla circum- scribe dorsal half of anterior margin; ante- ; a0. BB ; ~ ype A 9024 160. 6U Cc “TSKY ¥320 Fig. 7. Puparium of Procecidochares kristineae: (A) habitus, anterior to left; (B) anterior end, 1—in- vagination scar, 2—anterior thoracic spiracles; (C) cau- dal segment, posterior spiracular plates, |—rima, 2— interspiracular process. rior thoracic spiracles consist of three ovoid papillae (Fig. 8D); meso- and metathoracic lateral spiracular complexes consist of an open spiracle; abdominal lateral spiracular complex consist of an open spiracle (Fig. 8E-1) and a single verruciform sensillum (Fig. 8E-2); caudal segment bears posterior VOLUME 99, NUMBER | 15K A, \ b aid 15KU.x1600 ° ‘9019 Fig. 8. i9. au E Third instar of Procecidochares lisae: (A) habitus, anterior to left; (B) gnathocephalon, anteriolateral 79 @aa4 \ 16. aU SKY K1146 view, 1—dorsal sensory organ, 2—terminal sensory organ, 3—pit sensory organ, 4—lateral sensory organ, 5— supralateral sensory organ, 6—lateral sensillum; (C) gnathocephalon, anterior view, |—mouth hooks, 2—median oral lobe, 3—ventrolateral sensillum; (D) anterior thoracic spiracles; (E) first abdominal lateral spiracular com- plex, 1—spiracle, 2—verruciform sensillum; (F) posterior spiracular plate, 1—rima, 2—interspiracular process. spiracular plates; posterior spiracular plates with three ovoid rimae, ca. 0.025 mm in length (Fig. 8F-1), and four spiniform in- terspiracular processes, longest measuring 0.01 mm in length (Fig. 8F-2); compound sensilla ventrad of posterior spiracular plates consist of two verruciform sensilla. Second instar: Superficially smooth, cy- lindrical, rounded anteriorly and posterior- ly; gnathocephalon conical, smooth with few rugose pads; paired dorsal sensory or- gans consist of a dome-shaped papilla (Fig. 9A-1); anterior sensory lobe bear terminal sensory organ (Fig. 9A-2), pit sensory or- 80 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON aga2°7 18.8 \ 15SkKU X240o Fig. 9. Second instar of Procecidochares lisae: (A) gnathocephalon, anterior view, 1—dorsal sensory organ, 2—terminal sensory organ, 3—pit sensory organ, 4—lateral sensory organ, 5—supralateral sensory organ; (B) gnathocephalon, anteriolateral view, 1—stomal sense organ, 2—median oral lobe, 3—mouth hooks; (C) anterior thoracic spiracles; (D) caudal segment, posterior spiracular plates, 1—rima, 2—interspiracular process. gan (Fig. 9A-3), lateral sensory organ (Fig. 9A-4), and supralateral sensory organ (Fig. 9A-5); stomal sense organ ventrad of an- terior sensory lobe (Fig. 9B-1); mouth hooks tridentate, teeth conical (Fig. 9B-3); median oral lobe fleshy, laterally flattened (Fig. 9B-2); anterior thoracic spiracles on posterior margin of prothorax, consist of two-three ovoid papillae (Fig. 9C); lateral spiracular complex not observed; caudal segment bears posterior spiracular plates; posterior spiracular plates with three ovoid rimae, ca. 0.009 mm in length (Fig. 9D-1) and four spiniform interspiracular process- es, longest measuring 0.004 mm (Fig. 9D-2); compound sensilla ventrad of pos- terior spiracular plates were obscured in prepared specimens. First instar: Superficially smooth, cylin- drical, tapering anteriorly, rounded posteri- orly, minute acanthae circumscribe segmen- tal lines (Fig. 10A); gnathocephalon coni- cal, smooth, lacking rugose pads (Fig. 10B); dorsal sensory organs large, dome-shaped (Fig. 1OB-1, C-1); anterior sensory lobes bear terminal sensory organ (Fig. 10C-2), pit sensory organ (Fig. 10C- 3), lateral sensory organ (Fig. 10C-4), and supralateral sensory organ (Fig. 10C-5); stomal sense organs indistinct; mouth hooks bidentate, apical tooth with v-shaped frontal groove along midline (Fig. 1OB-2, D-1); ventrolateral sensillum ventrolaterad of mouth lumen (Fig. 1OB-3, D-2); anterior thoracic spiracles absent; lateral spiracular complexes not observed; caudal segment bears posterior spiracular plates (Fig. 10E- 1); posterior spiracular plates with two ovoid rimae, ca. 0.002 mm in length (Fig. 10F-1), and four, very reduced spiniform VOLUME 99, NUMBER | 15KY $1 Fig. 10. First instar of Procecidochares lisae: (A) habitus, anterior to left; (B) gnathocephalon, lateral view, 1—dorsal sensory organ, 2—mouth hooks, 3—ventrolateral sensillum; (C) gnathocephalon, 1—dorsal sensory organ, 2—terminal sensory organ, 3—pit sensory organ, 4 lateral sensory organ, 5—supralateral sensory organ; (D) gnathocephalon, ventral view, 1—mouth hooks, 2—ventrolateral sensillum; (E) caudal segment, 1—posterior spiracular plates, 2—verruciform sensillum, 3—compound sensillum, verruciform sensillum; (F) posterior spi- racular plates, 1—rima, 2—interspiracular process. interspiracular processes, longest measuring 0.001 mm (Fig. 1OF-2); two verruciform sensilla dorsolaterad of posterior spiracular plates (Fig. 1|OE-2); compound sensilla ven- trad of posterior spiracular plates consist of two verruciform sensilla (Fig. 1OE-3). Puparium: Superficially smooth, elon- gate-ellipsoidal (Fig. I1A); anterior end bears invagination scar (Fig. 11B-1), and anterior thoracic spiracles (Fig. 11B-2); caudal segment bears posterior spiracular plates, each with three ovoid rimae, ca. 0.04 82 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 15KV Ace 9 6665 166. 68U | & ISKY K266 Fig. 11. Puparium of Procecidochares lisae: (A) habitus, anterior to left; (B) anterior end, 1—invagi- nation scar, 2—anterior thoracic spiracle; (C) caudal segment, 1—rima, 2—interspiracular process. mm in length (Fig. 11C-1), and four thorn- like interspiracular processes, longest mea- suring 0.007 mm (Fig. 11C-2). Fifty-six pu- paria averaged 4.01 + 0.05 (range, 3.24- 4.60) mm in length; 1.83 + 0.03 (range, 1.40-2.35) mm in width. Procecidochares kristineae and P. lisae larvae are morphologically very similar. The only difference noted was the com- pound sensilla ventrad of the posterior spi- racular plates; in P. kristineae there was a verruciform sensillum and a stelex sensil- lum, but both sensilla in P. lisae were ver- ruciform. Thus, morphological differentia- tion between these two, possibly sibling species on separate, closely related hosts is minimal between their larvae, but distinc- tive between their adults. Similar morpho- logical differences between larvae and adults of two sympatric, nongallicolous, also probably sibling species, Trupanea ni- gricornis and T. bisetosa recently was re- ported (Knio et. al. 1996). In turn, Procecidochares kristineae and P. lisae are very similar to P. stonei (Green et al. 1993). The noticeable difference be- tween P. stonei and these two species in the third instar larva is that in the lateral spi- racular complex, the verruciform sensillum is anteriorad of the spiracle in P. stonei not posteriorad. In the first instar of P. stonei, the mouth hooks are not grooved dorsally and the interspiracular processes are multi branched and blade-like (Green et al. 1993), not ovoid or spiniform and unbranched as in P. kristineae and P. lisae, respectively. Procecidochares kristineae and P. lisae are the first species of Tephritidae described with a v-shaped frontal groove in the apical tooth (Goeden and Headrick 1992; Goeden and Teerink 1996a, b, c; Goeden et al. 1994b; Green et al. 1993; Headrick and Goeden 1990; Headrick et al. 1995; Knio et al. 1996). The nongallicolous, flower head infesting species P. flavipes differs from all three of the above gall-forming species in having thoracic segments cov- ered with small, polygonal rugose pads and the ventral intersegmental areas and dorsum of the caudal segment bearing minute acan- thae (Goeden et al. 1994). DISTRIBUTION AND Hosts Both Procecidochares kristineae and P. lisae are true monophages, each forming galls only on its sole host-plant species, Ambrosia dumosa and A. eriocentra, re- spectively. Procecidochares kristineae is VOLUME 99, NUMBER 1 widely associated with A. dumosa in the Colorado and Mojave deserts of southern California (Goeden and Ricker 1976a, Sil- verman and Goeden 1980). Accordingly, the geographic distribution of P. kristineae may coincide wholly or partly with that of A. dumosa in southern Nevada, southwest- ern Utah, and western and southwestern Ar- izona (Benson and Darrow 1981), and southward into the Sonoran Desert of Baja California and Sonora, Mexico (Shreve and Wiggens 1964). The distributions of P. li- sae and A. eriocentra in southern California are restricted to a small part of the north- eastern Mojave Desert (Goeden and Ricker 1976b), but probably coincides wholly or partly with the wider distribution of this na- tive ragweed in southern Nevada, south- western Utah, and northwestern and central Arizona (Benson and Darrow 1981). Estimating tephritid distribution from host-plant distribution is problematic be- cause some gall-forming tephritid flies are known to be less widely distributed than their hosts, e.g., A. bigeloviae (Cockerell) and A. trixa Curran are widely allopatric, sympatric, or absent on Chrysothamnus nauseosus (Pallas) Britton in different parts of Utah, Arizona, New Mexico, and Colo- rado (Dodson and George 1986). Similarly, A. trixa, but apparently not A. bigeloviae, occurs in California on C. nauseosus which is a widespread plant species in southern California (D. H. Headrick, RDG, and JAT, unpublished data). BIOLOGY The biology of P. kristineae was de- scribed in considerable detail by Silverman and Goeden (1980). The following account largely concerns P. lisae, which then serves as a basis for comparison with P. kristineae and two other taxonomically-close Proce- cidochares spp. studied previously in south- ern California, P. stonei (Green et al. 1993) and P. flavipes (Goeden et al. 1994a). Egg.—Most eggs are inserted singly and basally in axillary buds of the current sea- son’s branch growth. Some buds held two 83 or rarely three eggs, presumably oviposited by different females, judging by the slightly different degrees of development of two or three first instars found associated with sin- gle buds in field-collected branch samples (see below). Two eggs of P. kristineae per axillary bud also were reported as rare by Silverman and Goeden (1980). Adjacent axillary buds of infested branches bore eggs probably deposited in sequence by individ- ual females as they moved in sequence to the next more-proximal axils, as described for P. kristineae (Silverman and Goeden 1980). Larva.—Eggs hatch about a week after their deposition (Goeden and Silverman 1980, Green et al. 1993), but then the first instar larvae persist as the longest-lived stage in the life cycle of P. lisae within tiny open cavities of incipient galls (Fig. 12A). Single, aestivating first instars were found Overwintering in ovoidal cavities 0.74 + 0.03 (range, 0.54—0.93) mm long by 0.54 + 0.02 (range 0.37—0.74) mm wide formed within, beneath, or immediately laterad of 26 axillary buds. Four pairs each of these first instars were associated with single ax- illary buds, and three first instars with an- other bud, documenting again multiple ovi- positions in single buds. Each of these lar- vae continued to develop separately, as all galls examined of P. lisae, like P. kristineae (Silverman and Goeden 1980), contained only one larva in each. Thus, sometimes two or rarely three unilocular galls of P. lisae arose at the same axil; although, only one gall per axil was usual for both P. kris- tineae and P. lisae (Silverman and Goeden 1980). However, larvae of P. stonei develop gregariously in unilocular galls (Green et al. 1995); The aforementioned 26 galls were sam- pled on 7.1i1.1995, so that not only were these aestivating first instars the overwin- tering stage of P. lisae, but also the stage within which this species had persisted for 6 or 7 months since the previous April or May, when their parent adults emerged, mated, and oviposited. Both P. kristineae 84 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON i” ec Fig. 12. Life stages of Procecidochares lisae on Ambrosia eriocentra: (A) axillary bud gall containing aestivating first instar, (B) sagittal section of preceding axillary bud gall, (C) axillary bud gall containing second instar, (D) sagittal section of gall containing second instar, (E) full-size gall, (F) sagittal section through full- size gall containing a puparium, and showing exit channel excavated by late third instar, (G) lateral view of male, (H) lateral view of gravid female, (I) mating pair, dorsal view. Lines = 1 mm. VOLUME 99, NUMBER 1 and P. stonei also overwinter as first instars in incipient galls (Silverman and Goeden 1980, Green et al. 1993). The tiny, over- wintering galls of P. lisae were undetecta- ble in the field (Fig. 12A, B); although those of P. kristineae were readily dis- cerned (Silverman and Goeden 1980). The former galls caused only a slight axillary swelling along the leafless branches of dor- mant A. eriocentra, unlike overwintering galls of P. kristineae on A. dumosa which incorporated juvenile leaves (Silverman and Goeden 1980). The axils on branches where incipient galls of P. lisae were borne aver- aged 3.04 + 0.05 (range, 2.5—3.6) mm in diameter (n = 21) and were located an av- erage of 15.8 + 1.1 (range, 6—24.5) cm from branch apex. Following substantial late-winter/early- spring rainfall, host plants, galls, and larvae of P. lisae concurrently resume growth as also reported for P. kristineae (Silverman and Goeden 1980) and P. stonei (Green et al. 1993). Twenty-five galls of P. lisae con- taining second instars (Fig. 12C, D) aver- aged 5.72 + 0.35 (range, 2.41—11.40) mm in length; 5.44 + 0.34 (range, 2.28—11.40) mm in width. These spheroidal galls (Fig. 12C) contained single, ovoidal to spheroi- dal cavities (Fig. 12D) that averaged 2.53 * 0.21 (range, 1.14—-5.70) mm in length and 1.97 + 0.16 (range, 0.57—4.56) mm in width. The gall walls averaged 1.12 + 0.10 (range, 0.42—2.85) mm in thickness. The larvae of P. lisae quickly grew and molted into the third and last larval instar within mostly spheroidal galls (Fig. 12E, F), 21 of which averaged 8.0 + 0.4 (range, 4.3—11) mm in length and 7.1 + 0.3 (range, 4.2-10) mm in width. The central ovoidal or spheroidal cavities were enlarged by lar- val feeding to an average of 4.1 + 0.2 (range, 1.42—5.13) mm in length and 3.17 + 0.18 (range, 1.42—5.13) mm in width. The lateral wall thickness averaged 1.18 + 0.04 (range, 0.57—-2.15) mm. Prior to pu- pariation, the third instar chews an exit tun- nel through the gall wall (Fig. 12F) ending in a thin, circular to oval, epidermal win- 85 dow 1.4 + 0.04 (range, 1.2—1.7) mm (n = 13) in diameter. Pupa.—Fifty-six galls containing puparia (Fig. 12F), and thus of maximum size and final subspheroidal shape, averaged 10.2 + 0.4 (range, 3.9—16) mm in length; 8.9 + 0.3 (range, 3.5—16) mm in width. The ovoidal to spheroidal cavities of these mature galls averaged 4.8 + 0.2 (range, 3.1—3.4) mm in length; 4.0 + 0.1 (range, 2.6—6.2) mm in width. The lateral walls of these galls av- eraged 1.58 + 0.06 (range, 0.51—2.58) mm in thickness. Adult.—The adults (Fig. 12G, H) of P. lisae that emerged from excised, field-col- lected galls were sexually mature, or nearly so, each female containing many full-size ova. These proovigenic females mated in laboratory cagings from | to 7 days-old (n = 59). (ries 21). Jen aales, lived, Jed) = 0.8 (range, 7-16) days; 11 nonovipositing females, 8.7 + 0.7 (range, 5-12) days in insectary cagings. These mean longevities fell between the 1l-week averages for P. kristineae (Silverman and Goeden 1980) and the 3-week averages for P. stonei (Green et al. 1993) under similar insectary conditions. This compares with longevities that averaged 42 days for males and 39 and 102 days for two series of synovigenic fe- males (n = 11 and 20, respectively) of P. flavipes (Goeden et al. 1994a), which has an aggregative life cycle different from the circumnatal life cycles of P. kristineae, P. lisae, and P. stonei (Headrick and Goeden 1994). The reproductive behavior of P. lisae re- sembled that of P. kristineae described by Silverman and Goeden (1980) and by D. H. Headrick from field observations (see be- low), and that of P. stonei described in greater detail by Green et al. (1993). Fur- thermore, a lack of courtship behavior and exclusive exhibition of enantion type of wing movements by both sexes reported by Headrick and Goeden (1994) as character- istic of circumnatal species of Procecido- chares also was confirmed for P. lisae. Wing enantion as defined and described by 86 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Green et al. (1993) and Headrick and Goe- den (1994) was observed for P. lisae as males tracked females prior to copulation, during copulatory induction behavior, and following postcopulatory separation. Cop- ulation by only one pair of P. lisae was observed from start to finish for 62 min, but nine more partially observed matings all were of shorter duration. Disengagement by three different pairs of adults was observed, whereby the males turned and walked off and away from the females as they concur- rently pulled their genitalia from the fe- males’ cloacas in one continuous motion in 15, 18, and 30 sec, respectively. The wings of both sexes of P. lisae usually were parted at 45° and centered over their midlines dur- ing copulation; whereas, Silverman and Goeden (1980) reported that males of P. kristineae usually were only slightly parted during copulation, a position sometimes also adopted by P. lisae males. Field observations.—Field observations of P. kristineae behaviors on Ambrosia du- mosa were made by D. H. Headrick at Snow Creek, Riverside Co., 2-9 April 1992. Most observations were made for 1— 3 h daily on three large shrubs all bearing newly formed, immature inflorescences. The oviposition behavior of P. kristineae females was highly stereotypical. Each fe- male began by walking from the base of a current season’s branch to the base of the raceme, where she turned head-downward, recurved her abdomen and oviscape, and probed with the apex of the aculeus into a leaf axil. After ovipositing or probing, the female moved basally to the next leaf axil and probed or oviposited in the same man- ner. When the female reached the base of the new growth, she moved to another branch and repeated the above behaviors. Females probed many leaf axils, but ovi- posited into only 1 to 4 axils per branch. Oviposition times averaged 2 min 9 sec (n = 4; range, 2 min—2 min 20 sec). Females frequently stopped to rest and groom be- tween Ovipositions. Mating behavior of P. kristineae was similar to that reported by Silverman and Goeden (1980). Males encountered females either while perched on top of branches or while moving through the crown. Males tracked females slowly from behind. Males sat still in a passive position if the female turned to face him, or paused nearby. Males mounted females by jumping onto their dorsa, either head first or from behind. Once mounted, males clasped the costal margins of the female’s wings near the base with the foretarsal claws; the middle legs were wrapped around the middle of the ab- domen of the female and the hind legs were bent and crossed underneath the abdomen of the female. Once mounted, males im- mediately began rubbing their hind legs asynchronously anterior to posterior under- neath the abdomen of the female. Simulta- neously, the abdomen of the female was raised and the apex of the ovipositor was placed into the epandrium; exsertion of the aculeus followed (n = 5). Copulation in the field lasted 30 min (1 = 2). Seasonal history.—Procecidochares li- sae usually is univoltine, but occasionally, following substantial late-summer rainfall stimulating renewed host-plant growth, pro- duces a facultative second generation. Pro- cecidochares kristineae is normally bivol- tine throughout its much wider range in southern California, but as reported by Sil- verman and Goeden (1980) can _ locally show univoltine, biennial, and even trivol- tine reproduction (unpublished data), de- pending on whether rainfall locally triggers resumption of gall and larval development concurrent with host-plant regrowth. Both species, along with P. stonei, exhibit the circumnatal life cycle defined and described by Headrick and Goeden (1994). Natural enemies.—The following chal- cidoid Hymenoptera were identified as par- asitoids of P. lisae: Eurytoma sp. and E. veronia Bugbee (Eurytomidae), as primary, solitary parasitoids reared from mature galls; Halticoptera sp. (Pteromalidae), as a primary, internal, larval-pupal parasitoid; Tetrastichus sp. (Eulophidae), as a gregar- VOLUME 99, NUMBER 1 ious, puparial endoparasitoid; Torymus sp. (Bedeguaris group), Torymus sp. (Tubicola group), 7. capillaceus (Hiiber) (Torymi- dae), probably as primary, solitary, larval ectoparasitoids as reared from immature galls. Silverman and Goeden (1980) reported 10 species of parasitic Hymenoptera reared from fully formed galls of P. kristineae, an unidentified vertebrate predator of the lar- vae and identified spider predator of the adults, and two species of inquiline weevils. Green et al. (1993) subsequently reported four species of primary chalcidoid parasit- oids individually reared from puparia of P. stonei as well as a hyperparasitized larval parasitoid. The genera of parasitoids com- mon to all three Procecidochares hosts were Eurytoma, Halticoptera, and Tetras- tichus. ACKNOWLEDGMENTS We sincerely thank A. C. Sanders, Cu- rator of the Herbarium, Department of Bot- any and Plant Sciences, University of Cal- ifornia, Riverside, for identification of plants. The parasitoids variously were iden- tified by H. E. Andersen, Huntington Beach, CA; G. Gordh, now with the De- partment of Entomology, University of Queensland, Australia; or J. Luhman, now with the Minnesota Department of Agricul- ture, St. Paul, MN. We also are grateful to FL. Blanc, D. H. Headrick, A. L. Norrbom, J. D. Pinto, and G. J. Steck for their helpful comments on earlier drafts of this paper. LITERATURE CITED Benson, L. and R. A. Darrow. 1981. Trees and Shrubs of the Southwestern Deserts. University of Ari- zona Press, Tucson. Blanc, FE L. and R. H. Foote. 1961. A new genus and five new species of California Tephritidae. Pan- Pacific Entomologist 37: 73-83. Dodson, G. and S. B. George. 1986. Examination of two morphs of gall-forming Aciurina (Diptera: Te- phritidae). Ecological and genetic evidence for species. Biological Journal of the Linnean Society 29: 63-79. Foote, R. H., EF L. Blanc, and A. L. Norrbom. 1993. Handbook of the Fruit Flies (Diptera: Tephritidae) 87 of America North of Mexico. Cornell University Press, Ithaca, New York. Goeden, R. D. and D. H. Headrick. 1992. Life history and descriptions of immature stages of Neaspilota viridescens Quisenberry (Diptera: Tephritidae) on native Asteraceae in southern California. Proceed- ings of the Entomological Society of Washington 94: 59-77. Goeden, R. D., D. H. Headrick, and J. A. Teerink. 1994a. Life History and descriptions of immature stages of Procecidochares flavipes Aldrich (Dip- tera: Tephritidae) on Brickellia spp. in southern California. Proceedings of the Entomological So- ciety of Washington 96: 288—300. 1994b. Life history and description of im- mature stages of Paroxyna genalis (Thomson) (Diptera: Tephritidae) on native Asteraceae in southern California. Proceedings of the Entomo- logical Society of Washington 96: 612-629. 1995a. Life history and descriptions of im- mature stages of Valentibulla californica (Coquil- lett) (Diptera: Tephritidae) on Chrysothamnus nauseosus (Pallas) Britton in southern California. Proceedings of the Entomological Society of Washington 97: 548-560. 1995b. Life history and description of im- mature stages of Urophora timberlakei Blanc and Foote (Diptera: Tephritidae) on native Asteraceae in southern California. Proceedings of the Ento- mological Society of Washington 97: 780-791. Goeden, R. D. and D. W. Ricker. 1976a. The phy- tophagous insect fauna of the ragweed, Ambrosia dumosa, in southern California. Environmental Entomology 5: 45—SO0. 1976b. The phytophagous insect faunas of the ragweeds, Ambrosia chenopodiifolia, A. eri- ocentra, and A. ilicifolia. Environmental Ento- mology 5: 923-930. Goeden, R. D. and J. A. Teerink. 1996a. Life histories and descriptions of adults and immature stages of two cryptic species, Aciurina ferrugenia (Doane) and A. michaeli new species (Diptera: Tephritidae) on Chrysothamnus viscidiflorus (Hooker) Nuttall in southern California. Proceedings of the Ento- mological Society of Washington 98: 415—438. . 1996b. Life history and descriptions of adults and immature stages of Aciurina idahoensis Steyskal (Diptera: Tephritidae) on Chrysothamnus viscidiflorus (Hooker) Nuttall in southern Califor- nia. Proceedings of the Entomological Society of Washington 98: 681—694. . 1996c. Life history and descriptions of adults and immature stages of Aciurina semilucida (Bates) (Diptera: Tephritidae) on Chrysothamnus viscidiflorus (Hooker) Nuttall in southern Califor- nia. Proceedings of the Entomological Society of Washington 98: 752-766. Green, J. EF, D. H. Headrick, and R. D. Goeden. 1993. 88 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Life history and description of immature stages of Procecidochares stonei Blanc & Foote on Vigui- era spp. in southern California (Diptera: Tephrit- idae). Pan-Pacific Entomologist 69: 18-32. Headrick, D. H. and R. D. Goeden. 1990. Description of the immature stages of Paracantha gentilis (Diptera: Tephritidae). 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A Flora of Southern California. University of California Press, Berkeley. Shreve, FE and I. L. Wiggens. 1964. Vegetation and Flora of the Sonoran Desert, Vol. 2. Stanford Uni- versity Press, Palo Alto, California. Silverman, J. and R. D. Goeden. 1980. Life history of a fruit fly, Procecidochares sp., on the ragweed Ambrosia dumosa, (Gray) Payne, in southern Cal- ifornia (Diptera: Tephritidae). Pan-Pacific Ento- mologist 56: 283-288. Wasbauer, M. W. 1972. An annotated host catalog of the fruit flies of America north of Mexico (Dip- tera: Tephritidae). California Department of Ag- riculture, Bureau of Entomology, Occasional Pa- pers 19: 1-72. PROC. ENTOMOL. SOC. WASH. 99(1), 1997, pp. 89-100 LIFE HISTORY AND LABORATORY REARING OF OEDANCALA DORSALIS (SAY) (HETEROPTERA: LYGAEIDAE), WITH DESCRIPTIONS OF IMMATURE STAGES'! LANE J. LOYA AND J. E. MCPHERSON Department of Zoology, Southern Illinois University, Carbondale, IL 62901, U.S.A. Abstract.—The life history of the seed bug Oedancala dorsalis (Say) was investigated in southern [Illinois from June 1994—November 1995, and the immature stages were de- scribed. The bug also was reared from egg to adult under controlled laboratory conditions. This apparently bivoltine species fed and reproduced on Carex spp. Adults overwintered and became active in April. Eggs were found from mid-May to early September in the spikes of Carex crinita. Seasonal occurrences of adults and nymphs are discussed. Adults were last observed in mid-October. This species was reared on C. crinita in the laboratory at 26 + 0.5°C under a 16L:8D photoperiod. The incubation period averaged 13.32 days. The five nymphal stadia averaged 6.34, 5.28, 4.52, 4.02, and 5.52 days, respectively. Key Words: Seed bug, Lygaeidae, southern Illinois, life history, laboratory rearing, im- mature stages, descriptions, Carex The family Lygaeidae contains approxi- mately 4050 species worldwide (Slater and O’Donnell 1995). Next to the family Miri- dae, it is the second largest family of Het- eroptera in America north of Mexico, where it is represented by approximately 320 spe- cies (Ashlock and Slater 1988). Lygaeids are commonly called seed bugs, referring to their habit of feeding on dry, mature seeds (Sweet 1960). In the United States and Canada, the ly- gaeid subfamily Pachygronthinae contains the three genera Oedancala Amyot and Ser- ville, Pachygrontha Germar, and Phlegyas Stal (Ashlock and Slater 1988). Only five species of Oedancala occur within this geo- graphic area (Ashlock and Slater 1988, Bar- anowski and Slater 1989). Of these, O. dor- salis Say is the most widespread, ranging ' Part of a thesis submitted to Southern Illinois Uni- versity at Carbondale by the senior author in partial fulfillment of the requirements for the M.S. degree in zoology. from Ontario, Quebec, and Maine south to Florida, and west to South Dakota, Colo- rado, and Texas, and is the only species of the genus that has been reported from Illi- nois (Ashlock and Slater 1988, Baranowski and Slater 1989). Oedancala dorsalis has been little stud- ied. It has been swept from herbage in wet meadows, pastures, and waste places (Blatchley 1926), and reported to feed on Carex, Cyperus (Van Duzee 1888, Slater 1951), Ceanothus, and marsh grasses (Slat- er 1951). Van Duzee (1888) found adults from May to September “‘about” Buffalo, New York. Uhler (1887) stated it rarely was found later than early summer in eastern North America and felt it was “‘single brooded.”’ It overwinters under leaves and around the tussocks at the edges of swamps (Torre-Bueno 1925); small colonies have been found beneath logs along the borders of woodlands in late October (Blatchley 1926): 90 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON This lygaeid has been reared in the lab- oratory under unspecified conditions and the egg and nymphal instars, except the sec- ond, have been described (Slater 1951). In this paper, we present information on the field life history in southern Illinois and laboratory rearing of O. dorsalis and de- scribe the immature stages. MATERIALS AND METHODS Field life history.—During summer 1994, several adults were observed feeding and copulating on the spikes of Carex crin- ita Lamarck in the La Rue-Pine Hills Re- search Natural Area, Union County, IL. The site is located along the south side of Forest Road 345, approximately 1.25 miles (2.01 km) east of State Highway 3. The plants (approximately 80 individuals) were grow- ing at the bases of red maple (Acer rubrum L.) located at the edge of La Rue Swamp. The population of O. dorsalis, although small, seemed sufficiently large to permit a life history investigation of the bug. There- fore, a study was conducted from June 1994 to November 1995. Counts of all nymphs and adults observed in the field, and notes on their activity, were taken weekly from early April to early November, before the bugs emerged from and after they entered Overwintering sites, respectively. Speci- mens were collected by hand picking. Nymphs large enough to be identified to in- star, and adults, were released. Others were preserved in 70% ethanol and taken to the laboratory for closer examination. Also, five C. crinita staminate spikes were taken to the laboratory weekly and inspected for eggs. Species of Carex, other than C. crinita, that served as host plants were noted from Pine Hills; Oakwood Bottoms, Jackson Co.; and Lake on the Campus, Southern Illinois University at Carbondale, Jackson Co. Bugs were recorded as feeding if their rostra were inserted in seeds or if many individuals were on the same plant. Potential overwintering sites (e.g., leaf litter, under bark of fallen trees) were ex- amined periodically during November— March. Laboratory rearing.—Ten adults were collected on 17 May and on 24 May 1995, brought to the laboratory, and placed in two oviposition cages (5 males, 5 females/cage). Each cage consisted of a 1l-quart (approxi- mately 0.95-liter) Mason jar with a disc of moistened filter paper on the bottom. A pis- tillate spike of C. crinita (collected outside the study site) served as food and was in- serted in a vial (8.5 cm length, 2.3 cm diam at base) containing distilled water stoppered with cotton; the vial was placed upright in the bottom of the cage. The cage was closed with a disc of paper toweling and wire screening secured with the band of the two- piece Mason jar lid. A strip of paper tow- eling (approximately 3 XX 20 cm), which provided additional area for excrement ab- sorption, was placed inside the jar with one end over the lip and held in place by the band. The cages were examined daily for eggs, which were removed and placed on moist- ened filter paper in the bottom of a petri dish (approximately 9 cm diam., 2 cm depth) and covered with the lid. A thin lay- er of petroleum jelly was applied to the in- side of the rim to help prevent the first in- stars from escaping. Additionally, the lids were secured with two strips of masking tape to insure a tight fit. Nymphs (including the firsts) were kept in petri dishes prepared similarly to those for eggs (e.g., a ring of petroleum jelly); a section of a pistillate spike served as food. The nymphs were grouped by molting dates to accurately determine the stadia. Food and filter paper in the cages and jars were changed, and distilled water was added, approximately every 2—3 days; the exception to this was that once oviposition began, spikes in the cages were replaced daily because they were destroyed during examination for eggs. To obtain information on reproductive behavior and fecundity (fertility was deter- mined in laboratory rearing), field-collected VOLUME 99, NUMBER 1 fifth instars were brought to the laboratory and placed in Mason jars prepared similarly to oviposition cages. As they reached adults, they were segregated by sex in two additional Mason jars. At the end of a min- imum of two weeks, six pairs were placed in petri dishes (one male, one female/dish) prepared similarly to those used for nymphs but without the ring of petroleum jelly. Be- havioral observations were made for peri- ods of 1—2 hours. All specimens were kept in incubators maintained at 26+0.5°C and a 16L:8D pho- toperiod (approximately 2800 lux). Descriptions of immature stages.—The description of each stage is based on ten individuals. Eggs and first through fifth in- stars were selected from field collected and laboratory reared individuals; all had been preserved in 70% ethanol. Only those that had not swelled in the alcohol were used in the descriptions. Drawings were made with the aid of a camera lucida. Measurements, in mm, were made with an ocular microm- eter. Statistics —Averages are expressed as X + SE; standard errors of less than 0.005 are listed as 0.00. Total developmental periods for males and females were compared with Student’s t-test; level of significance was 0:05: RESULTS AND DISCUSSION Field life history—Oedancala dorsalis Overwintered as adults that became active in late April (Figs. 1, 2), began feeding and copulating on spikes of maturing Carex crinita, and reproduced shortly thereafter; it remained on the host into October. Adults and nymphs also fed on C. blanda Dewey, C. cephalophora Muhlenberg, C. comosa F Boott, C. conjuncta FE Boott, C. crus-corvi Shuttleworth, C. frankii Kunth, C. granu- laris Muhlenberg, C. squarrosa L., C. tri- buloides Wahlenberg, C. vulpinoidea Mi- chaux, and C. vulpinoidea var. ambigua F Boott (= C. annectens Bicknell) (Table 1). One adult and two nymphs were observed on C. lurida Wahlenberg and C. lupulina 9] Muhlenberg, respectively, but were not ob- served feeding. Eggs were found on C. crinita from mid- May to early September. They usually were laid singly, but, sometimes, they were de- posited side by side in clusters of 2—5. Gen- erally, they were inserted between the scales of staminate spikes; occasionally, they were found in pistillate spikes inserted between the scale and perigynium. Similar Ovipositional behavior has been observed in other species of Oedancala (Baranowski and Slater 1982, 1989). The first instars were found from early June to early September (Figs. 1, 2). They were observed most commonly near the bases of the staminate spikes with their yel- lowish orange abdomens protruding from between the perigynia. Second instars were found from mid-June to early September, third instars from mid-June to early Octo- ber, fourth instars from late June to mid- October, and fifth instars from early July to early October. Second instars through adults were observed most commonly clinging to the pistillate spikes, feeding on seeds. If disturbed, nymphs often would fall to the ground and remain motionless; adults sometimes would fly a short distance to an- other plant or to the ground. The mating position was end-to-end. No precopulatory behavior was observed in the field. Copulating adults were noted from late May to late June and from mid-July to early August. Other than mating, little in- teraction between individuals was ob- served. Although adults clearly overwinter, no individuals were located in the field be- tween November and March. Overwintered adults were most abundant from late May to early July and died off soon thereafter. New adults appeared in late July as evi- denced by the appearance of callow adults (N = 6) and rise in number of adults fol- lowing the first peak of fifth instars. Based on the peaks of abundance of the adults, eggs, and the first-fifth instars (Figs. 1, 2), and the two periods of copulation (see 92 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1st INSTAR PERCENT OF INDIVIDUALS 3rd INSTAR (N=74) 4th INSTAR (N=95) 5th INSTAR (N=139) | ever e during 1995 season in Union Co., IL. above), O. dorsalis apparently was bivol- tine in southern Illinois. Laboratory rearing.—Adults fed and copulated on the pistillate spikes of C. crin- ita. Of the 697 eggs laid during this study, 90.2% were inserted in the pistillate spikes between the scales and perigynia and 1.1% in the filter paper, and 7.8% were laid on the cotton and 0.9% on the sides of the jar. Most were laid singly (68.1%) or side by side in clusters of 2 or 3 (28.0%), less com- monly in clusters of 4—7 (3.9%). The incubation period averaged 13.32 days (Table 2). Eggs were pale green when laid, turning yellowish in 2—3 days. Pink eyespots were visible in 5 to 6 days. At this Field cycle of Oedancala dorsalis. Percent in each sample of total individuals of same stage collected time, the anterior half of the egg began to redden; the posterior half remained yellow- ish. The minute egg burster and the paired dorsal sclerites of the abdomen were visible in 8 days. The first instar emerged through a slit in the cephalic end of the egg. The ground col- or was yellowish just after emergence; the sclerotized areas were pink but darkened to brown within 3—5 hours. The first, second, third, fourth, and fifth stadia averaged 6.34, 5.28, 4.52, 4.02, and 5.52 days, respectively. The total develop- mental period was 39.00 days (Table 2). There was no significant difference between total developmental period for males (37.34 VOLUME 99, NUMBER | Cue PERCENT OF INDIVIDUALS ADULT (N=797) 1st INSTAR (N=91) 3rd INSTAR (N=74) 4th INSTAR (N=95) 5th INSTAR (N=139) Fig. 2: season in Union Co., IL. + 0.35 days, N = 125) and females (36.89 + 0.32 days, N = 122) (Student’s t-test = —0.9429, df = 245, p = .3467). Mortality during the nymphal stadia re- sulted from incomplete ecdysis and unnat- ural causes (e. g., drowning in water con- densation inside the dishes). Precopulatory behavior was observed be- tween three pairs of males and females and lasted 5-10 minutes. In two of the encoun- ters, the first contact was head-to-head with both individuals touching antennae in a slow, irregular, up-and-down motion. Sub- sequently, the male moved around to the side of the female while antennating her, continued until he was behind her, and then mounted her from the side and extended his genitalia downward until engaged. In one of the two pairings, the male tapped the fe- male with his legs during mounting. After 98 AUGUST |SEPTEMBER| OCTOBER —————_ Field cycle of Oedancala dorsalis. Percent of individuals of each stage per sample during 1995 5 minutes, while still copulating and re- maining attached, the male dismounted and rotated to about an 80° angle; shortly there- after, the pair assumed a 180° end-to-end position. During copulation, the female moved about the petri dish, pulling the male. Copulation lasted approximately an hour. In the third encounter, the female ini- tiated copulation. She antennated the male and crawled on and off his back repeatedly. The pair began copulating shortly thereaf- ter. The male cleaned its rostrum and an- tennae during copulation. This pair re- mained in copula for approximately 25 minutes. The six pairs of adults reproduced in the laboratory, with the females averaging 53.33 = 13.37 eggs (range, 10-108). The F, adults produced a second gener- ation. In addition, fifth instars collected in 94 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Southern Illinois food plants of Oedan- cala dorsalis. Stages Collection Host Taxon Collected* Site” Cyperaceae Carex blanda Dewey A PH C. cephalophora Muhlenberg N, A PH C. comosa F. Boott N,A Ee. PH C. conjuncta F. Boott A OB C. crinita Lamarck N, A ILCs 125 C. crus-corvi Shuttleworth A PH C. frankii Kunth N, A OB, PH C. granularis Muhlenberg A PH C. squarrosa L. N,A OB C. tribuloides Wahlenberg N,A OB C. vulpinoidea Michaux N, A OB; PH C. vulpinoidea Michaux var. ambigua F. Boott (= C. annectens Bicknell) N, A OB, PH 4N = nymph; A = adult. > LC = Lake on the Campus, Southern Illinois Uni- versity; OB = Oakwood Bottoms; PH = La Rue-Pine Hills/Otter Pond Research Natural Area. the field in late July and brought to the lab- oratory reached adults and subsequently re- produced. Both of these observations sup- port our statement that O. dorsalis is bi- voltine in southern Illinois. Descriptions of immature stages.—Egg (Fig. 3A—B): Length, 1.41 + 0.01; width, 0.34 + 0.01. Eggs usually laid singly, sometimes in small clusters of 2—7; each elongate and whitish to pale green at ovi- position, turning yellowish red during mat- uration. Chorion with longitudinal ridges; 3-7 micropylar processes in ring pattern at cephalic end. Nymphal instars: The first instar is de- scribed in detail, but only major changes from previous instars are described for sub- sequent instars. Comparative statements (e.g., more punctate) refer to previous in- stars. The length is measured from tip of tylus to tip of abdomen and the width, across the widest part of the body. Addi- tional measurements are given in Table 3. First instar (Fig. 4): Length, 1.59 + 0.02; width, 0.66 + 0.01. Body generally yellow- ish orange in live specimens, paler in pre- served ones. Sclerotized areas of head, tho- Table 2. Duration (in days) of each immature stage of Oedancala dorsalis under controlled laboratory conditions. Number Complet- ing Cumulative Stage Stadium Range x += SE mean age Egg? 621. 10416 19.324, 103 fiaee Nymph Ist instar 933 3-10 6.34 + .04 19.66 2nd instar 427 2-12 5.28 + .09 24.94 3rd instar 307 1-15 4.52 = 12 29°46 4th instar 263 2-11 4.02 + .07 33.48 Sth instar 247 2-8 5:52 = 104 739300 4697 eggs were laid. rax, abdomen, and legs dark brown. Head, thorax, and abdomen lightly setose; legs and antennae moderately setose. Body elon- gate, slightly pyriform, greatest width across abdominal segments 2-3. Head narrowed in front, lateral margins tapering before antennae. Tylus exceeding juga, apex broad. Compound eyes red; pale line extending from inner margin of each eye posteromedially, meeting at midline of vertex, then extending to posterior margin of head. Antennae four segmented; seg- ments | to 3 yellowish, segment 4 brown, segments white at incisures; segment | ex- tending beyond apex of tylus; segment 4 acute apically; ratio of antennal segment lengths approximately 9:10:10:18. Rostrum four segmented, brown, extending onto ab- dominal segment 1. Pro- and mesonota mostly sclerotized; sclerotized areas concolorous with head, in form of paired plates, plates separated me- dially. Pronotal plates subquadrate. Meso- notal plates subrectangular, posterior mar- gins generally straight, laterally bending posteriorly, plates approximately 0.4x length of pronotal plates at midline. Meta- notum more membranous; paired sclerites present, each subelliptical, approximately 0.7X length of mesonotal plates at midline. Thoracic pleura sclerotized, concolorous with corresponding notal plates; spiracles present on posterior margins of pro- and VOLUME 99, NUMBER 1 95 Fig. 3. Scanning electron micrographs of egg of Oedancala dorsalis. A, Egg. B, Micropylar region. 96 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 3. Measurements (mm)* of Oedancala dorsalis instars.” Nymph Ist Instar 2nd Instar 3rd Instar 4th Instar Sth Instar Body length* 1.59 + 0.02 22025003 2166)== O05 3.81 + 0.06 5.26 + 0.08 Antennal segs. Ist 0.18 + 0.00 0225 0:00 0327-2) 0/01 0.49 + 0.01 O75 = OO 2nd 0.20 + 0.00 0.26 + 0.00 0.34 + 0.00 0.45 + 0.01 0.61 + 0.01 3rd 0.20 + 0.00 0.25 + 0.00 0.34 + 0.00 0.44 + 0.00 0.59 + 0.01 4th 0.37 + 0.00 0.41 + 0.00 0.50 + 0.01 0.61 + 0.01 0.74 = 0.01 Head width at eyes 0.43 + 0.00 ODS72== 0101 0.73 + 0.01 0.93 + 0.01 1.14 + 0.01 Synthlipsis O32== 10:00 0.41 + 0.01 OD 27-0101 0.64 + 0.01 Oa 0101 Head length‘ 0.30 + 0.01 0.41 + 0.01 0.50 + 0.01 Om + O'02 0.86 + 0.02 Beak segments Ist 0.18 + 0.00 0.25 + 0.00 0.31 + 0.00 0.40 + 0.01 0.47 + 0.01 2nd 0.23 + 0.00 0.29 + 0.00 0.35 + 0.00 0.44 + 0.01 O52] OO 3rd On7- = 0100 OH23== 10:00 0.29 + 0.00 0.39 + 0.01 0.45 + 0.01 4th 0.24 + 0.00 0.28 + 0.00 0.34 + 0.00 0.40 + 0.00 0.46 + 0.00 Notal lengths‘ Pronotum 0.20 + 0.01 0.29 + 0.01 0.43 + 0.01 0.69 + 0.02 L.025-= {0101 Mesonotum 0.10 + 0.00 Ont 2== O10 0.31 + 0.01 0.59 + 0.02 0.93 + 0.02 Metanotum 0.10 + 0.00 0.11 + 0.00 0.10 + 0.00 0.10 + 0.00 OMI == 0100 Notal widths Pronotum 0.49 + 0.01 0.69 + 0.01 0.96 + 0.02 1S6)== 0:03 1.81 + 0.02 Mesonotum 053° 0:01 0.74 + 0.02 1.04 + 0.02 iL Sy Sea O08} 216 20102 Metanotum 0.60 + 0.01 0.80 + 0.02 O92 = 0102 1.00 + 0.04 1:27 = O106 Leg lengths Profemur 0.30 + 0.01 0.40 + 0.00 05670101 0.85 + 0.01 1.29) == O1O0 Protibia 0.31 + 0.01 OMI OlO (ON SI7/ se OO) 0.80 + 0.01 lipyl a= (0)C0)1 Protarsus 0.22 + 0.00 0.25 + 0.00 03370101 0.43 + 0.01 0.57 + 0.01 Mesofemur 0.29 + 0.01 0.38 + 0.01 OS50E2 0/01 OF 25-2010 ONE = (Ov Mesotibia 0.30 + 0.00 0.39 + 0.00 054220101 OWS == 0101 1O1F = 0:0) Mesotarsus 0.21 + 0.00 0.25 + 0.00 (32 22 (OMON 0:42 = 0:01 0.55 + 0.01 Metafemur 0.34 + 0.01 0.44 + 0.01 0.61 + 0.02 0.89 + 0.01 27-0101 Metatibia 0.39 + 0.01 0.51 + 0.01 0.68 + 0.01 0.97 + 0.01 1-355== 0102 Metatarsus 0.25 + 0.01 (AAS) BE (COMO O39==70'01 0.49 + 0.01 0.69 + 0.01 Abd. width 0.66 + 0.01 0.90 + 0.02 122 0104 1.64 + 0.03 1-95) 23)0105 ay + SE; values of <0.005 listed as 0.00. > Based on 10 individuals of each instar. © Measured at midline. mesopleura. Thoracic sterna membranous, concolorous with membranous areas of nota. Coxae, trochanters, and femora gen- erally brownish; tibiae brownish, yellowish apically; tarsi yellowish, last segment brownish in apical half; profemora slightly thickened, ventral margin usually with two small spines in distal half; tarsi two seg- mented. Abdomen with paired scent glands me- dially on intersegmental lines between seg- ments 4 and 5, and 5 and 6; each pair sur- rounded by circular, sclerotized plate; scent glands visible as median orange masses. Tergum 8 with pair of small, suboval scle- rites medially; tergum 9 with posterior two thirds sclerotized; terga 1-6 each with sparse, transverse row of setae, 2—8 each VOLUME 99, NUMBER 1 97 Figs. 4-8. Nymphal stages of Oedancala dorsalis. 4, First instar. 5, Second instar. 6, Third instar. 7, Fourth instar. 8, Fifth instar. 98 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON with single lateral seta, those of 5—7 lon- gest; terga 2—7 each with faint pseudointer- segmental line visible. Spiracles ventrolat- erally on segments 2—8. Sterna 5—9 each with single medial plate, plates of variable shape; sterna 5 and 6 each with pair of small sublateral sclerites; sternum 7 with sublateral sclerite each side on posterior margin; setae generally more numerous than those on corresponding terga. Second instar (Fig. 5): Length, 2.02 + 0.03; width, 0.90 + 0.02. Sclerotized areas of head and thorax sparsely and irregularly punctate. Ratio of antennal segment lengths ap- proximately 11:13:13:20. Thorax with notal plates larger, fused medially, forming single plates; pale, me- dian fusion line evident from near anterior margin of pronotum to posterior margin of metanotum. Pronotal plate covering prono- tum. Mesonotal plate nearly covering meso- notum, subrectangular, posterior margin concave, approximately 0.5 length of pro- notum at midline. Metanotum with plates fused; resulting plate subrectangular, cov- ering approximately half of metanotum, ap- proximately 0.3 length of mesonotum at midline. Paired, small, narrow, transverse sclerites on intersegmental line between metanotum and abdominal segment |. Pro- femora thicker, ventral margin dentate in distal half with sparse row of teeth, distal two most prominent. Abdomen, dorsally, sometimes with three poorly defined, reddish, longitudinal stripes, one medial and two lateral; segments 5—7 with lateral setae equal in length to those on remaining segments. Tergum 9 almost completely sclerotized. Otherwise, similar to first instar. Third instar (Fig. 6): Length, 2.66 + 0.05; width, 1.22 + 0.04. Sclerotized area of head and thorax more punctate. Ratio of antennal segment lengths ap- proximately 16:17:17:25. Mesonotal plate larger, covering meso- notum, posterior margin concave medially, approximately 0.8 length of pronotum at midline; wing pads visible, small, not ex- tending beyond posterior margin of meta- notal plate. Metanotal plate larger, covering more than two thirds of metanotum, poste- rior margin straight, approximately 0.3 X length of mesonotal plate at midline. Inter- segmental sclerites between metanotum and abdominal segment 1 sometimes hidden by metanotum. Thoracic spiracles obscured by sclerotized pleura. Profemora thicker, teeth on ventral margin more prominent. Abdomen with three stripes better de- fined; lateral setae often paired. Sterna 3—9 with medial sclerites, those on sterna 3—4 paired; intersegmental lines of sterna 1—5 occasionally with paired, transverse, linear, medial sclerites. Otherwise, similar to second instar. Kourth instar (Fig. 7): Length, 3-51 0.06; width, 1.64 + 0.03. Punctures more numerous. Body width usually greatest at abdominal segments 2-3. Head with sclerotized areas generally yellowish to yellowish brown; dorsum with posterior margin dark brown to black. Pair of red ocelli sometimes visible, posterior and mesad to eyes. Ratio of antennal seg- ment lengths approximately 10:9:9:12. Ros- trum extending onto metasternum. Thorax with sclerotized areas generally yellowish to brown; pleura with continuous, longitudinal, dark brown to black stripe, ob- scure to clearly defined. Pronotum usually with pair of longitudinal dark-brown to black stripes. Mesonotum with pair of stripes, concolorous and continuous with those of pronotum; mesonotum 0.9 length of pronotum at midline; wing pads longer, extending to abdominal tergum 2. Metano- tal plate larger, completely covering meta- notum, posterior margin arcuate; metano- tum approximately 0.2 length of meso- notum at midline; wing pads evident later- ally, extending to abdominal tergum 2, almost entirely covered by mesonotal wing pads. Meso- and metafemora and some- times profemora more yellowish; profemo- ra thicker, ventral margin dentate with three VOLUME 99, NUMBER 1 prominent subequal teeth distally, separated by smaller teeth. Abdomen, dorsally, with three, reddish, longitudinal stripes now clearly defined; ventrally, with pair of diffuse, sublateral, reddish, longitudinal stripes. Tergum 8 with sclerites larger. Otherwise, similar to third instar. Fifth instar (Fig. 8). Length, 5.26 + 0.08; width, 2.16 + 0.02. Punctures more nu- merous. Body width usually greatest across mesothorax. Head with sclerotized areas yellowish. Ocelli clearly visible. Antennal segment 1 brown on inner margin, remainder of seg- ment yellowish; segment 2 yellowish; seg- ment 3 yellow proximally, reddish brown to brown distally; segment 4 brownish; ra- tio of antennal segment lengths 5:4:4:5. Rostrum yellowish, extending onto poste- rior margin of mesosternum. Thorax with sclerotized areas yellowish to yellowish brown; ventrally with pair of brown markings on meso- and metathora- ces. Pronotum with stripes reduced, meso- notum with stripes still well defined; meso- notum 0.9X length of pronotum at midline; metanotum 0.1 length of mesonotum at midline; meso- and metanotal wing pads longer, extending onto abdominal tergum 3 or 4. Legs yellowish, ventral margin of pro- femora dark brown; profemora greatly swollen, ventral margin dentate with four prominent subequal teeth, separated by smaller teeth; tarsi apically, subacute dor- sally. Abdomen, ventrally, with two well-de- fined, longitudinal, sublateral red stripes; longitudinal red stripe present medially, varying from diffuse to well defined. Ter- gum 8 with paired sclerites fused to form plate almost entirely covering dorsal sur- face. Otherwise, similar to fourth instar. Diagnosis.—The five nymphal instars, in addition to size, can be separated readily by the relative lengths of the pro- and meso- nota; sclerotization of the metanotum; rel- ative lengths of the lateral abdominal setae; 99 presence or absence of dorsal abdominal stripes; presence or absence, and degree of development, of wing pads; and presence or absence, and number and density, of punc- tures. There is a progressive increase in the length of the mesonotum relative to the pro- notum through all instars. The first instar can be separated from later instars by the lack of a single sclerotized plate on the metanotum, the relatively longer lengths of the lateral setae on abdominal terga 5—7, lack of punctures, and lack of dorsal ab- dominal stripes. The second instar can be distinguished from later instars by the lack of distinct mesonotal wing pads; wing pad development becomes apparent in the third instar with progressive development in the subsequent instars. The third-fifth instars can be distinguished from each other by the length of the mesonotal wing pads, which reach the metanotum in the third instar, ab- dominal tergum 2 in the fourth, and abdom- inal tergum 3 or 4 in the fifth. Finally, there is a progressive increase in the number and density of punctures in the second-fifth in- stars. ACKNOWLEDGMENTS We thank the following faculty and staff members of Southern Illinois University at Carbondale: B. M. Burr and W. G. Dyer, Department of Zoology, for their critical re- views of the manuscript; S. C. Mueller, Re- search Photography and Illustration Facili- ty, for final drafts of the nymphal-stage il- lustrations; and S. J. Schmitt and R. D. Tin- dall, Center for Electron Microscopy, for the SEM photographs of the eggs. We also thank zoology graduate student C. S. Bundy for assistance in the field. We are grateful to the U. S. Forest Service for granting us permission to collect in the La Rue-Pine Hills Research Natural Area, Union Co., IL; and to M. A. Basinger, Department of Plant Biology, SIUC, and E. L. Shimp, U. S. For- est Service, for identification of the sedges. 100 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON LITERATURE CITED Ashlock, P. D. and A. Slater. 1988. Family Lygaeidae, pp. 167-245. In Henry, T. J. and R. C. Froeschner, eds., Catalog of the Heteroptera, or True Bugs, of Canada and the Continental United States. E. J. Brill, New York. 958 pp. Baranowski, R. M. and J. A. Slater. 1982. The Pach- ygronthinae (Hemiptera: Lygaeidae) of Trinidad with the description of a new species and notes on other sedge feeding lygaeids. Florida Ento- mologist 65: 492-506. . 1989. The utilization of grasses as host plants by a species of Oedancala (Hemiptera: Lygaei- dae) with the description of a new species from Florida and the West Indies. Florida Entomologist 72: 243-251. Blatchley, W. S. 1926. Heteroptera or True Bugs of Eastern North America. Nature Publishing Com- pany, Indianapolis. 1116 pp. Slater, J. A. 1951. The immature stages of American Pachygronthinae (Hemiptera: Lygaeidae). Iowa Academy of Science 58: 553-561. Slater, J. A. and J. E. O’Donnell. 1995. A Catalogue of the Lygaeidae of the World (1960-1994). New York Entomological Society, New York. 410 pp. Sweet, M. H. 1960. The seed bugs: a contribution to the feeding habits of the Lygaeidae (Hemiptera: Heteroptera). Annals of the Entomological Soci- ety of America 53: 317-321. Torre-Bueno, J. R. 1925. Some winter bugs. Bulletin of the Brooklyn Entomological Society 20: 70. Uhler, P. R. 1877. Report upon the insects collected by P. R. Uhler during the explorations of 1875, including monographs of the families Cydnidae and Saldae [sic], and the Hemiptera collected by A. S. Packard, Jr., M. D. Bulletin of the United States Geological and Geographical Survey of the Territories 3: 355—475, 765-801, plates 27—28. Van Duzee, E. P. 1888. Mimicry in Hemiptera. Psyche 5: 27-28. PROC. ENTOMOL. SOC. WASH. 99(1), 1997, pp. 101-109 TWO NEW GENERA AND THREE NEW SPECIES OF EUPLECTRINI (HYMENOPTERA: EULOPHIDAE) FROM THE NEW WORLD G. A. W. WIJESEKARA AND M. E. SCHAUFF (GAWW) Maryland Center for Systematic Entomology, Department of Entomology, University of Maryland, College Park, MD 20742, U.S.A.; (MES) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture % Na- tional Museum of Natural History, MRC 168, Washington, DC 20560, U.S.A. Abstract.—Two new genera (Alveoplectrus, type species A. floridanus W. & S. and Eurycephaloplectrus, type species E. columbianus W. & S.) and three new species (A. floridanus, A. truncatus, and E. columbianus) of the tribe Euplectrini (Hymenoptera: Eulophidae) are described. Euplectrus corumbae Ashmead is reclassified Alveoplectrus corumbae (Ashmead) comb. n. Alveoplectrus floridanus is a parasite of the larvae of Alarodia slossoniae (Lepidoptera: Limacodidae). Eurycephaloplectrus colombianus 1s pat- asitic on larvae of Sibine sp. (Lepidoptera: Limacodidae). The two genera are closely related and share some synapomorphies with the mainly Old World genus Platyplectrus. Key Words: Chalcidoidea, Eulophidae, New World, Alveoplectrus, Eurycephaloplectrus, Alarodia, Sibine, parasites, Limacodidae The tribe Euplectrini is unique within Eulophinae being the only cocoon weaving ectoparasites. Species in Euplectrini are united by the possession of an elongate hind tibial spur. This character has been ob- served to be somewhat variable. For ex- ample, the euplectrine genus Metaplectrus Ferriére has a relatively short hind tibial spur, and it appears very similar to the ge- nus Euplectrophelinus Girault (J. LaSalle personal communication), which is not placed in the Euplectrini. In the absence of a thorough phylogenetic analysis of all the included genera, we continue to define Eu- plectrini based on this tibial spur character. Members of the tribe are distributed mainly in the Old World tropics, although species of Euplectrus Westwood are found world- wide. The tribe consists of six genera of which only two, Euplectrus and Platyplec- trus Ferriére, are known to occur in the New World (Bouéek 1988). The tribe is represented in North America by 13 species of Euplectrus (Burks 1979) and a single species of Platyplectrus (Schauff and La- Salle 1993). In our study on Sri Lankan Eu- plectrini we reported specimens of Euplec- tromorpha Girault from the New World (Wijesekara and Schauff 1994). Although these specimens possess two submedian ca- rinae on the propodeum, the defining char- acteristic of Euplectromorpha, further anal- ysis led us to conclude that these species are not properly placed in Euplectromor- pha. In this paper, we erect two new genera for these species. Museum abbreviations are as follows: National Museum of Natural History, Washington, D.C. (USNM); The Natural History Museum, London (BMNH). Ter- minology for surface sculpture follows Har- ris (1979). Bouéek (1988) produced a key to the genera of Euplectrini. His key is modified 102 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON to include our new genera. Awara Boucéek (1988) is not included in this key as it does not properly belong in the Euplectrini (Z. Bouéek and J. LaSalle, in litt.). KEY TO THE WORLD GENERA OF EUPLECTRINI 1. Propodeum with single median carina posterior tosbasalkcupi(hicsglS) ee ane eee ieee 2 — Propodeum with two submedian carinae; me- dian carina lacking (Figs. 2, 6, 12, 14) ...... 5 2. Scutellum without sublateral groove (as in Fig. 10); pronotum with transverse carina Be ec R ES CP PRG ais seri oa dd Ruel Lae Euplectrus Westwood — Scutellum with distinct sublateral groove (Figs. 5, 13, 14); pronotum variable 3. Hind tibia with a single, relatively short, apical spur; hind basitarsus subequal in length to sec- onditarsomete se eee eee Metaplectrus Ferriére — Hind tibia with 2 distinctly elongate apical spurs; hind basitarsus much longer than second LATSOMETE 15 BS 5 ay Sac wae ce SSA S Ne pee wenchar ences 4 4. Submedian area of propodeum divided into more than four areolae; first tergum as long as half the length of metasoma ... .Aroplectrus Lin — Propodeum with undivided submedian area; first tergum of metasoma variable mags Es) Otte eet gece aap Platyplectrus Ferriere 5. Scutellum without sublateral groove (Fig. 10); head relatively broad, more than 1.6 wider than high; petiole distinct, plate-like in dorsal view (Fig. 12) Metter te att Oe Eurycephaloplectrus, new genus — Scutellum with distinct sublateral groove (Fig. 5); head less than 1.5 wider than high; petiole transverse, not visible dorsally ............ 6 6. Scutellum with sublateral groove broad, con- tiguous with the posterior margin (Fig. 5); hind tibia with a single elongate spur (Fig. 7); hind tibial apex projected beyond point of attach- ment of basitarsus (Fig. 7); head with distinct tentorial depression lateral to clypeus (Fig. 4); postoccipital carina present (Fig. 1); species predominantly brownish in color ....... badd Net Pt scence iP i ae aie Alveoplectrus, new genus — Scutellum with sublateral groove narrow, not contiguous with the posterior margin (Fig. 14); hind tibia with one or two spurs; hind tibial apex not extended beyond point of tarsal at- tachment; head without facial depressions ad- jacent to clypeus; species including their ap- pendages yellowish in color........... Tt A eee oe Euplectromorpha Girault Alveoplectrus Wijesekara and Schauff, new genus (Figs. 1-8, 15, 16) Type Species: Alveoplectrus floridanus Wi- jesekara and Schauff. Diagnosis.—Female with distinct tento- rial depression on face near lateral margin of clypeus (Fig. 4); postoccipital carina present (Fig. 1); forebasitarsus with distinct strigil (Fig. 3); scutellum with sublateral groves which continue along the posterior margin (Fig. 5); mesepimeron with a dis- tinct pit located closer to the posterior mar- gin; dorsellum with posteriorly directed cup-like carina (Fig. 6); propodeum with two sublateral carinae that diverge posteri- orly as a narrow triangle originating from basal cup (Figs. 2, 6); hind leg with single tibial spur which is as long as first two tar- someres together; hind tibial margin ex- tended beyond tarsal attachment point (Fig. aD: Description.—Female. Compound eye with numerous long silvery setae; malar su- ture present; clypeus distinct but not de- marcated by supraclypeal suture (Fig. 4); face convex; postoccipital carina present; POL 1.5X longer than OOL; frons and ver- tex uniformly covered with setae; occipital area flat compared to concave occiput of Platyplectrus, with scattered setae; mandi- bles reduced, not meeting medially and without teeth; first funicular segment and clava distinctly longer than other segment (Figs. 15, 16); scape not reaching level of anterior ocellus; pronotum uniformly cov- ered with setae, without transverse carina; mesoscutum similarly covered with setae; axilla smooth, without setae; sublateral scu- tellar grooves contiguous along posterior margin of scutellum; mesepimeron with a distinct pit close to posterior mesopleural margin (Fig. 8); dorsellum medially with carina in form of posteriorly directed cup (Fig. 6); propodeum with two submedian carinae (Figs. 2, 6); petiole transverse; me- tasoma longer than broad, first tergum cov- ering more than half length of metasoma; VOLUME 99, NUMBER 1 103 3 Figs. 1-4. 4, A. truncatus, head. forebasitarsus with distinct strigil (Fig. 3); hind tibia with a single elongate spur; apical margin of hind tibia elongate beyond point of tarsal attachment (Fig. 7); hind basitarsus longer than following tarsomeres. Discussion.—This genus can be distin- guished from Euplectromorpha and Platy- plectrus by the presence of the anterior ten- torial depression, extended hind tibial mar- gin, and cuplike median carina of the dor- sellum. In both Euplectromorpha and Platyplectrus the dorsellum does not pos- sess a cup-like carina, being either smooth in most species or irregularly carinated (few Platyplectrus), and their hind tibial margin does not extend beyond tarsal attachment point. The clypeal area is smooth with no demarcations or depressions in Euplectro- morpha, whereas in Platyplectrus distinct supraclypeal carinae demarcate the clypeus, 1-3, Alveoplectrus floridanus. 1, Head, posterior view. 2, Dorsellum and propodeum. 3, Strigil. and there is no distinct depression near the tentorial pits. The single hind tibial spur in Alveoplec- trus is very distinct. Bouéek (1988) consid- ered the presence of a single tibial spur as a good generic character to identify Meta- plectrus but we have observed some vari- ation among other species. For example, Euplectromorpha jamburaliyaensis has a single hind tibial spur, although most Eu- plectromorpha species have two hind tibial spurs (Wijesekara and Schauff 1994). Based on this, we treat this character as homoplasious within the tribe. The pres- ence of two submedian carinae on the pro- podeum has been regarded as autapomorph- ic for Euplectromorpha but is also charac- teristic of the two new genera. Euplectro- morpha differs from Alveoplectrus by having fine lateral furrows on the scutellum 104 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 5-8. sepimeron. which are not posteriorly contiguous, and lacking postoccipital and pronotal carinae and posterior pit of the mesepimeron. In all three species of Alveoplectrus the occipital carina is very distinctly developed as in some Platyplectrus, but we consider this character to be variable, as we also have examined single specimens of three additional undescribed species of Alveo- plectrus from Central and South America, two of which do not possess the occipital carina. The relationship of Alveoplectrus to other Euplectrini is difficult to assess. Overall, species of Alveoplectrus appear similar to those of Platyplectrus, and the posteriorly contiguous lateral carina on the scutellum, setaceous head, pronotum and mesonotum, and distinct strigil suggest a close relation- ship with that genus. In addition presence of a distinct pit on the mesepimeron closer to the posterior margin of the mesopleuron also suggest a closer relationship to Platy- Alveoplectrus truncatus. 5, Scutellum. 6, Dorsellum and propodeum. 7, Hindtibial spur. 8, Me- plectrus. Sharing of similar propodeal ca- rinae by Alveoplectrus and Euplectromor- pha is probably homoplasious. Etymology.—‘‘Alveo” comes from Latin ‘“‘alveus’” meaning cavity, and refers to the depressions on the face of species in the genus. KEY TO THE SPECIES OF ALVEOPLECTRUS 1. Mesepimeron divided by transverse sulcus (Fig. 8); frons below anterior ocellus without a transverse carinae o 2. 4). «4 se ee 2 — Mesepimeron not divided by a sulcus; frons below anterior ocellus with short transverse ca- LINACS edo Ramet ee corumbae (Ashmead) 2. Antennal clava truncated; distal 3 flagellom- eres distinct from others being whitish; occip- ital carinae without a lamina; posterior end of propodeum lacks alveole . .truncatus, new species — Antennal clava slightly asymetrical, not dis- tinctly truncated; all flagellomeres same color; occipital carinae with a lamina; posterior end of propodeum with lateral alveoli ....... Pia SR A eae er ecmee me floridanus, new species VOLUME 99, NUMBER | Alveoplectrus floridanus Wijesekara and Schauff, new species (Figs. 1-3, 15) Female.—Length 1.2—2.0 mm. Color: Head and mesosoma dark honey brown, an- tenna, legs except hind coxa, petiole, and metasoma except small area along lateral margin of first tergum yellow; hind coxa, petiole, and posterior lateral area of first ter- gum brownish but lighter than head and me- sosoma. Head: Width 1.2 height; intero- cellar distance 2.5 eye width; frons and vertex evenly covered with short white se- tae; occipital and postoccipital carinae pres- ent (Fig. 1); POL:OOL 5:3; antennal clava slightly asymmetric; single anellus present; first funicular segment 1.2 longer than re- maining funicular segments, which are sub- equal in length (Fig. 15). Mesosoma: Pro- notum rectangular, collarlike, uniformly se- tose; mesoscutum similarly covered with se- tae; scutellum sculpture asperous; scutellum longer than broad with distinct lateral fur- rows contiguous with posterior margin; ax- illa smooth; mesepimeron divided into lower and upper mesepimeron by transverse sulcus terminating as a pit close to posterior me- sopleural margin; propodeum smooth with two posteriorly diverging submedian carinae joining posteriorly with two lateral alveoli (Fig. 2). Metasoma: Longer than broad, pet- iole transverse, not visible from dorsal side; first tergum covers more than half metaso- mal length. Legs and wings: Hind coxa smooth; hind basitarsus 2 longer than re- maining tarsomeres; forewing with postmarginal vein longer than stigmal vein. Male.—Unknown. Host.—Larvae of Alarodia slossoniae (Lepidoptera: Limacodidae). Distribution._Florida, Monroe Co. Types.—Holotype 2 on point with data: Florida, Monroe Co., Crane Key, 16-IV-76. Coll. D. Simberloff, ex. larvae Alarodia slossoniae (USNM). Paratypes: same data as holotype. (2 2 USNM, 1 2 BMNH). 105 Etymology.—The species epithet refers to the type locality. Alveoplectrus truncatus Wijesekara and Schauff, new species (Figs. 4-8, 16) Female.—Length 1.7—2.0 mm. Color: head and mesosoma dark honey brown to black; first three funicular segments, coxae, and metasoma except lateral part of first ter- gite brownish; scape, pedicel, and legs ex- cept coxae yellowish; apical part of mid and hind femora and tibiae honey colored; an- tennal clava, fourth funicular, dorsal and ventral surface of first metasomal tergum whitish. Head: Width 1.2 X height; inter- ocular distance 2 eye width; occipital and postoccipital carinae present; POL:OOL 7: 4; first funicular segment 1.5 longer than second funicular segment; funicular seg- ments becoming shorter and broader toward apex of antenna; clava about 1.6 longer than the preceding segment and distinctly truncated (Fig. 16). Mesosoma: Pronotum rectangular and covered with short brown- ish setae; mesonotum also covered with similar setae; scutellum and axilla smooth (Fig. 5); mesepisternum and mesepimeron smooth, mesepimeron divided into lower and upper mesepimeron by a transverse sul- cus, which terminates as a pit close to pos- terior mesopleural margin (Fig. 8); propo- deum with diverging submedian carinae en- closing a coarse raised median area, other- wise smooth (Fig. 6); posterior margin of propodeum without lateral carinae forming posterior alveole. Metasoma: First tergum extending over half of the metasoma. Legs and wings: Hind coxa smooth; single elon- gate hind tibial spur as long as first two tarsal segments together; hind basitarsus 2 x longer than second tarsomere; postmarginal vein longer than stigmal vein. Male.—Unknown. Hosts.—Unknown. Distribution.—Costa Rica. Types.—Holotype 2 on point with data: Costa Rica, Puntarenas, Golfo Dulce 3 Km. 106 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON W. Piedras Blancas, 100m. 3-V-1989 coll. Hanson (BMNH). Paratypes: same data as holotype (1 2 USNM); Costa Rica, Puntar- enas, Gulfo Dulce 3 Km. S. Rincon, 10m. 2-III-1989. coll Hanson (2 2) Costa Rica, Heredia, Chilamate 75m. XII-1989, III- 1990, Hanson and Godoy (1 ¢), (BMNH). Etymology.—The species epithet refers to the truncated clava of the antenna. Alveoplectrus corumbae (Ashmead), new combination Euplectrus corumbae Ashmead, 1904: 517. Holotype ¢: Brazil, Corumba, H. H. Smith (USNM 60572). Female diagnosis.—Head, mesosoma, and metasoma excluding the dorsum of the first tergum honey brown; legs yellow; an- tennal clava, third and fourth funicular seg- ments brownish with a short transverse ca- rinae just below anterior ocellus; posterior margin of dorsal axiller surface with dis- tinct projection over scutoscutellar suture toward scutellum; basal cup of the propo- deum enlarged, with a laminated margin; mesepimeron not divided into two parts by a sulcus. Discussion.—Alveoplectrus corumbae can be easily distinguished from the Alveo- plectrus species described above by the presence of distinct projections of the pos- terior margin of axilla (straight in A. flori- danus and without any projection in A. truncatus), reticulate sculpture of the scu- tellum (smooth and asprous in A. floridanus and A. truncatus) and lack of a sulcus on mesepimeron dividing it into two parts (di- vided into upper and lower mesepimeron by a sulcus in both A. floridanus and A. trun- catus (Fig. 8)). Alveoplectrus corumbae was described by Ashmead (1904) from a single female specimen (USNM type no. 60572, exam- ined). Vidal Sarmiento and DeSantis (1979) described a male of A. corumbae reared from Spodoptera frugiperda (J. E. Smith) (Noctuidae) as “‘allotype’’. Eurycephaloplectrus Wijesekara and Schauff, new genus (Figs. 9-12) Type species: Eurycephaloplectrus colom- bianus Wijesekara and Schauff. Diagnosis.—Female head distinctly broader than high; occiput concave; head, pronotum and mesoscutum uniformly se- tose; scutellum smooth without lateral fur- rows and with two anteriorly contiguous distinct alveoli separating scutellum from axillae (Fig. 10); propodeum with posteri- orly diverging submedian carinae which originate from basal cup separately; median carina absent (Fig. 12); petiole wider than long and distinctly visible dorsally, dorsal surface smooth and expanded like a plate (Fig. 12); abdomen shorter than mesosoma. Description.—Female. Head more than 1.6X as broad as high; vertex and occiput uniformly covered with short setae; eye densely ciliated; malar sulcus distinct; clyp- eus not differentiated; occipital area con- cave; antennal scrobe distinctly marked; scape not reaching level of anterior ocellus. Pronotum and mesoscutum uniformly cov- ered with short setae; axilla and scutellum smooth; distinct alveoli between scutellum and axilla; scutellum without lateral furrow; each axilla with two pairs of short setae (Fig. 10); mesepimeron with distinct pit near posterior margin (Fig. 11); dorsellum smooth; propodeum with two posteriorly diverging submedian carinae, which reach posterior margin of propodeum between ab- domen and coxal foramina, closer to plica, making broader angle near basal cup when compared to Alveoplectrus (Fig. 12); peti- ole broader than long, platelike, and visible in dorsal view; metasoma shorter than me- sosoma; strigil of fore basitarsus not dis- tinct; hind tibia with two elongate spurs. Discussion.—Eurycephaloplectrus is closely related to Alveoplectrus. The syna- pomorphies for this relationship are 1) uni- formly ciliated head, pronotum, and meso- notum and 2) basally separated submedian VOLUME 99, NUMBER 1 Figs. 9-14. carina on the propodeum. The distinct pit close to the posterior margin of the mes- epimeron is similar to the state found in Platyplectrus and is considered homolo- gous to the similar structure in Alveoplec- trus. However this genus differs from Al- veoplectrus in lack of sublateral scutellar furrows, absence of a distinct depression on the face lateral to clypeus; lack of an elon- 9-12. Eurycephaloplectrus colombianus. 9, Head, posterior view. 10, Scutellum. 11, Mes- epimeron. 12, Dorsellum, propodeum, and petiole. 13, Platyplectrus sp., dorsum. 14, Euplectromorpha sp., dorsum. gate strigil; presence of two pairs of setae on each axilla; presence of a dorsally ex- panded petiole; and presence of two hind tibial spurs. Species of Alveoplectrus pos- sess distinct depressions lateral to the clyp- eus, posteriorly contiguous sublateral fur- rows on the scutellum, distinct strigil, no setae on axilla, transverse petiole, and a sin- gle hind tibial spur. Eurycephaloplectrus is 108 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 15-16. Female antenna. 15, Alveoplectrus floridanus. 16, A. truncatus. similar to Euplectrus in lacking lateral fur- rows on the scutellum but differs from it by having a distinct malar suture and lacking a median carina on the propodeum. Etymology.—‘‘Eurys’’ meaning broad and “‘cephalo”’ meaning head (both Greek) together refer to distinctly broad head of this genus, and is combined with the stem “*plectrus’’. Eurycephaloplectrus colombianus Wijesekara and Schauff, new species (Figs. 9-12) Female.—Length: 1.4—1.5 mm. Color: Head, ventral and lateral mesosoma, hind coxa, and petiole honey brown; mesosoma dorsally much darker than head, almost black; metasoma except anterior dorsal area, fore and hind coxae and hind femur light brown; antennae, legs except all coxae and hind femur and anterior dorsal area of metasoma yellow. Head: Width 1.6X height; interocular distance 2.5 eye width; mouth margin broad; antennal scrobe dis- tinctly depressed; vertex and occipital area uniformly covered with setae; eye densely setose; occipital and postoccipital carina ab- sent (Fig. 9); POL:OOL 9:7; occiput con- cave; first funicular segment 1.2 longer than second funicular segment; funicular segments 2—4 subequal in length; clava dis- tinctly longer than funicular segments. Me- sosoma: Pronotum collarlike and uniformly setose (Fig. 9); mesoscutum covered with similar white setae; mesepimeron not divid- ed (Fig. 11); dorsellum and propodeum smooth; basal cup distinctly developed; two submedian carinae originate separately from basal cup and diverge posteriorly; pro- podeal margin laterally with few white se- tae (Fig. 12). Metasoma: Petiole smooth, broader than long; dorsal surface of petiole distinctly expanded and plate-like; metaso- ma distinctly shorter than mesosoma. Legs and wings: hind tibia with two elongate spurs; hind basitarsus slightly longer than other tarsomeres; wings at rest extended be- yond apex of metasoma; postmarginal vein longer than stigmal vein. Male.—Unknown. Host.—Larvae of Sibine sp. (Lepidop- tera: Limacodidae). Distribution.—Colombia, Tibaitata. Types.—Holotype 2 on point with data: Colombia, Tibityata, 21-VIII-1974, Coll. J.A. Jimenez, “‘Acacias vet’’ ex. Sibine lar- vae (USNM). Paratypes; 3 females with same data as holotype (2 2 USNM, 1 2 BMNH). Etymology.—The species epithet refers to the type locality. ACKNOWLEDGMENTS We thank John LaSalle of the Interna- tional Institute of Entomology, London, for valuable advice, John Noyes of The Natural History Museum, London, for loan of ma- terial for this study and Gregory Zolner- wich of Texas A&M University and John Huber of The Canadian National Collecion for reviewing the manuscript. We appreci- ate the work of G. Venable on the plates of SEMs. LITERATURE CITED Ashmead, W. H. 1904. Classification of the chalcid- flies of the superfamily Chalcidoidea with descrip- a VOLUME 99, NUMBER 1 tions of new species in the Carnegie Museum, col- lected in South America by Herbert H. Smith. Memoirs of the Carnegie Museum, 1: i—xxi + 225-551 Bouéek, Z. 1988. Australasian Chalcidoidea (Hyme- noptera): A Biosystematic Revision of Genera of Fourteen Families, with a Reclassification of Spe- cies. CAB International Institute of Entomology, Wallingford. 832 pp. Burks, B. D. 1979. Family Eulophidae, pp. 967—1022. In Krombein, K. V., P. D. Hurd, Jr., D. R. Smith, and B. D. Burks, eds. Catalog of Hymenoptera in America North of Mexico. Vol. 1. Smithsonian Institution Press, Washington, D.C. 1198 pp. Harris, R. A. 1979. A glossary of surface sculpturing. 109 California Department of Food and Agriculture. Occasional Papers in Entomology 28: 31 pp. Schauff, M. E. and J. LaSalle. 1993. Nomenclatural notes on genera of North American Eulophidae (Hymenoptera: Chalcidoidea). Proceedings of the Entomological Society of Washington 95: 488- 503. Vidal Sarmiento, J. A. and L. DeSantis. 1979. Nuevas citas de Himenopteros para la Republica Argen- tina. II. Revista de la Sociedad Entomologica Ar- gentina 34: 19-23. Wijesekara, G. A. W. and M. E. Schauff. 1994. Re- vision of the tribe Euplectrini of Sri Lanka (Hy- menoptera: Eulophidae). Oriental Insects 28: 1— 48. PROC. ENTOMOL. SOC. WASH. 99(1), 1997, pp. 110-114 A NEW SPECIES OF GALL MIDGE (DIPTERA: CECIDOMYIIDAE) FROM ECUADOR ASSOCIATED WITH FLOWERS OF CLAVIJA (THEOPHRASTACEAE) RAYMOND J. GAGNE, CORNELIA OTT, AND SUSANNE S. RENNER (RJG) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture % National Museum of Natural History MRC-168, Washing- ton, DC 20560, U.S.A.; (CO, SSR) Institut fiir Spezielle Botanik und Botanischer Garten, Johannes Gutenberg University, D-55099 Mainz, Germany. Abstract.—A gall midge new to science, Charidiplosis clavijica Gagné, is described from specimens found standing on male flowers of Clavija weberbaueri (Theophrastaceae) in eastern Ecuador. The new species is notable for several unique characters involving the tarsal claws, the male antenna, and the female ovipositor. The new species is compared to the widespread and only other described congener, Charidiplosis triangularis (Felt), a species associated with flowers of cacao. Key Words: Clavija is a Neotropical plant genus of 50 species (Stahl 1991). The pollination and breeding system of Clavija species have not been studied previously in the field. Two of us (CO, SSR) are investigat- ing floral morphology in Clavija in an at- tempt to determine which species are an- drodioecious, as suggested by Stahl (1991). The reproductive biology of Clav- ija weberbaueri Mez. was studied during two Stays at the Jatun Sacha Biological Re- serve in eastern Ecuador in September, 1994 and October, 1995. This plant is an understory shrub of about 0.8-1.5 m height, frequent along trails at Jatun Sacha. Flowers are orange, bowl shaped, ca. 1 cm diameter, and emit a fruity scent, especially at night. The chemical composition of the floral scents of C. euerganea Macbr. and C. repanda Stahl have been investigated by Knudsen and Stahl (1994). Scents in these two species are very similar and con- sist almost exclusively of sesquiterpenes. In both years when C. weberbaueri was Neotropical, Charidiplosis, pollination studied at Jatun Sacha, numerous individ- uals were in bloom. One of us (CO) monitored six shrubs with bisexual flowers and ten male shrubs from October until December in both years. The only visitors seen on the male flowers during several observation periods, both during the day and at night until 11 p-m., were gall midges of the new species described here. The flies remained immo- bile on the flowers for long periods of time, except at dusk when some individu- als were observed to fly. Female flowering shrubs were extremely rare in both years, and we did not see any visitors on the few female flowers we found. Whether these gall midges pollinate the flowers is still un- clear. MATERIALS AND METHODS The new species was collected by one of us (CO) in September, 1994 and October, 1995, at the Jatun Sacha Biological Re- serve, Napo Province, Ecuador. The Re- VOLUME 99, NUMBER | serve is located 8 km E of Puerto Misa- hualli on the eastern slope of the Andes at about 400 m elevation (01°04’S, 77°36'W). Mean annual precipitation at Jatun Sacha is 3600 mm and the average yearly tempera- ture is about 24.5°C. Fresh-caught flies were killed and preserved in 70% ethanol and later slide mounted for scientific study using the method outlined in Gagné (1989, 1994). Specimens of the new species have been deposited in the Museo Ecuatoriano de Ciencias Naturales in Quito, Ecuador, and the National Museum of Natural His- tory, Smithsonian Institution, (USNM) in Washington, DC, U.S.A. Terminology for adult morphology follows usage in Mc- Alpine (1981). Charidiplosis clavijica Gagné, new species (Figs. 1—7) Adult.—Head (Figs. 1-2): Eye 9-10 facets long at vertex, connate; facets all closely adjacent. Vertex with narrow, dor- sal protuberance bearing 2 long setae. Frons with 3—4 setae per side. Labella elongate linear in frontal view, more or less ovoid in lateral view, with about 15 setae laterally and covered with fine, pliant setulae mesally. Labrum elongate, basally with several short, sensory setae, covered beyond base on dorsum with long, fine, pliant, setulae, and situated between the la- bellae. Hypopharynx wide, striated dorsal- ly and with elongate, setulose ligule ven- trally. Palpus 4 segmented, with basal pal- piger. Antenna with 12 flagellomeres, ped- icel enlarged, longer and wider than both scape and first flagellomere. Male flagel- lomeres (Fig. 2) gynecoid, cylindrical, with very short necks, and becoming suc- cessively narrower from base to apex; cir- cumfila appressed, as 2 rings encircling the flagellomeres, interconnected by 2 longi- tudinal strands. Female flagellomeres as in male but all of same width. Thorax: Scutum with 4, single, sparse longitudinal rows of setae. Anepisternum without vestiture. Epimeron with 4—6 setae 111 in longitudinal row. Wing length: d, 1.4— 1.6 mm (n = 5); 2, 1.5—1.6 mm (n = 4); R,; curved apically, joining C posterior to wing apex; Rs weak, oblique, situated prox- imal to mid distance between arculus and R, apex; M;,, evident on distal two-thirds length; Cu forked; CuP present before Cu fork. Fifth tarsomeres (Figs. 3, 7) with ac- ropod partly hidden from view; claws sim- ple, thin, curving near basal third; empodia and pulvilli rudimentary. Male abdomen: First through sixth ter- gites entire, rectangular, with mostly single, continuous, posterior row of setae, 1—3 lat- eral setae on each side near midlength, sparse setiform scales covering most of sclerites, and pair of trichoid sensilla on an- terior margin. Seventh tergite as for sixth but posterior row of setae interrupted mesally. Eighth tergite weakly sclerotized, anterior pair of trichoid sensilla the only vestiture. Genitalia (Fig. 5—6): cercus nar- rowing to rounded apex, with several pos- terior setae; hypoproct slightly longer than cercus, tapered gradually from base to very narrow apex, a pair of setae present apical- ly; aedeagus longer than gonocoxite, ta- pered evenly from base to narrow apex, gonocoxite with short, obtuse mesobasal lobe, cylindrical beyond; gonostylus elon- gate, cylindrical, barely tapered from base to apex, setulose near base, asetulose and ridged beyond. Female abdomen: First through seventh tergites as for male first through sixth. Eighth tergite square, shorter and narrower than seventh, with scattered setae on pos- terior half and anterior pair of trichoid sen- silla. Eighth sternite present, with scattered setae and anterior pair of trichoid sensilla. Intersegmental membrane with scattered se- tae on all surfaces. Ninth tergum (Fig. 4) dorsally with elongate, curved setae, later- ally and ventrally with short, scattered se- tae. Tenth tergum without setae. Cercus (Fig. 4) elongate-ovoid, completely setu- lose, with four apical to apicoventral sen- sory setae; hypoproct elongate, undivided. Pupa and larva.—Unknown. 112 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-4. Holotype.—