Journal of Hymenoptera Research

Volume 13, Number 1 April 2004

ISSN #1070-9428 CONTENTS

ENGEL, M. S., C. D. MICHENER, and M. G. RIGHTMYER. The cleptoparasitic bee tribe Rhathymini (Hymenoptera: Apidae): Description of a new genus and a tribal review

GIBSON, G. A. R A new species of Oozetetes De Santis (Hymenoptera: Chalcidoidea: Eupelmidae) attacking oothecae of Nyctibora acaciana Roth (Orthoptera: Blattellidae) 13

GONZALEZ, V. H. and C. D. MICHENER. A new Chilicola Spinola from Colombian Paramo (Hymenoptera: Colletidae: Xeromelissinae) 24

GRISSELL, E. E., K. KAMIJO, and K. R. HOBBS. Torymus Dalman (Torymidae: Hymenoptera) associated with coniferous cones, with descriptions of three new species 31

A. L. GRIXTI, J. C, ZAYED, and PACKER. Behavioral interactions among females of Acamptopoeum submetallicum (Spinola) and Nolanomelissa toroi Rozen (Hymenoptera: Andrenidae) 48

LANES, G. O., F. T. GOBBI, and C. O. AZEVEDO. Report on a collection of Bethylidae (Hymenoptera) from central Florida, USA, with description of a new species of Lepidosternopsis Ogloblin 57

PUCCI, T. and M. SHARKEY. A revision of Agathirsia Westwood (Hymenoptera: Braconidae: Agathidinae) with notes on mouthpart morphology 64

REINA, P. and J. LA SALLE. Two new species of Quadrastichus Girault (Hymenoptera: Eulophidae): Parasitoids of the leafminers Phyllocnistis citrella Stainton

(Lepidoptera: Gracillariidae) and Liriomyza trifolii (Burgess) (Diptera: Agromyzidae) 108

SMITH, D. R. and S. G. BADO. First food plant record for Lagideus Konow (Hymenoptera: Pergidae), a new species feeding on Fuchsia and Ludwigia (Onagraceae) in Argentina 120

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Title of Publication: Journal of Hymenoptera Research. Frequency of Issue: Twice a year. Location of Office of Publication, Business Office of Publisher and Owner: International Society of Hymenopterists, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C 20560-0168, U.S.A. Editor: E. Eric Grissell, Systematic Entomology Laboratory, USDA, % National Museum of Nat- ural History, 10th and Constitution NW, Washington, D.C 20560-0168. U.S.A. Managing Editor and Known Bondholders or other Security Holders: none. J. HYM. RES. Vol. 13(1), 2004, pp. 1-12

The Cleptoparasitic Bee Tribe Rhathymini (Hymenoptera: Apidae): Description of a New Genus and a Tribal Review

Michael S. Engel, Charles D. Michener, and Molly G. Rightmyer

Division of Entomology, Natural History Museum, and Entomology Program, Department of Ecology and Evolutionary Biology, Snow Hall, 1460 Jayhawk Boulevard, University of Kansas, Lawrence, Kansas 66045-7523, USA

— Abstract. The new genus Rhathymodes is proposed for Rhathymus acutiventris Friese [with its R. new synonym, fiiesei Ducke], and R. bertonii Schrottky; resulting in the new combinations: Rhathymodes acutiventris (Friese) and R. bertonii (Schrottky). A lectotype is newly designated for R. friesei. To accommodate the new genus, changes are suggested for a key to subfamilies and tribes of Apidae. The tribe Rhathymini and its two genera are characterized, as are the two species of Rhathymodes.

The neotropical cleptoparasitic apine Engel (2001); equivalents are indicated in tribe Rhathymini consists of moderate brackets in keys and descriptions. The ab- sized to large (13-28 mm body length) breviations T and S are for metasomal ter- species superficially resembling vespid ga and sterna; T3, for example, is the third wasps, especially Polistes, or suggesting in metasomal tergum. Antennal flagellar seg- form giant species of the bee genus No- ment is abbreviated F. Photomicrography mada. The impetus for the present paper was done using a Microptics ML-1000 was the discovery by Martin Cooper of Digital Imaging System. Lyme Regis, U.K., and almost simulta- Collections in which specimens are pre- neously by one of us (MSE), that some of served are indicated by names of cities or the smaller species hitherto placed in towns in brackets, with names of relevant Rhathymus do not run to the Rhathymini curators in parentheses, as below: in the to the subfamilies and tribes of key [Berlin] Museum fur Naturkunde, Michener Apinae by (2000: 571-574). Berlin, Germany (Frank These smaller species represent a second Koch). of until now genus the, monogeneric, [Budapest] Hungarian Natural His- It is a with some Rhathymini. genus prob- tory Museum, Budapest, able relative to plesiomorphies Rhathymus Hungary (Lajos Zambon). and therefore to some in- likely provide [Chamela] Instituto de Biologia, sight into relations between Rhathymini Universidad Nacional and other tribes of Apinae. We hope that Autonoma de Mexico, re- recognition of the new genus, named be- search station at Chame- low will the dis- Rhathymodes, encourage la, Jalisco, Mexico (Ricar- of its unknown and of its lar- covery hosts, do Ayala). val characteristics. [Heredia] Instituto Nacional de Bio- The morphological terminology used diversidad (InBio), Here- below follows that of Michener (2000) dia, Costa Rica (Carolina with some modifications as proposed by Godoy). Journal of Hymenoptera Research

habitus. Lateral of and Figs. 1-2. Khathymodes bertonii (Schrottky). 1, Dorsal 2, view head, mesosoma, anterior metasoma (note absence of pleural tubercle).

= [Lawrence] = Division of Entomology, [New York] American Museum of University of Kansas Nat- Natural History, New ural History Museum and York, New York, USA (Je- rome G. Biodiversity Research Rozen, Jr.). Center, Lawrence, Kan- [San Lorenzo]= Museo Nacional de His- sas, USA. toria Natural del Para- [Lyme Regis]= Martin Cooper collection; guay, San Lorenzo, Para- Lyme Regis, UK (Martin guay (Bolivar R. Garcete- Cooper). Barrett). Volume 13, Number 1, 2004

SYSTEMATICS than distance to from apex wing tip, apex rounded and separated from costal Tribe Rhathymini Lepeletier wing margin; pterostigma one-fourth to one- 1841: 539. BJw.thymites Lepeletier Type genus: fifth as long as marginal cell, border of Khathymus Lepeletier and Serville 1828. Com- pterostigma in that cell straight or gently bining stem: Rhathym—. concave; wing hairy, alar papillae absent — (Fig. 4). Hind with cu-a [cu-v] Description. Body usually without are- wing to shorter than as of dense pale appressed pubescence; oblique, longer slightly second abscissa of M + Cu; lobe mi- form elongate, pubescence short, so that jugal nute, rounded, about one-tenth as as habitus suggests Polistes wasps or giant long vannal lobe. Metasoma widest at T2 and Nomada (e.g., Fig. 1); coloration black to T3; Tl narrower than T2, lateral largely yellow, sometimes with metasoma markedly dorsal surface weak- red, or all red. Compound eyes slightly di- profile slanting, only differentiated and less than half as verging below. Clypeus protuberant to ly long as more surface. T7 of less than width of compound eye in lateral slanting, anterior, male to bidentate without view because lower part of compound eye tapering apex, and sometimes S3 quite broad. Mandible slender, simple. La- pygidial plate; S4, S5, of male and male brum as long as or longer than median strongly fringed; S7, S8, as illustrated length of clypeus. Proboscis long, in re- genitalia (Figs. 6-13), genitalia with both and lower pose reaching between or to apices of pro- upper gonostylar well rather coxae; labial palpus with first two seg- processes developed, upper slender with branched lower broad ments subequal in length, last two seg- hairs, and translucent; valve scler- ments minute, directed laterally; maxillary penis heavily but palpus absent. Antennal scape short, less otized, spatha largely membranous with than three times as long as wide; Fl about heavily sclerotized longitudinal bar half as long as F2. Epistomal sulcus absent at each side. Female without pseudopy- below anterior tentorial pits so that clyp- gidial area; pygidial plate present, taper- eus and lower paraocular areas are fused. ing to apical narrowly rounded point, lat- Lateral ocellus separated from median eral margins weakly concave to weakly ocellus by one-third ocellar diameter or convex. Sting well developed; gonoplac less; preoccipital area rounded. Scutellum [= gonostylus] long, slender, parallel-sid- grading from somewhat elevated to form ed. Comments.— transverse shining ridge to distinctly bi- Some of the characteristics tuberculate, posterior declivitous part lon- used to identify the Rhathymini by Mich- ger, sometimes much longer, than anterior ener (2000) turn out to be generic charac- subhorizontal part; axilla small, rounded, ters of Rhathymus rather than tribal fea- not produced to form tooth. Propodeal tri- tures. The key to subfamilies and tribes of angle hairy. Procoxa tapering, mesal api- Apidae (Michener 2000: 572) should be cal margin produced as flattened hairy changed so that couplet 17 omits the process that looks like slender hairy spine phrases about the mesepisternal tuber- in ventral view (Fig. 5). Protibia and me- cle. Furthermore, because of probable con- sotibia each with distinct outer apical fusion at couplet 16, Rhathymini should spine; tibial spurs unmodified. Claws each run out not only at couplet 17 but also with flattened basal tooth; arolia present. through couplet 20. Change the outcome Scopa absent. Forewing with three sub- of the second alternative of couplet 20 to marginal cells; marginal cell large, longer 20a and add a new couplet as follows: Journal of Hymenoptera Research

at all sulcus ab- 20a (20). Maxillary palpus absent and axilla small, not produced; epistomal areas are sent below anterior tentorial pit so that clypeus and lower paraocular fused Apinae, Rhathymini Maxillary palpus present, or if absent, then axilla produced to point {Odyneropsis in 21 Epeolini); epistomal sulcus usually complete

KEY TO GENERA OF RHATHYMINI

Mesepisternum with large submedian tubercle; vein cu-a [cu-v] of hind wing strongly oblique and distinctly longer than second abscissa of M+Cu; supraclypeal area strongly elevated, crested medially, not continuing convexity of clypeus; ocellocular area depressed below level of adjacent areas Rhathymus Lepeletier and Serville Mesepisternum without tubercle; vein cu-a [cu-v] of hind wing less strongly oblique and slightly shorter than or subequal to second abscissa of M+Cu; supraclypeal area with surface generally a continuation of convexity of clypeus although with small frontal tubercle at lower end of frontal line; ocellocular area not depressed Rhathymodes Engel, Michener, and Rightmyer

Genus Rhathymus Lepeletier and Diagnosis. —The principal characters of Serville this genus are included as contrasting par- Figs. 6, 7, 10, 11 enthetical notations in the description of Rhathymodes, below. Colax and Serville 1825: 213. Nomen Lepeletier 4, Comments.—The name Bureauella was nudum. synonymized by Michener (2000: 739). Its Rhathymus Lepeletier and Serville 1828: 448. type species, briefly described by Domi- Type species: Rhathymus bicolor Lepeletier nique (1898), was very large, yellowish and Serville 1828, monobasic. Lepeletier with dark metasomal bands. It is a 1841: 539. Dalla Torre 1896: 323. Michener possible senior of versicolor 2000: 739. Silveira et al. 2002: 129. synonym Rhathymus Friese 1906. Friese a to Colax Lepeletier and Serville 1828: 448. Nomen (1912) gave key the praeoccupatum [nee Hubner 1819 (Lepidop- species, including species now placed in tera); Wiedemann 1824 (Diptera); et Stephens Rhathymodes and in the genus Odyneropsis 1829 of the (Hymenoptera)]. Type species: Rhathy- Epeolini. About 16 species-group mus bicolor and Serville Lepeletier 1828, names have been proposed for Rhathymus monobasic. Established as a of but the synonym actual number of species is prob- Rhathymus and Serville 1828 and Lepeletier ably smaller; the genus is in need of re- not therefore available (ICZN 1999: Art. vision and comprehensive cladistic study. 11.6); see also Michener (1997). So far Biology.— as known, Rhathymus Liogastra Perty 1833: 146. Type species: Rhathy- species are cleptoparasites of mus bicolor Lepeletier and Serville 1828, Epicharis monobasic. (Apidae: Centridini), apparently deposit- their a small Rathymus Smith 1854: 278. Lapsus calami et ing eggs through opening in the host's brood-cell closure praeoccupatum [nee Dejean 1831 (Coleoptera), (Camargo et et Gistel 1848 (Echinodermata)]. al. 1975; Hiller and Wittmarm 1994- Rozen Bureauella 1898: 61. 1969, Dominique Type species: 1991, 2003). The hospicidal, first in- Bureauella star insignis Dominique 1898, mono- dispatches the host larva before feed- basic. on the ing provisions (Rozen 1969, 1991). Volume 13, Number 1, 2004

Figs. 3-5. Rhathymodes bertonii (Schrottky). 3, Frontal view of head. 4, Forewing. 5, Ventral-oblique view of head and mesosoma; arrow indicates short, setose extension of procoxa (note absence of pleural tubercle). Journal of Hymenoptera Research

to Larval stages were described by Rozen gins weakly concave, meeting apically translucent of T6 (1969, 1991), McGinley (1981), and Camar- form apex (margins of T6 go et al. (1975) who also described the straight or weakly convex, apex formed extension pupa. Rozen (2000, 2001) gave additional opaque and largely by of characters for distinguishing the mature of elevated discal part pygidial plate). larva and pupa of Rhathymus and Rozen 9. Apical process of second valvifer of fe- above base of (2003) described the eggs (as mature oo- male forming slender hook = robust and cytes) of two species. Raw (1991, 1992) gonoplac [ gonostylus] (more gave an account of the post-emergence not hooked). 10. Male S4 and S5 simple, well surfaces as as flight behavior of male Rhathymus as transverse, exposed long as some host data. those of adjacent sterna (male S4 and S5 broadly emarginate, much shortened me- Engel, Michener, and Rhathymodes dially so that only narrow margins are ex- new genus Rightmyer, posed, thus exposed part of S6 large). 11. 1-5, 8, 9, 12, 13 Figs. — Lateral extremities of male S5 not pro- Type species. Rhathymus acutiventris duced (strongly produced posterolaterally Friese 1906. and hairy, supporting hair tuft noticeable — 2000: 102- Diagnosis. The generic characters are from above, see Michener fig. listed below, each followed by the state of 2). 12. Male S3-S5 with apical fringes of the same character in Rhathymus, in paren- erect, curved or sigmoid hairs (fringes ap- theses. Body length 13-18 mm (16-28 mm pressed, not conspicuous, well developed in Rhathymus). 1. Supraclypeal-frontal area only on S4 and S5). 13. Genitalia and hid- convex, in profile continuing convexity of den sterna as in figures 8-9, 12-13 (cf. clypeus, frontal tubercle and carina above Figs. 6-7, 10-11 for Rhathymus); lateral it rather weak, see figures 2-3 (this area sclerotization of spatha arcuate and pro- strongly produced as crest, in profile ele- duced (not arcuate, not produced). vated above imaginary continuation of Included species. —Three names have clypeal convexity, sloping steeply at sides been provided for species of Rhathymodes, into depressions around antennal bases). as follows: Rhathymodes acutiventris 2. Ocellocular area not depressed (strongly (Friese), R. friesei (Ducke), and R. bertonii depressed below level of adjacent areas). (Schrottky). All are new combinations 3. Scutellum bituberculate, the two con- and, as indicated below, the first two are vexities (sometimes weak) forming line subjective synonyms. between dorsal and posterior declivitous Etymology. —The new genus-group surfaces (with shining ridge, sometimes name is a combination of Rhathymus (Gr. to form weak bituber- depressed medially rhathymos, meaning "carefree" or "lazy") on line between dorsal de- culation, and and the suffix -odes (Gr., an adjectival declivitous 4. surfaces). Mesepisternum rivative of eidos, meaning "resembling"). without anteromedian tubercle (with The gender is masculine. large, mostly impunctate and hairless, an- Distribution. —Jalisco, Mexico, to Para- teromedian 5. Mesobasitarsus tubercle). guay, essentially the same as for Rhathymus. shorter than mesotibia as meso- (as long Phylogenetic commentary. —Comparison 6. with second submar- with other tibia). Forewing tribes of Apinae suggests that cell lm-cu first recur- ginal receiving [= characters 1, 2, 4, 10, and 11 are plesiom- rent near middle or distal see vein] third, orphic relative to Rhathymus. Perhaps 4 (near of cell). 7. Hind characters 3 figure apex wing and more certainly 9 and the with cu-a [c-v] to or shorter than in 13 are subequal spatha derived (admittedly, the second abscissa of M+Cu 8. (longer than). sting and male genitalia are unknown in of female with lateral Pygidial plate mar- R. bertonii). Volume 13, Number 1, 2004

KEY TO SPECIES OF RHATHYMODES (based on females only)

1 Mandible longer than minimum distance between compound eyes; third submarginal cell as long as to longer than second submarginal cell; mesoscutal surface with four yellow stripes; propodeum yellow with black, median, longitudinal stripe on posterior surface (narrower and slightly fainter in specimens from Nicaragua); metasoma yellow or dusky yellow R. acutiventris (Friese) - Mandible shorter than minimum distance between compound eyes (Fig. 3); third submarginal cell markedly shorter than other submarginal cells (Fig. 4); mesoscutal surface appearing yellow with two black stripes (Figs. 1, 3); propodeum yellow without black, median, longitudinal stripe (Fig. 1); metasoma mostly black (Figs. 1-2) R. bertotiii (Schrottky)

Rhathymodes acutiventris (Friese), rior mesoscutal margin, submedian bands new combination sometimes partly fused; pretarsal claws; Figs. 8-9, 12-13 longitudinal median stripe on posterior surface of propodeum; apex of pygidial Rhathymus acutiventris Friese 1906: 120. Friese (sometimes brown rather than 1912: 226. Holotype [Budapest], examined. plate black). Ventral surface of Rhathymus friesei Ducke 1907: 458. Friese 1912: flagellum yel- 225. Silveira et al. 2002: 129. Lectotype [Ber- low-brown except yellow Fl. Tibial spurs lin], examined. New synonym. dark brown. Metasomal terga sometimes — dusky yellow with lighter yellow apical Description. Body length 13-18 mm. bands, although usually uniformly yel- Mandible of female longer than minimum low. Wings transparent brownish, veins distance between eyes. Third submarginal and pterostigma dark brown to blackish. cell measured on posterior margin nearly Pubescence golden, short except on distal as long as to longer than second. Body and part of labrum (where it forms two tufts), legs yellow (sometimes brownish yellow), genal area, sides of mesosoma and pro- the following areas black or blackish: api- of mesoscutum of cal half of mandible; labrum except for podeum; pubescence rather uniform shorter than ocellar two brownish spots near base; subanten- length, diameter, erect; of metasomal nal sulci and epistomal sulcus between pubescence terga appearing dark against anterior tentorial pits; upper surface of an- appressed, Punctation fine and tenna (sometimes brownish) except some- yellow background. dense so that most surfaces are dull but times yellow or yellow-brown on base and labrum, apex of flagellum; ocellar area extending clypeus (especially impunctate lower lobe and down on either side of supraclypeal ele- margin), pronotal hypoe- area because vation to antennal bases and usually ex- pimeral shining punctures less metasomal tending laterally on occiput behind sum- dense; terga especially dull because of fine mit of compound eye; transverse spots on uniformly punctuation and dense short hair. Male hidden anterior surface of pronotum; spot at an- sterna and as in 12-13. terior base of pronotal lobe; mesoscutum genitalia— Figs. 8-9, except for lateral marginal yellow band, Variation. Because of variation in scu- that the broadened anteriorly, and submedian lon- tellar form we at first believed gitudinal yellow band, thus four yellow specimens here placed in R. acutiventris bands on mesoscutum, none of them at- represented two or more species. Fre- taining anterior and posterior mesoscutal quently the convexities are prominent so margins or lateral bands attaining poste- that the declivitous surface below their Journal of Hymenoptera Research

and male 6, Ventral (left) and dorsal (right) views. Figs. 6-7. Rhathymus bicolor Lepeletier Serville, genitalia. = = seen in different views (a valve; b 7, Lateral view. Letters serve to indicate identical structures penis = = e = lower f = aedeagus; c upper gonocoxite; d upper gonostylar process; gonostylar process; spatha).

summits is about one and one half times Mexico and Honduras and find no differ- lat- ences between them. as long as the dorsal surface (seen in — eral view). Less commonly the convexities Material examined. MEXICO: Jalisco: are smaller. Thus, in a specimen from Su- 19, Chamela, 13 July 1990 (R. Ayala) of riname the declivitous surface is more [Chamela]. Chiapas: 16 , Agua Azul, N than twice as long as the dorsal surface Ocosingo, 23 April 1993 (F. Noguera) in a 1 9 and this condition is approached [Lawrence]; , Parque Laguna Belgica, specimen from Cerro Campana, Panama, 19.3 km N of Ocozocoautla, 1560 m, 12 the 1991 although another collected on same June (J. Ashe) [Lawrence]. GUATE- 3.5 of La day had larger convexities. From the few MALA: 19, Zarapa, km SE see 1500 23 1993 R. specimens available we no geograph- Union, m, Jun. (J. Ashe, ical significance in the scutellar variation. Brooks) [Lawrence]. HONDURAS: Atlan- It would have been desirable to examine tida: 19, Lancetilla Botanical Garden, male terminalia from all parts of the range Tela, 10 m, 15°46'N, 87°27'W, 22 June 1994 from south- R. but males are available only (J. Ashe, Brooks, methyl salicylate and ern Mexico, Honduras, and Panama; we eucalyptus oil attractants) [Lawrence]. have made dissections of specimens from Cortez: 36, 39, Parque Nacional Cerro Volume 13, Number 1, 2004

Figs. 8-11. Male internal sterna. 8-9, Khathymodes acutiventris (Friese): 8, Eighth sternum. 9, Seventh sternum. 10-11, Rhathyinus bicolor Lepeletier and Serville: 10, Eighth sternum. 11, Seventh sternum.

Azul-Meambar, Los Pinos, 800 m, 1995 (J. Ashe, R. Brooks) [Lawrence]. Co- 14°42.4'N, 87°54.7'W, 10-16 May 2002 (S. lon: 39, 14 km N of junction of Escobal Peck) [Lawrence]. NICARAGUA: Grana- and Pina roads, 30 m, 2 June 1996 (J. Ashe, da: 39, Volcan Mombacho, Santa Ana 2, R. Brooks) [Lawrence]; 26 , Barro Colora- malaise M. 29 1977 B. L. S. Kim- 15 May 1998, trap (J. Maes) do Island, May (R. & [Lawrence]. 1 9, Volcan Mombacho, Santa sey) [Lawrence]. SURINAME: Brokopon- 21 M. do: 1 Nature Witi Ana 2, June 1998, malaise trap (J. 9 , Brownsberg Preserve, Maes) [Lawrence]. COSTA RICA: Alajue- Creek Trail, 80 m, 4°56'55"N, 55°10'53"W, la: 19, Cano Negro, 20 m, 10-19 March 23-25 June 1999 (Z. Falin, A. Gandadin, H. 1993 (K. Martinez) [Heredia]. Gnanacaste: Hiwat) [Lawrence]. BRAZIL: Para: 16, 19, Maritza Biological Station, 550 m, 22 Estado do Para, Vbidos (Rio Branco), A. Pernambuco: May 1993 (J. & Ashe) [Lawrence]. 19, VIII.1912, Ducke [Berlin]. Parque Nacional Guanacaste, Est. Pitilla, 9 19, Caruaru, 900 m, April 1972 (M. Al- km S of Est. Cecilia, 700 m, 19 May-3 June varenga) [New York]. Minas Gerais: 16, 1993 (P. Rios) [Heredia]. Puntarenas: 19, Barbacena, 14.12.1905, Ducke [lectotype of or Fila Moras, Buenos Aires, A. C. Amistad, R. friesei, Berlin, see below]. Sao Paulo 1000m, 19 May 1993 (M. A. Zumbado, S. Parana: 16, Rio Parana, Siid-Brasil, 1904 Rojas) [Heredia]. PANAMA: Chiriqni: 19 [description and photograph, Friese 1912: [type of R. acutiventris, in Budapest, ex- 225, examined, Berlin]. — amined]. Panama: 29, Cerro Campana Lectotype designation. Several speci- Friese (Capira), 825 m, 8°44'N, 79°57'W, 1-5 June mens of R. friesei are located in the Journal of Hymenoptera Research 10

acutiventris male 12, Ventral (left) and dorsal (right) views. 13, Figs. 12-13. Rhathymodes (Friese), genitalia. Lateral view. Lettering as in Figs. 6-7.

— over half of the collection [Berlin] and many bear his char- Comments. Just speci- acteristic "typus" labels. As is unfortu- mens were collected by flight intercept set in forests. used nately not rare for specimens bearing such traps up Frequently by such to be useful labels, many are not part of the original coleopterists, traps prove were collected at in forest bees seen bee type series (e.g., several collecting rarely by localities not mentioned in Ducke's origi- collectors. nal account or, worse yet, were collected bertonii years after the publication of the species!). Rhathymodes (Schrottky), new combination However, among the material is at least 1-5 the specimen collected by Ducke at Bar- Figs. bacena in Minas Gerais (the other speci- Khathymus bertonii Schrottky 1920: 217. Holo- mens mentioned Ducke are not in Ber- by type [San Lorenzo], compared. and this is one which lin), assuredly upon — he based his description. For the express Diagnosis. Similar to R. acutiventris ex- purpose of nomenclatural stability we cept as follows: Mandible shorter than hereby designate and label this individual minimum distance between compound Lectotype: 1 Estado of male as lectotype. 6 , Brazil, eyes (Fig. 3), suggesting mandible de Minas Ger., Barbacena, 14.12.1905, of R. acutiventris. Third submarginal cell Ducke [Berlin]. markedly shorter than others (Fig. 4). Me- Volume 13, Number 1, 2004 11

information on from Drs. Bolivar R. Garcete- soscutum with mesal yellow stripes fused types Barrett, William L. Overal, Orlando Tobias Silveira, so that surface appears yellow with two Frank Koch, Charles Huber, and Elsa Obrecht. Im- black (as in the male) and very nar- stripes portant comments on the manuscript were provided row black anterior 1, margin (Figs. 3) by Drs. Jerome G. Rozen, Jr., Michael Ohl, and E. Eric (male also with minute black triangle at Grissell. We were able to examine the holotype of R. acutiventris thanks to Dr. Zambon of the Hun- posterior margin). Propodeum entirely Lajos garian Natural History Museum, Budapest, and Mr. yellow, without black median stripe (Fig. Gyorgy R. Makranczy who couriered the specimen. 1). T3-T6 black (with black bands in MGR was supported by a NSF Predoctoral Fellow- T3 and T4 with male), apical margins ship. This is contribution number 3323 of the Division brownish translucent (Fig. 1); preapical of Entomology, Natural History Museum and Biodi- Research of Kansas. part of pygidial plate reddish yellow, oth- versity Center, University erwise black. plate LITERATURE CITED The inner metatibial spurs appear broken alike on the two the R. S. F. apically, sides, Camargo, J. M. F., Zucchi, and Sakagami. 1975. Observations on the bionomics of truncate apices slightly darkened suggest- Epicharis rustica notes ing that the truncation is normal or that (Epicharana) flava (Olivier) including on its parasite Rhatkymus sp. (Hymenoptera, breaking occurred during the life of the Apoidea: Anthophoridae). Studia Entomologia 18: of the are insect. The spurs male holotype 313-340. not truncated, having typical, pointed api- Dalla Torre, C. G., de [K. W., von] 1896. Catalogus ces, furthering the suggestion that the Hymenopterorum hucusque descriptorum systemati- et X: truncation of the female is the result of ca synonymicus. Volumen Apidae (Anthophi- la). Engelmann; Lipsiae [Leipzig], Germany; 643 breakage. PP- Variation. —The feature is var- following Dejean, P. F. M. A. 1831. Species General des Coleopteres iable within R. acutiventris but is recorded de la Collection de M. le Comte Dejean [tome 5]. here for our specimen of R. bertonii: scu- Mequignon-Marvis; Paris, France; 8+883 pp. 1898. d'oeil sur les melliferes tellum strongly bituberculate with ascend- Dominique, J. Coup sud-americaines du museum de Nantes. Bulletin ing dorsal surface measured to apices of de la Societe des Sciences Naturelles de Vouest de la tubercles about two-thirds as as de- long France 8: 57-65. clivitous surface. — Ducke, A. 1907. Beitrag zur Kenntnis der Solitarbi- Material examined. Aside from the male enen Brasiliens (Hym.). Zeitschrift fur Systematis- holotype from Puerto Bertoni, Paraguay che Hymenopterologie und Dipterologie 7: 321-325, 361-368, 455-461.' [San Lorenzo], this species is known from M. S. 2001. A of the Baltic amber one female as follows: PARAGUAY: Par- Engel, monograph bees and evolution of the Apoidea (Hymenop- Nacional de aguari, Parque Ybycui [ca. tera). Bulletin of the American Museum of Natural 26°S, 57°W], 300 m elevation, December History 259: 1-192. H. 1906. Neue Schmarotzerbienen aus der neo- 13-18, 1989 (M. Cooper) [Lyme Regis]. Friese, Zeitschrift fur Comments.—This species was described tropischen Region. Systematische Hymenopterologie und Dipterologie 6: 118-121. from a male from Paraguay, preserved in Friese, H. 1912. Neue und wenig bekannte Bienen- the Museo Nacional de Historia Natural arten der neotropischen Region. Archiv filr Na- del In the Martin col- Paraguay. Cooper turgeschichte, Abteilung A 78(5): 198-226. Gistel 1848. des lection is one female that agrees in several [Gistl], J. Naturgeschichte Tierreichs,fiir hohere Schulen. Hoffmann; respects with the male and that we regard Stuttgart, Germany; xvi + 216+[4] pp., 32 pis. as R. bertonii. The above diagnostic re- Hiller, B., and D. Wittmann. 1994. Seasonality, nest- marks are based on the female with some ing biology and mating behavior of the oil-col- notes derived from the parenthetical orig- lecting bee Epicharis dejeanii (Anthophoridae, Porto 107- inal description of the male. Centridini). Biociencias I Alegre] 2(1): ACKNOWLEDGMENTS 124. Hiibner, J. 1816-1826. Verzeichniss bekannter Schmet-

We appreciate the loan of material from several tlinge [sic: Schmetterlinge]. Privately published; sources as indicated in the Introduction, as well as Augsburg, Germany; 431 + 72 pp. [Dating of sec- 12 Journal of Hymenoptera Research

177- A. 1992. Mate and size of tions: 1816, 1-16 pp.; 1819, 17-176 pp.; 1820, Raw, searching population 257-304 two univoltine, of the bee 208 pp.; 1821, 209-256 pp.; 1823, pp.; solitary species genus in Brazil with records of 1825, 305-431 pp.; 1826, Anzeiger 1-72 pp.] Epicharis (Hymenoptera) International Commission on Zoological Nomencla- threats to nesting populations. Entomologist 1-9. ture. 1999. International Code of Zoological Nomen- 111(1): 1969. The larvae of clature [Fourth Edition]. International Trust for Rozen, J. G., Jr. Anthophoridae Part 3. The Zoological Nomenclature; London, UK; (Hymenoptera, Apoidea), Melectini, Ericrocini and American Mu- xxix + 306 pp. [sic], Rhathymini. Na- seum Novitates 2382: 1-24. Lepeletier de Saint Fargeau, A.L.M. 1841. Histoire — 1991. Evolution of in turelle des Insectes Hymenopteres [tome second]. Rozen, J. G., Jr. cleptoparasitism of Roret; Paris, France; 680 pp. anthophorid bees as revealed by their mode A. L. G. Au- and first instars Lepeletier de Saint Fargeau, M., and J. parasitism (Hymenoptera: Apo- dinet-Serville. 1825, 1828. [Sections] In G. A. idea). American Museum Novitates 3029: 1-36. 2000. of some Olivier (ed.), Encyclopedic Mithodique, ou par ordre Rozen, J. G., Jr. Pupal descriptions clep- de matieres. Histoire naturelle. Insectes [vol. 10]. toparasitic bees (Apidae), with a preliminary ge- bees Agasse; Paris, France; part 1, 1825, 1-344 pp.; neric key to pupae of cleptoparasitic (Apo- Novitates 1-19. part 2, 1828, 345-832 pp. idea). American Museum 3289: R. 1981. of the Colletidae 2001. A taxonomic to mature lar- McGinley, J. Systematics Rozen, J. G., Jr. key based on mature larvae with phenetic analysis of vae of cleptoparasitic bees (Hymenoptera: Apo- apoid larvae (Hymenoptera: Apoidea). University idea). American Museum Novitates 3309: 1-27. ovariole and of California Publications in Entomology 91: 1-307. Rozen, J. G., Jr. 2003. Eggs, numbers, Michener, C. D. 1997. Genus-group names of bees modes of parasitism of cleptoparasitic bees, with and supplemental family-group names. Scientific emphasis on neotropical species (Hymenoptera: Papers, Natural History Museum, University of Kan- Apoidea). American Museum Novitates 3413: 1-36. sas 1: 1-81. Schrottky, C. 1920. Himenopteros nuevos o poco con- Michener, C. D. 2000. The Bees of the World. Johns ocidos sudamericanos. Revista do Museo Paulista Hopkins University Press; Baltimore, MD; 12: 179-227. xiv + [l]+913 pp. Silveira, F. A., G. A. R. Melo, and E. A. B. Almeida.

Perty, J. A. M. 1830-1833. Delectus Animalium Articu- 2002. Abelhas Brasileiras: Sistemdtica e Identificacdo. latorum, quae in itinere per Brasiliam annis Ed. IDMAR; Belo Horizonte, Brazil; 253 pp. MDCCCXVII-MDCCCXX jussu et auspiciis Maxi- Smith, F. 1854. Catalogue of the Hymenopterous Insects

miliani I. fosephi Bavariae regis augustissimi peracto in the Collection of the British Museum, part 2. Brit-

Dr. B. de . . et Dr. C. F. Ph. de collegerunt J. Spix. ish Museum; London, UK; 199^465 pp., vii-xii Martins. Digessit, descripsit, pingenda curavit Dr. pis.

Maximilianus . . est et edidit Carol. F. Perty. praefatus Stephens, J. 1829. The nomenclature of British insects;

Frederic. de Martins. . . Philip, Accedit dissertatio de being a compendious list of such species as are con- Insectorum in America meridionali habitantium vitae tained in the systematic catalogue of British insects, moribus et distributione genere, geographica. and forming a guide to their classification, &c, &c. Monachii Hiibschmann; [Munich], Germany; Baldwin and Cradock; London, UK; [2] +68 pp. + [viii] iii+44+224 pp., 40 pis. [Liogastra dates Wiedemann, C. R. G. [C. R.W.] 1824. Munus rectoris from 1833] in Academia Christiana Albertina aditurus analecta A. 1991. The circuitous trails Raw, used by males of entomologica ex Museo Regio Havniensi maxime con- the cuckoo bee, Rhathyiuus fulvus (Hymenoptera, gesta profert iconibusque illustrat. E regio typgra- in forest in Brazil. En- Anthophoridae) Salvador, phico scholarum; Kiliae [Kiel], Germany; 60 pp., tomologist 110(3): 110-113. lpl. J. HYM. RES. Vol. 13(1), 2004, pp. 13-23

A New Species of Oozetetes DeSantis (Hymenoptera: Chalcidoidea: Eupelmidae) Attacking Oothecae of Nyctibora acaciana Roth (Orthoptera: Blattellidae)

Gary A. P. Gibson

K. W. Agriculture and Agri-Food Canada, 960 Carling Avenue, Neatby Bldg., Ottawa, Ontario, Canada, K1A 0C6; email: [email protected]

Abstract. —Oozetetes nyctiboraphagus Gibson, new species, an egg predator in the oothecae of Nyctibora acaciana Roth, is described from Costa Rica and Nicaragua. The description of the male of Oozetetes and a is the first for the genus. The species is assigned to the bucheri species group Basal key is provided to distinguish females of the five described species of this group. regions of the forewing differentiated by convex or concave folds and sometimes setal lines are compared and a with putatively homologous forewing veins and cells of other Hymenoptera, comprehensive set of names is applied to these based on homology and historical usage in chalcid literature.

Oozetetes was established by De Santis ceicornis (Cameron, 1884) from Panama. (1970) for a single species, O. bucheri De Oozetetes compressicornis (Cameron, 1884) Santis, which was reared from oothecae of and O. gigas (Cameron, 1884), both from Pseudoischnoptera lineata (Olivier) (Orthop- Panama, are the only two species classi- tera: Blattellidae). Gibson (1995) subse- fied in the compressicornis-group. Males these quently transferred five species into Ooz- are unrecognized for any of species. etetes from other genera and recognized Host information is also lacking for any other than the two species groups, the bucheri- and com- described species type spe- I a female and male of pressicornis-groups, based on structures of cies, though saw ootheca the scape and face of females. The bucheri- an undescribed species from the group consists of species with females of an unidentified cockroach (Gibson was having a more or less tubular scape (Figs. 1995). The same parasitoid species of 13, 14), a distinct parascrobal region along subsequently reared from a new species the inner orbit so that the scrobal depres- cockroach and information concerning its behavior sion extends to the anterior ocellus (Figs, rearing and possible oviposition de- l-k), and the interantennal region flat or was given when the cockroach was acaciana Roth even slightly depressed above the level of scribed as Nyctibora (Orthop- tera: Deans and Roth the toruli (Fig. 1). The primary feature Blattellidae) by de- characterizing compressicomis-group fe- (2003). The rearing of two of seven cock- males is a strongly compressed scape, scribed species of Oozetetes from which is correlated with a different struc- roach oothecae suggests that all members as ture of the scrobal depression and paras- of the genus might use cockroaches crobal region than for bucheri-group fe- hosts. I have seen females representing from males (see discussion and figures in Gib- numerous species of the genus trop- son 1995). Classified in the bucheri-group ical and subtropical regions of the New are O. bucheri from Argentina, O. magni- World, as far north as Florida (Gibson clavatus (Ashmead, 1904) and O. splendens 1995). The purpose of this paper is to pro- (Walker, 1862) from Brazil, and O. testa- vide a name for the new species reared of Hymenoptera Research 14 Journal

concave folds. The from N. acaciana in order to facilitate on- convex and regions, to differ- folds and setal lines are analogous, if not going behavioral studies. A key to cells and distinct veins in entiate females of the five described buch- homologous, as illustrated Hub- is also as an other by

• several setal regions by lines or by fold) (Fig. 17, vnf), further differentiates Volume 13, Number 1, 2004 15 the cubital area from the posteriormost margins straight transverse or only very basal region of the wing, the vannal area slightly emarginate; syntergum with api- is cal into transverse (= claval area) (Fig. 17, vna), which re- margin reflexed flange, flexed so as to be subhorizontal or inclined Remarks. —Females of Oozetetes can be relative to the cubital area. Although only differentiated from other eupelmine fe- inconspicuously developed in Oozetetes, males using Gibson's (1995) key to genera many chalcids have another longitudinal based on females. As for other Eupelmi- convex fold near the posterior margin of nae, the sexes of Oozetetes are highly di- the wing. This fold apparently is not ho- morphic and the single male then known mologous with a vein, but can be termed was keyed together with males of A. {An- of their sim- the subcubital fold (Fig. 17, scf) because, astatus) Motschulsky because similar an- if present, it extends along the subcubital ilar bidentate mandibles and Anastatus is a setal line sensu Gibson (1997, fig. 5). The tennal structures. speciose, convex subcubital fold results in the pos- cosmopolitan genus whose members usu- terior margin of the forewing being ally are primary endoparasitoids of eggs curved down and it is this portion onto of several insect orders. Gibson (1995) sug- which the dorsally curved hamuli of the gested that apically broad, bidentate man- hind wing hook. Because of the various dibles in Eupelminae may be indicative of longitudinal folds the surface of the wing an endoparasitoid of eggs. Further, Ooze- is pleated behind the basal cell, often be- tetes may represent a monophyletic line- ing more or less M-like as if viewed in age having a specialized host relationship that renders Anasta- cross-section (Fig. 17, insert). with cockroach eggs tus paraphyletic. This latter possibility is Oozetetes De Santis indicated because the males of the two

^ „ ~„~ „~ „„ m genera are morphologically' very similar, Oozetetes Santis 1970: 32-33. . , / , De Type species: f L / °.,, , ~ ,, , , .^r, ,., but several features differentiate females Oozetetes bucheri De Santis, by monotypy and of the two taxa. Association of the sexes original designation. of species other than O. nyctiboraphagus, Diagnosis. —Mandible bidentate, with particularly those of the compressicornis small ventroapical tooth and broad dor- group, is necessary prior to determining soapical margin. Eye bare or superficially whether there are reliable features to dif- those of glabrous, at most only very sparsely and ferentiate males of Oozetetes from in inconspicuously microsetose (Figs. 1-4). Anastatus (see discussion Gibson 1995). Male antenna with pedicel only slightly Oozetetes nyctiboraphagus Gibson, new longer than broad and without row of se- species tae ventrally (Fig. 15); flagellum compact- ~ Fi s - l ' 2 ' 5 13' 15 ' 16 ' 18 ' 19) filiform with short seta and numerous ( § — multiporous plate sensilla in several rows Etymology. The species name is de- the and (Fig. 16); clava not conspicuously en- rived from Greek phagos (to eat) its host larged, only about as long as combined the generic name of only known Oozetetes is mascu- length of apical two funicle segments (Fig. cockroach, Nyctibora. Female with line and means searcher' or 'seeker', 15). propodeum (Fig. 7) plical 'egg— region conspicuously sculptured and with Type material. Holotype female: callar region convex and almost always "COSTA RICA: Guanacaste, Parque Na- largely or entirely sculptured and setose, cional Santa Rosa, Rd to Nicaragua, 1 km "A.R. coll. Female gaster (Fig. 9) dark or yellowish to N main Rd"; Deans, dry forest, orange, but without subbasal white band 30.VI.2001, em. 18.VII.2001, ex. Nyctibora tree" if light-colored; G^ with posterior margin sp. egg case on ant acacia (CNCI with No. male: same emarginate but Gt2-Gt 5 posterior Type 22872). Allotype Journal of Hymenoptera Research 16

Figs. 1-8. Oozetctes spp. 1-4, Head (9). 1 and 2, O. nyctiboraphagus. 3, O. splendens. 4, O. sp. nr O. nyctiboraphagus. 5-8, O. nyctiboraphagus (9). 5, Mesosoma (dorsal). 6, Mesosoma (dorsolateral) (arrow points to parapsidal band). 7, Apex of scutellum to base of gaster (dorsal). 8, Posterior half of acropleuron (lateral). Volume 13, Number 1, 2004 17

9-16. Figs. Oozetetes spp. 9-13, O. nyctiboraphagus. 9, Gaster (9, dorsolateral). 10, Head (6*, frontolateral). 11, Mesosoma (6, lateral). 12, Apex of scutellum to base of gaster (d, posterolateral). 13, Antenna (d). 14, O. testaceicornis, antenna (6). 15-16, O. nyctiboraphagus (6). 15, Flagellum. 16, Third flagellar segment. of Hymenoptera Research 18 Journal

and width: data as holotype (CNCI). Paratypes (108 9, tennal region clypeus; height = 19: in dorsal view as in 2, with 15c?): COSTA RICA: same data as holo- 16; Fig. ex- interorbital distance 0.32 times head type (169, 1 8); same data as holotype OOL 0.7 and POL 1.8 times mini- cept emerged 13.VII.2001 (139, IS), width; Guanacaste P.N. Santa Rosa Sect. Santa mum diameter of posterior ocellus; eye = Rosa, Acacia 005, 1.5 km E. Area Admin- height: eye width: malar space 11: 7.4: scrobal ex- istrativa, 25.VI.2001, Andy Deans, coll. un- 7.4; depression bell-shaped, ocellus and reticulate- der bird nest, emerged 13.VII.2001, oothe- tending to anterior to lower face ca of Nyctibora acaciana n. sp. Roth (22 9, rugulose strigose (Fig. 1); to microreticulate and 5

ex- under some of ulate-rugulose. Antenna dark brown apical tergum angles light, under some and coriaceous. cept scape with metallic luster shiny, only very finely — first funicular Variation and limits. Deans and angles of light and segment species ratio of Roth a lateral habitus yellowish; in lateral view (Fig. 15) (2003) provided funicu- of a female and male. scape (excluding radicle): pedicel: photograph (fig. 4E) = 4.2: Males I include in the series are all lar segments: clava 10: 2.6: 1.0: 5.0: type for one 3.2: 3.6: 3.0: 3.0: 6.6. Mesosoma in dorsal very similar to one another, except reared individual 13.VII.2001, view primarily dark with slight cupreous (emerged This individual has the head and luster under some angles of light except CNCI). antenna of a but the remainder of following more distinctly metallic green to female, the of a male the bluish: pronotum laterally, convex lateral body except mesopleuron is somewhat "feminized", a portion of mesoscutum, and propodeum; having less concave mesoscutum with inconspicuous whitish much larger acropleuron and setae and scutellar-axillar complex with femoral depression than other males. from the unidentified dark setae; propodeum with callus setose The female reared has F1 -F1 lateral to level of spiracle and with a few species of cockroach 6 8 yellowish setae between spiracle and bare plical re- and the other flagellar segments only light in lateral brown also gion (Fig. 12). Mesosoma view (left flagellum misshapen, near metallic green to bluish except following with Flg-FL fused and constricted dark without metallic luster: acropleuron, middle). All other females I include in the the holo- femoral depression, upper portion of me- type series are very similar to sepisternum and oblique transepisternal type, having the pedicel and flagellum line. Forewing hyaline with cc: mv: pmv: brown as well as the interorbital distance = = stv 19.6: 9.6: 8.0: 4.4; stigmal vein con- 0.30-0.33 times the width of the head (n = spicuously curved and tapered, similar to 10, x 0.31), a short but distinct OOL (Fig. female; costal cell setose ventrally and 2), and similar forewing setal and acro- bare dorsally except for 4 setae near an- pleural sculpture patterns. Several females terior margin medially; basal cell setose from Belize, Brazil, Costa Rica, Ecuador, and with continuous line of setae along Guatemala, Peru and Venezuela are simi- mediocubital and cubital folds, the two lar to females I recognize as O. nyctibora- folds only slightly angulate relative to phagus, but are excluded from the type se- each other; cubital and vannal areas bare; ries based on different combinations of disc with distinct lunate bare region be- features, including the relative width of basal cell yond but region closed posteri- the interorbital distance and OOL (e.g. line orly by of setae along cubital fold. Fig. 4), the extent to which the cubital and Legs dark brown with metallic green to vannal areas are setose, the length of the blue luster except the following white: tro- postmarginal vein relative to the stigmal chantelli, knees and apices of tibiae of fore vein, and sculpture of the acropleuron and middle legs, and basal 3 tarsal seg- posteriorly. I believe O. nyctiboraphagus ments of middle and hind legs (protarsi forms part of a species complex of several brownish-white). Metanotum overlain by very similar species that differ from each of scutellum apex (Fig. 11), band-like and other by different combinations of rela- coriaceous to reticulate (Fig. 12). Propo- tively subtle features. Additional rearings deum reticulate-coriaceous with inverted and specimens collected from diverse lo- Y-like median carinal complex and with a cations are required to test this hypothesis. few oblique rugae extending from poste- Biology. —Deans and Roth (2003) reared rior margin (Fig. 12). Metasoma in dorsal O. nyctiboraphagus from the oothecae of view primarily dark brown but with me- Nyctibora acaciana Roth. Based on the un- allic luster on Gt and on green basally x systematic arrangement of wasp pupae Volume 13, Number 1, 2004 21 within the oothecae, they suggested the Blattidae) while this is attached to the fe- wasp larva was mobile and feeds on many male and can not locate the ootheca if it is rather than just a single egg, that is, it acts covered by sand or sawdust. The ovipo- more like a predator than a true parasit- sition behavior and preferred time and oid. The wasps emerged about 2-3 place of oviposition for O. nyctiboraphagus months earlier than the cockroaches, from remains to be determined. Deans and mid-August to late September, with about Roth (2003) noted that other common 25-30 wasps emerging per ootheca cockroach oothecal inhabitants are un- through 1-3 holes chewed by emerging known for JV. acaciana and suggested this adults (Deans and Roth 2003, fig. 4D). species derives significant protection by Deans was unsuccessful in inducing developing on ant-acacias. However, if O. reared female wasps to oviposit into the nyctiboraphagus is the only species that at- oothecae of N. acaciana, either by confining tacks the oothecae, this may indicate it is females with free oothecae, females with the only species able to escape ant preda- oothecae still attached to female N. acaci- tion and successfully oviposit into oothe- ana, or by placing female wasps on a new- cae attached to acacias protected by ants. ly deposited oothecae on an ant-acacia in- Gibson (1986) commented on the prodi- habited by Pseudomyrmex spinicola (Emery) gious jumping ability of female Eupelmi- (Hymenoptera: Formicidae). In one repli- nae. He correlated this and the extreme cate of the latter experiment, a female an- sexual dimorphism that characterizes the tennated the ootheca but jumped away at subfamily with a highly derived mesoso- the first contact with an ant. Although ovi- mal skeletomusculature in females. Jump- position was never observed, the authors ing in female Eupelminae does not appear suggested O. nyctiboraphagus probably to be for movement from place to place oviposits into the ootheca while this is still and Gibson (1986) postulated it probably attached to the female cockroach so as to evolved as a rapid escape mechanism avoid the hostile ant-protected environ- from predators, possibly ants and spiders. ment. They noted that Anastatus floridanus The ant-wasp interaction noted by Deans Roth and Willis oviposits into the ootheca demonstrates the ability of O. nyctibora- of Eurycotis floridana Rehn (Orthoptera: phagus to escape ant predation.

KEY TO FEMALES OF DESCRIBED BI/CHER/-GROUP SPECIES

1. Flagellum conspicuously clavate (Fig. 14), funicle with apical 4 segments strongly transverse, and clava about as long as combined length of apical 6 segments and about 2.5 times as wide as Fl O. a magniclavatus (Ashmead) - Flagellum not conspicuously clavate (Fig. 13), funicle with apical 4 segments subquadrate to elongate, and clava shorter than combined length of apical 4 segments and less than twice as wide as FL_ 2

2. Fore- and middle legs beyond coxae yellowish orange; forewing (De Santis 1970, fig. 1) with following bare: costal cell dorsally except apically near parastigma, basal cell basally and posteriorly, cubital area, mediocubital notch, and vannal area (Fig. 17) O. bucheri De Santis - Fore- and middle legs largely dark beyond coxae, brown or with metallic luster; forewing with costal cell sometimes more extensively setose than above, but otherwise only vannal area bare 3

3. Antenna entirely dark, the scape with distinct bluish-purple luster; fore- and hind legs dark, the femora with distinct bluish-purple luster, but middle leg with femur and tibia yellowish-orange; lower face and parascrobal region smooth to finely coriaceous with Journal of Hymenoptera Research 22

O. 9 with schematic cross-section (insert) (abbre- Figs. 17-19. Oozetetes spp. 17, bucheri, holotype forewing, cubital of costal viations: be, basal cell; bf, basal fold; cc, costal cell; cua, cubital area; cuf, fold; lec, length suborbital fold; cell; mcf, mediocubital fold; men, mediocubital notch; mdf, medial fold; scf, smv, submarginal Setae of vein; vna, vannal area; vnf, vannal fold). 18-19, 0. nyctiboraphagus (9). 18, Base of forewing. 19, forewing in region around campaniform sensilla of uncus.

distance about 0.2 times distinct, scattered, setiferous punctures (Fig. 3); interorbital head width; costal cell dorsally with complete band of setae along anterior margin; bluish under some of gaster dark dorsally, but laterally bright metallic green to angles base of to light; forewing uniformly light brown from near parastigma apex O. splendeiis (Cameron) and Antenna either with scape or flagellum yellowish; all legs mostly dark; lower face parascrobal region reticulate-rugulose with only indistinct, shallow setiferous punctures cell either (Fig. 1); interorbital distance at least 0.3 times head width; costal entirely setose or mostly bare; gaster brownish with only slight metallic luster posteriorly (gaster missing beyond Gt, for O. testaceicornis); forewing obviously more hyaline apically so 4 as to have distinct medial brownish region vannal Forewing (Fig. 18) with costal cell largely bare dorsally except near parastigma, Volume 13, Number 1, 2004 23

area apically and mediocubital notch both setose, and basal cell with about apical half white interorbital at having setae; distance least 0.30 times head width (Fig. 2); acropleuron reticulate-strigose medially to distinctly reticulate posteriorly (Fig. 8) O. nyctiboraphagus Gibson Forewing with costal cell entirely setose dorsally, vannal area and mediocubital notch bare (hence with large bare region between basal cell and disc), and basal cell with dark setae except along extreme apical margin (also small region of white setae immediately beyond basal fold and recurved portion of cubital fold); interorbital distance 0.22 times head width; acropleuron coriaceous ventrally to punctulate medially or punctulate-co-

riaceous posteriorly . . . . '. O. testaceicomis (Cameron)

ACKNOWLEDGMENTS on ant-acacias. Transactions of the American Ento- mological Society 129: 267-283. I thank Andrew Deans, University of Illinois, for De Santis, L. 1970. Un nuevo eupelmido Argentino de ootecas the specimens that instigated this study and Michael parasito de cucarachas silvestres (Hy- Gates (USNM), Sue Lewis (BMNH) and Steve Hey- menoptera). Revista del Museo de La Plata (Nueva don (UCDC) for the loan of additional specimens. Ms. Serie) 11: 31-35. G. A. P. 1986. Mesothoracic skeletomuscula- Jennifer Read is gratefully acknowledged for the Gibson, ture and mechanics of and in Eu- plates of scanning electron micrographs and John flight jumping Huber and Michael Gates are acknowledged for re- pelminae (Hymenoptera, Chalcidoidea: Eupel- Canadian viewing the manuscript and suggesting several im- midae). Entomologist 118: 691-728. A. P. provements. Gibson, G. 1995. Parasitic wasps of the subfamily Eupelminae (Hymenoptera: Chalcidoidea: LITERATURE CITED Eupelmidae). Memoirs on Entomology International 5: i-v + 421 pp. Gibson, G. A. P. 1997. Chapter 2. Morphology and Ashmead, W. H. 1904. Classification of the chalcid in: terminology. Pages 16-44 Gibson, G A. P., J. flies, or the Chalcidoidea, with de- superfamily T. B. Huber, and J. Woolley (eds.), Annotated Keys scriptions of new species in the Carnegie Muse- to the Genera of Nearctic Chalcidoidea (Hymenop- um, collected in South America by Herbert H. tera). National Research Council Canada, NRC Smith. Memoirs of the Carnegie Museum 1: i-xi, Research Press, 794 pp. 225-551, xxxi-xxxix. pis. T. Huber, J. and M. J. Sharkey. 1993. Chapter 13. Bucher, G. E. 1948. The of Monodontomerus anatomy Structure. in: T. Pages 13-64 H. Goulet and J. dentipes Boh., an entomophagous chalcid. Cana- Huber (eds.), Hymenoptera of the world: An iden- dian journal of Research 26: 230-281. tification guide to families. Agriculture Canada Burks, B. D. 1938. A study of chalcidoid wings (Hy- Publication 1894/ E. Ottawa. 668 pp. menoptera). Annals of the Entomological Society of Nichols, S. W. 1989. The Torre-Bueno Glossary of En- America 31: 157-161. tomology. Revised edition of A Glossary of Ento- Cameron, P. 1884. Tenthre- Hymenoptera (Families mology by J.R. de la Torre-Bueno including Sup- Centrali-America- dinidae-Chrysididae). Biologia plement A by George S. Tulloch. The New York na. Insecta 1: 1-487 + 20 pis. Entomological Society. New York. 840 pp. Deans, A. R. and L. M. Roth. 2003. Nyctihora acaciana Walker, F. 1862. Notes on Chalcidites, and characters a (Blattellidae: Nyctiborinae), new species of of undescribed species. Transactions of the Ento- cockroach from Central America that oviposits mological Society of London (3) 1: 345-397. J. HYM. RES. Vol. 13(1), 2004, pp. 24-30

A New Chilicola Spinola from Colombian Paramo (Hymenoptera: Colletidae: Xeromelissinae)

Victor H. Gonzalez and Charles D. Michener

Entomology Program, Department of Ecology and Evolutionary Biology and Entomology Division, Natural History Museum and Biodiversity Research Center, University of Kansas, Lawrence, Kansas 66045, USA; email: [email protected] and [email protected]

Abstract. — Chilicola (Anoediscelis) paramo Gonzalez and Michener, n. sp., from a Paramo in the Eastern Andes of Colombia is described. The new species is not a close relative of other known species and is not considered a member of the primarily Andean group of C. ashmeadi. Cliilicola paramo differs from other species in such a way that Michener's (2002) key to Andean subgenera of Chilicola requires modification; a new key is therefore provided here. Aspects of the nesting biology of the new species are also given. — Resumen. Chilicola (Anoediscelis) paramo Gonzalez and Michener, n. sp., es descrita de un Paramo de la cordillera Oriental de Colombia. La nueva especie no esta relacionada a ninguna otra especie conocida y no es considerada un miembro del grupo principalmente andino de C. ashmeadi. Chilicola paramo es bien diferente de las otras especies del grupo de tal manera que la clave de Michener (2002) para los subgeneros andinos de Chilicola necesita ser modificada; una clave nueva es presentada aqui. Tambien se presentan notas sobre la biologia de nidificacion de la especie.

Recently, Michener (2002) revised the lar diameter, and metasomal tergum, re- Andean of Chilicola tropical species Spi- spectively. Photomicrographs were pre- that is nola, those that occur from Peru to pared by Prof. Michael S. Engel using a Venezuela above 1000 m. Such Andean Microptis ML-1000 Digital Imaging Sys- were in three species grouped subgenera: tem. Type specimens are deposited in the Anoediscelis Toro and Moldenke, Hi/laeo- following institutions: soma Ashmead and Oroediscelis Michener; the Andean species of the first consisting IAVH Instituto Alexander von Hum- of the group of C. ashmeadi (Crawford). boldt, Villa de Leyva, Boyaca, Co- This is a paper supplement to that revision lombia (J. E. Castillo). since the new species described herein dif- SEMC Entomology Division, Natural fers considerably from other known spe- History Museum, University of cies in such a way that Michener's (2002) Kansas, Lawrence, KS 66045- to Andean key subgenera requires modi- 7523, USA (Z. Falin). fication. Therefore, one objective is to cor- rect that The field work key. Aspects of the nesting biol- was done by V.G. with ogy for the new species are also discussed. the help of Paula Montoya. All nests of the new were MATERIAL AND METHODS species collected during cold, cloudy weather when the bees should follows have been in their Morphological terminology nests, at the locality in- Michener (2000, 2002). Abbreviations used dicated below after the description, on in are F, and T for fla- descriptions S, OD, August 23, 2003. This is during the tran- metasomal ocel- gellar segment, sternum, sition from the rainy to the dry season. Volume 13, Number 1, 2004 25

Figs. 1-2. Chilicola paramo. 1, Forewing (photo of a paratype). 2, Face, holotype male.

Mean values are given with standard er- long, swollen scape (Fig. 2) and the swol- rors. len hind femur of the male (Fig. 3). Male. —Body length 5.5 mm; forewing Chilicola (Anoediscelis) paramo length 4.0 mm. Coloration (paratype in Gonzalez and Michener, n. sp. parentheses): Black, clypeus with small 1-8 Figs. than ocel- — pale yellow spot (Fig. 2), larger Diagnosis. Except as indicated under lus, on middle of lower clypeal margin Group Characters below, this species (pale area extending nearly full width of agrees with the characterization of the C. lower margin of clypeus and in middle, ashmeadi group by Michener (2002). It dif- up as acute point to two fifths of length of fers from members of that group by the clypeus); mandible brown (pale yellow) distal stigmal perpendicular crossing sub- except for narrow black base and black marginal cells near first submarginal apex; flagellum black (under surface dark crossvein (Fig. 1) and especially by the brown); tegula with vague brownish black of Hymenoptera Research 26 Journal

antennal base, and lower mesepisternum; short setae pedicel covered with setae; rather long, 20D, on vertex, genal area, outer sur- propleura, coxae, hind femur, face of hind tibia, upper part of side of propodeum where setae are rather dense and plumose, sides of metasomal terga Hind male. Fig. 3. Chilicola paramo. leg, where setae are denser along posterior margins and form weak, scarcely notice- front femur able fasciae and on area posterolaterally; apex of apical laterally, pos- and area on distal third (half) of anterior terior margins of metasomal sterna; apical setae of median surface of front femur brownish yellow; fringe of S2 with part bent outer surface of front tibia brownish yel- coarse, distal parts strongly posteri- Structure: HEAD: Face low; small areas at apex of mid femur and orly and branched. base of mid tibia dark brown (brownish as long as broad; interocellar distance sub- to ocellocular distance, about two yellow); base of hind tibia blackish brown equal distance about 1.5 (brownish yellow); hind tibia with inner OD; ocelloccipital OD; swollen thicker than distal apical projection black (brown); tibial scape (Fig. 2), and about twice width spurs testaceous; front and middle tarsi part of flagellum of than and partly brownish; wings faintly smoky, F4, longer clypeus reaching veins and stigma black; posterior margins middle of anterior ocellus; pedicel subcy- of Tl to T5 translucent brownish black, lindrical, about 1.5 times as long as broad; Sculpturing: Surface of head and thorax flagellum much slenderer than in C. ash- throughout dulled by micropunctuation, meadi, Fl slightly shorter than pedicel, the same on yellow clypeal area as on ad- about 1.5 times as long as broad, F2 short- jacent black areas; metasoma dulled by er than others, F4 and subsequent seg- transverse lineolation. Punctures of clyp- ments progressively broader and longer, eus, scutum, scutellum, and sides of tho- all except Fll about 1.5 times as long as rax small, weak, mostly separated by two broad. THORAX: Legs robust, hind femur or three puncture widths, those of scutum, swollen, about twice as long as broad, scutellum, and sides of thorax consider- hind tibia with apical half somewhat en- ably smaller and weaker than in C. ash- larged (Fig. 3), with inner apical projection meadi; punctures coarser and closer on rest extending distad; hind tarsus longer than of head, on frons as close as possible; tibia, hind basitarsus parallel-sided. M£- punctures nearly absent or unrecognizable TASOMA: Sterna scarcely modified, S2 on metasoma; dorsal surface or basal area with basomedian tumescence, apical mar- of propodeum granular with several irreg- gin of S2 more convex than transverse ular longitudinal carinae on anterior half margins of S3 and S4; S6 with posterior (two fifths), posterior margin of dorsal margin rounded, apicomedian fringe con- surface marked by transverse arcuate sisting of erect setae much longer than reduced that is ridge, medially, largely shown in Fig. 4 because much foreshort- smooth and shiny; posterior part of tegula ened in drawing, large sublateral seta eas- smooth and partly shiny; marginal zones ily broken off; S7, S8, and genitalia as in of Tl to T5 than more shiny discs but none Figs. 5-8; S7 with distal lobe trifid, sug- the less transversely lineolate. Pubes- gesting that of C. venezuelnna Michener but cence: dull setae of- Short, sparse, whitish, with setae on only one branch (Fig. 5, ten when viewed dusky against white compare with Fig. 8a of Michener, 2002). background; setae longest, 30D or more, Female.—Agrees with description of »n scape, paraocular area and frons near male except for usual sexual characters Volume 13, Number 1, 2004 27

4-8. Chilicola Figs. paramo, male. 4, S6. 5, S7. 6, S8. 7 and 8, Genitalia. In divided figures, dorsal view is shown on left, ventral on right.

the and following: Coloration: Black ex- spicuous as to be easily described as mere- under side of cept flagellum dark brown; ly seta bases, on frons separated by about basal half of front tibia with brownish yel- one puncture width and micropunctua- low on outer and sometimes anterior sur- tion conspicuous. Pubescence: Setae of faces; extreme apex of front femur some- scape and adjacent face less conspicuously times brownish yellow; tibial spurs testa- long than in male, but generally setae lon- ceous. Sculpturing: Punctuation even ger and denser than in male, especially weaker than in male, punctures of scutum, scopal setae of hind femora and metaso- scutellum, and sides of thorax so incon- mal sterna, the latter not forming fringes of Hymenoptera Research 28 Journal

the characterization of the C. ashmeadi as in male but wide spread, as usual for Michener as fol- Chilicolinae those of S2 especially long group by (2002) except lows: Size (over 5 mm in body and plumose. Pedicel with only scattered larger HEAD: orbital of small setae. Structure: Face very slightly length). Upper tangent not male of me- longer than broad; scape normal, passing through upper part of inner swollen, about as wide as F2 and thinner dian ocellus; emargination eye than rather in male, not mar- than distal part of flagellum, shorter margin strong smooth lower interocular clypeus, not reaching anterior ocellus; gined by strip; Fl distance more than half of intero- pedicel nearly twice as long as broad; upper interalveolar distance near- and F2 about as long as broad, subsequent cular distance; to alveolocular distance; suban- segments progressively wider, broader ly equal sutures labrum about than long until F8 and F9 which some- tennal subparallel; as wide as times appear about as broad as long or four times long; maxillary pal- un- three fourths as as even slightly longer than broad. Legs pus nearly long pre- modified, not swollen. mentum, segments progressively longer from base to so Holotype.—Male. COLOMBIA: Boyaca: and more slender apex, Arcabuco Prov., Santuario de Fauna y Flo- that distal segment is the longest and most well be- ra de Iguaque, Camino de la Laguna, 5° slender; labial palpus extending 70' N, 73° 46' W, 3400-3600 m, in dry yond glossa, second segment nearly twice others flower stems of Espeletia argentea, 23-VIII- as long as broad, subequal. THO- 2003 (V. Gonzalez & P. Montoya) [IAVH RAX: Pronotal collar nearly as wide as No. 4117]. maximum width of flagellum; stigma Paratypes. —One male and four females about two thirds as long as length of mar- with same data as the holotype, the male ginal cell on costa; distal stigmal perpendic- and two females in SEMC, the other two ular crossing submarginal cells near first sub- females at IAVH. No. 4118-9 crossvein dorsal surface of — marginal (Fig. 1); Etymology. The specific name is a noun propodeum about as long as scutellum; in apposition; the word Paramo is the posterior tibia of male much longer than name for a well known vegetation type of femur, almost four times as long as apical high altitudes (3500-4100 m) in the north- width, widest at apex because of inner ern Andes, dominated by the asteraceous apical projection (Fig. 3). METASOMA: Tl genus Espeletia Mutis ex Humb. & Bonpl., slightly longer than broad in male, slightly which provided the nesting sites for Chil- broader than long in female; marginal icola paramo. zones of terga only narrowly smooth at extreme of otherwise trans- GROUP CHARACTERS apices terga, versely lineolate like tergal discs although The new species, C. paramo, is a member more shiny than discs; penis valves with of the subgenus Anoediscelis in the sense single dorsoapical diverging membranous of Michener It (2000, 2002). agrees with processes (Fig. 7) (as in C. ashmeadi).

KEY TO THE TROPICAL ANDEAN SUBGENERA OF CHILICOLA

The mentioned in the to the problem, above, key subgenera of the tropical Andes (Michener, 2002: 8), is a result of the venational character italicized in the section on Group Characters in listing differences between C. and the of paramo description the C. ashmeadi group. The distal stigmal perpendicular crosses the submar- cells near the first crossvein ginal submarginal (Fig. 1), so that according to the first character of the first key C. couplet, paramo would be a species of the subgenus Oroediscelis. Other characters, however, show that C. is not otherwise similar to Oroediscelis but is a paramo species of Anoediscelis. The following is a corrected key: Volume 13, Number 1, 2004 29

1 Malar space one third as long as broad or more; S4 of male with pair of tubercles or projec- tions; hind tibia and usually basitarsus of male swollen and modified Oroediscelis - linear = one third as as S4 of male Malar space ( absent), rarely nearly long broad; simple or nearly so; hind tibia and basitarsus of male slender, not modified, or distal part of tibia slightly enlarged and produced as in C. paramo (Fig. 3) 2 2 Head above antennal alveolus with depression (sometimes evanescent) extending up toward ocellocular region; S8 of male with apical process deeply bifid; body length usually 4.0 to 5.5 mm but as little as 3.0 mm in C. smithpardoi Michener Hylaeosoma - Head without depression above antennal alveolus; S8 of male with apical process truncate (Fig. 6); body length usually 3.0 to 3.8 mm, but 5.5 mm in C. paramo Anoediscelis

NESTING BIOLOGY whitish, and was in contact with the pith walls of the tunnel except where it formed As described for C. espeleticola Michener the ends of the cells and thus the parti- (Michener, 2002), nests of C. paramo were tions between cells. Cells in series were found in dead, dry, broken, pithy flower- separated only by such partitions; spaces ing stems of living plants of the Paramo between cells were absent. The semiliquid species, Espeletia argentea Humb. & Bonpl. food masses were dark brown and occu- (Asteraceae). The stems containing nests = pied about one third of the cell length; had diameters of 5.5 to 7.0 mm (X 6.4 of masses X = 2.2 mm ± ± n = in lengths pollen 0.6, 8) and were varying posi- = 0.3, n 4. Five of the collected nests con- tions from almost horizontal, lying on the tained 1, 3, 3, 5 and 6 cells each. The short- ground, to erect and with the nest en- est nest tunnel (8 mm) led to one cell. Two trance 70 cm above the ground level. All old nests were recognized by fragments of occupied stems had broken ends where cell membrane; each contained an adult the bees had entered the pith. Nests con- male C. paramo. Active nests (with cells) sisted of unbranched tunnels through the each contained also a adult female. axes of the stems. Tunnel diameters single = All the adults had unworn wings. ranged from 2.5 to 3.5 mm (X 2.8 ± 0.29, = One female of the subgenus Oroediscelis n 7); the variation in diameter suggests was found in the same area but in a dead that at least some nests were probably in stem of Rubus Linnaeus (Rosaceae). In the tunnels made by other insects. One tunnel, absence of males, the is not iden- not included in the above statistics and species tified. not considered a nest, was 1.9 mm in di-

it contained a female ameter; only single DISCUSSION probably resting during cold weather. Lengths of four nests measured from en- In view of the numerous characters dif- trances to the upper ends of the cells, i.e., ferentiating C. (Anoediscelis) paramo from lengths of open tunnels above the cells, the species placed in the C. ashmeadi were 8, 17, 50, and 60 mm. The cell series group, we believe that it should be ex- often occupied the lower ends of the tun- cluded from that group. It does not, how- nels, but in other cases the empty burrow ever, show a close relationship to any of extended below the cells. All the cells con- the remaining (Chilean) species of the sub- tained provisions or small larvae, no large genus; for illustrations and descriptions of larvae or pupae. The cells were cylindrical these species see Toro and Moldenke and averaged 6.4 mm in length (± 1.2, n (1979) and for commentary see Michener = 10) and about 3 mm in diameter. The (2002: 10, 11). cell membrane was similar to that of many The new species is another example of other colletids, translucent and slightly a probably apomorphic taxon from the 30 Journal of Hymenoptera Research

boldt, and El Sistema de Parques Nacionales Natur- high Andes which does not have an ob- ales de Colombia for their outstanding logistical sup- vious affinity to any other known species, port and for generously providing both permission the distinctiveness of the showing again to work in the park and lodging to V.G. during this the existence Andean fauna and perhaps study. Financial support for V.G. was provided by and NSF DBI-0096905 S. Ashe of more species groups within the Andean Ideawild by grant (to J. This is contribution Nr. 3349 of the Chilicola fauna. and M. S. Engel). Division of Entomology, Natural History Museum The nesting biology reported here for C. and Biodiversity Research Center, University of Kan- does not differ from paramo significantly sas. that of other Chilicola species. According to what is known from other species, it is LITERATURE CITED that C. nests in diverse likely paramo pithy Michener, C. D. 2000. The Bees of the World. Johns sticks and branches available in the Para- Hopkins University Press; Baltimore, MD; xiv + mo and is not likely to be specialized to [l]+913 pp. Michener, C. D. 2002. The bee genus Chilicola in the Espeletia argentea inflorescences. tropical Andes, with observations on nesting bi- ACKNOWLEDGMENTS ology and a phylogenetic analysis of the subgenera (Hymenoptera: Colletidae, Xeromelissinae). We are indebted to Paula Montoya for her help in Scientific Papers, Natural History Museum, The the field, Michael Engel for the photographic illustra- University of Kansas 26: 1-47. tions, Diana C. Arias and Jose E. Castillo from the Toro H. and A. Moldenke. 1979. Revision de los Xe- Seccion de Entomologi'a of the Instituto de Investi- romelissinae Chilenos. Anales del Museo de His- gation en Recursos Biologicos Alexander von Hum- toria Natural, Valparaiso 12: 95-182. J. HYM. RES. Vol. 13(1), 2004, pp. 31-47

Torymns Dalman (Torymidae: Hymenoptera) Associated with Coniferous Cones, with Descriptions of Three New Species

E. E. Grissell, K. Kamijo, and K. R. Hobbs

(EEG) Systematic Entomology Laboratory, PSI, ARS, USDA, % National Museum of Natural History, Washington, DC 20560-0168, USA, email: [email protected]; (KK) Nishi 2 Minami 6-2-41, Bibai, Hokkaido, 072-0025 Japan, email: [email protected]; (KRH) 74 673 Zircon Circle East, Palm Desert CA 92260 USA (deceased 2 August 2003)

— Abstract. Eight species of Torymus Dalman are associated with coniferous cones, including three new species described herein: Torymus pseudotsugae Hobbs, from cones of Pseudotsuga menziesii in the United States (California, Idaho) and Canada (British Colombia); Torymus hobbsi Grissell, from cones of Picea sitchensis (Bongard) Carriere in the United States (California, Oregon, Colorado); and Torymus ezomatsuanus Kamijo, from larvae of cecidomyiid in seeds of Picea glehnii (F. Schmidt) Masters and P. jezoensis (Siebold and Zuccarini) Carriere in Japan (Hokkaido). The previously described species are: T. azureus Boheman (Holarctic), T. caudatus Boheman (Hol- T. arctic), festivus Hobbs (Nearctic), T. janetiellae Graham and Gijswijt (Palearctic), and Torymus tsugae (Yano) (Palearctic). Of the eight known species, five are reported attacking Cecidomyiidae (Diptera) in coniferous cones, but specific hosts are unknown for the other three. Torymus tsugae is recognized and redescribed for the first time since its description in 1918. Torymus caudatus is for the first reported time from Japan (Hokkaido, ex cones of Picea jezoensis, new host plant) and from the New World (New York, USA, ex cones of Picea abies (L.) Karsten); T. azureus is reported for the first time from Japan (Hokkaido, ex seeds of Picea glehnii). An illustrated key to all species is given.

The hymenopterous family Torymidae control of cone-infesting cecidomyiids, contains a number of species associated which in some cases account for loses of with coniferous cones. Most of these be- up to 80% in seed production (Masters long to the genus Megastigmus Dalman 2003). and are phytophagous within seeds In this paper we review the world's (Grissell 1999). Less well known are spe- known cone-inhabiting Torymus, which cies of Torymus Dalman that inhabit total eight species, including three new cones. Although little specific host data ones described herein. What specific host are available for these species, it is prob- data are available are discussed under able that all are parasitoids of cone-in- each species and a summary host/parasit- habiting gall flies of the family Cecido- oid list is given at the end of this paper. myiidae (Fig. 31). It is possible, though This list encapsulates what is known unlikely based on known host records for about host plant, host insect, and the To- the genus, that some species may be seed- rymus associated with them. We provide feeders or might be attacking seed-feed- distribution and economic data for each ing Megastigmus. Basically little is known species as well as an illustrated key. We about these Torymus other than they oc- report the first New World occurrence of cur in coniferous cones. Though they ap- Torymus caudatus Boheman, and the first pear to be of little economic importance, occurrence of that species and T. azureus they likely play some role in the natural Boheman in Japan. of Hymenoptera Research 32 Journal Graham METHODS termined using Grissell (1995), Ce- and Gijswijt (1998), and Noyes (2002). Our concepts of described Palearctic checked cidomyiid host names were by are based on specimens deter- species Dr. Gagne, Systematic Entomol- and who Raymond mined by Graham Gijswijt (1998) all Laboratory. The authors' names for for several ogy designated neotypes species (T. hosts are in the plant and insect given azureus, T. caudatus) and to M. J. Gijswijt Host Plant/ Torymus list at the end of this who loaned specimens for examination (T. paper. caudatus, T. Graham and Gijs- janetiellae The abbreviations are used: and of following wijt). Examples of these specimens = ocellocular distance, POL = are OOL post- type material of T. festivus Hobbs ocellar distance, OD = lateral ocellus di- housed in the National Museum of Natu- brackets are used for la- ameter; square [ ] ral History, Washington, D.C. The senior bel data that the information is author has examined specimens of all indicating not on the label and has been added from taxa, including the new species described other sources. The technical herein, and those for the redesciption of T. key provides characters with which to tsugae Yano. morphological are often diffi- Plant nomenclature and distribution identify species, but these even with series of was checked using the Germplasm cult to see, specimens in excellent condition. Females are the Resources Information Network (GRIN 2000) and The PLANTS Database (USDA, NRCS, more reliable sex upon which to base de- 2001). Torymus names and hosts were de- terminations.

KEY TO CONE-INFESTING SPECIES OF TORYMUS DALMAN

sexes: 1 Female: basal cell open behind, cubital vein without setae (Fig. 28, arrow). Both metacoxa without setae along outer dorsal margin (as in Figs. 10, 11) and lateral ocellus Yano subequal in length to ocellocular length (Fig. 5); (ex Tsuga; Palearctic) tsugae - Female: basal cell partially (Fig. 27) to completely (Fig. 26, arrow) closed behind, cubital vein with 2 or more setae. Both sexes: either metacoxa with setae along outer dorsal margin (Figs. 8, 9) or metacoxa without setae (Figs. 10, 11) and lateral ocellus 0.7X or less length of ocellocular distance (Figs. 1, 23) 2

2 Metacoxa without setae along outer dorsal margin (Figs. 10, 11); frenal area of scutellum with (Fig. 3) or without setae 3 - Metacoxa with short setae along outer dorsal margin (Figs. 8, 9) [note that these are different from longer setae arising on posterior side of coxa in ventral half]; frenal area of scutellum with setae (as in Fig. 3) 5

3 Frenal area of scutellum without setae, glabrous and shiny; area polished compared to anterior of interocular distance 1 X scutellum; about eye height in facial view (as in Fig. 6); postgenal area in side view narrow, less than 0.3 X eye width. Female: metacoxa over narrowly elongate, nearly parallel-sided, 3X as long as wide (Fig. 10); club without area of ventral micropilosity on any segment; (ex Picca; Holarctic) azureus Boheman - Frenal area of scutellum with at least a sparsely setose, few setae (Fig. 3) or if setae not then entire scutellum apparent heavily reticulate; interocular distance about 1.5X eye in facial view height (Fig. 17); postgenal area in side view wide, 0.5-0.7X eye width Female: metacoxa not about 2.5-3X as (Fig. 21). elongate, long as wide (as in Figs. 8, 9, at least one club with ventral area of 11); segment micropilosity (as in Fig. 7) 4 4 Toruli 1.5-2X own diameter above ventral eye margin (as in Figs. 6, 16); venter of forewing basal cell essentially bare, with few or no setae. Female: club segment 3 with ventral area of (as in 7); (ex micropilosity Fig. Picea; Palearctic) ezomatsuanus Kamijo, n. sp. Volume 13, Number 1, 2004 33

T. dorsal view. Figs. 1-7. Torymus spp. 1-4, T. ezomatsuanus. 5-7, tsugae. 1, 5, Female, head, 2, Female, antenna. 3, Female, scutellum. 4, Male, antenna. 6, Female, head, front view. 7, Female, clava, ventral view.

- basal Toruli less than own diameter above ventral eye margin (Fig. 17); venter of forewing 2 and 3 with ventral cell evenly covered with short setae (Fig. 27). Female: club segments n. area of micropilosity; (ex Abies; Nearctic) hobbsi Grissell, sp.

5 Propodeal spiracle about 3 to 4X its own length from posterior margin of propodeum, ca. of metatibia 0.3X median length of propodeum (Fig. 13). Female: ovipositor 7x length (sometimes curling and distorting when dry so that it cannot be measured); (ex Picea; Holarctic) caudatus Boheman - of ca. Propodeal spiracle less than 3x its own length from posterior margin propodeum, not than 4x 0.5 to 0.7x median length of propodeum (Fig. 12). Female: ovipositor more length of metatibia 6 of 6 Funicle segments (Fig. 14) at apex without transverse row outstanding perpendicular as as setae (cf. Figs. 14, 15); female pro- and metafemora slender, over 4X long wide; profemur of male about 3.5 x as long as wide. Female: ovipositor less than 3x as long as metatibia. Male: setae on funicle segments recurved above surface, dense (ca. 3-4 7 rows), obscuring multiporous plate sensilla (Fig. 14) - at with transverse Funicle segments (Fig. 15; best seen in profile with backlighting) apex row of bristles clearly projecting at nearly right angles; female pro- and metafemora less than slightly enlarged, less than 3.5 X as long as wide; profemur of male enlarged, 3x as long as wide. Female: ovipositor at least 4x as long as metatibia (but usually distorted and difficult to measure). Male: setae on funicle segments appressed to surface, sparse (ca. 2 rows), multiporous plate sensilla apparent as white, flattened bands (Fig. n. 15); (ex Pseudotsnga; Nearctic) pseudotsugae Hobbs, sp. 34 Journal of Hymenoptera Research

5 or outer dorsal 7 Female: metacoxa (Fig. 9) with single row of setae (about fewer) along a between them and margin, reaching 0.2-0.3 x length of coxa and creating gap longer burnished similar to setae at apical 0.3 of coxa. Male: venter of scape barely sculptured, face, which is bright metallic green (ex Chamaecyparis; Palearctic) janetiellae Graham and Gijswijt - 10 or more setae outer Female: metacoxa (Fig. 8) with several setal rows (about total) along dorsal margin, reaching at least 0.5 x length of coxa and meeting longer setae at apical 0.5 of coxa. Male: venter of scape with granulose, coarse sculpture similar to that on face, which is matt blackish green; (ex Chamaecyparis, Thuja; Nearctic) .... festivus Hobbs

Torymus azureus Boheman tricts. Bakke (1955) summarized published Figs. 10, 26 records of rearings made from Picea cones containing Laspeyresia strobilella L. (Lepi- Torymus azureus Boheman 1834: 369-370. doptera: Olethreutidae) and proved con- that were the ac- Distribution and hosts. —Torymus azureus clusively cecidomyiids tual host. is known from western Russia (Ni- This was first in the kol'skaya 1952) and eastern Europe (Gra- species reported New World Grissell (1976) from cones ham and Gijswijt 1998). The species was by reared from Kaltenbachiola strobi and Ple- of endemic Picea engelmannii in Montana and New Mexico. The of T. azureus, meliella abietina (Diptera: Cecidomyiidae), origin whether Holarctic or introduced from one both from Picea abies (Norway spruce) to the other, is not known. It is cones (Bakke 1955, 1963; Graham and Gijs- region possible that T. azureus shifted from intro- wijt 1998). [Bakke (1955) did not specify duced to endemic the Picea species, but based on his sum- Norway spruce Engle- mann's but so far there are no re- mary work published in 1963 it may be spruce, cords of the from assumed that he worked only with Picea species Norway spruce in the abies in Norway.] Norway spruce is en- New World. The Palearctic cecido- demic to the Palearctic Region (USDA, myiid hosts are not known from the New NRCS We have seen World 2001). 16 from Swe- (Gagne,—pers. comm.). den reared from cones of Picea excelsa (= Discussion. This species would be P. abies). We report T. azureus here for the placed in the varians- group by Grissell first time from Japan (19, Ashoro, Hok- (1976) and in its own species-group by kaido, 1956, K. Kamijo; 1 9 Hokkaido (ex- Graham and Gijswijt (1998). Other species act locality unknown), 1956, K. Kamijo; known from coniferous cones are mem- 19, Tomakomai, Hokkaido, 1956, K. Kam- bers of the bedeguaris- or cingulatus-groups These of ijo). specimens were reared from Pi- Torymus. Torymus azureus is distin- cea glehnii (new host record), which is en- guished from species in these groups demic to the Palearctic (Japan and Russia; based on the following species-group GRIN 2000). characters: the frenal area is indicated by Bakke (1955) provided a detailed histor- an absence of setae (remainder of scutel- ical overview of Torymus azureus, includ- lum with sparse, elongate setae) and being both its ing taxonomic confusion with T. almost smooth compared to anterior of caudatus (see next species discussion), and scutellum and dorsum of mesosoma, its habits. biological According to Bakke which is feebly aciculate; there is no frenal T. azureus (1963) was common in the low- line. Additional characters that aid in lands and southern areas of identification Norway, are: the metacoxa distally whereas T. caudatus was more common in has long setae on the inner (flat) surface, the higher elevations and northern dis- but from an outer (i.e., normal) view these Volume 13, Number 1, 2004 35

11 v

16 17

Figs. 8-25. Ton/mus spp. 8-12, Metacoxa, outer view. 8, T. festmus. 9, T. janetiellae. 10, T. azureus. 11, T. hobbsi. 12-13, Propodeum. 12, T. pseudotsugae. 13, T. caudatus. 14-15, Funicle segment 5. 14, T. janetiellae. 15, T. pseudotsugae. 16-17, Head, frontal view. 16, T. pseudotsugae. 17, T. hobbsi. 18-21, Head, side view. 18, T. festivio. 19, T. pseudotsugae. 20, T. janetiellae. 21, T. hobbsi. 22-25, Head, dorsal view. 22, T. pseudotsugae. 23, T. hobbsi.

24, T. janetiellae. 25, T. festivus. of Hymenoptera Research 36 Journal

treated Grissell (1976) because the key are usually visible only distally (Fig. 10); western Nearctic It is distin- there are no setae along the dorsal margin; only species. from other cone-associated in females the metacoxa is elongate and guished spe- based on the set of charac- being 3 to 3.5 X as long as cies following parallel-sided but has the is about ters: a frenal area is not apparent wide (Fig. 10); and ovipositor and reticulate similar to 4.5 to 5.5 X as long as the meta tibial setae sculpture a frenal line color for both sexes is the anterior of the scutellum; length. The body with is indicated but is medially metallic blue green, sometimes pur- barely laterally obscure at some angles of view; plish reflections. except metacoxa has many short, recurved setae where the Torymus caudatus Boheman along the angled dorsal margin dorsal surface in Fig. 13 outer face meets the (as are a few setae Fig. 8) and there elongate Torymus caudatus Boheman 1834: 365-366. visible in the basal portion (in both sexes Distribution and hosts. —Torymus cauda- the metacoxa is not noticeably parallel- the is about 3 to tus is widespread in eastern Europe (Gra- sided); propodeal spiracle it been 4X its own from the ham and Gijswijt 1998) where has greatest length pos- about reared from Kaltenbachiella strobi in cones terior margin of the propodeum and of Picea abies (Bakke 1955). In Sweden it 1/4 the median propodeal length (Fig. 13); has been reared from cones of Picea excelsa and the ovipositor is about 7X as long as for the the metatibial but it is so (= P. abies). We report the species length, usually difficult to first time from Japan (29, 2d, Asahikawa, distorted that the length is Hokkaido, em. 27-VI-1990, K. Kamijo; measure. The body color is metallic green specimens in Laboratory of Systematic En- for both sexes. tomology, Hokkaido University, Sapporo, Torymus ezomatsuanus Kamijo, Hokkaido), where it was reared from new species cones of Picea jezoensis (new host record). Figs. 1-4 The species is herein reported in the New World for the first time reared from Female. —Body length 1.9-2.6 mm. Ovi- X of cones of Picea abies containing the cecido- positor sheaths about 2.4-2.5 length = myiid Dasineura [now Kaltenbachiola] metatibia. Dark blue to blue violet: pro- blackish. canadensis (Felt). The only known locality podeum often purplish. Antenna is New York (89 ,86, Syracuse, Onondage Coxae concolorous with mesosoma, re- Co., coll. 13-XII-1979, em. 12-1 to 25-111- mainder dark brown, with pro- and me- P. in metallic reflections. 1980, J. Sedwick; specimens National tafemora with Wings Museum of Natural History, Washington, almost hyaline with veins brownish yel- DC). Because Picea abies is a Palearctic tree low. Head in dorsal view (Fig. 1) about species introduced into the Nearctic 1.9X as wide as long; temples about 0.3 X (USDA, NRCS 2001), the single known as long as eye; occiptal carina not strong; population of T. caudatus from New York ocelli small, POL 1. 8-2.3 X OOL, OOL 1.3- was most introduced. 1.5X in 1.2-1.3 X as likely— OD. Head front view Discussion. Bakke (1955, 1963) re- wide as high, genae roundly converging viewed the taxonomic history and biology to mouth. Eyes separated by about 1.3 X of this species, especially with reference to their height, with inner orbits subparallel; its early and consistent confusion with T. malar space about 0.4 X eye height; in lat- azureus. This species is placed as a mem- eral view genal area about 0.5 X eye width. ber of the bedeguans-group as defined by Torulus slightly more than own diameter Grissell (1976) and Graham and Gijswijt above ventral eye margin. Clypeus with It cannot in (1998). be placed the key of lower margin almost truncate. Head irreg- Volume 13, Number 1, 2004 37 — ularly, finely reticulate, face with sparse, Male. Differs from female as follows: indistinct piliferous punctures. Antenna Body length 1.7-2.3 mm. Frons and face, (Fig. 2) weakly clavate; scape not reaching and sometimes mesosoma with coppery median ocellus, about 2.9 X as long as reflections. Legs more extensively darker, wide; combined length of pedicel and fla- Head in dorsal view fully twice as wide gellum 1. 2-1.3 X width of head; pedicel as long. Antenna (Fig. 4): scape 2.3X as nearly 1.5X as long as wide; anellus dis- long as wide; combined length of pedicel .4 X tinctly transverse; Fl much shorter than and flagellum 1.3-1 width of head; pedicel, slightly longer than wide to quad- pedicel slightly longer than wide; Fl a lit- rate; F2 as long as pedicel, a little longer tie longer than pedicel, slightly longer than wide; F4 subquadrate; F7 1.3-1.4X as than wide or quadrate; F2-F5 equal in wide as long; clava nearly as long as F5- length, quadrate to slightly longer than F7 combined; sensilla disposed in 1 row wide; F6-F7 usually slightly transverse; on each segment; C3 with a small tuft of clava shorter than F5-F7, C3 without ven- micropilosity beneath (as in Fig. 7). Me- tral micropilosity; flagellum with rather sosoma 1.6-1.8X as long as wide, in pro- short decumbent setae. Mesosoma more file propodeum sloping at about 60 de- slender, about 1.8X as long as wide. Scu- grees. Midlobe of mesoscutum densely re- tellum fully 1.3X as long as wide, with ticulate, transversely so anteriorly, with scattered setae all over scutellum except piliferous punctures minute. Scutellum on frenal area. Forewing less than 2.3 X as nearly 1.3X as long as wide, moderately long as wide. Metasoma shorter than me- convex, with sculpture as in posterior part sosoma. of mesoscutal midlobe, becoming weaker Etymology. —From "ezomatsu", the Jap- and shiny on frenal area, often smooth at anese name of Picea jezoensis, and the suf- extreme apex but without frenal line; setae fix -anus.

— : on scutellum (Fig. 3) sparse, usually ab- Type material. Holotype 9 Chitose sent medially, piliferous punctures a little (near Shikotsu-ko), Hokkaido, Japan, em. larger than on mesoscutum; flange very III-1992, ex cecidomyiid larva in seed of narrow, not trabeculate. Propodeum pol- Picea jezoensis, K. Kamijo (deposited in ished, with superficial alutaceous sculp- Laboratory of Systematic Entomology, ture, smoother medially and with a row Hokkaido University). Paratypes: 159, of very small fovea along base. Lower 10c? with same data as holotype; 2 9, Asa- mesepimeron small, 1.5X as high as wide, hikawa, Hokkaido, em. V-1983, ex ceci- Metacoxa twice as long as wide, dorsally domyiid larvae in seeds of P. jezoensis, F. without hairs in basal half, outer side Komai; 49, IS, Akan, Hokkaido, em. V- weakly reticulate; metafemur 4.9 X as long 1983, ex cecidomyiid larvae in seeds of P. as wide; metatibia with longer spur about glehnii, F. Komai (deposited in Laboratory as long as width of tibia, shorter spur of Systematic Entomology, Hokkaido Uni- about half of longer one. Forewing 2.4X versify, Biabi, Hokkaido, and National as long as wide; costal cell on upper sur- Museum of Natural History, Washington, face with a row of setae in apical half; bas- DC). al vein with about 6 setae; basal cell with Distribution. —Japan (Hokkaido). The 1-9 setae, widely open below; speculum of host tree is endemic to the Palearctic and moderate size, narrowly open below; rel- is found in China, Japan, and the Russian = ative lengths of M:PM:ST 31:9.5:4; ST Federation (GRIN 2000). petiolate. Metasoma a little longer than Host.—Reared from cecidomyiid larvae 0.7 to in of Picea and P. mesosoma; hypopygium reaching seeds — glehnii jezoensis. 0.8 of gaster. Ovipositor sheaths as long as Discussion. Torymus ezomatuanus be- metasoma plus half of mesosoma. longs to the cingulatus-group of Graham 38 Journal of Hymenoptera Research

2.7 to as and Gijswijt (1998) or to the bedeguaris- have the ovipositor about 3x group of Grissell (1976), whose definition long as the metatibia. In both sexes the includes the cingulatus-group. This species body color is metallic blue, belongs to the group of species without setae on the dorsum of the metacoxa and Torymus janetiellae Graham and is similar in size and color to T. azureus Gijswijt 9, 20, 24 from which it differs in having the frenal Figs- area setose (Fig. 3) and sculp- sparsely Torymus janetiellae Graham and Gijswijt 1998: tured about as for the remainder of the 115-116. scutellum (i.e., not nearly polished as in T. Distribution and hosts.—This was azureus); in females having the metacoxa species reared in the Netherlands from cones of less than about 2.5 X as long as wide (as lawsoniana in Fig. 11, not over 3X as in T. azureus, Chamaeajparis containing Jane- tiella Fig. 10) and antennal club segment 3 with siskiyou (Diptera: Cecidomyiidae) and an area of ventral micropilosity (as in Fig. (Graham Gijswijt 1998). Chamaecyparis lawsoniana is an endemic tree native to 7). Females have the ovipositor about 2.5 X southern and northern California the length of the metatibia. In both sexes Oregon the color is metallic blue to blue violet or (GRIN 2000) and the host cecidomyiid is native to purplish. Oregon (Gagne 1989). The fly is introduced and well established in Europe Torymus festivus Hobbs (Gagne 1989), as is the tree. It is likely that Figs. 8, 14, 18, 25 T. janetiellae is also introduced (Graham and t c i- it uu nncn ,-,, Anc Gijswijt} } 1998).' TorymusJ festivus was Torymus festivus Hobbs 1950: 173-175. . f ~ . / ^ ... reared from C. lawsoniana in California — and is similar to T. Distribution and hosts. This species was extremely janetiellae, an additional of lat- reared from Dasineura sp. (Diptera: Ceci- suggesting affinity the domyiidae) in seeds of nearctic endemic ter species to the Nearctic fauna. Graham and western red cedar {Thuja plicata) and Port Gijswijt (1998) also suggested that T. Orford cedar {Chamaecyparis lawsoniana) janetiellae could have been introduced from (Hobbs 1950). It is known from Oregon, Japan but— gave no reasons for this. Alaska, and the Northwest Territories Discussion. This species is listed as a (Grissell 1976). As these trees are native to member of the bedeguaris-group in Gra- the northwestern Nearctic (USDA, NRCS ham and Gijswijt (1998) and would key to that 2001), Torymus festivus should be consid- group as recognised by Grissell ered an It endemic Nearctic species. Cur- (1976). is distinguished from its most this rently species and T. pseudotsugae are phenotypic congenitor, T. festivus, by char- the only two species associated with co- acters given in the key. In females the ovi- niferous cones is that are thought to be en- positor less than 2.5 X the length of the demic to the New World. metatibial length. In both sexes the body Discussion. — is Torymus festivus is placed metallic green, in the bedeguaris-group based upon Gris- sell It is hobbsi new (1976). phenotypically similar to Torymus Grissell, species

Fi s - T. 11 ' 17 ' 21 ' 23 25 27 29 janetiellae in characters associated with § ' ' > > 30 the antenna, metafemur, and propodeum, Female.—Body length 2.0-2.5 mm; ovi- metacoxa, but be from may distinguished positor sheaths about 4X length of meta- it the by few characters given in the tibia. key. Metallic blue black except following Additionally, T. festivus is known only weakly brownish yellow: antenna, legs, from the Nearctic whereas T is janetiellae wing veins, ovipositor sheaths; wings hy- known from only the Palearctic. Females aline. Head in dorsal view (Fig. 23) about Volume 13, Number 1, 2004 39

1.8X as wide as long; temple about 0.5 X tae in basal half (Fig. 11), outer face retic- as long as eye; occipital carina weak; POL ulate; metafemur about 4X as long as 2.0 X OOL, OOL 1.5 X OD, lateral ocellus wide; metatibia with longer spur about as small, about 2.0 X own longest diameter long as width of tibia, shorter spur about from occipital carina (Fig. 23). Head in 0.6 X as long as longer one. Forewing front view (Fig. 17) about 1.1 X as wide as about 2.5 X as long as wide; upper and high, with genae roundly converging to lower anterior margin of costal cell (Fig. mouth; intermalar distance about 2.5 X 27) with setal row, apical half of costal cell malar distance. Eyes separated by about with 3 or 4 rows setal rows below; basal 1.4X own height, with inner orbits ven- vein with 1 to 2 rows of setae; basal cell trally diverging; malar space about 0.4 X below almost uniformly covered with wide- eye height; in lateral view eye appearing spaced setae, above with only a few setae, reduced (Fig. 21) with genal area at widest cubital vein beneath basal cell with few se- X point about 0.6 eye width. Torulus less tae apically (i.e., mostly open below); spec- than own diameter above ventral eye mar- ulum apparent on upper surface, but with gin (Fig. 17). Clypeus with lower margin setae on lower; approximate relative lengths essentially truncate. Head irregularly, of marginal : postmarginal : stigmal veins 30: finely reticulate. Antenna weakly clavate 9:4; stigmal vein petiolate. Metasoma slight- (Fig. 29); scape not reaching median ocel- ly longer than mesosoma; hypopygium lus, about 3x as long as wide; combined about 0.7x length of gaster. Ovipositor length of pedicel and flagellum about sheaths as long as body. equal to width of head; pedicel in lateral Male.—Body length 1.9-2.9 mm. Differ- view about 1.2X as long as wide; anellus ing from female as follows: Antenna (Fig. distinctly transverse; Fl slightly shorter 30): scape about 6x as long as wide; ped- than pedicel, quadrate; F2 slightly longer icel nearly 2x as long as wide; Fl shorter than pedicel, a little longer than wide; F4 than pedicel, about as long as width of slightly wider than long; F7 about 1.5X as pedicel; F2-F7 wider than long; clava ven- wide as long; clava equal to or longer than trally flattened, longer than F5-F7, C3 F5-F7 combined; sensilla arranged in 1 without ventral micropilosity; flagellum row on each segment; C3 without micro- with short decumbent setae. pilosity beneath. Mesosoma ca 1.8X as Variation. —This species appears to be long as wide, in profile propodeum slop- highly variable in sculpture. In one rearing at about 60 degrees. Entire dorsal sur- ing (Newport) 2 female and 2 males are face densely reticulate (transversely so just lightly sculptured and 2 males are heavily posterior to pronotum); lateral surfaces in sculptured. In an August rearing at Cres- most specimens nearly similarly sculp- cent City, 1 female and 1 male are lightly tured (including acropleuron, upper and sculptured and 2 females are heavily lower epimeron, and metapleuron), but sculptured, but an October rearing from some nearly smooth on lateral surfaces the same locality has 1 female and 4 males (see "Variation", below). Scutellum about all heavily sculptured. All other characters as long as wide, moderately convex, with- for these specimens appear to be consis- out frenal line; setae sparse (about as in tent. In all specimens the torulus varies Fig. 3) to absent; scutellar flange narrow from slightly to much less than its own (barely perceptable), with no trace of pits. diameter above the ventral eye margins Propodeum nearly as reticulate as scutel- (Fig. 17), but this variation appears to be lum, with row of small fovea along ante- due to slight distortions of the head. (See rior margin. Lower mesepimeron about also discussion section, below.) 1.8X as high as wide. Metacoxa about 3- Etymology.—This species is named in 3.5 X as long as wide, dorsally without se- honor of Kenneth R. Hobbs who, long Journal of Hymenoptera Research 40

../ill./ . . Volume 13, Number 1, 2004 41

about 0.6-0.7X the eye width in lateral view (Fig. 21); the interocular distance is about 1.5 X the eye height [also ezomatsuanus]; female and male T. hobbsi have the antennal club equal to or greater in length than the preceeding 3 segments (Figs. 29, 30); male T. hobbsi have the club somewhat flattened ventrally and distinctly acuminate at the apex (Fig. 30). Some specimens of T. hobbsi differ based upon the heavily sculptured acropleuron, upper and lower epimeron, metapleuron, and propodeum (all other species are essentially polished in these areas). Unfortu- nately the expression of sculpture in these areas is not consistent on all specimens, even in a single rearing, so that the above characters must be used in combination. In females, the ovipositor is about 4X as long as the metatibia. In both sexes the color is bluish black. Four specimens reared from the series collected in Crescent City appear to represent either extreme variations of Tory- mus hobbsi, or a different species entirely. These have setae on the dorsum of the metacoxa and on the frenal area, both characters that indicate assignment to a different species group. It will require collection and analysis of new material in excellent condition to determine if more than one Fig. 31. Pseudotsuga menziesii cone. Cross section in of Picea sitch- showing scale galls (white circles) of Contarinia ore- species is living the cones gonensis (Cecidomyiidae) (modified from Gagne 1989, ensis. Fig. 356, reprinted by permission). Torymus psendotsugae Hobbs, new species 12, 15, 16, 19, 22 1998) T. hobbsi runs to couplet 4 where it Figs. — fits neither of the two treated species. Holotype female. Body length 2.0 mm; Torymus hobbsi differs from other spe- ovipositor sheaths abut 4X length of me- cies of Torymus reared from cones based tatibia. Metallic blue with purple reflec- upon the basal cell and parastigmal areas, tions under indirect light, including pedi- which are nearly uniformly setose on the cel, anellus, head, thorax and abdomen; under side (Fig. 27). It also differs in part following light amber: scape, except apical in the following respects [exceptions are 2/3 of dorsal surface dark blue, trochan- noted in brackets]: the torulus is less than ters, femora at bases and apices, dorso-lat- its own diameter above the ventral eye eral surfaces of tibiae metallic blue fading margin (Fig. 17) [also ezomatsuanus, cau- to brown on ventral and mesolateral sur- datus, and azureus, especially when the faces; protarsal segments nearly uniform head is collapsed]; the postgenal area is brown, mesotarsal segments similar with Research 42 Journal of Hymenoptera

as at about 60 basitarsus lighter brown, metabasitarsus wide, propodeum sloping this area off white with each segment increasingly degree angle, smooth medially, anterior row of minute brown distally; wing veins light brown interrupting pits with ori- basally becoming darker distally, stigmal (Fig. 12); laterally longitudinally ented narrowed vein prominently petiolate; areas between reticulation; propodeum distance between inner postmarginal vein and stigmas vein light medially, margin brown. Face and dorsal aspects of thorax of spiracles 3X median propodeal length Mesosoma reticulate over bearing closely appressed white setae (Fig. 12). evenly about as long as distance between them; dorsal surface. Metasoma finely, distinctly twice reticulate and clypeal and apical scutellar hairs laterally dorsally except Mt2-Mt5 this length; metacoxa with dorsal setae. Mt2 smooth, medially deeply Head in dorsal view 1.7X as wide as long; emarginate [best seen on paratype speci- than temple 0.3 X as long as eye; occipital cari- mens], ovipositor longer body, na developed; POL 2.5 X OOL, OOL 1.6X Male.—Length 1.5-2.5 mm. Differing OD, lateral ocellus 1.7X own longest di- from female only in having more metallic ameter from occipital carina (Fig. 22). green color; ocellocular and ocelloccipital to lateral Head in front view (Fig. 16) 1.1 X as wide distances equal ocellus diameter; as high, with gena roundly converging to and epimeron about 0.7X as wide as high, mouth; intermalar distance 2.7 X malar Variation. —Females vary in length from distance. Eyes separated by about own 1.8 to 2.5 mm. with the ovipositor from 2.1 height, with inner margins ventrally to 2.3 mm. Females are uniform blue with slightly diverging; malar space 0.3 X eye some purple reflections. The males tend to height; in lateral view eye not appearing be more green, however, some are nearly reduced (Fig. 19; cf Fig. 21, hobbsi) with as blue as females. genal area at widest point 0.5 X eye width. Etymology. —This species is named for Torulus about 1.5-2X own diameter the association with its host plant, Pseu- above ventral eye margin (Fig. 16). Clyp- dotsuga menziesii. eus with lower margin truncate, slightly Type material. —Holotype 9: Oregon, recessed relative to ventral margin of head [Coos Co.], Two Mile Road west of High- (Fig. 16). Antenna cylindrical (as in Fig. 4), way 101 south of Bandon, 23-IX-1998, C. not scape reaching median ocellus, about Hobbs, B. Baugh and S. Brown coll., 4X as long as wide; combined length of reared from cones of Pseudotsuga menziesii, and pedicel flagellum about much greater emerged in lab 2-II-1999 (deposited in Na- than width of head; pedicel in lateral view tional Museum of Natural History, Wash- 2X as as anellus long broad, quadrate ta- ington DC). Paratypes: 739,596 (in USNM pering basally, ratio pedicel : anellus : Fl unless otherwise noted): 29, 86, same F7:club as through 7:3:4:5:5:5:5:6:3 (Fig. data as holotype. USA: California: 69, 29); anterior 3/4 of 1 pronotum transversely 6 , Santa Cruz Co., Bonnie Doon, 1 /2 mi. aciculate, posterior 1/4 finely reticulate as ne 4600 Smith Grade Road, 23-IX-1998, B. in remainder of dorsal surface of scutum Hobbs, C. Hobbs, K. Hobbs, reared from and scutellum; forewing (Fig. 27) relative- cones of Pseudotsuga menziesii, emerged 8- bare costal cell in basal 1 and I to ly basally, /4 2-II-1999; 39,16% [Siskiyou Co.], Hap- 1/3 with and apical upper lower setal py Camp, Klamath National Forest, 27- lower surface with rows, additional scat- VII-[19]54, Hopkins No. 34018d, Pseudo- tered setae behind setal basal cell = row, tsuga taxifolia (now menziesii); 19, Sis- with several setae paralleling submarginal kiyou Co., Etna, XI-1957, T. W. Koerber, vein, basal vein with 1 or 2 cubital setae, Hopkins No. 34097a, Pseudotsuga menziesii; vein setose; frenum and entirely frenal 4 9, 16, [Humboldt Co.], Orleans, 12-26- ne absent. Mesosoma about 1.6X as long VIII-[19]54, Hopkins No. 34091d, Pseudo- Volume 13, Number 1, 2004 43

= tsuga taxifolia (now menziesii); 139, 66, characters: the scutellum has no frenal Humboldt Co., Orleans, T. W. Koerber, area, is evenly covered with reticulate Hopkins No. 37500a, Pseudotsuga menziesii; sculpture and sparse setae, and has no fre- 49, Trinity Co., Salyer, IX-1957, T. W. nal line; the metacoxa is not elongate but Koerber, Hopkins No. 37504a, Pseudotsuga is parallel-sided, with a few long setae dis- menziesii. Idaho: Shoshone 9 and recurved setae 5$, 66 , Co., tally many short, along mi. nw Kellog, ex cones P. menziesii; 189, the angled dorsal margin where the outer 14c?, Idaho Co., ca. 1.8 mi sse Graves face meets the dorsal surface; the propo- Butte, US Hwy. 12, mi. marker 139. ex. deal spiracle is less than 3 X its own great- cones P. menziesii (specimens deposited in est length from the posterior margin of the Natural History Museum, London; Cana- propodeum and about 1/2 to 1/3 the median National Collection, Ottawa); 79, 5 6, dian propodeal length (Fig. 12). In males Boise Co., 1 mi. e Lowman, T9N, R&E, the funicular segements have 2 rows of Sec. 35, 19-VIII-1972, R. W. Clausen, ex appressed setae that do not obscure the cones P. sensilla in menziesii; 19, \6 , McCall, [Valley multiporous plate (Fig. 15), and Co.], 12-VIII-1971, J. Dale, Hopkins No. both sexes there is a single row of bristles 14280f, ex Douglas-fir cones. Oregon: 1 9, that project at nearly right angles (Fig. 15). 25-VIII- In is 76 , [Jackson Co.], Mistletoe, females the ovipositor about 4X as E. [19]16, J. Patterson, Pseudotsuga taxifolia long as the metatibial length. In both sexes = (now menziesii). CANADA: British Co- the body varies from metallic green, blue, lombia: 89, 96, southern Vancouver Is- or purple, with combinations of blue- land, VIII-1994, R. Bennetem. Spring 1995, green to bluish purple, = ex cones P. taxifolia (now menziesii). Interestingly Torymus pseudotsugae has Distribution. —This species is widely dis- not, until now, been recognized even tributed from northern California to though Douglas-fir seeds are a valuable southern—British Colombia. crop (Masters 2003) and have been the ob- Hosts. This species has been reared ject of much study (Johnson and Hedlin from seeds and cones of Pseudotsuga men- 1967, Hedlin et al. 1980, Hermann and ziesii. Douglas-fir is endemic to the west- Lavender 1990, Ministry of Forestry 2003). ern United States and Canada (GRIN According to Hermann and Lavender 2000) and so far has only a single known (1990) the most destructive insects in- cecidomyiid species (Contarinia oregonensis elude: the Douglas-fir seed chalcid (Me- Foote) that feeds in the cone scales (Fig. gastigmus spermotrophus Wachtl) (Hyme- 31). It is possible that a seed-feeding ceci- noptera), the Douglas-fir cone moth (Bar- domyiid could also be present, but none bara colfaxiana Kearfott) and the fir cone has been reported (Gagne 1989 and per- worm (Dioryctria abietivorella Grote) (Lep- sonal and the cone communication).— idoptera), Douglas-fir gall Discussion. Torymus pseudotsugae is a midge (Contarinia oregonensis Foote) and member of the bedeguaris species group as cone scale midge (C. washingtonensis John- defined by Grissell (1976) and Graham son) (Diptera). According to Hedlin et al. and Gijswijt (1998). Among western Ne- (1980) any of these insects may effectively arctic Torymus, the species would run to destroy a cone crop in a given location; T. coloradensis (Huber) in the key by Gris- Masters (2003) reported that cecidomyiid sell (1976); in the Palearctic Region it cone gall midges can cause a reduction of would run to T. hylesini Graham in Gra- up to 80% in seed production. There are ham and Gijswijt (1998) (neither of these at least 5 described species of Cecidomyi- species is associated with coniferous idae (Diptera) (Gagne 1989) on Pseudotsu- cones). It is distinguished from cone-as- ga, and it is one or more of these species sociated species by the following set of that is the likely host of Torymus pseudo- of Hymenoptera Research 44 Journal

is OOL 0.95-1 .2 X OD. Head in front tsugae. If Torymus pseudotsugae parasit- OOL, as view rounded, 1.16-1.18X as wide izing one or more of the cecidomyiids, (Fig. 6) cone-in- as 1.1 X their is most likely based on the other high. Eyes separated by in our with inner orbits Ma- festing Torymus examined study, height, subparallel. about one third of then its value as a biological control agent lar space height eye; towards has been overlooked completely. genae roundly converging mouth; in lateral view genal area about Torymus tsugae (Yano) 0.3 X eye width. Torulus about 1.5 X own Figs. 5-7, 28 diameter above ventral eye margin. Clyp- eus with lower margin truncate or very Callimome tsugae Yano [in Yano and Koyama] shallowly emarginate. Both mandibles tri- 1918a: 44-45; 1918b: 373. dentate. Vertex and frons irregularly, somewhat This species was originally described densely reticulate, granulate, from specimens reared from seeds of Jap- with sparse, rather distinct piliferous anese hemlock, Tsuga sieboldii. Yano did punctures. Face finely reticulate; setae on with indis- not give any other data for the types. face short, piliferous punctures not Nothing has been published on T. tsugae tinct. Antenna (Fig. 6): scape reaching X as since its description, and we have been median ocellus, 3.4 as long wide; ped- unable to locate specimens of this species icel plus flagellum about 1.3X width of for comparison. Information about types head; pedicel 1.7-1.9X as long as wide, a was requested from Dr. Akihiko Shino- little longer than Fl; flagellum distinctly hara, National Science Museum, Tokyo, clavate; anellus slightly transverse to al- and Dr. Tikahiko Naito, Kobe University, most as long as wide; Fl quadrate to Kobe; both replied that Yano type material slightly longer than wide; F2 and F3 equal is not housed at these institutions. Kamijo in length, a little longer than wide; F6 and (1962), writing of the type material of Me- F7 transverse; clava as long as F5-F7 com- gastigmus cryptomeriae Yano, M. inamurae bined, 2.0-2.25 X as long as wide: C3 with Yano, and M. thuyopsis Yano, stated that it a small tuft of micropilosity beneath (Fig. had been "destroyed by fire." Because To- 7); sensilla disposed in 1 row on each fu- rymus tsugae was described in the same nicle segment. Mesosoma nearly 1.7X as and paper (Yano Koyama 1918a) as these long as wide; in profile, propodeum slop- Megastigmus, it is likely that the types of ing at about 60 degrees. Mid lobe of me- this species were destroyed in the same soscutum transversely reticulate, more or fire. The following redescription is based less imbricate in anterior half; setae mod- on reared specimens from Japanese hem- erately dense, with piliferous punctures lock, the original host plant, which is na- shallow and indistinct. Scutellum 1.2X as tive to and Korea Japan (GRIN 2000). long as wide, flat in longitudinal axis, re- Female.— 1.8-2.3 Body length mm; ovi- ticulate, very densely so anteriorly; fre- sheaths about 3.9 X positor as long as me- num indicated by very weak, almost tatibia. Dark blue with a violet or a green- smooth sculpture but frenal line absent; ish in dark- tinge places. Scape testaceous, setae on scutellum about as in Fig. 3, with er and apically; pedicel flagellum blackish. piliferous punctures distinct. Propodeum testaceous: coxae and metafemur Legs longitudinally, weakly sculptured, with a dark blue; and mesofemora pro- and me- row of very small fovea along base. Lower tatibia medially darkened, usually with mesepimeron small, 1.5 X as high as wide. metallic reflections. with Wings hyaline, Metacoxa about 2.5 X as long as wide, veins testaceous. Head in dorsal yellowish very weakly sculptured, dorsally with a riew about 1.9X as (Fig. 5) wide as long; carina and without setae in basal half; me- carina not POL 2.2-2.27X cipital strong. tafemur 4.4-4.7X as long as wide, with Volume 13, Number 1, 2004 45 sparse, elongate piliferous punctures; me- 2.5 X as long as wide and not parallel-sid- tatibia with longer spur as long as width ed (about 3.5 X as long and parallel-sided of tibia, shorter spur longer than half of in azureus), and the absence of setae on the longer one. Forewing 2.5 X as long as basal vein and along the posterior margin wide; costal cell on upper surface with a of the basal cell (Fig. 28). In males, there row of setae apically; basal cell bare, open may be a few setae along the basal vein below (Fig. 28); basal vein with a few se- (as in male T. azureus), but the speculum tae; speculum moderately large, narrowly is open posteriorly (i.e., there are no setae open below; relative lengths of marginal : on the poximal part of the cubital vein, = postmarginal : stigmal veins 31:7:3.1; ST thus leaving a bare path from the specu- a little petiolate. Metasoma longer than lum to the posterior wing margin (Fig. 28). mesosoma, compressed; hypopygium Other species generally have setae in the about 0.7 of area reaching gaster. Ovipositor posteriad of the speculum (e.g., Figs. sheaths—slightly longer than body. 26, 27). Both sexes of T. tsugae are metallic Male. Differs from female as follows. blue to violet with some green tinges. Body length 1.6-2.1 mm. Antenna: scape about 2.4 X as long as wide; pedicel plus HOST PLANT/ TORYMUS LIST about 1.4X width of head; flagellum ped- (All species reared from coniferous icel 1.3X as as wide, a little shorter long cones. Insect hosts, when known, are Ce- than Fl; anellus distinctly transverse; Fl- cidomyiidae (Diptera) in cones or seeds.) F2 subquadrate; distal funicle segments weakly transverse; clava without visible Chamaecyparis lawsoniana (A. Murray) Par- tuft of micropilosity. Mesosoma slender, latore (Port Orford cedar) 1.8-1.9X X as long as wide. Scutellum 1.3 Dasineura sp.: Torymus festivus Hobbs as long as wide or more, with scattered Janetiella siskiyou Felt: Torymus janetiellae setae all over scutellum except on frenal Graham and Gijswijt area. Metasoma usually shorter than me- Picea abies (L.) Karsten (Norway spruce) sosoma. Kaltenbachiola canadensis (Felt): Torymus examined. — caudatus Specimens 69, 66 , Shiga-Ko- Boheman gen, Nagano Pref., Honshu, Japan, em. Kaltenbachiola strobi (Winnertz): Torymus 10.iv.-4.v.l957, ex seeds of Tsuga sieboldii, azureus Boheman, Torymus caudatus K. 5 in of Kamijo (59, 6 , Laboratory Sys- Boheman tematic Entomology, Hokkaido Universi- Plemeliella abietina Seitner: Torymus azur- ty; 19, 1<5, National Museum of Natural eus Boheman History, Washington,— DC). Picea engelmannii Parry ex Engelmann (En- Distribution. Japan (Honshu). The host gelmann spruce) tree is endemic to the Palearctic and is unknown host in cones: Torymus azureus known only from Japan and South Korea Boheman (GRIN —2000). Picea glehnii (F. Schmidt) Masters (Sakha- Host. Reared from seeds of Tsuga sie- lin spruce) boldii. cecidomyiid in seed: Torymus azureus Discussion. —This species would be Boheman, Torymus ezomatsuanus placed in the bedeguaris-group by Grissell Kamijo (1976) and Graham and Gijswijt (1998). Picea jezoensis (Siebold and Zuccarini) Car- Torymus tsugae is recognized in females by riere (ezo spruce) having a few setae on the frenal area (as cecidomyiid in seed: Torymus ezomatsu- in Fig. 3), the ovipositor sheaths less than anus Kamijo about 4X as long as hindtibia (4.5 to 5.5X Picea sitchensis (Bongard) Carriere (Sitka as long in azureus), the metacoxa about spruce) Journal of Hymenoptera Research 46

Network, National Plant Germplasm System, unknown host in cones: Torymus hobbsi ARS, USDA: http://www.ars-grin.gov/npgs/ Grissell tax/index. html. Franco western Nearctic Pseudotsuga menziesii (Mirbel) Grissell, E. E. 1976. A revision of Dalman To- (Douglas-fir) species of Torymus (Hymenoptera: Publications in unknown host in seeds and cones: To- rymidae). University of California + Hobbs Entomology 79: 1-120, 6 plates. rymus pseudotsugae Chal- Grissell, E. E. 1995. Toryminae (Hymenoptera: Donn ex D. Don (western red Thuja plicata cidoidea: Torymidae): a redefinition, generic cedar) classification, and annotated world catalog of Hobbs on International 2: 1- Dasineura sp.: Torymus festivus species. Memoirs Entomologxj, Tsuga sieboldii Carriere (Japanese hemlock) 470. E. E. 1999. An annotated of World unknown host in seeds: Torymus tsugae Grissell, catalog Megastigminae (Hymenoptera: Chalcidoidea: (Yano) the American Ento- Torymidae). Contributions of 1-92. mological Institute. 31 (4): ACKNOWLEDGMENTS Cibrian Hedlin, Alan F., Harry O. Yates III, David Cone and seed insects North Amer- We thank Dr. Raymond Gagne (Systematic Ento- Tovar. 1980. of information ican Environment Canada, Canadian mology Laboratory, Washington, DC) for conifers. hosts. Ottawa, ON; USDA Forest Ser- and help with the nomenclature of cecidomyiid Forestry Service, and W. for the Secretaria de We also thank M. J. Gijswijt Hogenes vice, Washington, DC; Agricultura from the Museum Am- Recursos Hidraulicos, Mexico. 122 gift of specimens Zoologisch y Chapingo, sterdam, Netherlands and Dr. Akihiko Shinohara pp. 1990. (National Science Museum, Natural History, Tokyo) Hermann, Richard K. and Denis P. Lavender. and Dr. Tikahiko Naito (Kobe University, Kobe), for Pseudotsuga menziesii (Mirb.) Franco, Douglas-Fir. of Barbara H. Honkala help in attempting to find type material Torymus In, Burns, Russell M., and Numbers was ob- 1990. Silvics North tsugae. Information for Hopkins' (Technical Coordinators). of file with the 654. tained from the Hopkins' cards on Sys- America: 1. Conifers. Agriculture Handbook National of tematic Entomology Laboratory, Museum U.S. Department of Agriculture, Forest Service, DC. For the available Natural History, Washington, reviewing Washington, DC. vol. 1, 675 pp. [Also we thank manuscript and making useful suggestions online as http://www.na.fs.fed.us/spfo/pubs/ John Noyes, the Natural History Museum, London, silvics_manual/ table_of_contents.htm] and Michael Gates, David Nickle, and David Smith, Hobbs, K. R. 1950. Notes on the classification of To- all of the Systematic Entomology Laboratory. Figure rymus with the biology and description of a new 31 is from (1989) (this paper) reprinted Gagne by per- species (Hymenoptera: Torymidae). Pan-Pacific mission of the Cornell Press. publisher, University Entomologist 26: 173-178. Johnson, N. E. and A. F. Hedlin. 1967. Douglas-fir LITERATURE CITED cone insects and their control. Canada Department of Forestry and Rural Development, Forestry Branch Bakke, A. 1955. Insects reared from spruce cones in Departmental Publication No. 1168: 1-11. northern Norway 1951. Norsk Entomologisk Kamijo, K. 1962. A revision of the species of the Me- Tidsskrift 9: 152-212. gastigminae occurring in Japan (Hymenoptera: Bakke, A. 1963. Studies on the spruce-cone insects Chalcidoidea) (Taxonomic studies on the Tory- Laspeyresia strobilella (L.) (Lepidoptera: Tortrici- midae of Japan), hisecta Matsumurana 25: 18-40. dae), Kaltenbachiola strobi (Winn.) (Diptera: Itoni- Masters, C. J. 2003. a clean of cone gall dae) and their parasites (Hymenoptera) in Nor- Making sweep duff removal shows as a non- way. Reports of the Norwegian Forest Research In- midge: promise chemical to seed orchard stitute 67 (19): 1-151. approach Douglas-fir Boheman, C. H. 1834. Skandinaviska Pteromaliner. IPM. Agrichemical and Environmental News No. 204 Kongliga Svoiska Vetenskapsakademiens Handlingar (http://aenews.wsu.edu) 54: 329-380 Ministry of Forestry, Government of British Colum-

( R. 1989. bia 2003. seed and related insects lagne, J. The plant-feeding gall midges of North (updated) Cone, America. Comstock Publishing Associates, Cor- in BC: http://www.for.gov.bc.ca/hti/IIG/ ni'll University Press, Ithaca and London. 356 pp. index.html. M. W. R. M. chalcid fauna of the Graham, de Vere and M. J. Gijswijt. 1998. Nikol'skaya, N. 1952. The Revision (if the European species of Torymus Dal- USSR. Jerusalem: S. Monson. 593 pp. [Translated man (s. lat.) (Hymenoptera: Torymidae). Zoolo- from the Russian by the Israel Program for Sci- he Verhai n 317: 1-202. entific Translation 1963.] sm Resources Information S. dial- Noyes, J. 2002. Interactive Catalogue of World Volume 13, Number 1, 2004 47

cidoidea —second (2001 edition). CD-Rom. Taxapad asitizing the seeds of coniferous trees.] Report of and The Natural History Museum, London. Forest Experiment, Forestry Bureau (Tokyo) 17: 39- JSDA, NRCS. 2001. The PLANTS Database, Version 58. [In Japanese.] 3.1. Plant National Data Center, Baton Rouge, LA Yano, S. and M. Koyama, 1918b. [On the wasps par- 70874-4490 USA: http://plants.usda.gov. asitizing the seeds of coniferous trees.] Insect *ano, S. and M. Koyama. 1918a. [On the wasps par- World (Gifu), 22: 372-376. [In Japanese.] J. HYM. RES. Vol. 13(1), 2004, pp. 48-56

Behavioral Interactions Among Females of Acamptopoeum toroi Rozen siibmetallicum (Spinola) and Nolanomelissa (Hymenoptera: Andrenidae)

Laurence Packer Jennifer C. Grixti, Amro Zayed, and

4700 Keele St., Toronto, Ontario, M3J 1P3, Canada; Department of Biology, York University, email: [email protected]

circle tube a Abstract—We present the results of experiments performed upon solitary panur- and the Nolanomelissa toroi Rozen. As gine bee Acamptopoeum submetallicum (Spinola) enigmatic females of A. submetallicum avoided one another. In expected for a solitary species, generally found bees with a re- contrast, N. toroi exhibited high levels of aggression, as generally among dissection and data on N. toroi are incom- productive division of labor. However, phenological or semisocial behaviors. Furthermore, no acts of were patible with either eusocial cooperation is The observed in the behavioral experiments suggesting that N. toroi not communal. extremely female biased sex ratio and low levels of mandibular wear among mated, reproductively active females in this species remain difficult to explain.

and bee Social behavior in bees varies greatly, ity to quickly discriminate classify without the need for de- ranging from solitary with one female per social behavior nest to eusocial with up to tens of thou- tailed and laborsome nest studies, is circle sands of individuals per nest (Michener tube experiments. This experimental 1974). A solitary bee constructs her own method, introduced by Breed et al. (1978), nest and provides stored food for her off- simulates nest tunnels using clear plastic spring, while eusocial bees (sensu Miche- tubing, where interactions among females ner 1974) have a reproductive division of can be observed. Circle tube arenas were labor, cooperative brood rearing, and originally used to study interactions more than a single generation of adults in among different castes of eusocial bees their nests at some point in the colony cy- (Breed et al. 1978; Pabalan et al. 2000) and cle. Communal behavior is a separate type to compare behaviors of species with dif- of social organization in which two or ferent social organizations (Kukuk 1992; more females share a nest without a re- Wcislo 1997). More recently it has been productive division of labor. To classify used to help predict the social organiza- the behavior of bees formally, nest exca- tion of species whose behavior is not vations at various times during the nest- known (Packer 2000; Packer et al. 2003). ing cycle accompanied by dissections of Interactions in the circle tube apparatus bees are usually needed to examine the ex- are initiated when two bees encounter one tent of their reproductive division of labor, another head-to-head within one body if any (Bell and Hawkins 1974; Brothers length of each other, referred to as a fron- and Michener 1974; Wcislo et al. 1993). tal encounter (FE). Behavioral interactions such studies are However, time consum- in circle tubes can usually be classified as and \g impossible perform in cases aggressive, cooperative, or avoidance. Ag- i (Packer et al. gressive interactions include one or both

. A have the abil- technique ay bees nudging, lunging or biting one an- 49 /olume 13, Number 1, 2004

Halictus )ther. Cooperative interactions involve dominance [e.g. ligatus (Pabalan ?ees passing one another, by rotating their et al. 2000) and Lasioglossum zephyrum ?ody in such a way that they pass venter- (Breed et al. 1978)]. of bees within :o-venter. Lastly, avoidance interactions The behavioral repertoire nvolve one or both bees turning away the circle tube apparatus has been com- rom a FE. The relative frequency of these monly reported for halictids (Breed et al. in a Smith and Weller Kukuk zategories of interactions species 1978; 1989; 1992; seems to correspond to its social organi- McConnell-Garner and Kukuk 1997; Wcis- ;ation (McConnell-Garner and Kukuk lo 1997; Pabalan et al. 2000; Packer 2000; L997; Paxton et al. 1999; Packer et al. 2003). Packer et al. 2003), as they are the most diverse bee ntraspecific interactions among solitary behaviorally family (reviewed Wcislo and Danforth :>ee species often result in avoidance in- by Packer 1997; eractions occurring at the highest fre- 1997). In comparison, the behavioral rep- Wcislo ertoire of bees within the Andren- }uency [e.g. Lasioglossum figueresi family Wcislo 1997), L. platycephalum (Rayment), idae, has been studied only once using et al. Al- L (Ctenonomia) sp. (McConnell-Garner these methods (Paxton 1999). and Kukuk 1997) and Penapis toroi Rozen though most andrenid species are solitary, Mich- Packer unpublished)]. In contrast, intra- some are communal (Paxton 1999; individuals of ener while none are known to be specific interactions among 2000), the circle a communal species result in a high fre- semisocial or eusocial. Using La- tube social interactions be- :mency of cooperative interactions [e.g. apparatus, females of two uoglossum hemichalceum (Cockerell) (Mc- tween pairs of conspecific ^onnell-Garner and Kukuk 1997), Ruizan- communal andrenids, Andrena scotica Per- v were heda mutabilis (Spinola) (Packer unpub- kins and Pamirgus calcaratus Scopoli iished) and Pamirgus calcaratus Scopoli described by Paxton et al. (1999). P. calcar- Taxton et al. 1999)]. These observations atus displayed highly cooperative behav- while A. scotica lower levels intuitively agree with the behaviors ex- ior, displayed but pected from the above two social organi- of cooperation both displayed very zations: females in a communal nest must low levels of aggression, share the nest entrance, and thus a high Nolanomelissa toroi Rozen, a recently the level of cooperation is required, whereas described genus and species from of the Atacama Desert in solitary bees generally do not interact with southern border other females in their nest tunnels, and Chile (Rozen 2003), has defied the at- find its thus may lack the behavioral repertoire tempts of several melittologists to it is abundant. Fur- needed for cooperation. Intraspecific inter- nest in places where actions among individuals of eusocial bees thermore, it has an extremely female bi- are more complex, as different castes ased sex ratio: only three males have been whereas (queens, guards, and foragers) interact dif- seen after extensive collecting, of have been observed, ferently with each other. For example, for- hundreds females to examine if ager-forager interactions often result in a This study was conducted estab- higher frequency of cooperation, than circle tube experiments could help — lish whether N. toroi is a communal guard guard interactions [e.g. Lasioglos- spe- sum zephyrum (Smith) (Breed et al. 1978)]. cies, with perhaps few nest entrances per While queen—queen interactions often re- female and intranidal mating, thereby ex- sult in a high frequency of cooperative plaining the biased sex ratio. Communal Halictus is well known in the and aggressive interactions [e.g. behavior subfamily Macrotera texana ligatus Say (Pabalan et al. 2000)]. Similarly, Panurginae [e.g. (Cres- queen-forager interactions may be coop- son) (Neff and Danforth 1992), Perdita por- Perdita erative, or aggressive presumably to assert talis Timberlake (Danforth 1991), of Hymenoptera Research 50 Journal

one are forced to repeatedly interact with opuntiae Cockerell (Custer 1928), Panurgus Panur- another et al. 1978). The bees were calcaratus (Scopoli) (Knerer 1980), (Breed and unmarked, as marking has been shown to ginus albopilosus Lucas (Rozen 1971) influence behavior (Packer submitted). Meliturgula braunsi Friese (Rozen 1968)], hem- The behaviors of two of bees were and also in halictids [e.g. Lasioglossum pairs for 15 = L. and simultaneously recorded minutes, ichalceum ( erythrurum) (Kukuk introduction into the tubes. Schwartz 1987; Kukuk and Crozier 1990) following their All trials were recorded a digital and some species of Agapostemon (Janjic using and interac- and Packer 2003)]. On the assumption that camera (Sony DCR-TRV25) were scored from the vid- N. toroi is communal, we hypothesize that tions resulting trial the tubes were discard- its behavior in circle tubes should be con- eo. After each new ones were used for the next sistent with this type of social organiza- ed and set of tion: interacting females should show trials, preventing possible phero- mone contamination of bees comparatively high levels of cooperation among pairs and Weller The bees and little aggression. Acamptopoeum sub- (Smith 1989). paired sol- were then in Kahle's solution metallicum (Spinola), a predominantly preserved for dissection. A total of 10 trials were con- itary andrenid (Rozen and Yanega 1999), behavioral in- was also studied for comparative purpos- ducted for each species. A recorded for those in- es. Acamptopoeum submetallicum occurs in teraction was only FE. Interactions the same subfamily as N. toroi, the Pan- teractions that followed a as if one or both urgine (Ruz 1987), with the former be- were classified avoidance, turned or backed from a longing to tribe Calliopsini Robertson, and bees away FE, if each other the latter belonging to the tribe Nolano- cooperative the bees passed melissini Rozen and Ascher, which ap- or aggressive if one or both bees nudged, or one another. pears to be the sister group to all other lunged, bit, fought Fights Panurginae (Rozen 2003, Ascher in Rozen involved continued contact for several sec- 2003). We expect avoidance, with little co- onds, and in some instances, several min- operation, to be the common mode of in- utes, during which a series of aggressive teraction between A. submetallicum fe- interactions occurred successively. Every males. FE ended either in a pass or avoidance interaction. In instances where a FE was fol- METHODS lowed by an aggressive interaction, and Acamptopoeum submetallicum females were ended in a pass, both interactions were collected over a nesting aggregation near scored separately. However, if the aggres- Parque National Fray Jorge, Region IV, sive interaction was ended by one or both Chile (S30°38'W71°36'), on Nov. 13, 2002, bees backing away, only the aggressive in- from 11:00 AM to 1:30 PM. Nolanomelissa to- teraction was scored. The proportion of roi females were collected as they visited the the three categories of behavioral interac- flowers of Nolaua rostrata (Lindley), located tions was calculated as the frequency of approximately 9 km north of Vallenar, Re- that behavior divided by the total number gion III, Chile (S28°31'W70°44'), on Nov. 17, of FEs during a circle tube trial. 2002, between 11:00 AM and 1:00 PM. Bees preserved in Kahle's solution were Circle tube experiments were conducted dissected using a dissection microscope on collected pairs of females of A. subme- (at 64 x magnification) to compare ovarian tallicum and N. toroi, within 5 minutes of development among females, and deter- their In an capture. outdoor shaded area, mine whether or not they had mated. bees were in 20 placed cm long clear plas- Ovarian development was established by tic tubes (internal diameter 5mm) joined scoring each of the six ovarioles as a frac- end-to-end to form a circle such that bees tion of a fully developed oocyte (a fully Volume 13, Number 1, 2004 51

in submetallicum and Table 1. Frequency of occurrence of different behavioral interactions Acatnptopoewn Nolanomelissa toroi. of Hymenoptera Research 52 Journal

solitary 3.6 communal 0.8 a semisocial .10 • eusocial X unknown .2 0.6 +» (0 k. Q. § 0.4 O

8 0.2

.11 12 —rX"

0.2 0.4 0.6 0.8 Aggression

of versus interactions, social or- Fig. 1. Plot of the average proportion cooperation aggressive grouped by 1. 2. L. (Ctenonomia) 3. L. hemichalceum, 4. ganization for the following species: Lasioglossum platycephalum, sp., 7. chloris L. pauxillum (Smith and Weller 1989), 5. Penapis toroi, 6. Ruizantheda mutabilis (Spinola), Corynura al. 9. A. 10. P. calcaratus (Spinola) (Packer unpublished), 8. Halictus lanei (Moure) (Packer et 2003), scotica, al. 11. A. 12. N. toroi (Paxton et 1999), submetaUicum , and (This study).

trial without a FE was excluded from fur- volving only one female backing out of a ther analysis. In total, 83 FEs were record- frontal encounter. 15% of all behavioral ined in 9 trials, with an average of 9.22 FEs teractions were aggressive, during which = per trial (SD 4.01). Females would often bites, lunges, nudges or fights were ob- stay in a FE for prolonged periods of time served. Only 2% of the interactions were before an interaction was observed, rang- cooperative passes (Fig. 1). ing from several seconds to well over a Mean ovarian development was 2.18 - = = minute and then either lunge, bite, or back (SD 0.60, n 16), and all females (n away from the opponent. The duration of 16) were mated (Table 2). 15 of the 16 fe- FEs was quite variable, ranging from a males contained at least one fully devel- second to over 3 minutes, in which case a oped oocyte within an ovariole, and the series of bites and a mandibular hold, remaining female contained developing where the mandibles of both bees are oocytes. The mean head width for A. sub- = clasped together in what appears to be a metaUicum females was 3.37mm (SD were observed. = prolonged bite, A sum- 0.14, n 16). Wing wear was highly var- of the occurrence of aggressive, co- iable: 8 females with < 2 nicks on the fore- e and avoidance interactions be- wing margin, 6 with a score of 20 (wherein ubmetallicum females is present- the margin of the wing was heavily erod- 1. Most of their interactions ed, and nearly half the wing was worn off) vv e (83%) (Fig. 1), usually in- and the remaining two each with 5 nicks. Volume 13, Number 1, 2004 53

= = Table 2. Dissection and body measurement averages for A. submetallicum (n 16), and N. toroi (n 20) : = = = = emales. Note: OD ovarian development, MW mandibular wear, WW wing wear, HW head width = md WL wing length.

Bee species OD Mated MW WW HW (mm) WL (mm)

4. submetallicum 2.18 ± 0.59 all mated 3.09 ± 3.27 8.5 ± 9.32 3.37 ± 0.14 7.17 ± 0.33

V. toroi 1.22 ± 0.46 all mated 0.3 ± 0.66 0.25 ± 0.64 3.03 ± 0.17 7.16 ± 0.36

for N. toroi fe- similarly, mandible wear was also highly ed. The mean head width = = variable, with 9 females with a score ^ 1, males was 3.03mm (SD 0.17, n 20). In i with a score of 5 (wherein half the man- contrast to A. submetallicum, wing wear dible was abraded), and 2 with a score of measurements showed that the majority 10 (the mandible worn to a short stump). of bees were unworn, with 17 females Differences in all the measured physiolog- with nicks on the forewing margin, and ical/morphological parameters between 3 with 1-2 nicks on the forewing margin. mteractants in a circle tube were not sig- Likewise, mandibular wear measurements that 16 females had nificantly correlated with the relative pro- revealed unworn their mandibles portions of aggressive or avoidance inter- mandibles, and 4 had actions (Spearman rank correlation, p ^ slightly worn. Similar to A. submetallicum, J. 129 for all tests). Note that two bees from differences in all the measured physiolog- in N. toroi separate trials escaped during transferring ical/morphological parameters attempts into Kahle's solution, and thus, females were not significantly correlated zould not be dissected and measured. with aggression or avoidance interactions Nolanomelissa toroi. —Frontal encounters in the circle tube (Spearman rank correla- > for N. toroi were frequent, occurring 193 tion, p 0.201, for all tests).— times in 10 trials, with an average of 19.3 Interspecific comparisons. Nolanomelissa = tube FEs per trial (SD 13.0). FEs would toroi females interacted in the circle av- quickly result in a bee either backing away more often than A. submetallicum. On from or acting aggressively towards its erage, N. toroi females had twice the num- opponent. Most females were very active, ber of frontal encounters as A. submetalli- = df = = consistently moving throughout the circle cum females (t -2.164, 17, p tube arena. A summary of the proportion 0.045). The circle tube behavior of N. toroi of behavioral interactions between pairs of was significantly more aggressive than N. toroi females is presented in Table 1. All that of A. submetallicum when using the of the interactions were either aggressive Mann-Whitney U test and the sequential = df = (62%) or avoidance (38%) (Fig.l). Aggres- Bonferroni adjustment (Z -3.056, = sive interactions included lunging, biting 8, p 0.002). However, the frequency of = = = or fighting. In five instances, prolonged avoidance (Z -0.287, df 8, p 0.774) = periods of fighting occurred, ranging from and cooperative interactions (Z —1.534, = = 1 to 3 minutes, during which a series of df 8, p 0.125) did not differ signifi- bites were observed. cantly between the two. Acamptopoeuui Of the 20 females used in circle tube tri- submetallicum females participating in the als, 14 contained at least one fully devel- circle tube experiments had significantly oped oocyte within an ovariole, and 6 con- higher levels of ovarian development = z = tained developing oocytes (Table 2). The (Wilcoxon signed-rank test, s 433, mean ovarian development was 1.22 (SD 4.356, p < 0.0001), mandibular wear (Wil- = = = = 0.46, n 20), and all females were mat- coxon signed-rank test, s 411, z 3.935, of Hymenoptera Research 54 Journal

all had ovaries it seems unlike- p < 0.0001), and wing wear (Wilcoxon developed = = = that it is a semisocial either. signed-rank test, s 403, z 3.802, p ly species The is a list of the facts 0.0001). following per- tinent to the biology of N. toroi. 1) It ap- DISCUSSION biased pears to have an extremely female The interactions of A. submetallicum fe- sex ratio, with approximately 100 females male en- males in the circle tube arena agree with being found on flowers for every countered. Its nests are dif- evidence of solitary nesting; more than 2) unusually 80% of frontal encounters were classified ficult to locate. 3) All females found on have at least some as avoidance, and very little aggression or flowers are mated and it ovarian that cooperation was observed (Fig. 1). Also development. 4) Considering seemed that A. submetallicum females females foraging for pollen with devel- are to have avoided initiating interactions in the circle oped oocytes expected already tube as supported by the comparatively excavated a nest, this species exhibits sur- low number of frontal encounters per trial prisingly little mandibular wear. 5) It exhibits levels of behavior in (avg. 9 FEs in 15 min.). On the other hand, high aggressive N. toroi interacted more frequently in the the circle tube apparatus. 6) It is active in there circle tube (avg. 19 FEs in 15 min.), with only in spring in years which has —60% of all interactions being aggressive, been sufficient winter rainfall. the so- and —40% avoidance (Fig. 1). Morpholog- What can we conclude regarding ical comparisons, with regard to wing and cial biology of N. toroi? First, based on ev- mandibular wear revealed that females of idence from circle tube experiments, N. to- N. toroi were younger, or at least had been roi does— not exhibit a communal organi- much less active in flight and nest exca- zation this species clusters on the oppo- vation than those of A. submetallicum. This site behavioral spectrum to known might suggest that bees are more aggres- communal andrenids and halictids (Fig. sive in early the nesting cycle. However, 1). Second, the presence of developed ova- no significant relationship was found be- ries accompanied by a short activity peri- tween the frequency of aggressive behav- od suggests that N. toroi does not have a ior and relative age in either species complex social organization despite the > [Spearman rank correlation: p 0.129 (A. high levels of observed. It is > aggression submetallicum) and p 0.201 (N. toroi) for worthwhile to note that N. toroi females all tests]. exhibit a peculiar pygidial plate and mod- Due to the difficulties associated with ified hind basitibial plates (Rozen 2003), nests and the finding highly female biased structures that are usually consistent in sex ratio throughout its activity period, we ground nesting bees (Rozen personal com- had that N. toroi hypothesized might be a munication). This fact, accompanied by communal species. However, the majority the extremely female biased sex ratio in of interactions among females of this spe- combination with low mandibular wear cies were with not one in- aggressive, and difficulty in finding nests are sugges- stance of cooperative behavior observed. tive of perhaps an unusual choice of nest Such levels of are high aggression more site and/or an unusual mating system. of a division of suggestive reproductive ACKNOWLEDGMENTS labor. However, this species is active at most once a year (only in years in which The research described here was carried out while there has for a funded a National been adequate winter rainfall) collecting samples project by a Geographic Research and Exploration Grant, to precluding standard eusocial colony cy- which we are grateful. All three authors are funded cle with spring gynes and summer work- by NSERC through graduate scholarships (J.C.G. and . As all females were mated and almost and a A.Z.) discovery grant (L.P.). We are grateful to Volume 13, Number 1, 2004 55

Horacio Larrain, Marta Pena, Alfredo Ugarte, and Lu- menoptera: Andrenidae). journal of the Kansas En- isa Ruz for their help in Chile. We would like to tomological Society 64: 394-405. E. differentiation in thank Dr. J.G. Rozen, Jr. and two anonymous review- Ordway, 1965. Caste Augochlorella ers for helpful comments on the manuscript. (Hymenoptera, Halictidae). Insectes Sociaux 12: 291-308. LITERATURE CITED Pabalan, N., K. G. Davey and L. Packer. 2000. Esca- lation of aggressive interactions during staged the encounters in Halictus ligatus Say (Hymenoptera: Ascher, J. S. 2003. Evidence for phylogenetic po- with a of circle tube be- sition of Nolanomelissa from nuclear EF-la se- Halictidae), comparison haviors with other halictine In- quence data. In Rozen, 2003. species. Journal of sect Behavior 13: 627-649. Bell, W. J. and W. A. Hawkins. 1974. Patterns of in- Packer, L. 1997. The relevance of phylogenetic sys- traspecific agonistic interactions involved in nest tematics to from medicine and defense of a primitively eusocial Halictine bee. biology: examples behavioral ecology. PI 1-29. In P. Grandcolas journal of Comparative Physiology 93: 183-193. (ed.). The Origin of Biodiversity in Insects: Phylo- Breed, M. D, J. M. Silverman and W. J. Bell. 1978. Tests Scenarios. Memoires du Agonistic behavior, social interactions, and be- genetic of Evolutionary Museum National d'Histoire Naturelle vol. 173. havioral specialization in a primitively eusocial bee. Insectes Sociaux 25: 351-364. 360pp. Packer, L. 2000. The of Thincohalitcus prog- Brothers, D. J. and C. D. Michener. 1974. Interactions biology nathic (Perez) Halictidae: Halic- in colonies of primitively social bees III. Ethom- (Hymenoptera: tini). Journal of Research 9: 53-61. etry of division of labor in Lasioglossum zephyrum Hymenoptera Packer, L., B. W. T. Coelho, S. Mateus and R. Zucchi. (Hymenoptera: Halictidae). Journal of Comparative 2003. Behavioral interactions females of Physiology 90:129-168. among Halictus (Seladonia) lanei Custer, C. P. 1928. The bee that works in stone; Perdita (Moure) (Hymenoptera: Halictidae). Journal of the Kansas So- opuntiae Cockerell. Psyche 35: 67-84. Entomological 76: 177-182. Danforth, B. N. 1991. Female foraging and intranest ciety Paxton, R. J., P. F. Kukuk and J. 1999. Effects behavior of a communal bee, Perdita portalis (Hy- Tengo. of and nestmate number on social in- menoptera: Andrenidae). Annals of the Entomolog- familiarity teractions in two communal bees, Andrena scotica ical Society of America 84: 537-548. calcaratus Andren- L. the bee and Panurgus (Hymenoptera, Janjic, J. and Packer. 2003. Phylogeny of Insectes Sociaux 46: 109-118. genus Agapostemon (Hymenoptera: Halictidae). idae). Rice, W. R. 1989. tables of statistical tests. Systematic Entomology 28:101-123. Analyzing Evolution 43: 223-225. Knerer, G. 1980. Panurgus calcaratus (Hymenoptera: Rozen, J. G., Jr. 1968. and immature of Andrenidae), a communal bee of Europe. Zoolo- Biology stages the aberrant bee American Mu- gischer-Anzeiger 204: 64-68. genus Meliturgula. seum Novitates 2331: 1-18. Kukuk, P. F. 1992. Social interactions and familiarity Rozen, G., Jr. 1971. and immature of in a communal halictine bee Lasioglossum (Chilal- J. Biology stages Moroccan bees. American Museum ictus) hemichalceum. Ethology 91: 291-300. panurgine Novitates 2457: 1-37. Kukuk, P. F. and M. P. Schwartz. 1987. Intranest be- 2003. A new and havior of the communal sweat bee Lasioglossum Rozen, J. G., Jr. tribe, genus, species of South American bee (Andrenidae: (Chilalictus) erythrurum (Hymenoptera: Halicti- panurgine on Nolana (Nolanaceae). dae). Journal of the Kansas Entomological Society 60: Panurginae), oligolectic 58-64. pp. 93-108. In G. A. R. Melo & I. Alves-dos-San- tos Lima aos Kukuk, P. F. and R. H. Crozier. 1990. Trophallaxis in (eds.). Apoidea Neotropica: Homenagem 90 Anos de S. Moure. Editora Cir- a communal halictine bee Lasioglossum (Chilalic- Jesus UNESC, Brazil. tus) erythrurum. Proceedings of the National Acad- ciuma, and D. 1999. emy of Sciences USA 87: 5402-5405. Rozen, J. G., Jr. Yanega. Nesting biology and immature of the South American bee McConnell-Garner, J. and P. F. Kukuk. 1997. Behav- stages Andreni- ioral interactions of two solitary, halictine bees genus Acamptopoeum (Hymenoptera: dae: Kansas Natural with comparisons among solitary, communal Panurginae). University of Publication 24: 59-67. and eusocial species. Ethology 103: 19-32. History Museum Special relation- Michener, C. D. 1974. The Social Behavior of the Bees. Ruz, L. 1987. Classification and phylogenetic Harvard University Press, Cambridge, Mass. ships of panurgine bees (Hymenoptera-Andren- Michener, C. D. 2000. The Bees of the World. The John idae). Ph.D. thesis, University of Kansas. Hopkins University Press, Baltimore and Lon- Smith, B. H. and C. Weller. 1989. Social competition don. among gynes in halictine bees: the influence of B. size on behavior. Neff, J. L. and Danforth, N. 1992. The nesting and bee and pheremones Journal of foraging behavior of Perdita texana Cresson (Hy- bisect Behavior 2: 397^111. 56 Journal of Hymenoptera Research

Wcislo, W. T., A. Wille and E. Orozco. 1993. Nesting context in a largely solitary bee, Lasioglossum biology of tropical solitary and social sweat bees, (Dialictus) figueresi (Hymenoptera: Halictidae). Lasioglossum (Dialictus) figueresi Wcislo and L. (D) Insectes Sociaux 44: 199-208. aeneiventre (Friese) (Hymenoptera, Halictidae). Wcislo, W. T. and B. N. Danforth. 1997. Secondarily Insectes Sociaux 40: 21^0. solitary: the evolutionary loss of social behavior. Wcislo, W. T. 1997. Social interactions and behavioral Trends in Ecology and Evolution 12: 468-474. J. HYM. RES. Vol. 13(1), 2004, pp. 57-63

Report on a Collection of Bethylidae (Hymenoptera) from Central Florida, USA, with Description of a New Species of Lepidosternopsis Ogloblin

Geane O. Lanes, Fernanda T. Gobbi, and Celso O. Azevedo

Universidade Federal do Espirito Santo, Departamento de Biologia, Av. Marechal Campos 1468, Maruipe, 29.040-090 Vitoria, ES, Brazil; (COA) email: [email protected]

Abstract. —Sixty species of Bethylidae of the genera Acrepyris Kieffer, Allobethylus Kieffer, Ape- nesia Westwood, Anisepyris Kieffer, Bakeriella Kieffer, Cephalonomia Westwood, Dissomphalus Ash- mead, Epyris Westwood, Goniozus Forster, Holepyris Kieffer, Laelius Ashmead, Lepidosternopsis Og- loblin, Plastanoxus Kieffer, Prorops Waterston, Prosierola Kieffer, Pseudisobrachium Kieffer, Wiabde- in Malaise at 12 sites in pyris Kieffer, and Sclerodermus Latreille were collected, primarily traps, Orange Co., 3 in Seminole Co., one in Osceola Co. and one in Volusia Co., Florida, USA. Lepidos- and irradiata ternopsis is recorded for the first time from the Nearctic Region Lepidosternopsis is described Lanes and Azevedo, sp. nov., is described and illustrated. The male of Lepidosternopsis for the first time. Taxonomic comments for some species are included.

There are about 2,000 species of Bethy- Our goal was to survey the Bethylidae lidae worldwide, 204 of which are record- of Florida, to establish range extensions variation of ed from the Nearctic Region. The few spe- and to analyze the taxonomic the studied The we cies whose biologies have been investigat- species. specimens studied 60 18 of which ed are idiobiont, or incipient koinobiont, represent species, are different from the 52 ectoparasitoids (Finnamore and Gauld recognized by Evans and Fullerton Five 1995). They parasitize larvae of Coleoptera (1997). genera: Wes- or Lepidoptera that live in cryptic situa- Allobethylus Kieffer, Cephalonomia Plastanox- tions. Most Bethylinae parasitize microle- twod, Lepidosternopsis Ogloblin, "s Kieffer, and Waterston were not pidopterans, whereas most Pristocerinae Prorops observed Evans and Fullerton and Epyrinae attack beetles, especially by (1997). Some occurred in remarkable those inhabiting wood or seeds (Evans species for 739 males of Pseu- 1964) numbers, example disobrachium (Kieffer). In the United States, Florida possibly flaviventre Lepidos- is recorded for the first time from has the richest bethylid fauna. Beginning ternopsis the Nearctic Region based on a new spe- in 1990, biologists at the University of cies describe in this a er " ' which we P P Central Florida surveyed the arthropod and Fullerton fauna of the campus (Evans TERMINOLOGY 1997) and other adjacent sites. Evans and Fullerton (1997) studied an assemblage of Terminology generally follows Evans 52 species of Bethylidae, based on approx- (1964). The nomenclature of integument imately 3,000 specimens from Central sculpture follows Harris (1979). The ter- Florida. We have studied an equally large minology of wing cells and veins follows second assemblage of Bethylidae in order Gauld and Bolton (1988). Abbreviations to better understand the fauna of this used for the description of the new species state. are: DAO, diameter of anterior ocellus, 58 Journal of Hymenoptera Research

Orlando. measured in frontal view; HE, height of 9. Orange Co., UCF, Mackay Marh. Red eye, measured in lateral view, across its Tract, Swgrass Maple. Orlando. maximum height; LFW, length of fore 10. Orange Co., UCF, Cypress wing; LH, length of head, measured in Forest. frontal view, from the vertex crest to the 11. Orange Co., Orlando. UCF, Long Leaf Pine Pine Oak. median apical margin of the clypeus; Sand Turkey OOL, ocello-ocular line, measured in la- 12. Orange Co., Orlando. UCF, Long Leaf tero-dorsal view, the shortest distance Pine Saw Palmetto. from the eye top to the posterior ocellus; 13. Orange Co., Orlando. UCF, Maiden- VOL, vertex-ocular line, measured in lat- cane Marsh. eral view, the distance from the eye top to 14. Orange Co., Orlando. UCF, Pond Pine vertex crest; WF, width of frons, measured Comm. Dahoon Holly. in frontal view, its minimum width; WH, 15. Orange Co., Orlando. UCF, Sand Pine width of head, measured in frontal view, Rosemary Scrub. its maximum width including the eyes; 16. Osceola Co. Walt Disney World. WOT, width of the ocellar triangle, mea- World Drive/US 192 SOI T25S R27E. sured in frontal view, the maximum Sand Pine /Rosemary Scrub. width, including the ocelli. 17. Sarasota Co., MCC-Venice Campus. Long Leaf Pine-Saw Palmetto. COLLECTION SITES 18. Volusia Co., Daytona Beach. Urban- Beachside, Halifax-River. The material examined is deposited at the Entomological Collection of the Uni- LIST OF SPECIES versity of Central Florida, Orlando (Stuart Eight species of Bethylidae are recorded M. Fullerton). It was collected using Mal- for the first time from Florida. These are aise traps, pit fall traps, UV light traps, indicated with an asterisk (*). of and sweeping between 1997 and 2000 in Listing genera follows Evans (1978), with species 18 sites in Central Florida in Orange, Sem- of each genus listed inole, Osceola, Sarasota and Volusia alphabetically. Counties. The collection sites are referred Subfamily Bethylinae to by number in the list that follows. Goniozus columbianus Ashmead. 56 9, 18. Sites 1, 6, 9, 10, 14. 1. Seminole 11, 13, Co., Econ Wilderness Area. * Goniozus complanatus Evans. 39. Site 15. Scrub Oak/Saw Palmetto (burned). Goniozus electus Fouts. 2 9 . Site 13. 2. Seminole Co., Econ Wilderness Area. Goniozus flavipes Fouts. 206. Sites 1, 2, 6, 9, 16. Scrub Oak/Saw Palmetto (unburned). Goniozus floridanus (Ashmead). 59. Sites 9, 10, 3. Seminole Co., Oviedo. 11. 4. Co., LK Tibet- Butler Pre- Orange Goniozus fratellus Evans. 16. Site 13. Flatwoods. serve/Scrubby Goniozus gracilicornis (Kieffer). 16. Site 11. 5. Orange Co., Orlando, Tibet Preserve Goniozus hortorum Brues. 1309, 2d. Sites 2, 6, 9, Myrtle Oak Scrub. 10, 11, 13, 14, 15, 16. Goniozus 6. Orange Co., Walt Disney World. C- 4 hubbardi Howard. 169,28. Sites 9, 13, 18. Stout Site SI 5, 16 T24S R27E. Xeric 14, Goniozus Oak/Flatwoods. nigrifemur Ashmead. 98 9, 66. Sites 1, 2, 6, 8, 9, 10, 11, 16. 7. Orange Co., Walt Disney Wolrd. MW- 13, 14, Prosierola bicarinata 39. Sites 4. 5 (unburned) S16 T24S R27E. Sand (Brues). 1, Pine/Oak Scrub. Subfamily Epyrinae 8. Orange Co., Walt Disney World. MW- Allobethylus floridanus Evans. 79. Sites 11. 7 S22 T24E. Sand 9, (unburned) Pine/ analis Anisepyris (Cresson). 1009, 39^. Sites 1, Oak Scrub. 2,6,8,9, 10,11,12,14. Volume 13, Number 1, 2004 59

Anisepyris columbianus (Ashmead). 349, 26 8. Acrepyris atra Klug. 3 c?. Sites 1, 2, 11. Sites 6, 8, 9, 10, 11, 12, 14. Acrepyris bridwelli Evans. 17c?. Sites 8, 11, 12, 14. Anisepyris grandis (Ashmead). 259, 211c?. Sites Acrepyris fraterna Evans. 2 9, 45

4, 5,' 6, 8, 9, 10, 11, 12, 14, 15, 16. ll,'l2, 14. Anisepyris subviolaceus Kieffer. 15 9. Sites 6, 8, Apenesia parapolita Evans. 5 9, 130c?. Sites 6, 8, 10, 11. 9, 10, 11, 12, 14, 16. Bakeriella mira Evans. 6 c?. Sites 9, 11. Dissomplialus apertus Kieffer. 311 c?. Sites 8, 9, 10, Cepmalonomia hyalinipensis Ashmead. 20 9, 2d. 11, 14. Sites 1, 9, 10^ 11, 14. Dissomplialus barberi Evans. 10c?. Sites 9, 11, 14. Evans. lie?. Sites Cephalonomia perpusilla 9, 11, Dissomplialus evansi Azevedo. 34 c?. Sites 6, 8, 9, 14. 10, 11, 12, 13, 14, 16. * Evans. 1 9 . Site 6. Ceplmlonomia quadiceps Dissomplialus kansanus Evans. 2c?. Sites 6, 10, 11. * Evans. 29. Site 9. Cephalonomia conophthori Dissomplialus krombeini Azevedo. 3

blue reflections, and some with reddish Laelius centratus (Say): females can have legs. legs light castaneous. ^ Cephalonomia conophthori Evans: the pro- Plastanoxus laevis (Ashmead): this wide- podeal disc is 1.25X as long as wide. spread species is recorded for the first Cephalonomia hyalinipennis Ashmead: some time from Florida. females are light castaneous and the Prorops obsoleta Evans: this species was head varies in length, some have the known only from 2 males and 1 female head longer or with side parallel behind from Trinidad. the eyes. Prorops nasuta (Say): in the Nearctic Re- Cephalonomia quadriceps Evans: this species gion this species was known only from was known from 3 males from Massa- 1 male from California. chusetts, Maryland and North Carolina, Pseudisobrachium arenarium Evans: the width USA. of the male frons ranges from 1.30 to 1.40X Epyris corticinus Evans: this species was height of eye and propodeal known from Virginia, Maryland and disc from 1.60 to 1.70X as long as wide. Pennsylvania, USA. In this series, some females have small Pseudisobrachium ashmeadi scutellar pits. Evans: length of Epyris deficiens Krombein: males can have propodeal disc in the males can be shorter than in poorly defined longitudinal striae. the type series, 1.26X as as wide. Epyris rufipes (Say): some females can have long the Pseudisobrachium propodeal disc evenly striate and flaviventre (Kieffer): in the median transverse males the head varies from dark casta- vein not strongly neous to the oblique. black, antennae from light Goniozus columbianus Ashmead: males can to dark castaneous, the diameter of the anterior have the head weakly coriaceous. ocellus varies from 0.17 to Goniozus 0.23 X the width of the and the complanatus Evans: this species frons, with was known only from Texas, USA. mesopleuron callus ill defined. Goniozus hortorum Brues: female heads Psedisobrachium rufiventre (Ashmead): this in species is similar to P. vary length, punctures size, frons tex- very flaviventre, ture, differing by the discoidal vein clypeus length, sculpture, and having of median slightly conspicuous, the antennae with height carina of clypeus. and Males can have the cubital vein 2.5X as segments slightly longer propodeal disc long as wide. slightly longer. Males can have the with callus well Goniozus hubbardi Howard: the third an- mesopleuron defined. Scleroderma Ashmead: the tennal segment can be 1.5X as lone as macrogaster pattern of coloration thick. body shows a large range of variation, but with the meta- Goniozus nigrifemur Ashmead: females can soma dark castaneous. The have the antennae as short as in G. emi- constantly head can be dark grants. Males can have the antennae castaneous, distinctly darker than the thorax; the thorax can reaching the vertex crest, the third an- be darker at the tenna 1 distinctly mesoscutum segment as long as thick, or the and and the posterior carina of the mesopleuron, head and propodeal disc thorax can be well-indicated. evenly light castaneous. catalinae Evans: Holepyris this species was Lepidosternopsis irradiata Lanes and known only from Arizona, USA. Azevedo, sp. nov. Holepyris graminis Evans: males can have Figs. 1-8 the posterior groove of the pronotal disc Male holotype.— 2.4 weakly Body length mm; developed. FW 1.6 mm. Color: Dark castaneous; an- Volume 13, Number 1, 2004 61

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dorsal. Man- Figs. 1-8. Lepidosternopsis irradiate sp. nov. 1—4, Male. 5-8, Female. 1, Head and mesosoma, 2, dible, frontal. 3, Forewing, dorsal. 4, Genitalia, ventral. 5. Head and mesosoma, dorsal. 6, Mandible, frontal. = 7, Forewing, dorsal. 8, Metasoma, ventral. (Scale bars 0.32 mm). Research 62 Journal of Hymenoptera

with volsella with tenna, mandible, legs and palpi casta- apex rounded; cuspis shorter than neous; wings hyaline, veins castaneous. laminar, very wide, slightly 4 ventral excavate in Head (Fig. 1): Mandible with apical paramere, margin api- teeth, the lower one relatively bigger and cal half; aedeagus bottle-shaped, progres- with truncate slender sharper (Fig. 2). Clypeus sively apically, apex emarginated, median lobe, with median carina concave basal margin rounded, in Female.— 2.8 mm; LFW 1.9 in profile. First four antennal segments Body length to male Color: Head ratio of 12:6:3:4; segment III 0.8X as long mm. Similar except: the as thick; segment XI 1.6 X as long as thick; and thorax castaneous, except prono- antennal sockets closed to each other, sep- turn light castaneous and propodeum yel- dark casta- arated by 0.3 X their diameter. Eye with lowish anteriorly; metasoma antennae and casta- sparse, short setae. Frons weakly coria- neous; palpi light ceous, shinning, with small, very sparse neous; mandibles and legs castaneous; vein castaneous. punctures, with shallow and small groove wings hyaline, light between the antennal sockets. LH 1.08X Head (Fig. 5): Mandible with 3 apical WH; WF 0.58 X WH; WF 1.12 X HE; OOL teeth, the higher very shorter than lowers 0.85X WOT; DAO 0.41 X WOT; frontal an- (Fig. 6). Clypeus very short, concave me- carina gle of ocellar triangle acute; posterior ocel- dian lobe, with median weakly lus distant from the vertex crest 0.5 X high and concave in profile. First four an- DAO. Vertex slightly convex with corner tennal segments in a ration of 25:8:4:4; seg- somewhat angled. VOL 0.85X HE. Head ment III as long as thick; segment XI 0.7X not flattened, its thickness 0.62 X LH. Me- longer than thick; antennal sockets close to sosoma (Fig. 1): Thorax slightly coria- each other, separated by 0.6X their diam- ceous. Pronotal disc 1.25 X as long as me- eter. Front very weakly coriaceous, shin- soscutum, with transverse carinae very ning. Head globoid, 0.58 X as high as long, weak. Notauli very weak, occupying an- LH 1.15X WH; WF 0.37X WH; WF 1.30X terior third of the mesoscutum; parapsidal HE; OOL 2.06 X WOT; DAO 0.3 X WOT; furrow absent; scutellar groove slender, posterior ocelli distant from the vertex 1.12X Propodeal disc as wide as long, 1.4X DAO. Vertex slightly convex, with weakly coriaceous, shinning; without me- corner somewhat angled. Temples slightly dian and posterior carinae, with lateral ca- converging anteriorly. VOL 1.20X HE. lateral rinae, margins straight; declivity Mesosoma (Fig. 5): Thorax coriaceous, without median carina; propodeal spiracle Pronotal disc 1.72X longer than mesoscu- totally directed outward. Mesopleuron turn; notauli very weak and inconspicu- weakly coriaceous, with wide, shallow fo- ous; parapsidal furrow, incomplete ante- veae all with a occupying nearly surface, riorly, occupying more than half of the small central Fore pit. wing (Fig. 3) with mesoscutum. Propodeal disc as long as veins Sc + Rs + + R, M, M Cu and lCu-a, wide, coriaceous, lateral margins concave one closed cell forming only (median), anteriorly. Mesopleuron slightly coria- Fore femur 2.1 X as as wide. long Median ceous, with large and shallow fovea oc- tibia not Claws Metaso- spinose. simple. cupying surface, with small central pit. ma: of sternites IV-VI Fore posterior margin wing as for male (Fig. 7). Fore femur with biemarginated. Hypopygium poste- 1.94X as long as wide. Metasoma: 1.73X rior narrow and margin concave. Genitalia as long as the mesosoma. Posterior margin (Fig. 4): Paramere divided in of sternites completely IV-VI biemarginated (Fig. 8). two arms, ventral arm with wide, apex Type material—Holotype 6, USA, Flor- somewhat truncate in ventral dorsal half, ida: Orange Co., Orlando, University Cen- ventral margin convex, margin straight, tral Florida, Cypress Forest, 2.VI.1999, dorsal arm shorter than ventral, slender, Malaise trap, P. Russell and S. Fullerton Volume 13, Number 1, 2004 63 col. (deposited in Entomological Collec- described the male of this genus for the tion, University of Central Florida, Orlan- first time. L. irradiata differs from all the do). Allotype 9, USA, Florida: Orange known species of the genus, because it is Co., Orlando, University Central Florida, macropterous with fore wings having MacKay Tract, Sawgrass Marsh, Red Ma- only the median cell closed and radial vein absent. ple, 14.VI.1999, Malaise trap, P. Russell and S. Fullerton col. Paratypes, USA, Flor- ACKNOWLEDGMENTS ida: Orange Co., Orlando, University Cen- We thank S. Fullerton for the loan of the material tral Florida: 16 , Marsh, Red Ma- Sawgrass and the duplicates retained. ple, 13.IX.1999, Malaise trap, P. Russell LITERATURE CITED and S. Fullerton col.; 16" and 19, LLP- Sand Pine, Turkey Oak, 7.XI.1997 and Azevedo, CO. 1999. Additions to the Neotropical Epyrinae (Hymenoptera, Bethylidae), with de- 2.VII.1997, Malaise trap, P. Russell and S. scriptions of a new species of Eepidosternopsis Fullerton col.; 19, Pond, Pine Comm., Da- from Brazil. Iheringia, serie Zoologia 87:11-18. hoon Malaise P. Holly, 11.V.1999, trap Evans, H.E. 1964. A synopsis of the American Bethy- 1 lidae Bulletin the Russell and S. Fullerton col.; 9 , Maiden- (Hymenoptera, Aculeata). of Museum 132(l):l-222. cane Marsh, 17.V.1999, Malaise trap, P. of Comparative Zoology Evans, H. E. 1978. The Bethylidae of America north Russell and S. Fullerton col. of Mexico. Memoirs of the American Entomological — refers to ab- Etymology. The name the Institute 27:1-332. sence of the radial vein in the fore wings. Evans, H. E. and S. M. Fullerton. 1997. Report on a collection of from Discussion.—The description of the ge- Bethylidae (Hymenoptera) Central Florida. Proceedings of the Entomological nus Lepidosternopsis was based on an ap- Society of Washington 99(1):174-179. terous female of L. kuscheliana Olgloblin, Finnamore, A. and I. D. Gauld. 1995. Bethylidae, p. 1953, from Masatierra Island (Chile). 470-479. In: Hanson, P. AND I. D. Gauld. The Evans (1964) described two species based Hymenoptera of Costa Rica. Oxford, Oxford Uni- on micropterous females from Australia. versity Press, 893p. Gauld, I. D. and B. Bolton. 1988. The Hymenoptera. Azevedo (1999) described the first mac- Oxford, Oxford University Press. 332 pp. of based ropterous species Lepidosternopsis, Harris, R.A. 1979. A glossary of surface sculpturing. on two females from Para, Brazil. Now we Occasional Papers in Entomology 28:1-31. J. HYM. RES. Vol. 13(1), 2004, pp. 64-107

Braconidae: A Revision of Agathirsia Westwood (Hymenoptera: Agathidinae) With Notes on Mouthpart Morphology

Thomas Pucci and Michael Sharkey

S-225 Science Center North, (TP) Department of Entomology, University of Kentucky, Ag. address: The Cleveland Lexington, Kentucky 40546-0091, USA; email: [email protected] (current Ohio Museum of Natural History, 1 Wade Oval Drive, Cleveland, 44106, USA); Center (MS) Department of Entomology, University of Kentucky, S-225 Ag. Science North, Lexington, Kentucky 40546-0091, USA; email: [email protected]

are Abstract. —A cladistic analysis of Agathirsia Westwood is performed and synapomorphies reduction of the ventral mandibular proposed for the first time for members of Agathirsia, i.e., tooth, and apicomedial pegs of the hind tibia flattened, and for members of its sister-group Cras- tibia and and tarsal claws somicrodus Ashmead, i.e., apicomedial pegs of the hind sharp seta-like, to the arid of Mexico simple, without basal lobes. Members of Agathirsia are restricted regions and the southwestern USA. Keys are provided to identify the genera of Agathidini and the species of nectar extraction of Agathirsia. The mouthparts of Agathirsia are described and three main types mechanisms are discussed, i.e., 1) sponge-like glossa that retracts within the galeae, 2) straw-like glossa that does not retract, and 3) labial palpi that form a drinking tube. The genus Agathirsia is revised to include 31 species, 23 of which are newly described, i.e., A. armandi, A. asterophila, A. bicolor, A. bifidilingna, A. campattisura, A. capillata, A. collini, A. davidi, A. foveiseries, A. heleni, A. kellyi, A. keni, A. jervisi, A. longigladia, A. longilingua, A. michelei, A. minuata, A. ninesevensi, A. papotii, A. parkeningi, A. real, A. rostrata, and A. tiro. Agathirsia rufiventris Westwood is considered a junior synonym of A. proxima Westwood. The monotypic genus Cenostomus is synonomized with Agathrisia. Cenostomus trichiosotmis (Cameron) is transferred to Agathirsia, and Crassoniicrodus pumilus (Szepligeti) is transferred to Bassus Fabricius.

Agathirsia Westwood is a member of the Crassoniicrodus from other agathidines Agathidinae, which is a well-established with which they may be confused. A cla- monophyletic group within the Braconi- distic analysis is performed with the aims dae (Sharkey 1992). Three autapomor- of testing the monophyly of Crassomicro- for the phies subfamily are the presence of dus and Agathirsia, revealing the relation- male on specialized tergal glands meta- ships among species of Agatliirsia, and un- somal 7 segments and 8 (Buckingham and covering the evolution of morphological Sharkey 1988), a wing fold between the characters such as those of the mouth- and the medial vein prestigma of the fore- parts. Previous to this study eight species and a narrow wing, marginal cell (Sharkey of Agathirsia were described but the single 1992). Agathidinae is a cosmopolitan sub- revision of the genus (Muesebeck 1927) re- of 52 family composed genera (Sharkey viewed only species found north of Mex- and members are found in 1997) most ter- ico. Neither Muesebeck's key to North restrial habitats. American agathidine genera, nor Shar- Here we describe and all key known key's (1997) key to the New World genera of members of species Agathirsia, which adequately distinguish between our con- are restricted to the arid of regions Mexico cepts of Crassoniicrodus and Agathirsia be- and the southwestern USA. We also pre- cause they rely on the shape of the mouth- sent a to and key distinguish Agathirsia parts—a variable character in both genera. Volume 13, Number 1, 2004 65

All known species of Agathidinae are vious defense mechanisms. Many species are or solitary koinobiont endoparasitoids of lar- of Agathirsia partly entirely orange, val Lepidoptera (Sharkey 1992) except for a common color for stinging Hymenop- one gregarious species recently discovered tera in the area. Other species of Agatliirsia by D. Janzen (unpublished). The few are entirely black with long, dense setae members of the Microdini and Agathidini and may mimic bees. with known biologies all attack the first The standard mandibular morphology instar larval stage of their lepidopteran for most species of Agathidinae is illus- are hosts, and although Agathirsia is a puta- trated in figure la. These mandibles tive member of the Agathidini we are not rather gracile, flattened and designed to sure if its members share this characteris- cut through silk, i.e., their own and/or In tic. Many species of Agathirsia and Cras- that of their host Lepidoptera. contrast, are somicrodus have very short ovipositors. the mandibles of species of Agatliirsia Within the Agathidinae this character cylindrical and much more robust (Figs. state is found almost exclusively in the Di- lb,c). Due to their similarity to the man- that nest sophrini, all members of which possess dibles of aculeate Hymenoptera the state. There are few observations of underground, we speculate that the shape members of Disophrini ovipositing but may be a modification that facilitates dig- these few indicate that they attack late in- ging. We suggest that species of Agathirsia stars of their hosts. We suspect that the are attacking larval Lepidoptera that are in the soil same is true for members of Agatliirsia and spending the daylight hours Crassomicrodus, at least those with short and /or that pupate in the soil. There is for ovipositors. only one host record Agathirsia; Bibby Little is known of the natural history of (1961) cites the larva of Acontia cretata a of and the members of Agathirsia. We speculate that (Noctuidae) as host Agathirsia, is consistent most or all species are diurnal based on natural history of this host their coloration and the absence of any la- with our hypothesis. This same mandib- bel data to indicate otherwise. Members of ular morphology, and perhaps the same in of Crassom- Agathirsia appear to be restricted to the biology, is present members arid regions of Mexico and the southwest- icrodus as their mandibles are even more of ern USA and most species are collected robust (Fig. Id). Many species Agathirsia from August through November, which is have the apex of the hind tibia flared, like The flared typically the end of a relatively wet period much bell-bottom pants. and the beginning of a dry period over region is highly sclerotized and we spec- modification most of the range. Many species have ulate that this may also be a elongate mouthparts and since most de- for digging. Relative to most Agathidinae, sert flowers produce nectar in daylight which have rather large, cylindrical pegs hours, this is further evidence to suggest on the hind tibia apico-laterally (Fig. 2a); that they are diurnal. those of members of Agathirsia are small value Most members of Agathirsia appear to and flat (Figs. 2b, c). The functional have mimetic coloration. E. G. Linsley not- of the flattened pegs is unknown but they ed the following behavior on a specimen may aid in soil excavation. "while tag of A. nigricauda: feeding [the] METHODS wasp spreads [its] wings and flashes ab- follows dominal and wing colors with each partial Morphological terminology rotation". This behavior is similar to that Sharkey and Wharton (1997). Glossa and the ba- of some stinging wasps and we speculate galea lengths were measured from that it is likely a case of Batesian mimicry ses of their respective palpomeres (see Fig. since members of Agatliirsia have no ob- 4b). Journal of Hymenoptera Research 66

mandibles for the that are Fig. 1. Mandibles of Agathidini: a, Agatliis sp., showing ground plan Agathidinae size of ventral twisted and function like scissors to cut through silk, b, Agathirsia testacea, showing maximum and tooth in the genus, c, Agathirsia nigricauda, showing typical mandible for Agathirsia, robust, cylindrical, for Crassomicrodus. lacking ventral tooth, d, Crassomicrodus divisus, showing typical shape of a mandible

The keys are designed for all known variation for both sexes. All measure- the species. Although ovipositor characteris- ments are in millimeters. Occasionally tics are included, both males and females glossae of short-tongued Agathirsia may be can be used. The percentages (%) indicat- partly to completely folded lengthwise ed in the keys represent the frequency along the midline. The glossae of these with which the user can expect to observe specimens often measure 15 to 30% short- the character state given a specimen that er than conspecifics because they are part- should key to that side of the couplet. We ly retracted. Measurements of these have weighted the common species to arrive at not been included in the descriptions ex- these figures. cept where noted. The number of hind tib- The key to Agathirsia was produced by ial pegs is a character that is often difficult editing a key generated using DELTA to observe; high magnification with dif- (Dallwitz et al. 1993). The species descrip- fused lighting is essential. ons of Agathirsia include total observed The holotypes of all described species of olume 13, Number 1, 2004 67

ig. 2. Hind tibial pegs: a, Bnssus ussitriensis, showing typical cylindrical pegs of the Agathidinae. b, Agathirsia igricauda, showing flattened pegs, c, Agathirsia collini, showing flattened pegs, d, Crassomicrodus divisus, show- ig hair-like spines.

{gathirsia were examined. If a reliably de- Florida (AEIC); American Museum of ermined species of Crassomicrodus was Natural History, New York, New York tot present in the original assemblage of (AMNH); Academy of Natural Sciences, naterial the holotype was examined to en- Philadelphia, Pennsylvania (ANSP); Univ- ure it was not a misplaced species of Aga- ersidad Autonoma de Tamaulipas, hirsia. — Ciudad Victoria, Tamaulipas, Mexico Depositories. The following are the con- (ATAM); British Museum of Natural His- ributing museums. Acronyms are from tory, London, England (BMNH); Califor- \rnett et al. (1993) except for ATAM nia Academy of Sciences, San Francisco, vhich is not included in their list: Amer- California (CASC); Canadian National can Entomological Institute, Gainesville, Collection of Insects, Ottawa, Ontario Research Journal of Hymenoptera 68

well basal lobe, b, bicolor, with basal Fig. 3. Claws: a, Agathirsia trichiosoma, showing developed Agathirsia with basal lobe absent, Head of rostrata, foveae lobe vestigial, c, Crassomicrodus divisus, d, Agathirsia showing below antennal insertions.

(CNCI); Deutshes Entomologisches Insti- mological Collections, Oxford, England tute Eberswalde, Germany (DEIC); Essig (OXUM); Snow Entomological Museum, Kansas Museum of Entomology, University of University of Kansas, Lawrence, California Berkley, California (EMEC); Los (SEMC); Texas A&M University, Depart- Angeles County Museum, Los Angeles, ment of Entomology Insect Collection, California (LACM); Michigan State Uni- College Station, Texas (TAMU); Universi- of Entomolo- versity, Department of Entomology Col- ty of Arkansas, Department lection, East Lansing, Michigan (MSUC); gy Collection, Fayetteville, Arkansas of Oregon State University, Department of (UADE); Bohart Museum, University Entomology Collection, Corvallis, Oregon California Davis, California (UCDC); Uni- OSUO); Oxford University Hope Ento- versity of Michigan, Division of Insects, Volume 13, Number 1, 2004 69

Fig. 4. Mouthparts: a, Agathirsia proxima, illustrating Type II mouthparts with the glossa adapted for straw- like sucking, b, A. nigricauda, illustrating Type I mouthparts in which the glossa is folded and enclosed by the galea in the drinking process.

Museum of Zoology, Ann Arbor, Michi- is described as "Elongated glossa, galea gan (UMMZ); United States National Mu- and labial palps; glossa exposed for much seum of Natural History, Washington, of its length in dried specimens; labial D.C. (USNM). palps closely aligned with glossa and as- in nectar extraction /conduction". MOUTHPART MORPHOLOGY sisting Agatliirsia sp.l (probably A. longilingua Pucci and as an A recurring adaptation of parasitoid Sharkey n.sp.) was given wasps is the presence of elongate mouth- example of this category. Jervis also as- then of parts to facilitate nectar extraction from signed a, undescribed, species Aga- thirsia = A. to his VI which deep corollas (Jervis 1998). This is espe- ( jervisi) Type consists of with cially apparent in the Braconidae and wasps "Elongated glossa and labial inner surfaces of labial Aculeata. In Braconidae, elongate mouth- palps; parts are present in members of five of the palps concave" and covered with long and dense setae. 29 subfamilies (Jervis 1998). Elongate glos- the three of sae are common in Agatliirsia but there is We recognize same types nectar extraction in a great deal of interspecific variation. Jer- mechanisms Agathir- vis (1998) categorized the nectar feeding of sia. Species with Jervis' Type I mouthparts parasitoid wasps into six types and placed have glossae that vary in length from ap- species of Agathirsia into three of these. proximately 0.25 to 1.8 mm. The glossae His Type I consists of "Elongated glossa are covered with dense setae that appar- and galea, glossa shallowly bilobed/ ently are specialized for liquid uptake. forked and concealed by the galea for The morphology of these setae is variable. most of its length in dried specimens". Figure 8a shows the spoon-like setae of A. = Agathirsia nigricauda (Viereck) and A. cres- testacca (glossa length 0.48 mm) and fig- soni Muesebeck & Walkley are cited as ex- ure 8b shows the scale-like setae of A. ni- = amples. Another category (Jervis' Type IV) gricauda (glossa length 1.4 mm). After of Hymenoptera Research 70 Journal

Fig. 5. Agathirsia spp., thoracic sculpture: a, A. davidi, showing foveate (crenulate) notaulus and sternaulus. b, c, A. proximo., showing smoothly impressed notauli and mostly absent sternaulus. d, A. testacea, showing posterior lobes of mesopleura.

the glossa is loaded with liquid we sup- liquid towards the mouth opening. The pose it is retracted into the galea. Dried galea likely acts as a template forcing the specimens, in which the glossa are partly two halves of the glossa together as they retracted, show that the glossa folds pos- are retracted. As with all other drinking teriorly along its midlength as it is retract- types, the cibarial and pharyngeal pumps ed. This results in the posterior surfaces of provide the negative pressure necessary to each half of the glossa being pushed suck the liquid into the oral cavity. nst each other which would squeeze Jervis' Type IV mouthparts are manifest Volume 13, Number 1, 2004 71

A. Fig. 6. Agathirsia spp. a, b, Propodea. a, A davidi. b, cressoni. c, d, Abdominal terga. c, A trichiosoma, d, A davidi

in specimens in which the galea is signif- teriorly along most of its length, thereby of the icantly shorter than the long, exposed forming a tube. The outer surface scale-like se- glossa (Fig. 4a). All of these species pos- tubular glossa is covered by out- sess glossae that are more than 2 mm in tae (Fig. 8d). The presence of setae on length. Because it seems apparent that the er surface of the glossal tube has two pos- glossa cannot fold or otherwise be con- sible explanations. Jervis (1998) posited that tained within the galeae, nectar must trav- that these setae are hydrophylic and el from the apex to the base of the glossa the glossa is retracted into the galeae itself. The glossa is strongly curved pos- where the nectar is extracted. He suggest- Journal of Hymenoptera Research 72

short seta, Fig. 7. Agathirsia spp., antennal setae, a, A. asteropliila, showing long seta, b, A. testacea, showing c, A. sericans, showing seta of medium length, d, A. testacea, showing foveae around ocelli.

ed that the tube itself was used primarily glossa to be folded into shorter sections. to secrete salivary juices to dilute nectar in Therefore, contrary to Jervis (1998), we order both to pre-orally digest nectar and suggest that the scale-like, hydrophilic, se- to optimize its viscosity. We cannot imag- tae on the outer surface of the glossa direct ine how a glossa, that is frequently many liquid toward the postero-medial longitu- times the length of the galeae, could be dinal fissure where they are sucked into a retracted to such degree. There are no the lumen of the glossa and subsequently dried that show retrac- specimens partial towards the oral cavity. This hypothesis and there are no tion, apparent morpho- requires that the glossa be able to open such as transverse stri- logical structures, and close, at least part of its length, to al- ations the that 1 on glossa, would allow the liquid to enter. There are interesting Volume 13, Number 1, 2004 73 of Hymenoptera Research 74 Journal

petiole \2nd submarginal cell

A. terminal and scales, b, A, Fig. 9. Agathirsia spp., glossae. a, nigricauda, showing 1) long setae, 2) glossal A. proxima, showing tube-like morphology and specialized setae, c, Forewing of davidi

of the species. One of its maxillary palps has transport liquid along the length only 4 palpomeres instead of the usual 5; proboscis aided by negative pressure ap- th th the 4 and 5 are fused. The other palp plied by the pharyngeal and cibarial has 5 palpomeres, the last being extremely pumps. small. the labial are Incredibly, palpi very CLADISTIC SECTION long, 1.7 mm, and 5 segmented, instead of 4, which is ground-plan for the Hymenop- Methods.—The data matrix for the cla- tera. The last palpomere is very small and distic analysis is presented in Table 1. globose. Palpomeres 2-4 (and possibly 1) Characters 1 and 13-18 are treated as or- have the inner sides concave and hirsute. dered. Character 13 is meristic and char- The glossa is hirsute, 1 .2 mm, and sits be- acters 14-18 are continuous. Median val- tween the labial palpi. The galea is much ues were used when there was variation smaller than the glossa and we suspect in these characters. A heuristic search was that the setae on the glossa and inner sur- performed using PAUP* version 4.0M0 face of the labial palpi are hydrophilic and (Swofford 2002). Five hundred random Volume 13, Number 1, 2004 75

Table 1. Character matrix for species of Agathirsia and outgroups, including multiple species of Crassomkrodus. of Hymenoptera Research 76 Journal

hair-like or if are lost, additions were carried out, each followed if the pegs are they The con- Some of the "hairs" are thicker than oth- by TBR and unlimited maxtrees. us to that tinuous characters were all scaled between ers and this lead suggest they are modified but this is not a well 10 states (0-9) and could have a minimum pegs the contin- corroborated conclusion. It may be more of 9 steps. We down-weighted the character state uous characters so that the total weight of appropriate to describe as "without thickened each continuous character (99) was ap- in Crassomicrodus setae" but the makes proximately equivalent to the total weight peg-like terminology difference in the results of the of a discrete binary character (100). This no practical was accomplished by giving all continu- analysis. 1- with basal lobe ous characters a weight of 11 and all bi- 3. Foreclaw: (Fig. 3a), 2- without basal lobe nary characters a weight of 100. Trees (Fig. 3c). Very tiny basal lobes are coded were rooted with Pselaphanus trogoides (vestigial) (Fig. 3b) as uncertain. Szepligeti due to its purported and sup- 4. Two carinae on first ter- ported sister-group relationship with longitudinal 1- Agathidinae (van Achterberg 1990). Before gum extending past spiracles: present; of 2- absent. This character is variable within starting this revision we were uncertain of members of the monophyly of both Crassomicrodus and Agathis and the phylogeny will have to be resolved before Agathirsia because testable synapomor- Agathis can be confidence in our of phies had never been proposed for either there scoring genus. To test for the monophyly of Aga- this taxon. thirsia and Crassomicrodus and to strength- 5. Quotient of galea divided by glossa: 2- less than en our outgroup analysis we included 10 1- more than 0.55 (Fig. 4b); the Mor- species of Crassomicrodus as well as mem- 0.40 (Fig. 4a). See Mouthpart a dis- bers of the genera Agatiiis and Bassus. The phology section for more complete IV undescribed species of Bassus (Bassus sp.) cussion. State 2 represents Jervis' Type was included because it has flattened pegs mouthpart morphology described earlier, as do all species of Agathirsia and we were 6. Hind coxal cavity: 1- open; 2- closed, not sure to which genus it belonged. It 7. Relative position of lateral ocelli and was included to test the question of its hind margin of eyes: 1- tangent with pos- placement in either— Bassus or Agathirsia. terior margin of eyes; 2- anterad posterior Character List. margin of eyes. State 2 appears to be syn- 1. Mandible: 1- with well-developed apomorphy for members of Agathis. This second tooth (Figs, la, d); 2- with vestigial state is associated with the forward migra- second tooth (Fig. lb); 3- without second tion of the ocelli due to invagination of the tooth (Fig. lc). Twelve species of Agathirsia back of the head. The invagination allows are polymorphic for states 2 and 3. Further the head to rotate anterodorsally into a observations of rare species will no doubt prognathous position without the occiput taxa yield more that possess both states, hitting the pronotum. The prognathous The second author suspects that the sec- position facilitates probing into deep floral ond mandibular tooth may be present in nectaries. specimens of most species but that it is 8. Shape of the labrum: 1- oval; 2- cir- worn off in the process of digging through cular. The elongation of the face in mem- soil, bers of Agathis includes the elongation of 2. Apical pegs of the hind tibia: 1- flat- the labrum which is therefore rather more tened 2- rounded dis- distally (Figs. 2b, c); circular than those of typical agathidines, 3- hair-like tally (peg-like) (Fig. 2a); (Fig. which are significantly wider than long. 2d). Although we have described state 3 This character state is also found in (the 5 "hair-like" we do not know pegs really appropriately named) Agathirsia rostrata, /olume 13, Number 1, 2004 77 is well as most members of the Cremnop- consensus tree of 32 minimum length trees ini. The state is also scattered throughout (branches with a minimum length of zero he Microdini and Disophryini. collapsed), each with a weighted length of 9. Mandible: 1- dorsoventrally flattened 4266 steps. The minimum length trees values: in- Fig. la); 2- thick and relatively cylindrical have the following consistency Figs. lb-d). See Biology section for more dex 0.47, retention index 0.82, and re- nformation on mandible morphology. scaled consistency index 0.38. The char- 10. Medial setae on tergum 3: 1- at most acters mapped onto the consensus tree are i fringe distally; 2- present distally as a those that are unequivocal over all pri- that are vide, complete band (Figs. 6c, d). As men- mary trees, with two exceptions ioned previously, the extensive setosity discussed below. In the following discus- in bees. sion the numbers in brackets re- nay be effective mimicking square [ ] 11. Second submarginal cell: 1- quad- fer to character states. ate; 2- triangular (Fig. 9c). A quadrate cell The monophyly of Agathirsia is sup- s found in the outgroups Pselaphanus and ported by one unequivocal autapomor- zarinus. The character was included to phy, i.e., the reduction of the second man- idd some resolution to the outgroup anal- dibular tooth (Figs. 1 b, c) (character 1[2]). ysis. Most species lack a second mandibular still a 12. Rs + M vein: 1- complete; 2- not tooth (Fig. lc), although some have of the inner zomplete (Fig. 9c). A complete Rs + M slight indication (ventral) a small /ein is found in the outgroups Pselaphanus tooth (Fig. lb). The presence of md Earinus. The character was included ventral tooth appears as a reoccurrence of :o add some resolution to the outgroup the tooth from an ancestral species of Aga- malysis. thirsia in which the tooth was absent. The The following six characters are meristic presence of flattened pegs on the apico- Dr continuous and the absolute values are medial surface of the hind tibia (Figs. 2b, scaled between and 9. c) (character 2[1]) is an autapomorphy im- 13. Number of distally flattened pegs plied by DELTRAN optimization only. 3n hind tibia. The number of pegs varies The flattened pegs of the hind tibia are between 0-21. found convergently in the undescribed 14. Quotient of basal width of tergum 1 species of Bassus included in the analysis 1. divided by length of tergum Quotients (Bassus sip.). are between 0.40-1.13. The sister-group relationship of Cras- 15. Quotient of glossa length divided somicrodus + Agathirsia is supported by the man- oy foretibia length. Quotients are between the robust cylindrical shape of 120-3.10. Because there are only a few dibles (character 9[2]), which appears to in the soil. species with exceptionally long tongues, be a modification for digging most species clump between states and The results of the cladistic analysis sug- I. gest two synapomorphies for members of 16. Quotient of ovipositor length divid- Crassomicrodus. The first, which is implied ed by foretibia length. Quotients are be- exclusively by DELTRAN optimization, is tween 0.54-6.30. that the apical pegs of the hind tibia are unlike 17. Quotient of malar space length di- hair-like (Fig. 2d) (character 2[3]), in vided by eye height. Quotients are be- the standard agathidine pegs illustrated tween 0.22-0.59. figure 2a and unlike the flattened pegs of 18. Quotient of length of labial palpom- members of Agathirsia (Figs. 2b, c). A ere 2 divided by length of labial palpom- weakness with this proposed autapomor- ere 4. Quotients are between 0.9-2.5. phy is that it rests on the assumption that Results. —Figure 10a shows the strict the flattened pegs of Agathirsia and the 78 Journal of Hymenoptera Research

Earinus limitaris Pselaphanus trogoides Agathirsia testacea A. collini A. fulvocastanea A. heleni

A. kellyi A. jervisi A. minuata tiro bifidilingua campanisura michelei armandi A. reai A. ninesevensi A. parkeningi A. rostrata A. davidi keni sericans foveiseries asterophila A. trichosoma A. capillata A. papoui A. rufula A. longilingua longigladia proxima bicolor cressoni A. nigricauda Crassomicrodus nigriceps C. sp. C. medius C. muesebecki C. apicipennis nigrithorax fenestratus fulvescens divisus pallens Agathis malvacearum arida semiaciculata Bassus sp. B. brooksi b Earinus Pselaphanus Agathirsia Crassomicrodus Agathis Bassus

Agathirsia spp a Strict consensus tree of and species outgroups all characters, b, Strict consensus tree employing using only discrete, non-meristic, characters. Volume 13, Number 1, 2004 79 hair-like spines of Crassomicrodus evolved tent then the randomized characters that independently from a peg-like ancestor. It replace them should add significantly to is equally parsimonious to suggest that the length of the tree. Characters 13-18 the flat pegs were derived from the hair- were randomized with "true random like spines or vice versa. The second pro- numbers" a program found at posed autapomorphy for Crassomicrodus is www.random.org. Rather than being re- the loss of lobes at the base of the tarsal placed with completely random numbers, :laws (character 3[2]) (Fig. 3c). The ple- the states were rearranged to maintain the omorphic condition, with a well-defined character state frequency, for example, if :>asal lobe, is shown in figure 3a. The loss there were 3 cells with a score of 7 in the ji this lobe is also found in some members original matrix the random matrix would Df the Earinini, and sporadically in other also contain 3 cells scored as 7. One hun- agathidines including some species of dred randomized datasets were generated Agatliirsia. and analyzed in Paup using TBR, unlim- Autapomorphies for AgatJiis are the an- ited Maxtrees, 50 random addition-se- terior placement of ocelli (character 7[2j) quence replicates with a time limit of 120 and shape of the labrum (character 8[2]). seconds on each replicate on a Pentium 4 rhe closed hind coxal cavities appear as 1.8GHz processor. Taken as a whole, the an autapomorphy for Bassus (character characters are 43% longer when random- 5[2]). This result is a product of the abbre- ized than they are in the non-randomized viated nature of this analysis. The closed results and they make the resulting mini- :oxal cavities are far more widespread in mum length trees an average of 21% lon- the Agathidinae, occurring in most Micro- ger. iini, Cremnoptini and Disophryini. At Within Agatliirsia several major clades present there is no evidence for the mono- supported by the continuous characters ohyly of Bassus though the two included are worth discussing. A. testacea is the sis- species are part of a much larger mono- ter-taxon of the remainder of the species shyletic group. of Agatliirsia. A. testacea is particularly in- Figure 10b illustrates the strict consen- teresting because it is the sole species with sus tree generated when the meristic and a large sclerite between the foramina of continuous characters (characters 13-18) the metasoma and the hind coxa (charac- are excluded. In this simplified tree all po- ter 6[2]). This putatively derived state is lytomies have been collapsed. The discrete found convergently in the vast majority of :haracters resolve the genera but do noth- Bassus, as well as most Microdini, Diso- ing to resolve the relationships within phryini, and Cremnoptini. The monophy- Agatliirsia. All of the resolution within ly of the remainder of the species is sup- Agatliirsia illustrated in figure 10a is the ported by four continuous characters. result of the addition of the meristic and Moving up the tree there is a large poly- continuous characters. Due to the limited tomy of seven species and one large number of discrete characters, and to the monophyletic group united by the com- nature of ordered multistate characters, plete loss of the second mandibular tooth characters 13-18 would add much reso- and three continuous characters. Further lution even if they were totally random. up the tree there is a polytomy composed To test whether or not the meristic and of four lineages. One of these clades con- continuous characters have information tains all of the species with long glossae content we randomized the meristic and (Jervis Type IV). Because the indepen- continuous characters while holding the dence of some of the characters is suspect, character state frequencies constant. If the monophyly of the group and the characters 13 to 18 have information con- unique derivation of the elongate glossa Journal of Hymenoptera Research 80 '

other not consid- be distinguished from all that this topology suggests are may observa- the following combination ered well supported, though the Agathidinae by complex tion that other characters do not imply of characters: Labio-maxillary longer than other groupings indicates some support, usually (95%) elongate (galea labial What is interesting, and better supported, mandible) (Figs, la, 4a, b); palpom- A. rela- more than 0.5 of is the distant placement of jervisi ere 3 no t reduced, length some confidence 4 tarsal claws not tive to this clade, giving pa ip0mere (Figs. 4a, b); IV , 1- to the conclusion that the Jervis Type i without two teeth; c e^ e ^ sharp VI are inde- and Jervis Type mouth-parts Rs+M vem f forewing incomplete (Fig derived from type I mouth- carina of pendently ^ posterior transverse propo- parts. deum absent; hind coxal cavity usually

DESCRIPTIONS AND KEYS o/o not from metasomal fo- ( 80 ) separated nar- Agathidini Sharkey 1992 ramen, sometimes (20%) separated by row sclerite. Diagnosis—Members of the Agathidini

WORLD KEY TO THE GENERA OF AGATHIDINI + BASSUS

The Bassus to the tribe Notes: The limits of the Agathidini are based on Sharkey (1992). genus belongs well defined and can sometimes be confused with members of Microdini (Sharkey 1992) but the genus is not a are included in this If no characters in side of couplet the Agathidini; it is therefore key. any particular characters. definitive then the reader should choose that alternative with the most applicable

hind coxae 1. Wide transverse carina, i.e. more that % as wide as long, between usually wide sclerite, (90%) present; hind coxa and metasoma usually separated by rarely shorter than mandible; with a very narrow sclerite or none at all; galea usually (90%) third labial palpomere usually (90%) less than half the length of the fourth; propo- Fabricius deum often (70%) areolated Bassus Wide transverse carina between hind coxae rarely (1%) present, sclerite usually absent

i.e. less that as wide as (90%) or very narrow, Y8 long, (9%); galea usually (99%) longer than mandible; third labial palpomere more than half the length of fourth (95%); propodeum not areolated 2 (Agathidini) tibial RS of 2 (1) Mandible dorso-ventrally flattened (Fig. la); hind spines peg-like (Fig. 2a); not forewing usually (90%) straight; second submarginal cell usually (90%) petiolate; head excavated posteromedially and ocelli shifted anterad such that line drawn over as lateral ocelli meeting the compound eyes; labrum usually (90%) almost as high wide Agathis Latreille flat- Mandible not flattened as above (Figs, lb-d); hind tibial spines either somewhat tened or hair-like (Figs. 2b-d); RS of forewing usually (90%) sinuate (Fig. 9c); second submarginal cell usually (90%) petiolate (Fig. 9c); head not excavated posteromedially and ocelli not shifted anterad such that a line drawn over lateral ocelli is approximately tangent with posterior margin of compound eyes; labrum usually (97%) 3 distinctly wider than high 3 (2) Glossa usually (90%) at least as long as mandible; hind tibial pegs somewhat flattened (Figs. 2b,c); claws with distinct to vestigial basal lobe (Figs. 3a, b) Agathirsia Westwood Glossa shorter than mandible; hind tibial pegs hair-like (Fig. 2d); claws with basal lobe absent (Fig. 3c) or sometimes (20%) vestigial Crassomicrodus Ashmead Volume 13, Number 1, 2004 81

Agathirsia Westwood TRANSFER OF CRASSOMICRODUS PUMILUS TO BASSUS PUMILUS Agathirsia Westwood 1882: 20. Type species Westwood, Agathirsia rufula designated by One of the species of Crassomicrodus that Viereck, 1914: 6. [Examined]. we investigated is clearly misplaced and Agathona Westwood 1882: 22. Type species Aga- is here transferred to Bassus. The fact that thona sericans Westwood, monobasic. Syn. by it is from South Africa was the first clue Szepligeti 1904: 128. [Examined]. that it was to Paragathis Ashmead 1889(1888): 638. Type spe- suggesting misassigned = Crassomicrodus. in the cies Microdus thoracicus [ cressoni] Cresson, Originally placed monobasic. Syn. by Ashmead 1900: 128. [Ex- genus Epimicrodus by Szepligeti (1913), it amined]. was transferred to Crassomicrodus when Cenostomus Cameron 1905: 387. Type species, Brues (1924) synonymized Epimicrodus un- Cenostomus trichiosomus mono- Cameron, by der Crassomicrodus. The name Bassus pum- typy. Syn. n. [Examined]. ilus is available. The specific name is oc- in Description. —Head: Gena lacking flange cupied Agathis, i.e., Agathis pumila and rationale for its posteroventrally, posterolateral corner (Ratzeburg), placement in to are rounded to acute; mandible robust and cy- Agathis (as opposed Bassus) discussed lindrical in cross-section (Figs. lb,c); sec- by Sharkey (1985, 1992). ond tooth of mandible much reduced or absent (Figs. lb,c); third (penultimate) la- Bassus pumilus (Szepligeti), n. comb. bial palpomere more than half as long as 1913: 385; last (distal) palpomere (Fig. 4a,b); frons Epimicrodus pumilus Szepligeti not bordered with carinae; face not elon- (DEIC). [Examined]. Crassomicrodus Brues 1924: 144. gated into rostrum except for A. rostrata. pumilus Mesosoma: Notauli always impressed, fo- The following description is not meant veolate to smooth (Figs. 5a,b); propodeum to be exhaustive but rather to support the not evenly areolated, usually rugose to placement of the species in Bassus. varying degrees (Figs. 6a,b); tarsal claws Penultimate labial (number usually with basal lobe but sometimes palpomere 3) half the length of number 2; mandible lobe vestigial (Figs. 3a,b); hind tibia with narrow and scissor-like; not dor- small flattened pegs distally (Figs. 2b,c); occiput excavated as in second cubital cell of the forewing trian- sally Agathis; propodeum declivous, with gular and usually petiolate (Fig. 9c); wings sharply strong posterior transverse carinae; hind coxal transparent to infumate (Fig. 9c); hind cox- cavity 1 metasomal al cavities open, sharing common opening closed; tergum striate; terga hind tibial claws with with the metasoma except for A. testacea. wide; pegs peg-like; basal lobes and Metasoma: First median tergite smooth, present sharply angled; in Bassus con- lacking sculpture, or pair of well-devel- forewing 3Rs decurved as hind 2Cub tu- oped longitudinal carinae (Figs. 6c,d); ovi- spicuus (Wesmael); wing positor (when fully exerted) from 0.1 to bular; ovipositor approximately 0.8X 1.3X as long as the body. body length.

KEY TO FEMALES AND MALES OF AGATHIRSIA

Note: An interactive version of this key with color illustrations is available at www.uky.edu/~mjshar0, it is much simpler to use primarily due to its use of color characters, nonetheless the following key is effective in distinguishing species. 1 Labial palpi elongate, concave, and encasing glossa A. jervisi 2 Labial palpi not as above, not encasing glossa (as in Fig. 4b) of Hymenoptera Research 82 Journal

3 than in 4a) 2 (1) Glossa more than 2x longer galea (as Fig. • 6 Glossa less than 2x longer than galea (as in Fig. 4b) black; entirely black; notauli 3 (2) Hind femur black; metasomal terga entirely propodeum foveate (as in Fig. 5a) or en- Hind femur orange; metasomal terga not entirely black; propodeum partially 5 foveae in 5b) tirely orange; notauli lacking (as Fig. flattened on of hind tibia (as in Figs. 2b,c) six or 4 (3) Number of thick, apically pegs apex in more less; tarsal claws with vestigial basal lobes (as Fig. 3b); ovipositor length

A - than 6X hind basitarsus length longigladia of hind tibia in 2b,c) more Number of thick, apically flattened pegs on apex (as Figs. in less than six; tarsal claws with distinct basal lobes (as Fig. 3a); ovipositor length A. than 6X hind basitarsus length longilingua for at least one third of (as in Fig. 5 (3) Foveae of sternaulus present length mesopleuron A. nifula 5a); glossa length less than 3mm Foveae of sternaulus restricted to extreme posterior of mesopleuron (Fig. 5c); glossa

A - length at least 3mm proxima A. testacea 6 (2) Pair of lobes between midcoxae (Fig. 5d) 7 Without pair of lobes between midcoxae of characters: first black color 7 (6) With the following combination tergum lacking basally; flattened hind tibial hind tibia lacking black color; at least 9 thick, apically pegs 8 (as in Fig. 2b) 9 Not exactly as above black for distal one third; 8 (7) Dorsal metasoma usually orange basally and completely x A. hind femur orange; glossa length 0.68-0.95 foretibia length nigricauda dorsal metasoma with transverse Tergum 1 black distally; remaining usually orange x black bars; hind femur usually (-90%) black; glossa length 0.60-0.69 foretibia A. cressotti length 10 9 (7) Hind femur entirely to mostly black 20 Hind femur entirely to mostly orange 11 10 (9) Metasomal terga entirely black 15 Metasomal terga with at least some orange or yellow A. rostrata 11 (10) Foveae present between tentorial pit and antennal insertion (Fig. 3d) 12 Foveae absent between tentorial pit and antennal insertion 12 (11) Setae on propodeum present along midline; ovipositor length greater than 3X hind basitarsus length A. parkeningi Setae on propodeum absent along midline (as in Figs. 6a,b); ovipositor length less than 3X hind basitarsus length 13 13 (12) Setae on tergum 3 dense distally (as in Fig. 6c); ovipositor shorter than length of hind basitarsus A. capillata Setae on tergum 3 not dense distally (as in Fig. 6d); ovipositor longer than hind basitarsus length 14 14 (13) Propodeum with pronounced rugosity near midline (as in Fig. 6b); distal half of hind tibia entirely black A. keni

Propodeum without pronounced rugosity near midline (Fig. 6a); distal half of hind tibia not entirely black A. davidi 15 (10) Mesoscutum entirely black 16 Mesoscutum partially to entirely orange 19 16 (15) Setae on propodeum present along midline 17 Setae on propodeum absent along midline (as in Figs. 6a,b) 18 17 Glossa X (16) length greater than 0.75 foretibia length; basal lobe of claw not large (smaller than Fig. 3a) A. sericans Glossa less X than 0.75 foretibia . length length; basal lobe of claw large (Fig. 3a) ... A. trichiosoma Volume 13, Number 1, 2004 83

18 (16) Setae on tergum 3 dense (as in Fig. 6c) and present on at least distal 0.75 ... A. sericans Setae on tergum 3 not dense and present on less than distal 0.75 (as in Fig. 6) A. asterophila 19 of for at (15) Foveae sternaulus present least one third length of mesopleuron (as in Fig. 5a); ovipositor length equal or shorter than length of hind basitarsus; setae on tergum 3 dense (as in Fig. 6c) and present on at least distal 0.75 A. sericans of Foveae sternaulus restricted to extreme posterior of mesopleuron (as in Fig. 5c); ovipositor more than 2x length of hind basitarsus; setae on tergum 3 not dense and present on less than distal 0.75 (as in Fig. 6d) A. bifidilingua 20 (9) Mesoscutum entirely black 21 Mesoscutum partially to entirely orange or brownish orange 28 21 (20) Metasomal terga entirely black 22 - Metasomal terga partially to entirely orange or brownish orange 25 22 Labial 2 at least (21) palpomere equal to combined length of palpomeres 3 + 4 (as in Fig. 4b) 23 - Labial palpomere 2 shorter than combined length of palpomeres 3 + 4 24 23 (22) Glossal length more than 0.75 X foretibia length; ovipositor length more than 3x hind basitarsus length A. michelei Glossal length less than 0.75 X foretibia length; ovipositor length less than 2x hind basitarsus length A. paponi 24 (22) Notauli lacking foveae (as in Fig. 5b); longest setae at midlength of antenna approximately equal to antennal width (as in Fig. 7a) A. kellyi Notauli foveate (as in Fig. 5a); longest setae at midlength of antenna less than 0.5 X antennal width (as in Fig. 7b) A. collini 25 (21) Labial palpomere 2 shorter than combined length of palpomeres 3 + 4 26 Labial palpomere 2 approximately equal to combined length of palpomeres 3 + 4 (as in Fig. 4b) A. michelei 26 (25) Claws with basal lobe distinct (as in Fig. 3a); propodeum largely sculptured and setae present except along midline (as in Figs. 6a,b); ovipositor distinctly longer than mesosoma A. campauisura Claws with basal lobe vestigial (as in Fig. 3b); propodeum largely smooth and glabrous medially; ovipositor shorter than mesosoma 27 27 (26) Sternaulus with foveae restricted to extreme posterior of mesopleuron (as in Fig. 5c); glossal length more than 0.5 X foretibia length; longest setae at midlength of antenna less than 0.5 X antenna width (as in Fig. 7b) A. ninesevensi Sternaulus completely foveate for entire length of mesosoma; glossal length less than 0.5 X foretibia length; longest setae at midlength of antenna approximately equal to antenna width or longer (as in Fig. 7a) A. tiro 28 (20) Area between tentorial pit and antennal insertion with line of foveae (as in Fig. 3d) A. foveiseries - Area between tentorial pit and antennal insertion without line of foveae 29 29 (28) Propodeum entirely black 30 Propodeum partially or entirely orange 32 30 (29) Propodeum smooth along midline; tarsal claws with basal lobes distinct (as in Fig. 3a) A. heleni - Propodeum sculptured along midline (as in Fig. 6a); claw with basal lobe variable ... 30 31 (30) Labial palpomere 2 approximately equal to combined length of palpomeres 3 + 4 (as in Fig. 4b) A. armandi Labial palpomere 2 shorter than combined length of palpomeres 3 + 4 A. bicolor 32 (28) Head entirely orange; glossal length shorter than 0.5 X foretibia length; ovipositor less than 1.5x hind basitarsus length A. minuata Face and frons at least partially black; glossal length more than 0.5 X foretibia length; ovipositor longer than 1.5X hind basitarsus length 33 Journal of Hymenoptera Research 84

first orange 33 (32) Forecoxa and midcoxa orange; tergum A. fulvocastaneafufrncastanea Forecoxa and midcoxa black; first tergum black basally

widths in 9c). Metaso- armandi Pucci and Sharkey, than 2 vein (as Fig Agathirsia5 0.76- n ma: Posterior width of first tergum setae on ter- 82 X length of first tergum; Distribution Known only ii ter- (Fig. llj).— 3 gparse dista y (as m Fig. 6d); Mexico. gum from the type locality in Puebla, m 3 with transverse groove distinct or Males.—Color: and black, orange Orange vestigial antenna maxil- except as follows: black; Female—Unknown, and labial partly black; o\ Mex- lary palpomeres Specimens examined.—Holotype: head except clypeus black; pronotum ueD 3 mi. N .^' p la, Petalcingo, black; mesos- black ventrally; propleuron viii 2 1.1963 (Parker & Stange) (USNM). black cutum sometimes slightly anteriorly; i j same data as paratype: r holotype black mesopleuron mostly except slightly (tjsnm) sometimes orange anteriorly; metanotum Remarks.—Similar to A. heleni and A. bi- black; with black laterally; metapleuron cdor_see characters in the key and re- black; hind tarsus sometimes propodeum markg under thoge species< dark first black; remaining au- orange; tergum Etymology.—In honor of the senior dark with black mottling; ; terga orange thor g father ex- wings slightly infumate. Body length: 6.6-7.0. Head: Longest cluding ovipositor Agathirsia asterophila Pucci and seta at of antenna approximate- midlength Sharkey, sp. n. less than half antenna ly half or slightly Distribution Southwestern width (as in Figs. 7b,c); labial palpomere (Fig. lib).— + in Mexico. 2 subequal to palpomeres 3 4 (as Fig. Females and males.—Color: black 4b); mandible without indication of sec- Mostly as follows: and labial ond tooth (as in Fig. lc); glossa fork length except maxillary pal- fore and midfemur 0.33-0.40; glossa length less than 2x galea pomeres partly pale; with small area foretibia length; glossa length 0.83-0.90; glossa orange distaiiy; or length 0.64-0.83 X shorter than foretibia sometimes entirely partly pale yellow/ X shorter foretarsus sometimes or length; malar space 0.41-0.42 orange; entirely midtibia than eye height; face above clypeus with partly pale yellow; partly pale hind distinct or vestigial longitudinal carinae; yellow; midtarsus sometimes pale; fur- area anterior to ocellus with pits forming tibia pale basally, pale color extending hind tarsus sometimes V-shape (as in Fig. 7d); area between ten- ther laterally; part- first and second torial pit and antennal insertion without ly or entirely pale yellow; line of foveae. Mesosoma: Notauli foveate terga both yellow basally; wings slightly and complete (as in Fig. 5a); propodeum infumate. Body length: excluding oviposi- mid- mildly rugose, setae present except along tor 8.2-10.3. Head: Longest seta at midline and posterior central area (as in length of antenna subequal to antenna 2 Fig. 9a,b); sternaulus foveate (as in Fig. width (as in Fig. 7c); labial palpomere 5a); mesepisternum without lobes be- subequal to palpomeres 3 + 4 (as in Fig. tween midcoxae; basal lobe of claws small 4b); mandible with or without indication

(cf. Figs. 3a,b); number of thick, apically of second tooth (as in Fig. lb,c); glossa flattened apical pegs on hind tibia 0-4; fork length 0.13-0.17; glossa length less 0.57- hind basitarsus straight; 1-cu-a and 1-M of than 2X galea length; glossa length irewing separated by a distance greater 0.61; glossa length 0.38-0.42 X shorter than olume 13, Number I, 2004 85

X — oretibia length; malar space 0.23-0.29 Remarks.—Similar to A. sericans and A. horter than eye height; face above clype- trichiosoma —see characters in—the key. is with or without median longitudinal Etymology. Lover of aster referring to arinae; area anterior to ocellus with pits the Asteraceae pollen commonly attached to Pollen identified Gretch- orming V-shape (as in Fig. 7d); area be- specimens. by en ween tentorial pit and antennal insertion Jones USDA, ARS, APMRU, College without line of foveae. Mesosoma: Notauli Station, TX. oveate and complete (as in Fig. 5a); pro- Agathirsia bicolor Pucci and Sharkey, •odeum rugose posteriorly, foveate ante- sp. n. iorly, setae present except along midline Distribution —Known from nd posterior central area (as in Fig. 9a,b); (Fig. llh). two localities in Chihuahua, Mexico. ternaulus foveate (as in Fig. 5a); mesepis- — ernum without lobes between midcoxae; Females and males. Color: Black and or- variable, black as follows: asal lobe of claws distinct (as in Fig. 3a); ange; except and labial lumber of thick, apically flattened apical maxillary palpomeres mostly sometimes ven- »egs on hind tibia 3-8; hind basitarsus ta- orange; clypeus orange trally; entirely to slight- >ered distally; 1-cu-a and 1-M of forewing pronotum orange black sometimes eparated by a distance greater than 2 vein ly ventrally; propleuron black and orange; mesoscutum and scu- widths (as in Fig. 9c). Metasoma: Posterior tellum black and or- /idth of first tergum 0.40-0.50 X shorter orange; mesopleuron ange; metanotum partly to entirely or- tian length of first tergum; setae on ter- ange; propodeum black or black and or- ;um 3 not dense (as in Fig. 6d), present ange; metapleuron black or black and or- n distal one third; tergum 3 with trans- ange; legs orange except tarsi orange to erse groove distinct or vestigial; ovipos- pale yellow; forewing clear in basal half, :or length 1.0-1.2; ovipositor length 0.67- infumate in distal half; first tergum orange .92 X shorter than hind basitarsus length. distally; remaining terga orange, some- Specimens examined. —Holotype: 9, Mex- times with dark orange mottling; hind :o, Morelos, 3.8 mi. W Yautapec, 3800', wing clear to slightly infumate in basal iii.17.1962 (Ordway) (SEMC). Paratypes: half and distinctly infumate in distal half. /lexico: Guerrero: 16, 10.3 mi. S. Iguala, Body length: excluding ovipositor 7.7-8.5. ii.23.1981 (Bogar, Schaffner, & Friedlan- Head: Longest seta at midlength of anten- ier) (TAMU); Jalisco: 16, 16 km N Autlan, na approximately half or < half antenna ii.31.1978 (Plitt & Schaffner) (TAMU); width (as in Figs. llb,c); labial palpomere 6, 16 km N Autlan, vii.31-viii.2.1978 2 shorter than combined length of 3 + 4; Plitt 28 mi. E & Schaffner) (TAMU); 19, mandible with indication of second tooth Guadalajara, viii.15.1962 (CNCI); Morelos: vestigial (as in Fig. lb) or absent (as in Fig. 6, Cuernovaca [Cuernavaca], viii.15.1954 lc); glossa fork length 0.32; glossa length Dreisbach) 16, as (MSUC); previous less than 2X galea length; glossa length 16, 12 mi. E Cuernavaca, elev. AEIC); 0.74; glossa length 0.53 X shorter than for- viii.12.1954 200', (Dreisbach) (AEIC); 19, etibia length; malar space 0.40-0.43X 2 mi. E elev. Cuernavaca, 4300', shorter than eye height; face above clype- 'hi. 14.1954 Kans. Mex. (Univ. Expedition) us without or with vestigial longitudinal 11 mi. S SEMC); 19, Tlaltizapan, carinae; area anterior to ocellus with pits •hi. 16. 1962 (Roberts & Marston) (SEMC); forming V-shape (as in Fig. 7d); area be- as '9, 16, same data holotype (SEMC); tween tentorial pit and antennal insertion 9, 2mi. W. Zacatepec, vii.28.1967 (R.H. & without line of foveae. Mesosoma: Notauli LM. Painter) (AEIC); Puebla: 19, Chietla, foveate and complete (as in Fig. 5a); pro- iii.13.1991 (Pena) (ATAM). podeum mildly rugose, setae present ex- of Hymenoptera Research 86 Journal

Monlh Month

distributions of selected Fig. 11. Agathirsia spp., and phenologies species.

in cept along midline and posterior central distance greater than 2 vein widths (as of first area (as in Fig. 9a,b); sternaulus foveate Fig. 9c). Metasoma: Posterior width of (as in Fig. 5a); mesepisternum without tergum 0.75-0.91 X shorter than length lobes between midcoxae; basal lobe of first tergum; setae on tergum 3 sparse dis- trans- claws vestigial (Fig. 3b); number of thick, tally (as in Fig. 6d); tergum 3 with apically flattened apical pegs on hind tibia verse groove absent or vestigial; oviposi- 1.8- 11-15; hind basitarsus tapered distally; 1- tor length 2.1-2.3; ovipositor length cu-a and 1-M of forewing separated by a 1.9X longer than hind basitarsus length. Volume 13, Number 1, 2004 87

111 IV V VI VII Vlll IX X XI Month

Fig. 11. Continued.

examined. — 9 Mex- number of measured. basal Specimens Holotype: , specimens The ico, Chihuahua, 16 km N Cd. Jimenez, lobe of the claw is vestigial (Fig. 3b) unlike viii.26.1991 (Griswold) (CNCI). Paratypes: the other two species. 19, 16, same data as holotype (CNCI); Etymology. —Two colors—referring to 1$, same data as holotype except the orange and black coloration. viii.27.1991 (CNCI). Pucci and Remarks.—Similar to A. heleni and A. ar- Agathirsia bifidilingua — Sharkey, sp. n. mandi see characters in the key and re- — marks under those species. A. bicolor is Distribution (Fig. 11a). Known only longer than A. heleni and A. armandi but from type locality —in Arizona. overlap is possible considering the limited Holotype female. Color: Mostly black 88 Journal of Hymenoptera Research

in is much earlier than most with some orange; orange as follows: date April of pronotum dorsally; mesoscutum; scutel- species Agathirsia. —Forked — lum partly orange; mesopleuron with or- Etymology. tongue referring ange area under sternaulus; fore and mid- to glossa shape. legs with some orange; forewing slightly , . . . _ . , Agathirsia Pucci and infumate in basal half, infumate in distal campantsura^ar eY' P" half; hind wing clear in basal half, infu- Distribution. —Known mate in distal half. Body length: excluding (Fig. 111). only in Mexico, ovipositor 9.4. Head: Longest seta at mid- from type locality Morelos, —Color: Black and or- length of antenna approximately half an- Holotype female. Black as follows: max- tenna width (as in Fig. 7c); labial palpom- ange. except orange or- ere 2 subequal to palpomeres 3 + 4 (as in illary and labial palpomeres mostly Fig. 4b); mandible without indication of ange; clypeus partially orange ventrally; or- second tooth (as in Fig. lc); glossa fork forecoxa and foretrochanter partially length 0.50; glossa length less than 2X ga- ange; remainder of foreleg orange; midleg lea length; glossa length 1.0; glossa length orange; hind leg orange except hind tarsus 0.77 X shorter than foretibia length; malar black and orange; terga orange except space 0.54 X shorter than eye height; face black mottling; wings slightly infumate. above clypeus with vestigial median Ion- Body length: excluding ovipositor 7.3. Head: gitudinal carinae; area anterior to ocellus Longest seta at midlength of antenna < without pits forming V-shape; area behalf antenna width (as in Fig. 7b); labial tween tentorial pit and antennal insertion palpomere 2 shorter than combined length without line of foveae. Mesosoma: Notauli of 3 + 4; mandible with indication of sec-

very weak and absent anteriorly; propo- ond tooth (as in Fig. lb); glossa fork length deum mildly rugose, midline mostly 0.30; glossa length less than 2X galea smooth, setae present except along midlength; glossa length 1.0; glossa length line (similar to Fig. 6a); sternaulus slightly 0.77X shorter than foretibia length; malar with impressed only a few foveae poste- space 0.46 X shorter than eye height; face in riorly (as Fig. 5c); mesepisternum with- above clypeus without median longitudi- out lobes between midcoxae; basal lobe of nal carinae; area anterior to ocellus with- claws in vestigial (as Fig. 3b); number of out pits forming V-shape; area between flattened thick, apically apical pegs on tentorial pit and antennal insertion with- hind tibia 7-12; hind basitarsus tapered out line of foveae. Mesosoma: Notauli fo- 1-cu-a and 1-M in distally; forewing par- veate anteriorly, barely impressed as or Metasoma: tially nearly overlapping. smooth groove posteriorly; propodeum Posterior width of first 0.80 X tergum foveate, rugose posteriorly to spiracles, shorter than of first length tergum; setae shallow medial longitudinal furrow par- on 3 in tergum sparse distally (as Fig. 6d); tially smooth, setae present except along 3 without transverse tergum groove; ovi- midline and posterior central area (as in 4.3; 3.9 X positor length ovipositor length Figs. 9a,b); sternaulus foveate (as in Fig. than hind longer — basitarsus length. 5a); mesepisternum without lobes be- Males. Unknown. tween midcoxae; basal lobe of claws dis- examined.— Specimens Holotype: 9, tinct (as in Fig. 3a); number of thick, api- USA, Arizona, Pima 12 mi. E Co., Tucson, cally flattened apical pegs on hind tibia 5- iv. 13.1957 hind (USNM). 8; basitarsus tapered distally; 1-cu-a Remarks.—The combination of an or- and 1-M of forewing separated by a dis- mesoscutum and black is tance ange legs greater than 2 vein widths (as in Fig. to this enough distinguish species from all 9c). Metasoma: Posterior width of first ter- other known The collection X Agathirsia. gum 0.78 shorter than length of first ter- Volume 13, Number 1, 2004 89

gum; setae on tergum 3 sparse distally (as without line of foveae. Mesosoma: Notauli in 3 with Fig. 6d); tergum transverse foveate and complete (as in Fig. 5a) or ab- groove vestigial; ovipositor length 4.3; ovi- sent posteriorly; propodeum mildly ru- 4.6 X positor length longer than hind bas- gose posteriorly mildly foveate anteriorly, itarsus length.— shallow furrow along midline mildly Males. Unknown. sculptured, setae present except along examined.— 9 midline and central area in Specimens Holotype: , Mex- posterior (as ico, Morelos, Yautepec, vii.31.1963 (Parker Figs. 9a,b); sternaulus foveate (as in Fig. & Stange) (USNM). 5a); mesepisternum without lobes be- Remarks. —Similar to A. ninesevensi—see tween midcoxae; basal lobe of claws dis- characters in the key. These two species tinct (as in Fig. 3a); number of thick, api- also differ in the distance 1-cu-a and 1-M cally flattened apical pegs on hind tibia 7- hind are from each other, however, intraspecific 11; basitarsus tapered distally; 1-cu- variation, when it is better understood, a and 1-M of forewing separated by a distance than 2 in may make this character of limited value. greater vein widths (as Fig. — Metasoma: Posterior Etymology. Bell calf—referring to the 9c). width of first ter- dilated hind tibia and narrow basitarsus. gum 0.61-0.74 X shorter than length of first tergum; setae on tergum 3 long and capillata Pucci and Agathirsia Sharkey, dense on distal half (as in Fig. 6c); tergum n. sp. 3 with transverse groove; ovipositor — 0.68- Distribution (Fig. 11m). South-central length 0.90-0.98; ovipositor length Mexico. 0.89 X shorter than hind basitarsus length. examined.— 9 Mex- Females ami males. —Color: Mostly black Specimens Holotype: , 12 mi. E with some orange to yellow markings. ico, Morelos, Cuernavaca, 4300', viii.14.1954 Black except as follows: forefemur, tarsus, (Chillcott) (AEIC). Paratypes: Mexico: Guerrero: 33 mi. N and tibia with some orange or pale yellow; 19, Taxco, elev. 5700', viii.29.1963 (Scullen & Bolin- midfemur mostly black, slightly orange (OSUO); Morelos: 19, 3 mi. NW Te- distally; midtibia black and pale yellow to ger) viii.16.1962 (Roberts & orange; midtarsus black and pale yellow; quesquitango, 1 9 same data as hind femur black or with some orange dis- Martson); , holotype (AEIC); 16 [metasoma missing] same data tally; hind tibia pale yellow basally and as collector (Univ. Kans. medially, hind tarsus black and orange to holotype except Expedition) (SEMC); 19, 45 mi. S Cuer- pale yellow; wings clear to slightly infu- navaca, elev. 4300' ix.12.1957 (Scullen) mate. Body length: excluding ovipositor (CNCI); Puebla: IS, 5 mi. S Izucar de Ma- 7.2-8.0. Head: Longest seta at midlength of tamoros, viii.1.1963 (USNM). antenna approximately half antenna Remarks. —Similar to A. davidi, A. keni width (as in Fig. 7c); labial palpomere 2 and A. parkeningi—see characters in the subequal or shorter than combined length key and remarks under A. parkeningi. In of 3 + 4; mandible without indication of addition, A. capillata has longer setae second tooth (as in Fig. lc); glossa fork throughout the body and reduced sculp- length 0.17-0.18; glossa length less than ture on the propodeum and pronotum 2X galea length; glossa length 0.49-0.64; compared to A. davidi (Fig. 5a) and A. keni. glossa length 0.36-0.47 X shorter than for- Etymology. — Hairy— referring to the etibia length; malar space 0.34-0.38X long and extensive setae. shorter than eye height; face above clype- collini Pucci and us with or without median longitudinal Agathirsia Sharkey, n. carinae; area anterior to ocellus with pits sp. — forming V-shape (as in Fig. 7d); area be- Distribution (Fig. lie). South-central tween tentorial pit and antennal insertion Mexico. Research 90 Journal of Hymenoptera

2.1- Females and males. —Color: Mostly black itor length 2.2-2.3; ovipositor length or brown as fol- 2.3 X than hind basitarsus length. with some orange light longer — examined. 9 , Mex- lows: maxillary and labial palpomeres Specimens Holotype: ix.8-9.1948 mostly orange; foretrochantellus orange or ico, Puebla, Matamoros, (Wag- Mexico: Puebla: brown; forefemur orange or orange and ner) (AEIC). Paratypes: as or brown; foretibia and foretarsus orange; 79, 16, same data holotype midtrochanter, midtrochantellus, and ix. 10. 1948 (AEIC); 26, Petalcingo, midfemur orange or orange and brown; viii.3.1963 (Parker & Stange) (USNM); 19, 3 mi. & midtibia entirely orange or with small 16, N Petalcingo (Parker Stange) 5 mi. S Izucar de Matamo- black area basally; midtarsus orange; hind (USNM), 36, trochanter and hind trochantellus orange; ros, viii. 1.1963 (Parker & Stange) (USNM). — to A. A. michelei hind femur usually orange; hind tibia en- Remarks. Similar kellyi, and A. —see characters in the tirely orange or slightly black basally; papoui key. — au- hind tarsus black and orange; wings clear Etymology. In honor of the senior thor's cousin. or slightly infumate. Body length: excluding ovipositor 5.9-7.4. Head: Longest seta Agathirsia cressoni (Muesebeck and at midlength of antenna < half antenna Walkley) width (as in Fig. 7b); labial palpomere 2 + Microdus thoracicus Cresson 1872: 181 shorter than combined length of 3 4; (preoc-— mandible with or without indication of cupied by Nees von Esenbeck 1834: 143 Earinus thoracicus). [Examined]. second tooth (as in Fig. lb,c); glossa fork currently thoracicus 1904: 129. length 0.20-0.30; glossa length less than Agathirsia Szepligeti Agathirsia cressoni Muesebeck and Walkley 2X galea length; glossa length 0.48-0.59; X 1951: 116. glossa length 0.44-0.53 shorter than for- — etibia length; malar space 0.31-0.46X Distribution (Fig. 11a). Southeastern shorter than eye height; face above clype- Texas to northeastern Mexico. us with or without median longitudinal Females and males. —Color: Orange and carinae; area anterior to ocellus with pits black; black except as follows: antenna or- forming V-shape (as in Fig. 7d) or wide ange basally; maxillary and labial palpom- and vestigial V-shape; area between ten- eres mostly orange; pronotum orange dor- torial pit and antennal insertion without sally; mesoscutum orange except some- line of foveae. Mesosoma: Notauli foveate times black medially or entirely black; scu- and complete (as in Fig. 5a); propodeum tellum black and /or orange; metanotum mildly rugose, inverted V-shape anterior- black or black and orange; propodeum setae ly, present except along midline and black or with some orange; foretrochanter posterior central area (as in Figs. 9a,b); and foretrochantellus black, or black and sternaulus foveate (as in Fig. 5a); mesepis- orange; forefemur black and orange; for- ternum without lobes between midcoxae; etibia and foretarsus orange to pale yel- basal lobe of claws distinct (as in Fig. 3a); low; midtrochantellus black or black and number of thick, apically flattened apical orange; midfemur mostly black to mostly on hind tibia pegs 2-6; hind basitarsus ei- orange; midtibia and midtarsus orange to ther straight distally or very slightly ta- pale yellow; hind trochanter and hind tro- 1-cu-a and 1-M of pered; forewing sepa- chantellus black to orange; hind femur rated a distance than 2 vein by greater mostly black except slightly orange distal- widths in Metasoma: Posterior (as Fig. 9c). ly, sometimes mostly orange; hind tibia width of first 0.71-0.92X tergum shorter pale yellow, usually somewhat orange in than of first length tergum; setae on ter- distal quarter; hind tarsus orange to pale 3 in tergum sparse distally (as Fig. 6d); yellow; first tergum orange basally, black 3 without transverse gum groove; ovipos- distally; remaining terga orange with to Volume 13, Number 1, 2004 91

6 transverse black bands especially on ter- characters in the key and remarks under 2 and ga 3, bands sometimes broken me- A. nigricauda. dially, dark mottling also occurs on some davidi Pucci and specimens; forewing shaded yellow to in- Agathirsia Sharkey, S n- fumate in basal half, infumate in distal P* — half; hind wing infumate or slightly infu- Distribution (Fig. 111). Southern Arizo- mate. na and Body length: excluding ovipositor Texas to northern— Mexico. 9.7-11.2. Head: Longest seta at midlength Females and males. Color: Mostly black of antenna < half antenna width (as in except as follows: fore and midfemora with small Fig. 7b); labial palpomere 2 subequal to orange area distally; fore and midtibiae black palpomeres 3 + 4 (as in Fig. 4b); mandible and pale yellow /orange; with or without indication of second tooth a U tarsi pale yellow to black; hind tibia a le (as in Figs. lb,c); glossa fork length 0.21- P yellow basally laterally; forewing clear to infumate in basal in- 0.32; glossa length less than 2x galea slightly half, fumate in distal hind clear length; glossa length 1.2-1.3; glossa length half; wing to X infumate. 0.60-0.69 shorter than foretibia length; slightly Body length: excluding 6.8-9.6. Head: seta malar space 0.38-0.45 X shorter than eye ovipositor Longest at of antenna half height; face above clypeus with median midlength approximately antenna width in labial longitudinal carinae; area anterior to ocel- (as Fig. 7c); pal- omere 2 (as in 4b) or less lus with pits forming vestigial or distinct P subequal Fig. to 3 + 4; mandible without in- V-shape (as in Fig. 7d); area between ten- palpomeres dication of second tooth (as in torial pit and antennal insertion without Fig. lc);

lossa fork len th - 2°-°- 30 line of foveae. Mesosoma: Notauli foveate § § ; glossa length less than 2X §alea len§th; §lossa len§th and complete (as in Fig. 5a); propodeum °-54^-71 lossa len th 0.44-0.58X shorter foveate in- ' g 8 rugose posteriorly, anteriorly, - than foretlbia lenS th; malar sPace °' 24 verted V-shape anteriorly, setae present 30X short than e e hei face above midline and central §ht; except along posterior °; f / us Wlth °r without median area longitu- (Fig.° 6b); sternaulus foveate (as in Fig. v . . dinal .,, i ^ carinae; area anterior to ocellus with c u u . , . , 5a); mesepisternum without lobes be- L T , , . „.

. lts in .j u i i i_ r i j- P forming V-shape (as Fig. 7d); area tween • • midcoxae; basal lobe of claws dis- f . i ,

. , , . . tentorial . , between and antennal ™ „ , , , rpit mser- tinct (as in Fig. 3a); number of thick, api- .., ,_,. c , • tlon without line of foveae. „ a Y i. , ., Mesosoma: No-

flattened . i cally apicalr hpeesa on hind tibia ,- r . ,,.„.,_, . . , . tauli foveate . and in -. i^ , ,. „ , complete (as Fig. 5a); hind basitarsus 1- ,, 14-21; taperedr distally; A •, ,-,,., ,, propodeum mildly rugose, setae present cu-a and 1-M in forewing separated by ap- t al midHne and terior centra] the width of proximately 1-M, sometimes . area (Fi 6a); sternaulus foveate (Fi 5a) more. Metasoma: Posterior width slightly meSepisternum without lobes between of first 0.75-0.94X shorter than tergum midcoxae; basal lobe of claws small to dis- length of first tergum; setae on 3 tergum tinct; number f thick, apically flattened not dense (as in 6d), on distal _ . Fig. present apical pegs on hind tibia 5 9; hind basi one third or less; 3 with transverse tergum tarsus tapered distally; 1-cu-a and 1-M of 1.8-2.2; groove; ovipositor length ovipos- forewing separated by a distance greater itor 1.0-1. 2 X than hind bas- length longer than 2 vein widths (Fig. 9c). Metasoma: itarsus length. Posterior width of first tergum 0.56-0.68 X examined.— Specimens Holotype: 9, shorter than length of first tergum; setae No. 1728.1 Ho- USA, Texas, type (ANSP). on tergum 3 not dense (as in Fig. 6d) pre- 9 6 AT motypes: (44 ) AM, AEIC, ANSP, sent for approximately distal 0.4; tergum CNCI, TAMU, USNM. 3 with transverse groove distinct or ves- Remarks. —Similar to A. — nigricauda see tigial; ovipositor length 1.5-2.1; ovipositor 92 Journal of Hymenoptera Research

of Ft. viii.23.1969 length 1.4-1.8 X longer than hind basitar- N Rockpile, W Davis, sus length. (Board & Hafernik) (TAMU). — Remarks. —Similar to A. A. keni Specimens examined. Holotype: 9, — capillata, USA, Arizona, Parker Canyon Lk., and A. parkeningi see characters in the A. and A. viii.20.1974 (H. & M. Townes) (AEIC). key and remarks under capillata Paratypes: Mexico: Chihuahua: 1$, 16, parkeningi. —In of the senior au- 118 km N Chihuahua, viii.29.1991 (Gris- Etymology. honor thor's brother. wold) (CNCI); 1(5, General Trias, viii.20.1991 (Griswold) (CNCI); Sonora: Agathirsia foveiseries Pucci and 19, Alamos, elev. 1200', ix.22.1963 (Scul- Sharkey, sp. n. len & Bolinger) (OSUO); 1 9, 14 mi. W Al- Distribution —Known amos, elev. 800', ix.22.1963 (Scullen & Bol- (Fig. llh). only from in Arizona. inger) (OSUO); 1 9, 17 mi. E Navajoa, elev. type locality —Color: 700', ix.22.1963 (Scullen & Bolinger) Holotype female. Mostly orange and black; as follows: an- (OSUO); 1(5, road 35 km SE Cancoba, orange except tenna black and labial viii.23.1971 (Rozen & Favreau) (AMNH); distally; maxillary black and 1(5, road 35 km SE Cancoba, viii.29.1984 palpomeres orange; propleuron mesoscutum and scutellum (Pulawski) (CASC); USA: Arizona: 29, black; orange; black me- 2<5, Cochise Co., Chiri. Mrs. Pinery Cyn., mesopleuron orange, ventrally; tanotum black and 10 mi. NW Onion Saddle, viii.16.1965 orange; propodeum black and fo- (Ballmer) (USNM); 16, Cochise Co., orange; metapleuron orange; recoxa black; foretrochanter black and or- Apache, viii.18.1964 (Rozen) (AMNH); 19, foretarsus to testaceous; mid Cochise Co., Apache, viii. 20.1971 (Rozen ange; orange and hind trochanter black and & Favreau) (AMNH); 16, Cochise Co., orange; hind coxa with a little black; hind trochan- Willcox, viii.18.1958 (Hurd) (USNM); 19, tellus black and orange; first tergum yel- Canelo, viii.22.1974 (H. & M. Townes) lowish basally; remaining terga orange ex- (AEIC); 29, Nogales, viii.24.1939 (Cran- cept dark mottling present; wings slightly dall) (USNM); 36, Nogales, viii.22.1974 infumate in basal half, infumate in distal (H. & M. Townes) (AEIC); 19, 36, Santa half. Body length: excluding ovipositor 6.8. Cruz Co., 5 mi. N Lochiel, ix.6.1971 (Gris- Head: Longest seta at midlength of anten- sell & Denno) (UCDC); 19, 1<5, Santa na subequal to antenna width (as in Fig. Cruz Co., 17 mi. NE Patagonia, elev. 4950', 7c); labial palpomere 2X longer than com- viii.27.1955 (Scullen) (USNM); 16, Santa bined length of 3 + 4 (similar to Fig. 4b); Cruz Co., Pena Blanca, viii.30.1963. (Park- mandible with indication of second tooth er & Stange) (USNM); 39 same data as (as in Fig. lb); glossa fork length 0.41; holotype (AEIC); 1<5, Sonoita, viii.21.1974 glossa length less than 2x galea length; (H. & M. Townes) (AEIC); 19, 16, Pima glossa length 0.78; glossa length 0.62X Co., Sycamore, elev. 4000', viii.15-16.1993 shorter than foretibia length; malar space (Sharkey) (CNCI); Texas: 19, Jeff Davis X 0.53 shorter than eye height; face above Co., Madera Co., W Ft. Davis, viii.23.1969 clypeus with median longitudinal carinae; & 1 9 (Board Hafernik) (TAMU); , Jeff Da- area anterior to ocellus with pits forming vis Co., 20 mi. S Toyahvale on SR 17, V-shape (as in Fig. 7d); area between ten- viii.23.1974 (Greenbaum) (TAMU); 16, torial pit and antennal insertion with line Jeff Davis Co., 11.3 mi. W SR 17 on SR 166, of foveae (as in Fig. 3d). Mesosoma: Notauli viii.21.1974 (Greenbaum) (TAMU); 4c5, foveate and complete (as in Fig. 5a); pro- Jeff Davis Co., 2.3-4.1 mi. Limpia Cyn., W podeum rugose, longitudinal oval shape Davis Mrs., State Park on SR viii. 118, 18- medially, setae U974 present except posterior (Greenbaum) (TAMU); 2c?, 1 mi. central area and somewhat along midline; Volume 13, Number 1, 2004 93 sternalus foveate (as in Fig. 5a); mesepis- excluding ovipositor 8.4. Head: Longest seta ternum without lobes between midcoxae; at midlength of antenna < half antenna basal lobe of claws vestigial (as in Fig. 3b); width (as in Fig. 7b); labial palpomere 2 number of thick, apically flattened apical subequal to (as in Fig. 4b) or shorter than pegs on hind tibia 6-8; hind basitarsus ta- combined length of 3 + 4; mandible with indication of second tooth in pered distally; 1-cu-a and 1-M of forewing (as Fig. lb); fork separated by a distance greater than 2 vein glossa length 0.51; glossa length less than 2x widths (as in Fig. 9c). Metasoma: Posterior galea length; glossa length 1.1; X width of first tergum 0.46 X shorter than glossa length 0.79 shorter than foretibia malar 0.53 X shorter than length of first tergum; setae on tergum 3 length; space eye face above with median present distal half; tergum 3 with trans- height; clypeus carinae; area anterior to ocel- verse groove vestigial; ovipositor length longitudinal lus with in 1.7; ovipositor length 1.4X longer than pits forming V-shape (as Fig. 7d); area between tentorial and anten- hind basitarsus length. pit Males. —Unknown. nal insertion without line of foveae. Meso- soma: Notauli foveate and in Specimens examined. —Holotype: 9, complete (as Fig. 5a); propodeum mildly to fo- USA, Arizona, Douglas, viii.27.1979 (Bo- rugose veate, setae present midline hart) (UCDC). except along (similar to sternaulus foveate Remarks. —The combination of facial Figs. 9a,b); (as in Fig. 5a) for posterior half to three sculpture and coloration is distinctive for quarters, absent this Due to the variation of anteriorly; mesepisternum species. pro- without lobes between midcoxae; basal podeal sculpture found in Agathirsia and lobe of claws distinct (as in Fig. 3a); num- the fact that only one specimen has been ber of thick, apically flattened apical pegs examined, the oval-shaped areola of the on hind tibia 3-6; hind basitarsus tapered propodeum, noted above, may not be a re- distally; 1-cu-a and 1-M of forewing sepa- liable character. rated by a distance greater than 2 vein —Line of foveae— to Etymology. referring widths (as in Fig. 9c). Metasoma: Posterior the lines of foveae on the face. width of first tergum 0.81 X shorter than length of first tergum; setae on tergum 3 Agathirsia fulvocastanea Westwood sparse distally (as in Fig. 6d); tergum 3 with transverse Agathirsia fulvo-castanea Westwood 1882: 22. groove; ovipositor length 2.7X than [Examined]. 4.0; ovipositor length longer hind basitarsus length. Distribution. — from the examined. — 9 Only known Specimens Holotype: , Mex- type locality which is an unknown site in ico, 75 (Coffin) (OXUM). Mexico. Remarks.—Similar to A. proxima, A. real and A. —see characters in the Holotype female. —Color: Mostly orange to rufula key. In A. has foveate brownish orange; parts with other colora- addition, fulvocastanea notauli and an indication of a second tion are as follows: Antenna black distally; tooth unlike A. and A. The head with black areas medially on face and proxima rufula. of should not frons; pronotum slightly black ventrally; presence brownish-orange be used to this propleuron black; mesopleuron black ven- decisively identify species considering the similarity to what is called trally; propodeum with small black area orange elsewhere. posteriorly; metapleuron black posteriorly; forecoxa black; midcoxa black; hind coxa Agathirsia heleni Pucci and Sharkey, with small black area ventrally; first ter- sp. n. black infu- — gum basally; forewing slightly Distribution (Fig. lib). Known only mate in basal half, infumate in distal half; from type locality in San Luis Potosi, Mex- hind wing slightly infumate. Body length: ico. Journal of Hymenoptera Research

shorter male.—Color: width of first 0.74-0.77X Holotype female and paratype tergum antenna than of first tergum; setae on ter- Mostly orange except as follows: length 3 (as in Fig. 6d); ter- black; head entirely black except clypeus gum sparse distally 3 with transverse sometimes dark orange; pronotum black gum groove; ovipositor X 1.4; length 1.75 longer ventrally; propleuron black; propodeum length ovipositor than hind basitarsus and metapleuron black; forecoxa black; length, 2 foretrochanter and fore-trochantellus Specimens examined.—-Holotype: ,Mex- Luis Metehuala, x.25.62 black or black and orange; midcoxa black ico, [San Potosi], or black N.L. (AEIC). Paratype: 16, same or orange; midtrochanter black (Townes) data as (AEIC). and orange; midtrochantellus, midfemur, holotype Similar to A. armandi and A. midtibia, and midtarsus black; hind coxa Remarks.— characters in the and re- black or black and orange; hind trochanter bicolor—see key marks under those The in black or orange; hind trochantellus black species. glossa A. armandi and A. bicolor seems to be about or dark orange; hind femur black; hind to black twice the of A. heleni but tibia black distally, dark orange length proper first measurement of A. heleni has not been at- basally; hind tarsus black and orange; due to the condition of the tergum black; tergum 2 black basally, retained speci- with mens. maining terga dark orange or orange -In honor of the senior au- black mottling; wings slightly infumate. Etymology.— 5.4-6.7. thor's mother. Body length: excluding ovipositor of anten- Head: Longest seta at midlength . Pucci and ^ u ir •.,. / T3in tux. Agathtrsia* 'jervtsi Sharkey,J na < half antenna width (as in rig. 7b); "' labial palpomere 2 subequal to (as in Fig. — from the 4b) or shorter than combined length of 3 Distribution. Known only in Mexico, + 4; mandible with very weak indication type locality Guerrero, Color: black ex- of second tooth; glossa fork length -0.10 Holotype female.— Mostly as follows: antenna fuscous (?, difficult to measure on specimen); glos- cept distally, sa length less than 2x galea length; glossa pedicel orange distally; maxillary palp fuscous length 0.48? (glossa folded); glossa length brown, lighter distally; labial palp dis- 0.33-0.49 X shorter than foretibia length; basally and gradually turning yellow of brownish- malar space 0.43 X shorter than eye height; tally; ventral margin clypeus ba- face above clypeus without median Ion- orange; tibiae with small orange area tarsi gitudinal carinae; area anterior to ocellus sally and distally; fore- and mid with pits partially forming V-shape (sim- mostly orange; hind tarsus partly orange; in ilar to Fig. 7d); area between tentorial pit forewing clear in basal half, infumate and antennal insertion without line of fo- distal half; hind wing clear in basal half, veae. Mesosoma: Notauli foveate and com- slightly infumate in distal half. Body plete (as in Fig. 5a); propodeum mildly ru- length: excluding ovipositor 5.8. Head: Lon- gose and setae present except smooth and gest seta at midlength of antenna approx- glabrous along midline and posterior cen- imately half antenna width (as in Fig. 7c); tral area (setae as in Fig. 6a); sternaulus labial palpomere 2 subequal to combined foveate (as in Fig. 5a); mesepisternum length of 3 + 4; mandible with indication fork without lobes between midcoxae; basal of second tooth (as in Fig. lb); glossa lobe of claws distinct (as in Fig. 3a); num- length unknown; glossa length more than ber of thick, apically fattened apical pegs 2X galea length; glossa length 1.2; glossa on hind tibia 3-5; hinc nsitarsus tapered length 1.0X longer than foretibia length; distally; 1-cu-a and 1- f forewing sep- malar space .25X shorter than eye height; I by a distance gr ater than 2 vein face above clypeus without longitudinal as in Fig. 9c). Metasoma: Posterior carinae; area anterior to ocellus with pits Volume 13, Number 1, 2004 95

in forming V-shape (as Fig. 7d); area be- 0.36 X shorter than eye height; face above tween tentorial pit and antennal insertion clypeus without median longitudinal ca- without line of foveae. Mesosoma: Notauli rinae; area anterior to ocellus without pits foveate and complete (as in Fig. 5a); pro- forming V-shape; area between tentorial podeum rugose, setae present except pos- pit and antennal insertion without line of terior central area; sternaulus foveate (as foveae. Mesosoma: Notauli complete, non in without Fig. 5a); mesepisternum lobes foveate (as in Fig. 5b); propodeum foveate, between basal lobe of midcoxae; claws inverted V-shape anteriorly, setae present distinct in (as Fig. 3a); number of thick, except along midline and posterior central flattened apically apical pegs on hind tibia area (as in Fig. 9a,b); sternaulus foveate hind basitarsus 3; tapered distally; 1-cu-a (as in Fig. 5a); mesepisternum without and 1-M of forewing separated by a dis- lobes between midcoxae; basal lobe of tance than 2 vein in greater widths (as Fig. claws distinct (as in Fig. 3a); number of Metasoma: Posterior 9c). width of first ter- thick, apically flattened apical pegs on .85 X shorter than first gum length of ter- hind tibia 1-2; hind basitarsus straight dis- setae on 3 gum; tergum sparse distally (as tally; 1-cu-a and 1-M in forewing separat- in Fig. 6d); tergum 3 without transverse ed by a distance subequal to width of 1- groove; ovipositor length 1.0; ovipositor M (see Fig. 9c). Metasoma: Posterior width X length .98 longer than hind basitarsus of first tergum 0.68 X shorter than length length. of first tergum; setae on tergum 3 sparse Males. — Unknown. distally (as in Fig. 6d); tergum 3 with examined. — 9 Mex- transverse Specimens Holotype: , groove vestigial; ovipositor ico, Amula, Guerrero, 6000 ft. Aug. (H.H. length unknown. Smith) Godman-Salvin Coll. (BMNH). Females. —Unknown. Remarks. —This is the with examined. — only species Specimens Holotype: 6 , Mex- elongated labial palpi that are convex me- ico, Puebla, Huegotzingo, elev. 5-6000', dially and form— a drinking tube. viii.17.1962 (Milliron) (CNCI). Etymology. The specific name is in Remarks. —Similar to A. collini, A. mich- honor of Mark Jervis for his interesting re- elei and A. papoui—see characters in the search on hymenopteran mouthpart mor- key. phology. Etymology. —In honor of the senior author's aunt. Agathirsia kellyi Pucci and Sharkey, sp. n. Agathirsia keni Pucci and Sharkey, Distribution —Known sp. n. (Fig. Hi). only — from type locality in Puebla, Mexico. Distribution (Fig. llg). Known from 2 Holotype male. —Color: Mostly Black ex- localities in Jalisco, Mexico. cept as follows: all femora and tibiae or- Holotype female and paratope male. —Color: ange; all tarsi mostly orange; wings slight- Mostly black except as follows: foretibia ly infumate. Body length: excluding ovi- black or orange to brown; foretarsus positor 7.1. Head: Longest seta at mid- sometimes with some orange distally; length of antenna subequal to antenna midtibia black or orange to brown; mid- width (as in Fig. 7c); labial palpomere 2 tarsus orange to mostly black; hind tibia shorter than combined length of 3 + 4; mostly pale yellow basally; forewing clear mandible with indication of second tooth or slightly infumate in basal half, slightly (as in Fig. lb); glossa fork length 0.13; infumate to infumate in distal half; hind glossa length less than 2X galea length; wing clear to slightly infumate. Body glossa length 0.62; glossa length 0.52 X length: excluding ovipositor 7.8-8.3. Head: shorter than foretibia length; malar space Longest seta at midlength of antenna ap- 96 Journal of Hymenoptera Research

as follows: and la- proximately half antenna width (as in Fig. Black except maxillary with color fem- 7c); labial palpomere 2 shorter than com- bial palpomeres some pale bined length of 3 + 4; presence of 2nd ora and tibiae of fore and midlegs with mandibular tooth not seen due to speci- small orange area distally; hind tibia with men position; glossa fork length 0.43; glos- small orange area basally; forewing infu- sa length less than 2x galea length; glossa mate or slightly infumate; hind wing clear X length 0.70-0.83; glossa length 0.51-0.61 to infumate in basal half, slightly infumate shorter than foretibia length; malar space to infumate in distal half. Body length: ex- X face 0.33-0.37 shorter than eye height; cluding ovipositor 7.3-7.4. Head: Longest above clypeus with or without median seta at midlength of antenna < half anten- area anterior to ocel- longitudinal carinae; na width (as in Fig. 7b); labial palpomere lus with in pits forming V-shape (as Fig. 2 shorter than combined length of 3 + 4; 7d) or vestigial V-shape; area between ten- mandible without indication of second torial and antennal insertion without pit tooth (as in Fig. lc); glossa fork length line of foveae. Mesosoma: Notauli foveate 0.37-0.45; glossa length more than 2x ga- and (as in 5a); complete Fig. propodeum lea length; glossa length 3.4-3.5; glossa setae midline rugose, present except along length 2.6-2.7X longer than foretibia (similar to 6a); sternaulus foveate (as Fig. length; malar space 0.29-0.33 X shorter in Fig. 5a); mesepisternum without lobes than eye height; face above clypeus with- between midcoxae; basal lobe of claws out or with vestigial longitudinal carinae; distinct (as in Fig. 3a); number of thick, area anterior to ocellus without pits form- apically flattened apical pegs on hind tibia ing V-shape; area between tentorial pit 5-8; hind basitarsus 1-cu- tapered distally; and antennal insertion without line of fo- a and 1-M of a dis- forewing separated by veae. Mesosoma: Notauli foveate and com- tance greater than 2 vein widths (as in Fig. plete (as in Fig. 5a); propodeum mildly ru- 9c). Metasoma: Posterior width of first ter- gose posteriorly, foveate and longitudinal gum 0.79-0.92 X shorter than length of medial furrow anteriorly, setae present ex- first; setae on tergum 3 not dense (as in cept along midline and posterior central Fig. 6d), present on distal half; tergum 3 area (as in Figs. 9a,b); sternaulus foveate with transverse groove; ovipositor length (as in Fig. 5a); mesepisternum without 2.1; ovipositor length 1.6X longer than lobes between midcoxae; basal lobe of hind basitarsus length. claws (as in 3b); number of Specimens examined.—Holotype: 9, Mex- vestigial Fig. thick, flattened on ico, Jalisco, 18 mi. SW Guadalajara, apically apical pegs hind tibia 2-3; hind basitarsus dis- ix.6.1971 (Villegas & Kane) (UCDC). Para- straight 1-cu-a and 1-M of type: Mexico: Jalisco: 16, Lagos de Mo- tally; forewing separated a distance than 2 vein reno, elev. 6300', viii.12.1954 (Dreisbach) by greater (USNM). widths (as in Fig. 9c). Metasoma: Posterior Remarks. — width of first 0.80 X shorter than Similar to A. capillata, A. dav- tergum of first idi and A. parkeningi—see characters in the length tergum; setae on tergum 3 and in key remarks under A. capillata. sparse distally (as Fig. 6d); tergum 3 — with Etymology. In honor of the senior au- transverse groove vestigial; oviposi- thor's uncle. tor length 8.2; ovipositor length 8.2 X longer than hind basitarsus length. Pucci and Agathirsia longigladia examined.— Specimens Holotype: 9 , Mex- Sharkey, sp. n. ico, Morelos, Cuernavaca, 3i. 19-21. 1987 Distribution (Fig. llg).—Known only (Parker) (CNCI). Paratype: 16, same data type locality in Morelos, Mexico. as holotype (CNCI). and male. —Color: — Dlotype female paratype Remarks. The combination glossa Volume 13, Number 1, 2004 97

length and coloration make this species first tergum; setae on tergum 3 sparse dis- distinctive. — — tally (as in Fig. 6d); tergum 3 with trans- Etymology. Long sword referring to verse groove distinct to absent; ovipositor the elongate ovipositor. length 5.3-6.9; ovipositor length 4.4-4.7X longer than hind basitarsus length. Agathirsia longilingna Pucci and Specimens examined. —Holotype: 9, Mex- Sharkey, sp. n. ico, Jalisco, San Fandia [San Fandila?], Distribution —Central Mexico. (Fig. llj).— ix.24.1963 (Michelbacher) (EMEC). Para- Females and males. Color: Mostly black types: Mexico: Jalisco: 16, same data as except as follows: maxillary and labial pal- holotype (EMEC); Queretaro: 39, 9 mi. N pomeres mostly orange; clypeus some- Queretaro, ix.21.1977 (Chemsak & A.&M. times partially orange; fore and midfe- Michelbacher) (EMEC); Zacatecas: 19, 8 or mora orange orange and black; all tib- mi. NW Rio Grande ix.27.1975 (Villegas) iae and tarsi orange; hind femur black and (UCDC). — orange, orange may be restricted to ex- Remarks. The combination of glossa treme distal portion; forewing slightly in- length and coloration distinguishes this fumate or infumate in basal half, infumate species from —all others in the genus. in distal half; hind wing slightly infumate. Etymology. Long tongued—referring Body length: excluding ovipositor 8.0-9.3. to the elongate glossa. Head: Longest seta at midlength of anten- Agathirsia michelei Pucci and Sharkey, na < half antenna width (as in Fig. 7b); n. labial palpomere 2 shorter than combined sp. + — length of 3 4; mandible with or without Distribution (Fig. 111). Known only indication of second tooth in from in (as Fig. lb,c); type locality — Jalisco, Mexico. glossa fork length 0.63-0.72; glossa length Holotype female. Color: Mostly black more than 2X galea length; glossa length and orange; black except as follows: max- 4.5-5.5; glossa length 2.8-3.1 X longer than illary and labial palpomeres mostly or- foretibia length; malar space 0.50-0.59 X ange; all trochanters black and orange; all shorter than eye height; face above clype- trochantelli femora and tibiae orange; ter- us with or without median longitudinal ga with 3 dark reddish-brown transverse carinae; area anterior to ocellus with pits bands on terga 2 and 3; wings slightly in- forming V-shape (as in Fig. 7d); area be- fumate. Body length: excluding ovipositor tween tentorial pit and antennal insertion 8.2; Head: Longest seta at midlength of an- without line of foveae. Mesosoma: Notauli tenna approximately half antenna width foveate and complete (as in Fig. 5a); pro- (as in Fig. 7c); labial palpomere 2 subequal podeum foveate except smooth or mostly to palpomeres 3 + 4 (as in Fig. 4b); man- smooth central area and rugosity sur- dible without indication of second tooth rounding and below spiracles, setae pre- (as in Fig. lc); glossa fork length 0.49; glos- sent except narrowly along midline (sim- sa length less than 2x galea length; glossa ilar to Fig. 6a); sternaulus foveate (as in length 1.4; glossa length equal to foretibia Fig. 5a); mesepisternum without lobes be- length; malar space 0.49 X shorter than eye tween midcoxae; basal lobe of claws dis- height; face above clypeus without medi- tinct (as in Fig. 3a); number of thick, api- an longitudinal carinae; area anterior to cally flattened apical pegs on hind tibia ocellus with pits forming vestigial V- 10-12; hind basitarsus tapered distally; shape (similar to Fig. 7d); area between distance between 1-cu-a and 1-M in fore- tentorial pit and antennal insertion with- wing subequal to the width of 1-M (see out line of foveae. Mesosoma: Notauli fo- Fig. 9c). Metasoma: Posterior width of first veate and complete (as in Fig. 5a); pro- tergum 0.80-0.93 X shorter than length of podeum foveate, setae present except 98 Journal of Hymenoptera Research

area anterior to ocellus along midline and central posterior area gitudinal carinae; in in (as in Figs. 9a,b); sternaulus foveate (as with pits forming V-shape (as Fig. 7d); area tentorial and antennal in- Fig. 5a); mesepisternum without lobes be- between pit tween midcoxae; basal lobe of claws dis- sertion without line of foveae. Mesosoma: foveate in tinct (as in Fig. 3a); number of thick, api- Notauli and complete (as Fig. setae cally flattened apical pegs on hind tibia 7- 5a); propodeum foveate, sparse pre- in 8; hind basitarsus tapered distally; 1-cu-a sent laterally; sternaulus foveate (as Fig. and 1-M of forewing separated by a dis- 5a); mesepisternum without lobes be- tance greater than 2 vein widths (as in Fig. tween midcoxae; basal lobe of claws ves- 9c). Metasoma: Posterior width of first ter- tigial (as in Fig. 3b); number of thick, api- tibia 1- gum 0.74 X shorter than length of first ter- cally flattened apical pegs on hind gum; setae on tergum 3 sparse distally (as 2; hind basitarsus straight distally; 1-cu-a in Fig. 6d); tergum 3 with transverse and 1-M of forewing separated by a dis- groove; ovipositor length 5.4; ovipositor tance greater than 2 vein widths (as in Fig. length 4.9 X longer than hind basitarsus 9c). Metasoma: Posterior width of first ter- X first length. — gum 0.71 shorter than length of ter- Males. Unknown. gum; setae on tergum 3 sparse distally (as examined. — 9 in 3 Specimens Holotype: , Mex- Fig. 6d); tergum without transverse ico, Jalisco, 12 mi. S Encarnacion de Diaz groove; ovipositor length 0.54; ovipositor & P. 0.73 X shorter than basitarsus (C. Vaurie)— (AMNH). length hind Remarks. Similar to A. collini, A. kellyi length. and A. —see in — papoui— characters the key. Male. Unknown. Etymology. In honor of the senior au- Specimens examined. —Holotype: 9, thor's sister-in-law. [USA, New Mexico] Hidalgo Co., Rodeo, viii.21.1958 (Bohart) (UCDC). Agathirsia minuata Pucci and Sharkey, — Remarks. The combination of glossa sp. n. — length and coloration distinguishes this Distribution (Fig. Ilk). Known only species from all others in the genus. from — type locality. Etymology. Small—referring to the rel- —Color: Holotype female. Mostly orange atively small glossa, ovipositor and over- as follows: except antenna black distally; all length. mesosoma orange except propodeum somewhat darkened; midleg somewhat Agathirsia nigricauda (Viereck) hind tibia darkened, darkened distally; Crassomicrodus nigricaudus Viereck 1905: 288. hind tarsus black; large brown area on ter- [Examined]. ga 2 and 3 the result of stains (likely Microdus caused nigricaudus Withington 1909: 329. by internal fluids); wings very Agathirsia nigricauda Muesebeck 1927: 14. infumate. slightly Body length: excluding 5.7. Head: seta at mid- ovipositor Longest Distribution (Fig. Hi). —Common in of length antenna approximately half an- northern Mexico and southern Arizona, tenna width in labial (as Fig. 7c); palpom- New Mexico, and Texas; occasionally 2 ere shorter than combined length of 3 + found in states somewhat north of these. 4; mandible with weak indication of sec- Females and male. —Color: Quite variable, ond tooth; fork glossa length unknown; black and orange: black except as follows: glossa less than 2X length galea length; Antenna orange basally; maxillary and la- glossa bial length approximately 0.20; glossa palpomeres mostly orange; rarely or- 0.20 X shorter than foretibia length length; ange behind ocelli and lateral area of face; malar space 0.55 X shorter than eye height; clypeus sometimes orange; pronotum or- ice above without median lon- clypeus ange dorsally; mesoscutum orange, some- Volume 13, Number 1, 2004 99 times with black area anteriorly; scutellum wing separated by a distance greater than black usual- 2 and/or orange; mesopleuron vein widths (as in Fig. 9c). Metasoma: ly black, rarely mostly orange to orange; Posterior width of first tergum 0.65-0.85 X metanotum sometimes black and orange; shorter than length of first tergum; setae black or black and propodeum orange, on tergum 3 not dense (as in Fig. 6d), pre- rarely entirely orange; metapleuron rarely sent on distal one third or less; tergum 3 with some orange; legs orange except: with transverse groove distinct or vesti- coxae of fore and black midlegs and /or gial; ovipositor length 2.2-3.2; ovipositor coxa orange, hind orange or black and or- length 1.3-2.1 X longer than hind basitar- fore ange, and midtibiae orange to pale sus length. hind tibia usu- examined. — Col- yellow, pale yellow except Specimens Holotype: 9 , at least somewhat in distal ally orange orado, Colorado Springs, 5015 ft., 643, viii quarter, tarsi orange to testaceous; terga (Tucker) (SEMC). Homotypes: (200 9 6) orange basally, black distally, rarely with AEIC, AMNH, CASC, CNCI, EMEC, black area basally and orange distally; LACM, MSUC, OSUO, SEMC, TAMU, forewing shaded yellow in basal half, UCDC, USNM. slightly infumate to infumate in distal Remarks.—Similar to A. cressoni—see half; hind wing slightly infumate to infu- characters in the key. The characters we mate. Body length: excluding ovipositor used to separate these either have rare ex- 9.1-10.4. Head: seta at Longest midlength ceptions, e.g., terga coloration, hind femur of antenna < half antenna width (as in color, notauli sculpture, and relative glos- labial 2 to sa or Fig. 7b); palpomere subequal length, can be very similar, e.g., rel- 3 + 4 palpomeres (Fig. 4b); mandible ative length of ovipositor. Although these without indication of second tooth (as in species overlap in time and space, A. ni- Fig. lc) or very weak; glossa fork length gricauda has a greater geographic range 0.19-0.34; glossa length less than 2x galea and an earlier flight period. Specimens of length; glossa length 1.2—1.5; glossa length both species have been collected within a 0.68-0.95 X shorter than foretibia length; few days from each other, late September malar space 0.35-0.47X shorter than eye to early October, in Pearsall, Texas. These height; face above clypeus with or without specimens are housed in TAMU. median longitudinal carinae; area anterior Agathirsia ninesevensi Pucci and to ocellus with pits forming V-shape (as in n. or area between tentorial bnarkey, sp. Fig. 7d) vestigial; — pit and antennal insertion without line of Distribution (Fig. 11a). Known only foveae. Mesosoma: Notauli from in foveate, usually type locality— Texas, incomplete anteriorly (—75%) otherwise Holotype female. Color: Mostly black ex- weakly or totally complete; propodeum cept as follows: antenna fuscous basally; foveate to rugose, less sculpture anterior- maxillary and labial palpomeres black and ly, often with inverted V-shape anteriorly, orange; foretrochantellus, forefemur, for- central area usually depressed with sculp- etibia, and foretarsus orange; midtrochan- ture differentiated from surrounding are- tellus, midfemur, midtibia, and midtarsus as, setae present except along midline and orange; hind leg orange except hind coxa central posterior area (as in Figs. 9a,b); black; terga orange except tergum 1 and sternaulus foveate (as in Fig. 5a); mesepis- distal tip of metasoma black; wings infu- ternum without lobes between midcoxae; mate. Body length: excluding ovipositor basal lobe of claws distinct (as in Fig. 3a); 8.2. Head: Longest seta at midlength of an- number of thick, apically flattened apical tenna > half antenna width (as in Fig. 7a); pegs on hind tibia 11-18; hind basitarsus labial palpomere 2 shorter than combined tapered distally; 1-cu-a and 1-M of fore- length of 3 + 4; mandible without indi- 100 Journal of Hymenoptera Research cation of second tooth (as in Fig. lc); glos- mate. Body length: excluding ovipositor seta at of sa fork length 0.23; glossa length less than 7.8-8.6. Head: Longest midlength to half or less antenna 2X galea length; glossa length 0.94; glossa antenna subequal length 0.67 X shorter than foretibia length; width (as in Figs. llb,c); labial palpomere + 4 in malar space 0.54 X shorter than eye height; 2 subequal to palpomeres 3 (as Fig. face above clypeus without median lon- 4b); mandible with or without indication in gitudinal carinae; area anterior to ocellus of second tooth (as Figs. lb,c); glossa less without pits forming V-shape; area be- fork length 0.11-0.24; glossa length 0.49- tween tentorial pit and antennal insertion than 2x galea length; glossa length without line of foveae. Mesosoma: Notauli 0.60; glossa length 0.36-0.46 X shorter than foveate anteriorly, present as smooth foretibia length; malar space 0.28-0.33 X groove medially and posteriorly; propo- shorter than eye height; face above clype- deum with large smooth and glabrous us with median longitudinal carinae; area area medially, weak rugosity and sparse anterior to ocellus with or without pits setae elsewhere; sternaulus foveate (as in forming V-shape; area between tentorial Fig. 5a) posteriorly, smooth groove ante- pit and antennal insertion without line of riorly; mesepisternum without lobes be- foveae. Mesosoma: Notauli foveate and tween midcoxae; basal lobe of claws ves- complete (as in Fig. 5a); propodeum mild- tigial (as in Fig. 3b); number of thick, api- ly rugose, central area depressed, setae cally flattened apical pegs on hind tibia 3- present except narrowly along midline 4; hind basitarsus tapered distally; 1-cu-a (similar to Fig. 6a); sternaulus foveate (as in and 1-M forewing partially or nearly in Fig. 5a); mesepisternum without lobes overlapping. Metasoma: Posterior width of between midcoxae; basal lobe of claws first 0.61 X tergum shorter than length of distinct (as in Fig. 3a); number of thick, first tergum; setae on tergum 3 sparse dis- apically flattened apical pegs on hind tibia in 3 tally (as Fig. 6d); tergum without 9-15; hind basitarsus tapered distally; 1- transverse groove; ovipositor length 3.4; cu-a and 1-M of forewing separated by a X ovipositor length 2.8 longer than hind distance greater than 2 vein widths (as in basitarsus length. Fig. 9c). Metasoma: Posterior width of first Male.—Unknown. X — tergum 0.67-0.73 shorter than length of Specimens examined. Holotype: 9, first tergum; setae on tergum 3 not dense Pecos x.17.1973 USA, Texas, Co., (Bohart) (as in Fig. 6d), present on distal third to (UCDC). half; tergum 3 with transverse groove; ovi- Remarks. —Similar to A. — campanisura positor length 0.83-0.88; ovipositor length see characters in the key and remarks un- 0.69-0.73 X shorter than hind basitarsus der A. campanisura. length. —In honor of Etymology. the musical Specimens examined.—Holotype: 9, Mex- Old 97's. group ico, Hidalgo, 13.5 mi. NE Tizayuca, elev. 7700', viii.28.1962 (SEMC). Paratypes: Agathirsia paponi Pucci and Sharkey, Mexico: Guanajuato: lc?, 10 mi. NW Leon, sp. n. viii.19.1954 (Chillcott) (CNCI); Hidalgo: Distribution Central (Fig. lib).— Mexico. 19, 36, same data as holotype (SEMC); Females and males. —Color: Most black ex- 16, same data as holotype except 3.5 mi. as follows: and labial cept maxillary pal- NE Tizayuca (SEMC); Jalisco: lc? [meta- sometimes with fem- pomeres pale color; soma missing], 16, Guadalajara, [6 date], ora orange; tibiae orange except hind tibia (McClendon) (ANSP). may be darkened tarsi or — distally; orange Remarks. Similar to A. collini, A. kellyi mostly hind tarsus be — orange except may and A. michelei see characters in the key. dark clear to orange; wings slightly infu- We have viewed a damaged specimen re- Volume 13, Number 1, 2004 101

A. sembling papoui except that the length gum 3 with wide shallow area; ovipositor of the glossa appears to be subequal to the length 5.6; ovipositor length 5.1 X longer foretibia. It collected in was Mexico, Jalis- than hind basitarsus length. co, Largos de Moreno, elev. 6400', August Males. —Unknown. 1954 and is housed in SEMC. — 21, — Specimens examined. Holotype: 9, Mex- Etymology. In honor of the senior au- ico, Jalisco, Zapotlanejo, x.3.[19]66 (G.E. & thor's grandfather. A.S. Bohart)—(CNCI). Remarks. Similar to A. capillata, A. dav- Agathirsia parkeningi Pucci and — idi and A. keni see characters in the key. Sharkey, sp. n. In addition, glossa length and malar space Distribution — (Fig. lie). Known only are distinctly longer in A. parkeningi than from in type locality— Jalisco, Mexico. in the others. Holotype female. Color: Mostly black ex- Etymology. —In honor of the musician cept as follows: maxillary and labial pal- Christopher Parkening. pomeres with some pale color; all femora with small orange area distally; all tibiae Agathirsia proxima Westwood with some orange basally; foretarsus with Agathirsia proxima Westwood 1882: 22. [Exam- some pale color; midtarsus mostly black; ined]. hind tarsus pale yellow basally, black dis- Agathirsia rufiventris Westwood 1882: 21. [Ex- infumate. exclud- tally; wings Body length: amined]. Syn. n. ing ovipositor 7.1. Head: Longest seta at midlength of antenna approximately half Distribution (Fig. llh). —Central to or < half antenna width (as in Fig. llb-c); southern Mexico. labial palpomere 2 shorter than combined Eemales and males. —Color: Orange and length of 3 + 4; mandible without indi- black, color quite variable; antenna orange cation of second tooth (as in Fig. lc); glos- basally black distally; maxillary and labial sa fork length 0.49; glossa length less than palpomeres mostly orange; head orange 2X galea length; glossa length 1.2; glossa except at least central area of face black, length 0.86 X shorter than foretibia length; often frons and face black, rarely entirely malar space 0.43 X shorter than eye height; black; pronotum varies from orange, to face above clypeus without median lon- black with small orange area dorsally; gitudinal carinae; area anterior to ocellus propleuron black or orange and black; me- with pits forming V-shape (as in Fig. 7d); soscutum orange; scutellum orange or area between tentorial pit and antennal in- black and orange; mesopleuron mostly or- sertion without line of foveae. Mesosoma: ange to black; metanotum black or black Notauli foveate and complete (as in Fig. and orange; propodeum black to orange; 5a); propodeum foveate, setae present metapleuron black to mostly orange with throughout; sternaulus foveate (as in Fig. some black; (rarely mesosoma entirely 5a); mesepisternum without lobes be- black); legs orange except sometimes tween midcoxae; basal lobe of claws ves- black on basal portion of coxae, tarsi or- tigial (as in Fig. 3b); number of thick, api- ange to pale yellow; terga orange except cally flattened apical pegs on hind tibia 3- often with black or dark orange mottling; 4; hind basitarsus tapered distally; 1-cu-a forewing slightly infumate in basal half, and 1-M of forewing separated by a dis- infumate or slightly infumate in distal tance greater than 2 vein widths (as in Fig. half; hind wing slightly infumate. Body 9c). Metasoma: Posterior width of first ter- length: excluding ovipositor 7.3-9.0. Head: gum 0.60 X shorter than length of first ter- Longest seta at midlength of antenna ap- gum; setae on tergum 3 not dense (as in proximately half or < half antenna width Fig. 6d), present on distal one third; ter- (as in Figs. llb,c); labial palpomere 2 102 Journal of Hymenoptera Research

shorter than combined length of 3 + 4; A. rufiventris. The head and mesosoma are mandible without indication of second completely black. tooth (as in Fig. lc); glossa fork length Agathirsia reai Pucci and Sharkey, 0.56-0.69; glossa length more than 2X ga- sp. n. lea length; glossa length 3.1-3.9; glossa — length 2.2-2.8 X longer than foretibia Distribution (Fig. Ilk). South-central Mexico. length; malar space 0.38-0.49 X shorter — than eye height; face above clypeus with- Females. Color: Mostly orange except as follows: out or with vestigial longitudinal carinae; antenna black or with some or- and labial area anterior to ocellus with pits forming ange basally; maxillary palpomeres head black or- vestigial V-shape (similar to Fig. 7d); area mostly orange; except and behind between tentorial pit and antennal inser- ange clypeus ocellus; propo- tion without line of foveae. Mesosoma: No- deum and metapleuron with some black; tauli clear in basal half, infumate in dis- complete, non foveate (Fig. 5b); pro- wings tal half or infumate podeum weakly sculptured, usually with slightly throughout. small depression along anterior midline, Body length: excluding ovipositor 6.5-7.4. Head: seta at of anten- smooth below, setae present except along Longest midlength na < half antenna width in midline and central posterior area (as in (as Fig. 7b); labial 2 to Figs. 9a,b); sternaulus with few foveae at palpomere subequal palpom- extreme base eres 3 + 4 (as in 4b); mandible with (Fig. 5c); mesepisternum Fig. indication of second tooth without lobes between midcoxae; basal (as in Fig. lb); lobe of claws glossa fork 0.31; less distinct (as in Fig. 3a); num- length glossa length ber of than 2X galea 0.89- thick, apically flattened apical pegs length; glossa length on hind tibia 1.0; glossa length 0.71-0.86 X shorter than 3-6; hind basitarsus straight 1-cu-a foretibia length; malar 0.45-0.53 X distally; and 1-M of forewing sep- space shorter than face above arated by a distance greater than 2 vein eye height; clypeus with distinct or widths (as in Fig. 9c). Metasoma: Posterior vestigial longitudinal carinae; area anterior to ocellus with width of first tergum 0.59-0.82 X shorter pits forming (as in area be- than length of first tergum; setae on ter- V-shape Fig. 7d); tween 3 tentorial pit and antennal insertion gum sparse distally (as in Fig. 6d); ter- without line of foveae. Mesosoma: Notauli gum 3 with transverse groove distinct to foveate and complete (as in Fig. 5a); pro- absent; ovipositor length 6.0-7.1; oviposi- podeum foveate, with or without inverted tor length 5.0-5.6 X longer than hind bas- V-shape anteriorly, setae itarsus length. present except midline and central area examined.— along posterior Specimens 9 , Mex- Holotype: in (as Figs. 9a,b); sternaulus foveate (as in ico, 74 (Coffin) (OXUM). Coffin #25 "Also 5a); without at Los Fig. mesepisternum lobes be- Barros. Hovering in the neighbour- tween midcoxae; basal lobe of claws dis- hood of plants inhabited larvae." by Ho- tinct (as in Fig. 3a); number of thick, motypes: (33 9 6) AMNH, CNCI, EMEC, api- OXUM, UCDC. cally flattened apical pegs on hind tibia 1- 2; hind basitarsus Remarks.— tapered distally; 1-cu-a Very similar to A. rufula—ihe and 1-M of forewing separated a dis- characters in the are the by key only ones tance than 2 greater vein widths (as in Fig. found to distinguish these Also species. 9c). Metasoma: Posterior width of first ter- similar to A. fulvocastanea and A. reai—see X gum 0.73-0.82 shorter than length of characters in the key and remarks under first tergum; setae on 3 ter- A. tergum absent; fulvocastanea. We have viewed three 3 with gum shallow indentation; oviposi- specimens that we consider to be melanic tor length 5.4-6.3; ovipositor length 5.4- one of orms, these being the of 5.7X than holotype longer hind basitarsus length. Volume 13, Number 1, 2004 103

Males. —Unknown. without lobes between midcoxae; basal examined.— Specimens Holotype: 9.. Mex- lobe of claws distinct (as in Fig. 3a) or ves- Lk. ico, Morelos, Tequesquitengo, approx. tigial (as in Fig. 3b); number of thick, api- ix.13.1957 5000', (Scullen) (CNCI). Para- cally flattened apical pegs on hind tibia 1- Mexico: Puebla: type: 19, Chietla, Aten- 5; hind basitarsus tapered distally; 1-cu-a elev. 1098 cingo, m, ix.25.1989 (Pena) and 1-M of forewing separated by a dis- (ATAM). tance than 2 vein widths in — greater (as Fig. Remarks. Similar to A. fulvocastanea, A. 9c). Metasoma: Posterior width of first ter- A. — proximo, and rufula see characters in gum 0.63-0.65 X shorter than length of the key and remarks under A. fulvocasta- first tergum; setae on tergum 3 not dense nea. — (as in Fig. 6d), present on distal half; ter- Etymology. In honor of the senior au- gum 3 without transverse groove; ovipos- thor's grandmother. itor length 2.8-3.1; ovipositor length 2.5- 2.8 X longer than hind basitarsus length. Agathirsia rostrata Pucci and Sharkey, Specimens examined. —Holotype: 9, Mex- sp. n. — ico, Puebla, 30 mi. SW Tehuacan, elev. Distribution (Fig. llg). Northeastern to 6800', x.13.1968 (R.H. & E.M. Painter) south-central Mexico. (AEIC). Paratypes: Mexico: Nuevo Leon: Females and males. —Color: black 1 9 Mostly , San Pedro Iturbide, 32 km W Linares, except as follows: antenna sometimes dark x.6.1962 (H. & M. Townes) (AEIC); Puebla: orange basally; fore and midfemora some- 1 9, same data as holotype (AEIC); 16", nr. times with small orange area distally; for- Puebla, x.15.1962 (Townes) (AEIC). etibia completely fuscous or orange basal- Remarks. —The combination of facial ly; foretarsus orange or fuscous basally; sculpture and coloration is diagnostic for midtibia orange or fuscous basally; mid- this species. In addition, the head in fron- tarsus orange basally; hind tibia orange tal view appears somewhat like a rostrum hind tarsus black and similar to that basally; orange; (Fig. 3d), — of Agathis species. wings infumate. Body length: excluding Etymology. Beaked, with a muzzle—re- ovipositor 5.8-6.8. Head: Longest seta at ferring to the head shape. midlength of antenna < half antenna Westwood width (as in Fig. 7b); labial palpomere 2 Agathirsia rufula shorter than combined length of 3 + 4; Agathirsia rufula Westwood 1882: 21. [Exam- mandible with indication of second tooth ined]. in (as Fig. lb) or vestigial; glossa fork — length 0.10-0.14; glossa length less than Distribution (Fig. lid). South-central 2X galea length; glossa length 0.48-0.53; Mexico. glossa length 0.45 X shorter than foretibia Females and males. —Color: Mostly orange length; malar space 0.50-0.59 X shorter except as follows: antenna black distally; than eye height; face above clypeus with- maxillary and labial palpomeres with out median longitudinal carinae; area an- some orange; frons and face black medi- terior to ocellus with or without pits formally; pronotum black ventrally; propleu- ing V-shape (as in Fig. 7d); area between ron black or black and orange; mesopleu- tentorial pit and antennal insertion with ron mostly black to mostly orange; metan- line of foveae, rarely vestigial (Fig. 3d). otum orange or black and orange; propo- Mesosoma: Notauli foveate and complete deum black and orange; metapleuron (as in Fig. 5a); propodeum mildly rugose, black or black and orange; tarsi sometimes sometimes with medial furrow anteriorly, pale yellow; terga usually orange to dark long, dense setae throughout; sternaulus orange except black mottling often pre- foveate (as in Fig. 5a); mesepisternum sent; wings infumate to slightly infumate. Journal of Hymenoptera Research 104

6.8-8.2. Agathirsia sericans (Westwood) Body length: excluding ovipositor of anten- Head: seta at midlength Westwood 1882: 23. [Lecto- Longest Agathona sericans half or < half antenna na approximately type. Examined]. labial 1904: 129. width (as in Fig. llb-c); palpomere Agathirsia sericans Szepligeti shorter than 3 + 4; mandible 2 palpomeres to Distribution (Fig. lie).—Southwestern without indication of second tooth (as in south-central Mexico. fork length 0.45-0.51; glos- Fig. lc); glossa Females and males. —Color: Mostly black sa more than 2X length; glos- length galea as follows: antenna sometimes or- 1. 6-2.0 X except sa length 2.1-2.6; glossa length and labial ange basally; maxillary palpom- than foretibia length; malar space longer eres with some pale color; mesoscutum 0.48-0.51 X shorter than eye height; face and mid sometimes partly orange; fore or without median above clypeus with to black; hind fe- legs from entirely orange area anterior to ocel- longitudinal carinae; mur sometimes with small orange area lus with forming wide V-shape or with some hind pits distally; hind tibia orange; wide to 7d); vestigial V-shape (similar Fig. tarsus with varying degrees of orange; between tentorial and antennal in- area area pit first tergum with yellow or orange sertion without line of foveae. Mesosoma: basally; tergum 2 yellow basally; forewing com- Notauli not foveate (as in Fig. 5b), infumate but usually darker in distal half; plete or not complete; propodeum weakly hind wing slightly infumate. Body length: anterior Head: Lon- sculptured, depression along excluding ovipositor 10.0-11.6. mid- of antenna midline, setae present except along gest seta at midlength approx- or shorter line and central posterior area (as in Figs. imately half antenna width (as labial 2 9a,b); sternaulus foveate (as in Fig. 5a) in Fig. llb,c); palpomere subequal to combined of 3 + 4 in 4b); posteriorly, absent anteriorly; mesepister- length (as Fig. num without lobes between midcoxae; mandible with or without indication of tooth in fork basal lobe of claws distinct (as in Fig. 3a); second (as Figs. lb,c); glossa 0.38-0.47; less than number of thick, apically flattened apical length glossa length 2X 1.5-1.8; pegs on hind tibia 2-6; hind basitarsus galea length; glossa length as foreti- of fore- length 0.85-1 .2 X as long straight distally; 1-cu-a and 1-M glossa bia malar space 0.31-0.39 X shorter separated by a distance greater than length; wing with than eye height; face above clypeus 2 vein widths (as in Fig. 9c). Metasoma: or without median carinae; Posterior width of first tergum 0.69-0.84 X longitudinal area anterior to ocellus with pits forming shorter than length of first tergum; setae in 7d); area between ten- on 3 in V-shape (as Fig. tergum sparse distally (as Fig. 6d); without torial pit and antennal insertion tergum 3 with transverse groove; ovipos- line of foveae. Mesosoma: Notauli foveate itor length 4.4-5.8; ovipositor length 4.0- and complete (as in Fig. 5a); propodeum 6.1 X longer than hind basitarsus length. V- foveate to rugose, often with inverted Specimens examined.—Holotype: 9, Mex- shape anteriorly and carina extending ico, 72 (Coffin) (OXUM). Coffin # 405 Sept. from medial posterior end to center of 1840, nr. Chapultepec. (8$ 6) Homotypes: propodeum, setae present throughout or AEIC, ATAM, CNCI, OSUO, OXUM. glabrous along midline and /or posterior- Remarks.— similar to A. — me- Very proxima ly; sternaulus foveate (as in Fig. 5a); the characters in the are the ones key only sepisternum without lobes between mid- found to distinguish the two species. Also coxae; basal lobe of claws small to vesti- similar to A. and A. real—see fulvocastanea gial (as in Fig. 3b); number of thick, api- in 8- characters the key and remarks under cally flattened apical pegs on hind tibia A. fulvocastanea. 13; hind basitarsus tapered distally; 1-cu- Volume 13, Number 1, 2004 105 a and 1-M of forewing separated by a dis- dian longitudinal carinae; area anterior to tance greater than 2 vein widths (as in Fig. ocellus with pits forming distinct V-shape 9c). Metasoma: Posterior width of first ter- (Fig. 7d) or vestigial; area between tento- X gum 0.43-0.53 shorter than length of rial pit and antennal insertion without line first tergum; setae on tergum 3 dense on of foveae. Mesosoma: Notauli complete, distal 0.75 to throughout and colored yel- varying from smoothly impressed to fo- low except rarely white; tergum 3 with veate; propodeum rugose, setae present transverse groove distinct to absent; ovi- laterally; sternaulus foveate (as in Fig. 5a); positor length 1.0-1.6; ovipositor length mesepisternum with pair of lobes between .63-.87X shorter than hind basitarsus midcoxae (Fig. 5d); basal lobe of claws distinct in number of length. — (as Fig. 3a); thick, api- Specimens examined. Lectotype: 9, cally flattened apical pegs on hind tibia 6- Mexico, 76 Vi (Coffin) (OXUM): designated 14; hind basitarsus tapered distally; lcu-a by van Achterberg, 1980. Homotypes: and 1-M of forewing almost contiguous, (19 9(5) AEIC, ATAM, EMEC, CASC, separated by the width of a vein or less CNCI, LACM, MSUC, OSUO, OXUM, (Fig. 9c). Metasoma: Posterior width of first SEMC. tergum 0.85-1. 13 X as long as length of Remarks. —Similar to A. trichiosoma and first tergum; setae on tergum 3 sparse dis- — in A. asteropJiila see characters in the key. tally (as Fig. 6d); tergum 3 without transverse groove; ovipositor length 3.2- Agathirsia testacea Muesebeck 3.6; ovipositor length 2.4-3.0 X longer than hind basitarsus length. Agathirsia testacea Muesebeck 1927: 13. [Exam- Specimens examined. —Holotype: 9, ined]. — USA, Mesilla [New Mexico], xi.6 (USNM). Distribution (Fig. llj). Central Califor- Homotypes: (285 9(5) AEIC, AMNH, nia to southeast Texas south to central CASC, CNCI, EMEC, LACM, MSUC, Mexico. OSUO, SEMC, TAMU, UADE, UCDC, Females and males. —Color: Mostly orange USNM. — the except as follows: antenna black distally; Remarks. The lobes between mid- maxillary and labial palpomeres partly coxae are distinctive for this species (Fig. black; mesosoma may contain black areas; 5d). hind tibia usually black distally and hind A. testacea is the most commonly col- tarsus usually somewhat darkened; black lected species of Agathirsia and has one of mottling usually present on terga; fore the largest geographic ranges. The range and hind wings from completely infumate of collection dates extends well into to clear in basal half and infumate in distal spring, which is unusual for Agathirsia. cites the larva of Aeon tin half. Body length: excluding ovipositor 6.7- Bibby (1961) A. testacea 8.4 (one exceptional specimen 5.5). Head: crctata (Noctuidae) as a host of Longest setae at midlength of antenna < from Yuma Arizona, viii.19.1953. half antenna width (as in Fig. 7b); labial Agathirsia tiro Pucci and Sharkey, sp. n. palpomere 2 shorter than combined length — Texas of palpomeres 3 + 4; mandible with in- Distribution (Fig. Ilk). Southern and northeast Mexico. dication of second tooth (Fig. lb), rarely — distinct or weak indication; glossa fork Females. Color: Black and orange, or- follows: antenna length 0.12-0.20; glossa length less than ange except as black; 2X galea length; glossa length 0.43-0.53; maxillary and labial palpomeres partly fore glossa length 0.35-0.42 X shorter than for- black; head black; mesosoma black; hind tibia black etibia length; malar space 0.48-0.65X and midcoxa partly black; tarsus black shorter than eye height; face without me- to dark orange basally; hind Research 106 Journal of Hymenoptera

re- School or Carneton Co. [Cam- to orange; first tergum black or orange; Cemetery?], eron iv. 13.1956 maining terga with some black mottling; Co.?], (Beamers, Stephen, to infumate in Michener & forewing slightly infumate Rozen) (SEMC). in dis- Remarks.—The combination of basal half, clear to slightly infumate glossa hind and coloration makes this tal half but lighter than basal portion; length species in basal clear distinctive. wing slightly infumate half, half. —Recruit, —refer- to slightly infumate in distal Body Etymology. beginner 5.7-5.9. Head: to the collec- length: excluding ovipositor ring relatively early (spring) Longest seta at midlength of antenna > tion dates. half antenna width (as in Fig. 7a); labial trichiosoma (Cameron), n. palpomere 2 shorter than combined length Agathirsia comb. of 3 + 4; mandible with indication of second tooth in lb); fork (as Fig. glossa length Cenostomus trichiosomus Cameron 1905: 387. unknown; less than 2X glossa length galea [Examined]. 0.25; length; glossa length approximately Agathis trichiosoma Shenefelt 1970: 362. glossa length approximately 0.22-0.32 X — to shorter than foretibia length; malar space Distribution (Fig. llf). Southwestern 0.46-0.52 X shorter than eye height; face south-central Mexico.— above clypeus without median longitudi- Females and males. Color: Mostly black nal carinae; area anterior to ocellus with except as follows: maxillary and labial pal- or without pits forming V-shape (as in Fig. pomeres partly pale; head black; mesoso- 7d); area between tentorial pit and anten- ma black; foretibia partly pale; foretarsus nal insertion without line of foveae. Me- pale yellow to orange to fuscous; midtibia sosoma: Notauli weakly foveate basally, partly pale yellow; midtarsus sometimes barely impressed as smooth groove else- orange to pale yellow; hind tibia pale yel- where; propodeum largely smooth medi- low/orange basally extending distally on ally, foveate elsewhere, setae present ex- lateral side; hind tarsus sometimes with cept along midline; sternaulus foveate (as orange or pale yellow; first tergum some- in Fig. 5a); mesepisternum without lobes times with small yellow /orange area ba- between midcoxae; basal lobe of claws sally; tergum 2 with some yellow or or- vestigial (as in Fig. 3b); number of thick, ange basally; forewing slightly infumate apically flattened apical pegs on hind tibia in basal half, infumate in distal half; hind hind 2-4; basitarsus straight distally; 1-cu- wing slightly infumate. Body length: ex- a and 1-M of forewing separated by a dis- cluding ovipositor 8.5-12.0. Head: Longest tance than 2 greater vein widths (as in Fig. seta at midlength of antenna subequal to Metasoma: first 9c). Posterior width of ter- 0.5-0.75 antenna width (as in Figs. lla,c); gum 0.73-0.87X shorter than length of labial palpomere 2 subequal to palpom- first setae on 3 tergum; tergum sparse dis- eres 3 + 4 (as in Fig. 4b); mandible within 3 tally (as Fig. 6d); tergum without out indication of second tooth (as in Fig. transverse groove; ovipositor length 1.2- lc) or weak; glossa fork length 0.17-0.23; 1.5-1.6X 1.3; ovipositor length longer than glossa length less than 2x galea length; hind basitarsus length. glossa length 0.60-0.76; glossa length 0.40- Males. —Unknown. 0.51 X shorter than foretibia length; malar examined. — Specimens Holotype. 9, Mex- space 0.22-0.27 X shorter than eye height; Nuevo 50 mi. ico, Leon, S Nuevo Laredo, face above clypeus with or without me- ii.22.1972 (Parker & Miller) (AEIC). Para- dian longitudinal carinae; area anterior to 29, same data as types: holotype (AEIC); ocellus without pits forming distinct or USA: Texas: 19, Del iv.18.1984 Rio, (Bo- vestigial V-shape (as in Fig. 7d); area be- Southmost (UCDC); 19, [Ranch, tween tentorial pit and antennal insertion Volume 13, Number 1, 2004 107 without line of foveae. Mesosoma: Notauli mous reviewer for their critical reviews of the manuscript. Research was supported by NSF grants DEB- foveate and complete (as in Fig. 5a); pro- 9972024 and DEB-0205982. Support was also provid- rugose, longitudinal furrow an- podeum ed through Kentucky Agriculture Experiment num- teriorly, long, dense setae throughout; ber 02-08-164. sternaulus foveate (as in Fig. 5a); mesepis- LITERATURE CITED ternum without lobes between midcoxae; C. 1990. Revision of the subtribe Me- basal lobe of claws distinct (as in Fig. 3a); Achterberg, van, socoelina Viereck Braconidae). number of thick, apically flattened apical (Hymenoptera: Zoologische Mededelingen (Leiden) 64: 31-57. pegs on hind tibia 6-10; hind basitarsus Arnett, R. H., G. A. Samualson, and G. M. Nishida. 1-cu-a and 1-M of fore- tapered distally; 1993. The insect and spider collections of the world. wing separated by a distance greater than Sandhill Crane Press. Gainesville, FL. G. R. and M. 1988. Abdom- 2 vein widths (as in Fig. 9c). Metasoma: Buckingham J. Sharkey. inal exocrine in Braconidae Posterior width of first tergum 0.43-0.59 X glands (Hymenop- tera). Adimnces in Parasitic Hymenoptera Research. shorter than of first setae length tergum; 1: 199-242. on 3 dense on distal 0.66-0.75; ter- tergum Dallwitz, M. J., Paine, T. A., and E. J. Zurcher. 1993. 3 transverse gum with groove; ovipositor User's guide to the DELTA system: a general system length 0.87-0.97; ovipositor length 0.54- for processing taxonomic descriptions. 4th edition, uno.edu/delta/ 0.78 X shorter than hind basitarsus length. http: //biodiversity. Harder, L. D. 1983. Functional differences of the examined. — pro- Specimens Holotype: 9 , Mex- boscides of short- and long-tongued bees (Hy- ico, B.M. HYM. 3.C.974 Homo- (BMNH). menoptera, Apoidea). Canadian Journal of Zoology. types: (23 9 6) AEIC, ATAM, BMNH, 61: 1580-1586. CASC, CNCI, MSUC, SEMC, UADE, Jervis, M. A. 1998. Functional and evolutionary as- UCDC, USNM. pects of mouthpart structure in parasitoid wasps. Journal of the Linnaean 63: 461- Remarks.—Similar to A. sericans—see Biological Society. 493. characters in the key. Muesebeck, C. F. W. 1927. A revision of the parasitic We recognize two forms that may be wasps of the subfamily Braconinae occurring in America north of Mexico. the Unit- separate species. None of the following Proceedings of ed States National Museum. 69: 1-73. characters used to distinguish them are Bassus Fa- Sharkey, M. J. 1985. Notes on the genera without exceptions and the putative forms bricius and Agathis Latreille, with a description in time and I (holo- overlap space. Type of Bassus arthurellus n. sp. (Hymenoptera: Bra- type form): tergum 2 completely yellow or conidae). The Canadian Entomologist. 117: 1497- orange behind gently sloping groove, 1502. M. 1992. Cladistics and tribal classification black hind basitarsus, median body length Sharkey, J. of the Agathidinae (Hymenoptera: Braconidae). 10.8. Type II: tergum 2 partially yellow or Journal of Natural History. 26: 425-447. behind 1997. orange V-shaped groove, pale yel- Sharkev, M. J. Subfamily Agathidinae, pp. 69- low hind basitarsus, median body length 84. />;: Manual of the New World Genera of the 9.2. Family Braconidae. R.A. Wharton, P.M. Marsh, and M.J. Sharkey (Eds.). Special Publication of the International Number ACKNOWLEDGMENTS Society of Hymenopterists. 1, Washington D.C. R. 1997. We thank all the curators from the institutions list- Sharkey, M. J. and A. Wharton. Morphology 19-38. /;/: Manual of the ed previously, Peter Southgate who helped with most and terminology. Pp. the Braconidae. of the SEM images, Dan Crowdus who helped with New World Genera of Family R.A. P.M. and Sharkev many technical aspects of the manuscript and wrote Wharton, Marsh, M.J. a program to create the datasets and automate the (Eds.). Special Publication of the International Soci- Number repeated PAUP analyses that incorporated the ran- ety of Hymenopterists, 1, Washington domized characters, Mads Haahr for creating the true D.C. D. L. 2002. Us- random number generator used (from www. random, Swofford, PAUP* Phylogenetic Analysis Other Methods) Version 4.0 org), Chris O'toole for information on Westwood ing Parsimony (*and blO. Sinauer Massachu- types, Don LaFontaine for host information, and Eric Associates, Sunderland, Grissell, Mark Jervis, Donald Quicke, and an anony- setts. J. HYM. RES. Vol. 13(1), 2004, pp. 108-119

Two New Species of Quadrastichns Girault (Hymenoptera: citrella Eulophidae): Parasitoids of the Leafminers Phyllocnistis Stainton (Lepidoptera: Gracillariidae) and Liriomyza trifolii (Burgess) (Diptera: Agromyzidae)

Placido Reina and John La Salle

of via S. 98- (PR) Dipartimento di Scienze e Tecnologie Fitosanitarie, University Catania, Sofia, 95124 Catania, Italy; email: [email protected]; (PR, JLS) CSIRO Entomology, GPO Box 1700, Canberra, ACT 2601, Australia; email: [email protected]

La Salle and Reina Abstract.—The parasitoids Quadrastichus citrella Reina and Q. plaquoi and La Salle (Hymenoptera: Eulophidae) are described. Quadrastichus citrella, native of South-East citrella Stainton Asia, has been used as a biological control agent of the citrus leafminer Phyllocnistis the has invaded. (Lepidoptera: Gracillariidae) in several countries that pest recently Quadrastichus of Characters plaquoi, from India, is a parasitoid Liriomyza trifolii (Burgess) (Diptera: Agromyzidae). of the of these new species are discussed, as well as their relationship to other members "anysis are group" of species. Distributional and biological details of these wasps given.

Leafmining insects can be severe pests SW USA to South America in the last de- that reduce plant metabolic activities and cade (Hoy and Nguyen 1997, Gates et al. can lead to desiccation, and premature fall 2002). of the leaves. If leaves are seriously at- In newly infested citrus-growing re- tacked, crops can be reduced or seedling gions, the CLM indigenous antagonists plants even totally destroyed (Spencer (mainly parasitoids) have never been able 1990). The leafmining habit is found in to reduce damage below economic thresh- several species of Lepidoptera and Diptera old, as reported from Florida (Hoy and and also in some Coleoptera and Hyme- Nguyen 1997), Israel (Argov and Rossler noptera. The main antagonists of leafmi- 1996), Italy (Siscaro et al. 2003), Spain ners are parasitic Hymenoptera, especially (Garrido Vivas 1995) and Turkey (Uygun those belonging to Chalcidoidea, which et al. 1996). However in the native range, penetrate the mines with their ovipositors pest population can be controlled below to lay eggs in or on the body of the mining the economic threshold by natural ene- larvae or to feed on the host's body fluids mies, which represent its main biological (Askew and Shaw 1974). mortality factor (Morakote and Nanta Among Lepidoptera, the citrus leafmi- 1996, Tan and Huang 1996, Wang et al. ner (CLM) Phyllocnistis citrella Stainton 1999). Therefore, in order to achieve nat- is an (Gracillariidae) important pest which ural control of this pest, several parasit- has recently spread throughout all citrus oids have been introduced to the newly areas around the world. This species, na- infested countries from the native range of five of South-East Asia, entered Africa and P. citrella. Among these, Quadrastichus ci- Australia in the 1900's early and has trella sp.n. (Hymenoptera: Eulophidae) the Mediterranean Ba- spread throughout has been used in Cyprus, Israel, Italy, Mo- md in the New World from SE and rocco and Spain (as Q. sp. "A" in Schauff Volume 13, Number 1, 2004 109 et al. 1998; as Q. sp. in Smaili et al. 1999, might be used in future sustainable con- Argov 2000, Barbagallo et al. 2000). How- trol programs. ever, the only evidence of establishment Species of Quadrastichus are often en- for this species is in Spain, where the spe- doparasites of Cecidomyiidae (Diptera); cies overwintered in the Valencia area less commonly, they parasitize Cynipidae (Garcia Mari et al. 2000). (Hymenoptera), Buprestidae and Curcu- Among Diptera, agromyzid leafminers, lionidae (Coleoptera), Agromyzidae and and particularly the species belonging to Tephritidae (Diptera). Also, Q. sajoi (Sze- the genus Liriomyza Mik, are considered lenyi) larvae are predatory of eriophid damaging pests of numerous vegetable mites within galls (Graham 1991, La Salle and floricultural crops throught the world. 1994, Hansson and La Salle 1996). Several of these species can cause exten- Within the Tetrastichinae, QuadrasticJius sive economic damage to a large range of is characterized by having the following host plants under both field and green- characters: a single seta on submarginal house conditions (Spencer 1989). Knowl- vein; mesoscutum usually with a single edge about agromyzid natural enemies adnotaular seta; propodeum without Y- has become increasingly important as a shaped paraspicular carina; cereal setae key element to biological control strategies unequal in length, with one being distinct- of these pests. A large number of parasit- ly longer and sinuate; antenna with all fu- oids have been recorded in the New and nicular segments longer than wide; scu- Old World, especially species from the tellum with submedian and sublateral families Eulophidae and Pteromalidae lines; propodeal spiracles close or fairly (Chalcidoidea) and, less commonly, spe- close to metanotum, with their rim ex- cies of the families Braconidae (Ichneu- posed; ovipositor sheaths not or only monoidea) and Eucoilidae (Cynipoidea) slightly projecting beyond last tergite of (Konishi 1998, Murphy and La Salle 1999). gaster (Graham 1991). Nevertheless, the majority of these species This genus was treated under the name are generalists and care must be taken Cecidotetrastichus Kostjukov (Kostjukov when deciding to introduce exotic natural 1977, Graham 1987), but Boucek (1988: enemies. Biological control strategies ap- 677) remarked on the similarity between propriate for agromyzid leafminers in Quadrastichus and Cecidotetrastichus and field vegetables often include the intro- Graham and La Salle (1991) placed Ceci- duction of appropriate exotic natural en- dotetrastichus in synonymy with Quadras- emies or conservation and enhancement tichus. Discussions on differentiating of local natural enemies. However, these Quadrastichus from related genera, such as strategies are not mutually exclusive, as it Aprostocetus Westwood, Citrostichus Bou- is clear that any introductions should take cek, Oomyzus Rondani and Tetrastichus into account the existing local natural en- Haliday in particular, are available in lit- emy community (La Salle 1993, La Salle erature (Graham 1987, 1991, La Salle 1994, and Gauld 1993). Recently, Murphy and Schauff et al. 1998, Reina and La Salle La Salle (1999) recommended that, due to 2003). At the moment, keys to Quadrasti- the prevalence and often general nature of cJius species are available only for Europe leafminer parasitoids, effort should be put (Graham 1991). into and understanding conserving indig- ABBREVIATIONS enous leafminer parasitoids rather than re- Insect Col- lying solely on the introduction of exotic ANIC, Australian National parasitoids. Quadrastichus plaquoi sp.n. lection, CSIRO Entomology, Canberra, L. in The Natural found parasitising trifolii (Burgess) Australia; BMNH, History India is an example of a species which Museum, London, UK; DISTEF, Diparti- of Hymenoptera Research 110 Journal

First funicular mento di Scienze e Tecnologie Fitosanitar- One anellus. segment De- shorter than other two which are ie, University of Catania, Italy; EMBT, slightly Thai- in about 0.7-0.8 X as partment of Agriculture, Bangkok, subequal length (Fl F2 or F3 Club land; INPC, National Pusa Collections, In- long as length). 3-segment- 5.0-5.5 X as as wide, lon- dian Agriculture Research Institute, New ed, long slightly than F2 and F3 combined; distinct ter- Delhi, Haryana, India; IZCAS, Institute of ger minal and suture between Zoology, Chinese Academy of Sciences, spine oblique National Museum 2 Mesosoma (Figs. 6-7): Beijing, China; USNM, apical segments. Pronotum of Natural History, Washington D.C., Slightly sculptured. medially USA. about 0.3 X as long as mesoscutum. Mid- lobe of mesoscutum 1.5-1.6X longer than Reina and La Quadrastichus citrella scutellum, with very weak to indistinct nov. Salle, sp. median line and single adnotaular seta 2, 4-11, 14, 15) (Figs. 1, placed in posterior half. Scutellum about — with 0.6 X as as broad. Dorsellum Diagnosis. Body mainly yellowish long very metanotum brown, transverse dark band short (0.2-0.3 X as long as broad), not ex- th with over 0.2- on 3-4 gastral segments. Frons tending posteriorly propodeum, broad median area rather than distinct 0.3 X as long as scutellum. Propodeum median line. Malar sulcus curved, without more strongly sculptured than thorax and distinct median carina and distinct fovea beneath eye. Antennal scape not with encloses a con- reaching above top of vertex. First funic- paraspicular carina which area Fore ular segment slightly shorter than other cave surrounding spiracle. wing 2-2.2 X as as broad. Sub- two which are subequal in length. Midlo- (Fig. 10): long be of mesoscutum with single adnotaular marginal vein with a single dorsal seta. X than seta placed in posterior half. Anterior seta Marginal vein 6.2 longer stigmal on scutellum longer than posterior one. vein. Postmarginal vein present, about Dorsellum very short (0.2-0.3 X as long as 0.9 X as long as stigmal vein. Fringe on broad). Propodeum well sculptured, with marginal vein 0.1-0.2X width of wing. distinct median carina and anterior mar- Speculum present and extending below gin not covered by dorsellum. Forewing marginal vein for 0.2-0.3 X its length; bare with distinct speculum and bare area be- area extending behind full length of mar- hind marginal vein. ginal vein. Metasoma: Ovate and slightly Female. —Length 0.9-1. 3mm. Body shorter than head and mesosoma com- mainly yellowish (Fig. 1); following parts bined. Cercus (Fig. 8) with one seta dis- brown to black: ocellar triangle, basal por- tinctly longer than the remaining setae tion of pedicel (about half pedicel length), and sinuate. flagellar segments, mesosternum, metan- Pupa.—Yellow-orange with dark longi- th otum, transverse band on 3^4 gastral seg- tudinal stripe for entire length (Fig. 2). ments. Head (Fig. 4): Frons with broad me- Meconium usually can be found outside dian area rather than distinct median line. the pupa near the caudal region. Malar space about 0.5 X eye height. Malar Male. —Length 0.7-1. lmm. Similar to fe- sulcus (Fig. 5) without fovea, distinctly male except darker markings: pronotum, curved especially beneath eye. Anterior axilla partially and costula, mesosternum, of margin clypeus slightly bilobate, with- mesopleura, mesoscutum especially in an- out distinct teeth. Mandible 3-dentate. An- terior half, propodeum, dorsellum and fi- tenna (Fig. 14): Torulus placed slightly nally gaster in posterior half. Fore wing: above lowest margin of eye. Scape 5.0- Fringe distinct, setae on marginal vein 5.5 X as as than about 0.3X long wide, slightly longer width of wing (Fig. 11). Anten- and the of vertex. height, reaching top na (Fig. 15): Basal whorls of setae present Volume 13, Number 1, 2004 111

*t y

Figs. 1-3. 1-2, Quadrastichus citrella, female. 1, Ovipositing on Phyllocnistis citrella larva. 2, Pupa. 3, Quadras- tichus plaquoi, female. of Hymenoptera Research 112 Journal

Figs. 4-9. 4-8, Quadrastichus citrella, female. 4, Head, frontal view. 5, Malar space. 6, Thorax. 7, Propodeum. 8, Cereal setae. 9, Quadrastichus citrella, male, scape. Volume 13, Number 1, 2004 113

on all funicular at X segments and least 0.5 guished from both Quadrastichus liriomyzae as as Funicle long flagellum length. seg- Hansson and La Salle and Q. plaquoi by ments ratio F1/F2/F3/F4: 5/12/15/16. having dorsellum very short (0.2-0.3 X as Club slightly longer than Fl and F2 com- long as broad), fore wing with a distinct bined. Ventral plaque (Fig. 9) on scape speculum (extending below marginal vein small ovate, very (about 0.1-0.2X as long for 0.2-0.3X its length) and the area be- as scape), and placed about slightly above hind marginal vein asetose for its entire the middle. length; males can also be recognized by —This is Etymology. species named for having a distinctly smaller plaque on the specific name of its host, Phyllocnistis scape: it is oval in shape, about 0. 1-0.2 X citrella Stainton. as as and — long scape length, placed slight- Distribution. Quadrastichus citrella is rely above the middle. Quadrastichus lirio- corded as a native of China, Japan, Taiwan myzae and Q. plaquoi have a longer dor- and Thailand (as Q. sp. "A" in Schauff et sellum (at least 0.4 X as long as broad), al. 1998). — fore wings have speculum almost indis- Biology. Quadrastichus citrella is an tinct, the area behind marginal vein setose idiobiont ectoparasitoid of second and for its entire length, and males have also third citrella Phyllocnistis instar larvae, the a longer ventral plaque on the scape: only host recognized by now. Its devel- about 0.6 (liriomyzae) or 0.7 (plaquoi) X as opmental cycle takes about 20 days at long as scape length. 20°C and R.H.>80%; at the same temper- Quadrastichus plaquoi Reina and La ature, the adults survive up to 40 days (as S - Salle ' P' nov Q. sp. in Argov and Rossler 1998, Llacer

Fi s - ( 3 ' 12 ' 13 ' 16-22) et al. 1998). — 8 Type material. Holotype 9: Thailand, Diagnosis. —Body yellowish with dark 1996, Y. Rossler, ex Phyllocnistis citrella spot in middle of pronotum and with th Stainton (BMNH). Paratypes 22d, 319, transverse dark band on 4 gastral seg- deposited as follows: same data as holo- ment. Frons with broad median area rath- type (Id, 59 ANIC; Id, 39 BMNH; Id, er than distinct median line. Malar sulcus 59 DISTEF; Id, 39 EMBT; Id, 19 INPC; curved and without fovea beneath eye. 19 IZCAS;ld,2 9 USNM); Israel, Bet Da- Antennal scape reaching slightly higher gan, Jaffa Corp., first delivery from Israel than top of vertex. First and second funic- to iv. Catania (Italy), 18. 1996, coll. by E. ular segments equal in length. Midlobe of Swirski, ex P. citrella (26, 19 ANIC; 28, mesoscutum with single adnotaular seta 1 9 DISTEF); Italy, University of Catania, placed in posterior half. Anterior seta on mass rearing on P. citrella, 26.vi.1996, coll. scutellum longer than posterior one. Pro- by G. Siscaro (26, 29 ANIC; 26, 19 podeum quite shiny, without a distinct BMNH; 4d,39 DISTEF; 2d, 19 EMBT; median carina. Dorsellum rounded pos- ld, 19 IZCAS; Id, 19 USNM). teriorly, at least 4.0X as long as broad. Comments. —Several specimens (with Forewing with speculum very small, area the same data as the type material) have just distal to basal vein almost completely been examined and the only variation we covered with setae, and area behind mar- could recognize is with the coloration on ginal vein—setose. mesoscutum and gaster. The anterior half Female. Length 0.9-1. 4mm. Body of midlobe of mesoscutum can sometimes mainly yellowish (Fig. 3) with following have some brownish markings, while the parts brown to black: basal portion of ped- th transverse band on 3-4 gastral segments icel (about 0.5 its length), ocellar triangle, may rarely be either larger or indistinct. flagellar segments, medial 0.3-0.4 of pron- Quadrastichus citrella may be distin- otum, mesoscutum anteriorly, median 114 Journal of Hymenoptera Research

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ISj Volume 13, Number 1, 2004 115 area on the propodeum, and transversal male but with darker markings as follows: 4 th band on gastral segment. Head (Fig. 18): anterior half of mesoscutum, metanotum, Frons without a distinct median line and dorsellum, propodeum and almost entire with broad median area. Malar only space basal half of gaster. Fore wing with fringe about 0.6 X as as Malar long eye height. distinct, and setae on marginal vein about sulcus 19) curved and 0.3 X (Fig. distinctly width of wing (Fig. 13). Antenna (Fig. without fovea. truncate anterior- Clypeus 17): Basal whorls of setae present on all without distinct teeth. Mandible ly, 3-den- the funicular segments and about 0.4-0.5 X tate. Antenna Torulus (Fig. 16): placed as long as flagellum length. Funicular seg- slightly above lowest eye margin. Scape ments ratio F1/F2/F3/F4: 6/10/11/9. 5x as as wide, long reaching slightly Club longer than F2 and F3 combined, above of vertex. One anellus top present. which are longest funicular segments. First and second funicular segments equal Ventral plaque (Fig. 22) on scape placed in length, third one about 0.8 X as as long medially and large: about 0.7x long as other two. Club 4.0-4.5 X as 3-segmented, scape length. long as wide, longer than Fl and F2 com- — Etymology. Males of this species have a bined, with distinct terminal spine and su- large ventral plaque on the scape in com- ture between apical 2 segments slightly parison with the much smaller one pre- oblique. Mesosoma (Figs. 20, 21): Pronotum sent in Q. citrella; therefore, the name pla- uniformly lineolate, medially 0.2-0.3 X as qnoi refers to this character. long as mesoscutum. Midlobe of mesos- Distribution. —Known only from Hima- cutum 1.6-1.7X slightly sculptured, longer chal Pradesh, India. than scutellum, and with a very weak to — Biology. Quadrastichus plaqnoi has been indistinct median line; single adnotaular recorded only on Liriomyza trifolii (Bur- seta present in posterior half. Scutellum gess). about 0.7X as long as broad. Dorsellum at Type material. —Holotype 9: India, Su- least 0.4 X as long as broad, evenly round- lam, Himachal Pradesh, 2000, ex Liriomyza ed posteriorly, 0.3-0.4 X as long as scutel- trifolii (Burgess) (BMNH). Paratypes 5cT, lum and slightly extending posteriorly 9$, deposited as follows: data as holotype over propodeum. Propodeum shiny and (IcT, 39 ANIC; 16, 19 BMNH, 2 9 DIS- without distinct median carina; distinct TEF;1<3,19 INPC;1<5,19 IZCAS; 1^,19 paraspicular carina present which enclos- USNM). es a concave area associated with spiracle. Comments.—No distinguishable varia- Fore wing (Fig. 12): 2.2-2.4X as long as tion has been recognized from the exam- broad. Submarginal vein with single dor- ined material. sal seta. Marginal vein about 4x as long It be difficult to Q. as stigmal vein. Postmarginal vein slightly may distinguish pla- from The fe- shorter than stigmal vein: about 0.9 X as qnoi Q. liriomyzae. Q. plaqnoi male has Fl and F2 in and long as stigmal vein. Fringe on marginal equal length above the of vein 0.2-0.3 X as long as wing width. Spec- scape reaching slightly apex vertex; moreover, both sexes have ulum very small, area just distal to basal pronotum with a median dark vein almost completely covered with se- mainly yellow area. female Fl tae, and area behind marginal vein setose. Q. liriomyzae has slightlv Metasoma: Oval in shape in dorsal view shorter than F2, scape not reaching above the of vertex and both sexes have and slightly shorter than head and meso- apex soma together. Cercus with one seta dis- pronotum dorsally entirely dark. Q. pla- tinctly longer than remaining setae and qnoi can be distinguished from Q. citrella sinuate. Tip of ovipositor slightly exserted. as suggested above in Q. citrella descrip- Male.—Length 0.7-0.9mm. Similar to fe- tion, and in the key below. 116 Journal of Hymenoptera Research

Figs. 18-22. 18-21, Quadrastichus female. plaquoi, 18, Head, frontal view. 19, Malar space. 20, Thorax. 21, Propodeum. 22, Quadrastichus plaquoi, male, scape. Volume 13, Number 1, 2004 117

DISCUSSION not foveate, pronotum uniformly sculptured and scutellum without an offset Within the genus, Q. citrella and Q. pla- strip along its hind edge. Other species quoi belong to the (see Gra- "anysis group" within this group are the European Q. ci- ham with which share exten- 1991), they trinus (Thomson) and Q. xanthosoma (Gra- sive on the frons yellow markings body, ham), the North American Q. flora (Gi- with a median area rather a usually than rault) and the Asian Q. liricvin/zae Hansson median carina, malar sulcus curved and and La Salle.

KEY TO OLD WORLD SPECIES OF THE "ANYSIS GROUP" OF QUADRASTICHUS

1. Both sexes: mesoscutum completely dark brown to black; speculum distinct and extending almost half of vein . marginal length. Male: ventral plaque about 0.5 x scape length . . Q. anysis (Walker) - Both sexes: mesoscutum to partially' totally yellowish. Other characters variable 2 2. Female: Fl > 1.3X F2 3 - Female: Fl < F2 4 3. Female: 1.5X than hind gaster longer tibia; thorax and gaster mainly blackish; last tergite 0.8-1.5X as long as broad (see Graham 1974, 1991) Q. citrimis (Thomson) - Female: 2.0 X gaster longer than hind tibia; thorax and gaster mainly yellow; last tergite 1. 3-2.0 X as long as broad (see Graham 1974, 1991) Q. xanthosoma (Graham) = 4. Both sexes: pronotum mainly yellow with a dark spot medially. Female: Fl F2. Male: ventral plaque (Fig. 22) placed medially and about 0.7X long as scape length Q. plaquoi, n. sp. - Fl < Female: F2; pronotum different, either entirely dark dorsally (lirionn/zae) or entirely yellow (citrella). Male: pronotum entirely dark; ventral plaque on scape variable 5 5. Both sexes: speculum indistinct, area just distal to basal vein almost completely setose; area behind marginal vein setose for entire its length. Female: pronotum dark dorsally. Male: ventral plaque on scape about 0.6X as long as scape length Q. liriomyzae Hansson and La Salle - Both sexes: speculum present and extending below marginal vein for 0.2-0.3 X its length; area behind marginal vein asetose for entire its length. Female: pronotum yellow dorsally. Male: ventral plaque (Fig. 9) about 0.1-0.2X as long as scape length Q. citrella, n. sp.

ACKNOWLEDGMENTS Argov, Y. and Y. Rossler. 1998. Rearing Methods for the Citrus Leafminer, Phyllocnistis citrella Stain- We are particularly grateful to Yoram Rossler of ton and Its Parasitoids in Israel. Biological control the Israel Cohen Institute for who Biological Control, 11: 18-21. kindly gave us specimens of Q. citrella from Thailand. Argov, Y. 2000. Biological control of Phyllocnistis ci- We also thank Gaetano Siscaro of DISTEF, who pro- trella in Israel. Proceedings of International Society vided us with both specimens of Q. citrella and pic- of Citriculture, Florida 2000, Poster n°48. tures of the and female. We also wish to thank pupa Askew, R. R. and M. R. Shaw. 1974. An account of Suzanne Lewis and John (BMNH) for lending Noyes the Chalcidoidea (Hymenoptera) parasitising us Q. lirionn/zae paratypes. leaf-mining insects of deciduous trees in Britain. LITERATURE CITED Biological journal of the l.innean Society 6: 289-335. Barbagallo, S., S. Longo, G. Siscaro, P. Reina and L. Argov, Y. and Y. Rossler. 1996. Introduction, release Zappala. 2000. Status of biological control of the and recovery of several exotic natural enemies Citrus leafminer (Phyllocnistis citrella Stainton) for biological control of the citrus leafminer, in Italy. Proceedings of the XXI International Con- Phyllocnistis citrella, in Israel. Phytoparasitica 24: gress of Entomology, Brazil, August 20-26, 2000, 33-38. 1:375. 118 Journal of Hymenoptera Research

1993. Parasitic Boucek, Z. 1988. Australasian Chalcidoidea (Hyme- La Salle, J. Hymenoptera, biological of of control and In: and Bio- noptera). A Biosystematic Revision Genera biodiversity, Hymenoptera and I.G. Gauld Fourteen Families, with a Reclassification of diversity (La Salle, J. (eds), Walling- ford, CAB International: 197-215. species. CAB International, Wallingford, Oxon, UK; La 1994. North American of Tetrasti- 832 pp. Salle, J. genera C. chinae Garcia Mari, F., R. Vercher, J. Costa Comelles, (Hymenoptera: Eulophidae). Journal of Granda, C. Marzal, C. Alfaro, D. Castrillon, V. Natural History 28: 109-236.

and I. D. Gauld. 1993. their Bueno and M. Villalba. 2000. Importation and La Salle, J. Hymenoptera: their on the of oth- establishment of exotic parasitoids for the biodiversity and impact diversity er In: and (La logical control of the Citrus Leaf Miner Phylloc- organisms. Hymenoptera Biodiversity in and I.G. Gauld nistis citrella (Lepidoptera: Gracillariidae) Salle, J. (eds), Wallingford, UK; Spanish citrus orchards. Proceedings of the Inter- CAB International: 1-26. A. and A. Garrido. 1998. national Society ofCitricidture, Florida 2000 Poster Llacer, E., Urbaneja, J. Jacas n. 54. Ciclo biologico de Quadrastichus sp., parasitoide Garrido Vivas, A. 1995. Pln/llocnistis citrella Stainton, exotico del minador de las hojas de los citricos citrella en laboratorio. Bol- aspectos biologicos y enemigos naturales encon- Pln/llocnistis Stainton,

I etin de Sanidad 24 669-678. trados en Espana. Levante Agricola, trim.: 13-21. Vegetal, Plagas (4): E. D. L. R and. P. Nanta. 1996. the citrus Gates, M. W., J. M. Heraty, M. Schauff, Wag- Morakote, Managing B. D. B. Wahl. 2002. leafminer in Thailand. the Interna- ner, J. Whitfield and Survey Proceedings of of the Parasitic Hymenoptera on Leafminers in tional Conference "Managing the Citrus Leafminer", 1996: 30-33. California. Journal of Hymenoptera Research 11 (2): Orlando, Florida, April 23-25, S. Salle. 1999. 213-270. Murphy, and J. La Balancing biological Graham, M. W. R. de V. 1974. A new species of Te- control strategies in the IPM of New World in- trastichus Walker (Hym., Eulophidae) from Brit- vasive Liriomyza leafminers in field vegetable ain. Entomologist's Gazette 25 (2): 106-110. crops. Biocontrol Neivs and Information 20, 91N- Graham, M. W. R. de V. 1987. A reclassification of the 104N. to European Tetrastichinae (Hymenoptera: Eulo- Reina, P and J. La Salle. 2003. Key the World Gen- phidae), with a revision of certain genera. Bullet- era of Eulophidae Parasitoids (Hymenoptera) of tin of the British Museum (Natural History). Ento- Leafmining Agromyzidae (Diptera). http:// mology Series, 55: 392 pp. www.ento.csiro.au/science/eulophid-key/eulo- Graham, M. W. R. de V. 1991. A reclassification of the phids.htm Tetrastichinae European (Hymenoptera: Eulo- Schauff, M.E., J. La Salle and G.A. Wijesekara. 1998. phidae): revision of the remaining genera. Mem- The Genera of Chalcid Parasitoids (Hymenop- oirs of the American Entomological Institute, 49: 1- tera: Chalcidoidea) of Citrus Leafminer Pln/lloc- 322. nistis citrella Stainton (Lepidoptera: Gracillari- M. W. R. de V. La Salle. Graham, and J. 1991. New idae). Journal of Natural History 32: 1001-1056. synonymy in European Tetrastichinae (Hyme- Siscaro, G., V. Caleca, P. Reina, M.C. Rizzo and L. noptera: Eulophidae), including designation of Zappala. 2003. Current status of the biological some and neotypes, lectotypes, new combina- control of the citrus leafminer in Sicily. Proceed- tions. Entomologist's Gazette 42: 89-96. ings of Citrus Working Group "Integrated control C. and La Salle. 1996. Hansson, J. Two new eulophid in citrus fruit crops", Valencia (Spain) 6-8 Novem- parasitoids (Hymenoptaera: Chalcidoidea) of Lir- ber 2002. OILB/SROP, Bulletin de V Organisation iomyza trifolii (Burgess) (Diptera: Agromyzidae). Internationale Lutte Biologique et Integree, 26(6): Oriental Insects 30: 193-202. 29-36. M. A. and R. Hoy, Nguyen. 1997. Classical biological Smaili, C, M. Afellah, A. Aarab and L. Zrida. 1999. control of the Citrus Leafminer ci- Pln/llocnistis Biologie et ecologie de Pln/llocnistis citrella (Lep.: trella Stainton (Lepidoptera: Gracillariidae). Trop- Gracillariidae) et initiation de la lutte biologique ical Lepidoptera 8 (Suppl. 1): 1-19. sur clementinier dans le Gharb au Maroc. Pro- Konishi, K. 1998. An illustrated to the key hymenop- ceedings of 51st International Symposium on crop terous of in parasitoids Liriomyza trifolii Japan. protection, Gent, Belgium, 64 (3a): 121-131. Miscellaneous Publication of the National Institute of Spencer, K.A. 1989. Leafminers. In Kahn, P.R. (ed) Sciences 22: 27-76. Agro-Environmental [In Japa- Plant protection and quarantine Vol. II. Selected nese]. pests and pathogens of quarantine significance. Boca V. V. Kostjukov, 1977. Stravnitel'naya morfologiya Raton, FL, USA; CRC Press: 77-98.

khal'tsid Tetrastichinae i sistema roda podsem. Spencer, K.A. 1990. Host specialization in the world Tetrastichus 1844 Eulo- Haliday (Hymenoptera, agromyzidae (Diptera). Khmer Academic Publish- phidae). Entomologicheskoe Obozreniye 56: 177- ers, 444 pp. 194. [In Russian] Tan, B. and M. Huang. 1996. Managing the citrus Volume 13, Number 1, 2004 119

leafminer in China. the International Proceedings of ternational Conference "Managing the Citrus Leaf- Conference "Managing the Citrus Leafminer", Or- miner", Orlando, Florida, April 23-25, 1996: 101. Florida, 1996: 49-52. Undo, April 23-25, Wang, L.D., M.S. You, J. Wang and Q. Wu. 1999. Uygun, N., M. Kersting, N.Z. Elekcioglu, I. Karaca, Niches of citrus leafminer and its natural ene- R. L. Yumruktepe and Erkilic. 1996. Status of the mies. Journal of Fujian Agricultural University, 28 citrus leafminer in Turkey. Proceedings of the In- (3): 319-324. J. HYM. RES. Vol. 13(1), 2004, pp. 120-124

First Food Plant Record for Lagideus Konow (Hymenoptera: Pergidae), a New Species Feeding on Fuchsia and Ludwigia (Onagraceae) in Argentina

David R. Smith and Silvina G. Bado

U. S. (DRS) Systematic Entomology Laboratory, PSI, Agricultural Research Service, Department P.O. Box of Agriculture, % National Museum of Natural History, Smithsonian Institution, 37012, MRC-168, Washington, DC 20013-7012, USA; email: [email protected]; Av. (SGB) Catedra Zoologia Agricola, Facultad de Agronomia, Universidad de Buenos Aires, San Martin 4453, C1417 DSQ, Cap. Fed., Argentina; email: [email protected]

Abstract. —Lagideus badoae Smith, n. sp. (Pergidae: Syzygoniinae), from Argentina and Uru- guay is described and illustrated. Adults were reared from larvae feeding on Fuchsia sp. and Ludwigia peploides (Kunth) Raven (Onagraceae) in Buenos Aires, Argentina. This is the first food plant record for a species of Lagideus. The larva, bearing two long apical filaments, is similar to the larva of Syzygonia cyanocephala Klug of Brazil and resembles larvae of the Australian genus Philomastix Froggatt (Pergidae: Philomastiginae).

Lagideus Konow currently includes 22 and upper margins of the antennal crests species and occurs from southern Mexico and usual carinate hind margin of the pos- to northern Argentina. Smith (1990) re- tocellar area, long apical hind tibial spines vised this exclusively Neotropical genus, (usually more than half the length of the separated it from the other three genera of basitarsi), mostly sclerotized basal plates, Syzygoniinae, added 18 new species, and and a small mesoscutellum. The base of gave a key to the 22 species. Food plants vein M in the forewing (near Sc+R) is usu- for the genus have remained unknown ally distinctly swollen. Sexual dimorphism until adults were reared from larvae feed- in Lagideus species is especially evident in the antennae as It is dif- ing on Fuchsia sp. and Ludwigia peploides described above. (Kunth) Raven (Onagraceae) in Buenos ficult to associate sexes, and it is possible Aires, Argentina, by SGB. Members of a few of the described species represent Lagideus are not commonly collected, and opposite sexes of the same species. This new and L. townesi Smith are the most species are known only from a few species two for which both sexes have been specimens. Considering the scarcity of only associated. study material for Lagideus, its potential great diversity, and probable occurrence of badoae Smith, new many more undescribed species, it is Lagideus species not surprising that these reared specimens (Figs. 1-9) a new represent species. Female. —Length (holotype and para- The is genus Lagideus distinguished types), 6.0 mm. Antenna and head black. from other of the genera Syzygoniinae by Thorax black with pronotum orange. Legs 8-segmented antennae which are filiform black with basal half of hind femur, ex- or serrate in females and bipectinate or treme bases of fore- and midfemora, basal in unipectinate males, presence of three halves of tibiae, and entire fore- and mid- cubital cells in the forewing, carinate inner tarsi mostly white. Abdomen black with Volume 13, Number 1, 2004 121

Figs. 1-4. Lagideus badoae. 1, Head, dorsal view. 2, Female antenna. 3, Male antenna. 4, Female lancet.

behind orange lateral stripes. Wings hyaline; stig- from above narrowing eyes (Fig. ma and veins black. Antenna (Fig. 2) 8- 1); distances between eye and hind ocel- hind segmented, segments 3-7 slightly serrate, lus, between hind ocelli, and from

1 ocellus to of head as 25: each slightly expanded at apex; antenna posterior margin length 1.8X head width; first and second 25:15; postocellar area 3.7X broader than behind. Hind basitarsus segments each slightly longer than broad; long, carinate third segment longer than fourth segment, shorter than length of remaining tarsal segments 4-8 gradually decreasing in segments combined, as 6:8; inner hind tibial of hind basitar- length. Malar space as broad as diameter spur about half length of front ocellus. Lower interocular dis- sus. Sheath (Figs. 5, 6) rounded in lateral concave on tance slightly shorter than eye length; eyes view, slightly ventro-apical intero- in dorsal thick at base and slightly converging below; upper margin, view, cular distance, to lower interocular dis- tapering evenly to acute apex, with long tance, to eye length as 100:80:85. Head backward projecting hairs, many longer of Hymenoptera Research 122 Journal

of sheath in dorsal than greatest breadth view, and many curved at their apices. annular hairs Lancet (Fig. 4) with short evenly distributed on annuli, hairs present on each annulus (except at extreme apex); apex blunt, nearly truncate; serrulae center pointed, those at base and deeper than those near apex; each serrula with 6- 9 fine anterior and posterior subbasal teeth; margin at apex with very fine teeth. Male.—Length, 5.8 mm. Color like fe- antenna male but legs mostly white and brownish. Antenna (Fig. 3) with segments 3-6 unipectinate; ramus of third segment of rami long, subequal to length segment; of segments 4-6 each shorter than length 7 of respective segment; segment triangu- at lar; segment 8 elongate, not expanded with apex. Genitalia (Figs. 7, 8) with harpe long, stiff hairs on inner surface; parapenis about as long as broad, rounded at apex; penis valve oval. Larva. —Flattened on leaf; thoracic legs directed laterally; abdominal segments with lateral lobes; abdominal segments 3- annulate; apical segment with two long filaments. (From photo, Fig. 9; specimens not saved for study.) — 9 labeled Fed. Types. Holotype , "Cap. Arg. 05/2003," Deposited in Ciencias Na- turales Museo, Universidad Nacional de La Plata, La Plata, Argentina. Paratypes: ARGENTINA: Same data as holotype (2$, 16). URUGUAY: Montevideo, Rec'd 47, in the National HL Parker (1 $ ). Deposited Museum of Natural History, Smithsonian Institution, Washington, DC, and Univer- sidad de Buenos Buenos Ar- Aires, Aires, lateral Figs. 5-8. Lagideus badoae. 5, Female sheath, Male gentina. — view. 6, Female sheath, dorsal view. 7, genital Etymology. Named after Ing. Agr. Sil- capsule, ventral view of right half. 8, Male genitalia, vina G. Bado, the co-author of this paper lateral view of penis valve. and discoverer of the larva and food — plant. Food plants and life history. Adults were bred from larvae feeding on Fuchsia sp. after pupation. Rearing conditions were 25 and Ludwigia peploides (Kunth) Raven (On- ± 3° C and 50-60% relative humidity. Lar- agraceae). Larvae were collected in early vae were gregarious feeders on the lower April when they were about 1 cm long. leaf surface, skeletonizing the leaves of Pupation occurred about 10 days after col- both food plants. lection, and adults emerged about a week Remarks. —Of the 22 species treated by Volume 13, Number 1, 2004 123

Smith (1990) 15 are known only from the peal area, clypeus, and labrum whitish, female and six only from the male. The lateral spots on the mesoprescutum or- for single species which both sexes are ange, and an entirely black abdomen. The is L. townesi known Smith. genitalia of L. badoae most closely resem- The female of badoae is in the bles that of L. Lagideus umncatus (Smith 1990: fig. group that has hairlike annular armature 308). rather evenly distributed the full length of Some other species described from each annulus of the lancet (Smith 1990: southern South America are L. albitarsus figs. 313-319, 323-324) (not clusters of Malaise (Santa Catarina, Brazil, and Uru- long, stout spines on the dorsal half of the guay), L. crinitus (Konow) ("Argentina"), lancet as in figs. 318-322). From the ten L. luticus Smith (Tucuman, Argentina), species with annular hairs, the female of and L. townesi Smith (Tucuman, Argenti- L. badoae is separated by having annular na). The male of L. albitarsus has consid- hairs on each annulus of the lancet except erable orange on the thorax and the ramus the extreme apex, the annuli subparallel, of the third segment is much longer than and the of the lancet the of the apex nearly truncate, length segment (Smith 1990: fig. and by the black thorax with the prono- 285); the female of L. crinitus has much of turn orange and black abdomen with a lat- the thorax and abdomen (except apex) or- eral orange stripe. The second half of cou- ange, the antenna only 1.5 X the head plet 2 in Smith's (1990) key can be modi- width, the hind basitarsus longer than the fied to include "abdomen black with lat- length of the remaining tarsal segments eral orange stripes." This would take L. combined, and the antenna with only seg- badoae to couplet 12, the second half of ments 5-7 serrate; the females of L. luticus which can be modified to add "abdomen and L. townesi have a cluster of long spines orange with orange lateral stripes and tho- on the dorsal half of the lancet (Smith rax black with pronotum orange." Thus, 1990: figs. 318, 319), and the male of L. L. badoae will key to L. wygodzinskyi Mai- townesi has the ramus of the third segment aise, known from Sao Paulo and Rio de much longer than the length of the seg- it in Janeiro, Brazil, from which can be sep- ment (as Smith 1990: fig. 285). arated by the subparallel annuli and trun- A photo of the larva was taken (Fig. 9), cate apex of the lancet. The lancet of L. though no larvae were saved for further wygodzinskyi has the basal and apical an- description. It is unusual, mainly by the nuli divergent and the apex acute (Smith presence of two long apical filaments at the of the 1990, fig. 313). apex abdomen; otherwise, the The male of Lagideus badoae will key to laterally protruding legs and lateral lobes couplet 20 which includes L. wuncatus of the abdominal segments resemble some Smith and L. yantuus Smith. Both these North American Acordulecera Say (Pergi- species have the hind basitarus longer dae: Acordulecerinae) larvae. The only than the length of the remaining tarsal other known larvae in the Neotropics with segments combined and are different in such long apical filaments is Syzygonia color. Lagideus wuncatus (described from cyanocephala Klug which feeds on Tibouch- Buenos Aires, Argentina), has dark orange ina spp. (Melastomataceae) in Brazil (illus- antennae, the upper half of the mesepis- trated by Azevedo Marques 1933). Smith ternum, tegula, lateral spots on the meso- (1990) placed Syzygonia Klug and Lagideus prescutum, and mesoscutellum orange, in the same subfamily, the Syzygoniinae, and the abdomen black with the first and based on adult characters. The similarity second terga mostly orange. Lagideus yan- of the larvae may help support that con- tuus (described from Sao Paulo, Brazil) elusion, has the antenna dark orange, the supracly- Other sawfly larvae with similar long of Hymenoptera Research 124 Journal

badoae on Fig. 9. Larva of Lagideus feeding Ludivigia sp.

of Philomastix LITERATURE CITED apical filaments are those in Austra- Azevedo L. A. de. 1933. Tenthredinidae sp. (Pergidae: Philomastiginae) Marques, conhecida por "Mosca de Serra" cuja larva, ou lia (Froggatt 1901, fig. 6; Naumann 1991, "falsa lagarta" e nociva a varias especies do ge- 42-13 A). In Philomastix, the long fila- fig. nero Tibouchina (Biologia de Bergiana cyanocephala from the ninth da ments protrude segment, (Klug, 1824) Konow, 1899). Ministerio Agricul- de Rio de not the apical one; in the photo it appears tura, Institute Biologico Defesa Agricola, 11 7 that the filaments of L. badoae protrude Janeiro. pp., pis. W. W. 1901. The pear and cherry slug (Er- from the Froggatt, apical segment. iocampa limacina, Retz), generally known as Selan- dria cerasi, with notes on Australian sawflies. Ag- ricultural Gazette New South Wales, Miscella- ACKNOWLEDGMENTS of neous Publication No. 497, 11 pp. Naumann, I. 1991. 42. Hymenoptera (wasps, bees, We thank Cathy Apgar, Systematic Entomology ants, sawflies), pp. 916-1106 In The Insects of Aus- Laboratory, U. S. Department of Agriculture, for tak- tralia, A Textbook for Students and Research Work- of the adults and arranging the plates. ing pictures ers. Division of Entomology, Commonwealth Sci- Maria Pannunzio took the of the larva Julia picture entific and Industrial Research Organisation, and assisted in the of adults. We extend rearing Melbourne University Press. Second Edition, thanks to the for the following reviewing manuscript: Volume II, pp. 543-1137. N. M. U.S. Forest and Schiff, Service, Stoneville, MS, Smith, D. R. 1990. A synopsis of the sawflies (Hy- G. L. Miller and E. E. Grissell, Systematic Entomology menoptera: Symphyta) of America south of the Laboratory, USDA, Beltsville, MD, and Washington, United States: Pergidae. Revista Brasileira Ento- DC, respectively. mologia 34: 7-200. J. HYM. RES. Vol. 13(1), 2004, pp. 125-133

Description of the Antistrophns rufus (Hymenoptera: Cynipidae) Species Complex, Including Two New Species

John F. Tooker, Andrew R. Deans, and Lawrence M. Hanks

Department of Entomology, University of Illinois at Urbana-Champaign, 505 S. Goodwin Urbana Avenue, 61801, [(JFT) current address: Department of Entomology, 501 ASI Building, Pennsylania State University, University Park, PA 16802]; (LMH) email: [email protected]

Abstract. — describe the We Antistrophus rufus species complex of gall wasps, including a rede- of A. scription rufus Gillette and descriptions of two new species: A. meganae Tooker and Hanks and A. jeattae Tooker and Hanks. Larvae of the three species develop in stem galls of different, but congeneric asteraceous plant species that are endemic to tallgrass prairies of midwestern A. North America, rufus in Silphium laciniatum L., A. meganae in S. terebinthinaceum Jacquin, and A. jeanae in S. perfoliatum L. Adults of the three species are very similar morphologically, but differ in structure of the antennae, length of ovipositors, depth of galls in plant tissues, and mass of mature larvae. An allozyme study confirmed that wasps from the three plant species are reproductively isolated from one another.

Antistrophus Walsh 1869 (Cynipidae: iting stems of S. laciniatum and S. terebin- Aylacini) is a Nearctic genus currently thinaceum due to phenological differences comprising at least eight species, all of between host plants (Tooker et al. 2002). which form galls in asteraceous plants Wasps from the two plant species mate as- (Burks 1979, Nieves-Aldrey 1994). Six spe- sortatively when brought into contact (un- cies form galls in species of Silphium and published data), and males preferentially Lygodesmia that are endemic to tallgrass respond to plant volatiles associated with prairies of midwestern North America their natal host species (Looker et al. (Burks 1979, Gleason and Cronquist 1991). 2002). Allozyme studies confirmed that Larvae of Antistrophus rufus Gillette feed wasp populations inhabiting S. laciniatum in small, single-chambered, ellipsoid galls and S. terebinthinaceum were reproductive (~3 mm in length) in flowering stems of isolated from one another, and a Nei's ge- Silphium species that do not affect the stem netic distance of 0.56 further indicated that surface and so are not discernible exter- the populations actually represent differ- nally (Gillette 1891, Beutenmuller 1910, ent species (Looker et al. 2002). Looker et al. 2002, Looker and Hanks In this paper, we extend our studies of 2004a, b, c). Antistrophus species by including a third Antistrophus rufus was originally de- population associated with Silphium per- scribed from specimens reared from Sil- foliatum. Differences in phenology be- phium laciniatuni L. (Gillette 1891), but the tween this population and those inhabit- species name also has been applied to ing S. laciniatum and S. terebinthinaceum specimens from S. terebinthinaceum Jac- (see below), again associated with host quin, S. perfoliatum L., and S. integrifolium plant phenology, suggest limited gene Michaux (Beutenmuller 1910). In our stud- flow. We confirm reproductive isolation of ies of the ecology of A. rufus in prairies of the S. perfoliatum population with allo- central Illinois, we found evidence of re- zyme studies, and also report that wasps productive isolation of populations inhab- from the three Silphium species differ in of Hymenoptera Research 126 Journal

view from of stemmaticum to ventral the morphology of the antennae and ovi- top area is of in host of clypeus. Supraclypeal positors, depths galls plants, margin between and to- and mass of mature larvae. On the basis the medial area clypeus of these morphological, and eco- ruli. genetic, mass conclude that An- To how gall dimensions and logical differences, we study a of at of larvae varied across plant species, tistrophus rufus comprises complex wasp Gillette in S. we dissected stems of S. col- least three species, A. rufus perfoliatum, 2002 from BRP laciniatum, A. meganae Tooker and Hanks lected in early Spring and A. where all three species co-oc- n. sp. in S. terebinthinaceum, jeanae Silphium extracted mature Tooker and Hanks n. sp. in S. perfoliatum. curred. We gall wasp and mor- larvae and them, and measured We provide here descriptions weighed the surface of for each species. the depth from stem galls phological diagnoses = with a microscope micrometer (N 22 METHODS Data for in galls in twenty stems). wasps with To compare the morphology of Antistro- S. perfoliatum were compared pub- adult in S. laciniatum and phus populations, we reared gall lished data for wasps and Hanks wasps from Silphium stems we collected S. terebinthinaceum (Tooker 2001- during the winters of 1998-1999 and 2004c). Illinois. between An- 2002 from prairie sites in central We compared differences from Fithian in for mor- We collected S. laciniatum tistrophus populations means N larval Railroad Prairie (FRP, Vermilion Co., phological variables, gall depth, and Rail- 40° 06.78, W 87° 54.10) and Buckley mass with analysis of variance (ANOVA; road Prairie (BRP, Iroquois Co.; N 40° Wiley 1981). Differences between individ- 34.88, W 88° 02.70), S. terebinthinaceum ual means were tested with the LSD Railroad Prairie from East St. Joseph means separation test (Sokal and Rohlf 88° and Pax- (ESJRP; N 40° 06.77, W 00.48) 1995). Railroad Prairie Ford N 40° ton (PRP; Co.; For the allozyme analysis, we reared 88° and S. from 26.17, W 06.36), perfoliatum Antistrophus adults from stems of S. per- BRP and PRP. We measured dimensions all three foliatum collected at BRP, where of heads and antennae of females using plant species co-occurred. We conducted digital photographs (e.g., Fig. 1) produced cellulose acetate electrophoresis for six with electron- and microscopy (scanning loci using methods described by Tooker et in compound microscopes) conjunction al. (2002). We compared allele frequencies with software image analysis (Image-Pro* for the S. perfoliatum population with fre- Plus Version 4.5, Media Inc., Cybernetics, quencies for the S. laciniatum and S. tere- Silver MD). We dissected Spring, ovipos- binthinaceum populations (from Tooker et itors from ten female from each wasps al. 2002) using RxC contingency table and measured with a micro- plant species tests (software ver. 2.1, Bill Engles, Uni- scope micrometer the length from the tip versity of Wisconsin, Madison). to the second valvifer (see Fig. 2). We present means ± 1 SE throughout. Terminology relating to morphology and wing venation follows Nieves-Aldrey Antistrophus rufus species complex (1994) and descriptions of sculpturing fol- (Figs. 1-8) low Harris (1979). Post-Ocellar Line (POL) — in cambi- is the distance between inner margins of Diagnosis. Galls developing lateral ocelli; Ocell-Ocular Line (OOL) is um and pith of flowering stems of Sil- the distance from outer edge of a lateral phium laciniatum, S. terebinthinaceum, and ocellus to inner margin of the compound S. perfoliatum. Second flagellomere longer eye. Head height is measured in frontal than first. Notauli evident, but faint ante- Volume 13, Number 1, 2004 127

Figs. 1-5. Antistrophns species. 1, Second flagellomere of female A) A. rufus; B) A. meganae; C) A. jeanae (scale = bar 100 (xm); lines 1 and 2 indicate length and width measurements, respectively. 2, Ovipositors of A) A. = rufus; B) A. meganae; C) A. jeanae (scale bar 1 mm); arrows indicate length measurement. 3, Head of A. rufus = = female (scale bar 200 |xm). 4, Mesonotum and metanotum of A. rufus female (scale bar 200 |xm). 5, Head = and mesosoma of A. rufus male (scale bar 200 (xm). 128 Journal of Hymenoptera Research

6. = Fig. Antistrophus rufus female (scale bar 1 mm). Volume 13, Number 1, 2004 129 riorly. Distinct medial ridge separating MALE. Differs from female in shape and scutellar fovae. size of metasoma (Fig. 8). Antennae lon- — 2.6 Description. FEMALE. Length ± ger, with 14 antennomeres (Fig. 7). 0.05 to mm (Fig. 6). Light brown amber GALL. Small (~3 mm in length), ellip- often with darker (even black) areas in soid, and monothalamous, hidden in cam- vertex, mesoscutum, and dorsum of me- bium and pith of flowering stems of Sil- tasoma. Antennae and legs same color as phium, not evident in external view, body. Head with fine areolate sculpturing Material examined. —We examined the and sparsely setose (Fig. 3, 5). Head in holotype in the Insect Collection of the II- dorsal view 1.8X broader than long; in linois Natural History Survey, Cham- frontal view, 1.4X broader than high; head paign, IL (INHS), which was reared from height 2x compound eye height. POL S. laciniatum, as well as 10 females and 5 slightly shorter than OOL and 2X greatest males that we reared from stems of each diameter of lateral ocellus. Malar space of three Silphium species (S. laciniatum, S. 0.8 X eye height. Face with laterally radi- terebinthinaceum, and S. perfoliatum) col- ating striae. Supraclypeal area with two lected from prairies in central Illinois (see pronounced tentorial pits. Antennae 13- above). Depositories for specimens are segmented and imbricate, distal flagello- INHS and the National Museum of Nat- mere 2X longer then flagellomere 10 (Fig. ural History, Smithsonian Institution, 7). First flagellomere 0.75 X as long as sec- Washington, DC (NMNH). ond. Pronotum finely areolate, setose, Comments. —Previous work revealed prominent lateral expansions, and distinct that no alleles were shared between gall submedial pits (Fig. 4). Mesoscutum finely wasps reared from S. laciniatum and S. ter- areolate; anterior fourth of notauli faint, ebinthinaceum (Table 1, Tooker et al. 2002). moderately convergent posteriorly. Medi- New data (Table 1) show that gall wasps an mesoscutal impression faint to promi- reared from S. perfoliatum had alleles at all nent, but most evident in posterior third, six loci that were not represented in wasps Two faint submedial impressions in ante- from either S. laciniatum or S. terebinthin- rior fourth and parallel to median mesos- aceum, and in fact, 48 and 71% of alleles at cutal impression. Two additional mesos- loci MDH and ME, respectively, were cutal impressions lateral to notali and unique to S. perfoliatum wasps. Moreover, most evident in posterior third (Fig. 4). locus IDH was fixed for unique alleles in Scutellum finely areolate. Scutellar fovae all three wasp populations (Table 1). R-C prominent and rounded, separated from contingency tables tests for each locus each other by distinct medial ridge. Me- were highly significant (P < 0.001), sug- sopleuron setose and finely areolate (Fig. gesting a lack of gene flow between pop- 5). Propodeum finely areolate and densely ulations of gall wasps inhabiting different setose laterally. Propodeal carinae mod- Silphium species. These data strongly sup- erately divergent posteriorly. Metasoma port the morphological evidence that nitid. In lateral view, length of metasoma these populations represent different spe- approximately equal to or slightly shorter cies. than head + mesosoma + propodeum Gillette (Fig. 6). Metasomal tergites I and II occupy Antistrophus rufus more than half of abdomen. Hypopygidial (Figs. 1-8) short. Ovipositor 2.8 ± 0.08 spine . very Antistrophus mfus Gillette 1891: 195 mm long. Wings hyaline with pale brown venation: Rl and Rs of forewing not ex- Diagnosis. —Distinguished from other tending to wing margin, radial cell open, species in complex by the following char- in in Rs curved slightly anteriorly (Fig. 6). acters: 1) Larvae developing galls 130 Journal of Hymenoptera Research

Table 1. Allelic frequencies of A. rufus species complex reared from stems of Silphium laciniatum ("S. lac") and S. terebinthinaceum ("S. to:") and Silphium perfoliatum ("S. per.") from three prairies in central Illinois. Sample sizes are in parentheses. The most anodal band was assigned "A" with electrophoretic mobilities of other bands relative to this band on starch and cellulose acetate gels. Populations are named after their prairies in S. laciniatum S. terebinthinaceum are (see Materials and Methods). Data from gall wasp populations and from Tooker et al. (2002).

Locus/ Volume 13, Number 1, 2004 131

Table 2. of and first Comparison width, length, width/length ratio of and second flagellomeres (Fig. 1) for three of the A. members rufus species complex and ANOVA results. We measured flagellomeres of 20 female reared from each of the wasps three plant species, averaging the values from each antennae to generate mean measurements for each individual. Values within column with different letters are significantly different (LSD < "*" "***" P 0.05). Significant P values for ANOVA indicated by (P < 0.05), (P < 0.0001). 132 Journal of Hymenoptera Research

—Similar to A. but larvae the most significant source of mortality for Biology. rufus, within in stems of A. rufus larvae with rates of parasitism of- develop galls flowering Adults ten exceeding 90% (Tooker and Hanks S. terebinthinaceum. begin emerging in and continue to 2004a). Species of parasitoids reared from from stems mid-June for 20 A. rufus galls include eurytomids (Euryto- emerge approximately days (Tooker ma lutea Bugbee and an unidentified Eur- and Hanks 2003a). Females oviposit in internodes of S. terebinthinaceum after ytoma species), an ormyrid (Ormyrus labo- stem tus Walker), eupelmids (Eupelmus vesicu- bolting. Similar to A. rufus, A. meganae lar- laris [Retzius] and two unidentified Brase- vae can suffer high parasitism rates and ma species), and a pteromalid the same guild of parasitoids appears to (unidentified Homoporus species; Tooker attack both species (Tooker and Hanks and Hanks 2004a). Populations of gall 2004a). are decimated when are wasps prairies Tooker and but recolonize Antistrophus jeanae Hanks, burned, quickly (Tooker new and Hanks 2004b). species (Figs. 1C, 2C) Antistrophus meganae Tooker and — Diagnosis. Larvae developing in galls Hanks, new species in cambium and pith of flowering stems (Figs. IB, 2B) — of S. perfoliatum. See description for fur- Diagnosis. Larvae developing in galls ther details. in cambium and pith of flowering stems Description. —FEMALE. Generally same of S. terebinthinaceum. See description for as others in species complex, but width/ further details.— length ratio of first flagellomere signifi- Description. FEMALE. Generally same cantly larger, and that of second flagello- as others in species complex, but width/ mere intermediate to other species in com- length ratio of first and second flagello- plex (Table 1, Fig. 1C). Ovipositor signifi- meres significantly smaller than in both A. cantly shorter than that of A. rufus (Table A. rufus and jeanae (Table 2, Fig. IB). Ovi- 3, Fig. 2C). GALL: Intermediate in depth positor significantly shorter than in A. ru- to other species in the complex (Table 3). fus (Table 3, Fig. 2B). GALL: Significantly LARVAE: Mass of mature, overwintering closer to the stem surface than other spe- larva significantly lower than in A. rufus cies in complex (Table 3). LARVAE: Mass and A. jeanae (Table 3). of — mature, overwintering larvae interme- Types. Holotype reared in May 2002 diate to that of A. A. rufus and jeanae (Ta- from stems of S. perfoliatum collected at ble 3). — BRP and deposited with INHS. We depos- Types. Holotype reared in June 2002 ited nine female and five male paratypes from stems of S. terebinthinaceum collected reared from stems of S. perfoliatum collect- at ESJRP and deposited in INHS. We de- ed in Spring 1999 from PRP (four females, nine posited female and five male para- three males) and Spring 2002 from BRP reared types from stems of S. terebinthin- (five females, two males) in INHS. We de- aceum collected in Spring 1999 from PRP posited five female paratypes (three from (four females, three males) and Spring PRP and two from BRP) and three male 2002 from ESJRP (five females and two paratypes (two from PRP and one from males) in INHS. We deposited five female BRP) in NMNH. from paratypes (three ESJRP, two from Etymology. —Named in honor of Jean and three male PRP) paratypes (two from Michelle Hanks, spouse of third author ESJRP and one from PRP) in NMNH. LMH. —Named in honor of — Etymology. Megan Biology. Similar to A. rufus, but larvae Weaver Tooker, of the first author. spouse develop within flowering stems of S. per- Volume 13, Number 1, 2004 133

Table 3. of Comparison ovipositor length, gall depth, and mass of larvae for the A. rufus species complex, and ANOVA results. Values within column with different letters are significantly different (LSD P < 0.05). "*" "**" "***" Significant P values for ANOVA indicated by (P < 0.01), (P < 0.001), (P < 0.0001).

Species Ovipositor length (mm) Gal! depth (mm) Larval mass (mg)

A. rufus 3.19 ± 0.07 a 2.20 ± 0.06 a 2.29 ± 0.13 a A. meganae 2.54 ± 0.15 b 1.05 ± 0.03 c 1.66 ± 0.09 b A. jeanae 2.68 ± 0.09 b 1.76 ± 0.10 b 0.88 ± 0.16 c = F, „ 9.44* F, M = 57.9*** F, 6.77*

Natural Bulletin the Illinois State Labo- foliatum. Adults emerge from stems for a History. of of Natural History 3: 191-197. period of about 20 days beginning in mid- ratory Gleason, H.A. and A. Cronquist. 1991. Manual of Vas- and females in stem inter- May oviposit cular Plants of Northeaster)i United States and Ad- after to nd nodes bolting. Similar other spe- jacent Canada, 2 Edition. The New York Botani- cal The New York. cies in the complex, parasitoids can inflict Garden, Bronx, Harris, R. A. 1979. A glossary of surface sculpturing. high levels of mortality on A. jeanae larvae. Occasional Papers in Entomology, California Depart- ACKNOWLEDGMENTS ment of Food and Agriculture No. 28: 1-31. L. 1994. Nieves-Aldrey, J. Revision of west-European of the tribe Ashmead We thank D. Voegtlin (Illinois Natural History Sur- genera Aylacini (Hyme- Re- for F. L. noptera, Cynipidae). journal of Hymenoptera vey) providing specimens, Ronquist, J. Ni- search 3: 175-206. eves-Aldrey, A. Valerio, and W.-Y. Choi for advice SAS Institute. 2001. SAS/STAT User's Guide for Per- on morphological characters and taxonomy, J. Whit- sonal Computers, release 8.01. SAS Institute, Cary, field for comments on the manuscript and use of mi- North Carolina. croscopic and photographic equipment, S. H. Ber- Sokal, R. R. and F. J. Rohlf. 1995. Biometry. 3" edition. locher, for guidance with allozymes, and S. Robinson W. H. Freeman, New York. and the staff of the at Imaging Technology Group A. L. 2002. Tooker, J. F., W. Koenig and M. Hanks. University of Illinois at Urbana-Champaign for assis- Altered host plant volatiles are proxies for sex tance with the Environmental Scanning Electron Mi- pheromones in the gall wasp Antistrophus rufus. We also assistance in the field croscope. appreciate the National Sciences ProceedingsC of Academy of and lab E. J. Kron, A. L. Crumrin, J. A. provided by U.S.A. 99 ; 15486-15491. K. R. F. and M. Mohler, Rinkenberger, J. Westberg, F. L. M. 2004a. in- Tooker, J. and Hanks. Endophytic W. Tooker. sect communities of two prairie perennials (As- teraceae: Silphium spp.). Biodiversity and Conser- LITERATURE CITED vation (in press). F. L. 2004b. Tooker, J. and M. Hanks. Impact of pre- Beutenmuller, W. 1910. The North American species scribed burning on endophytic insect communi- of and their Bulletin American Mu- Aylax galls. of ties of prairie perennials (Asteraceae: Silphium seum Natural 28: 137-144. History spp.). Biodiversity and Conservation (in press). Burks, B.D. 1979. 1045- F. L. Superfamily Cynipoidea, pp. Tooker, J. and M. Hanks. 2004c. Trophic position 1107. /// K. V. P. D. D. R. Krombein, Hurd, Jr., of the endophytic beetle Mordellistena aethiops of Smith, and B. D. Burks (eds), Catalog Hyme- Smith (Coleoptera: Mordellidae). Environmental noptera in America North of Mexico. Volume 1, Entomology (in press). Smithsonian Institution Press, Washington, DC. Wiley, E. O. 1981. Phylogenetics: the Theory and Practice Gillette, C.P. 1891. Descriptions of new Cynipidae in of Phylogenetic Systematics. John Wiley and Sons: the collection of the Illinois State Laboratory of New York. J. HYM. RES. Vol. 13(1), 2004, pp. 134-148

The Neotropical Species of Deuterixys Mason (Hymenoptera: Braconidae)

James B. Whitfield and M. Teresa Oltra Moscardo of Urbana, IL 61801, (JBW) Department of Entomology, 320 Morrill Hall, University Illinois, USA; email: [email protected]; Universidad de Valencia, (MTOM) Instituto Cavanilles de Biodiversidad y Biologia Evolutiva, [email protected] Apartado 22085, 46071 -Valencia, Spain; email:

Mason are revised for Abstract—The neotropical species of the braconid wasp genus Deuterixys of which are new to science: D. colombiana, D. the first time. Seven species are recognized, four D. The three D. bennetti Whit- erythrocephala, D. hansoni, and tehuantepeca. remaining species, and D. Whitfield, were known from the Nearctic field, D. pacifica Whitfield quercicola previously into the An illustrated identification is region but are found to range Neotropical. key presented for the New World species.

the of Our taxonomic knowledge of parasitoid sented here for species Deuterixys in- Mason in the wasps in tropical parts of the world, Neotropical Region began no information eluding the Neotropical Region, is notori- with absolutely published of the in the area under ously incomplete. It is especially so for about any species less well parasitoids whose hosts are also study. understood, for example the microlepi- The genus Deuterixys was erected by to house the doptera as a group. Small caterpillars tend Mason (1981) species previ- to be less conspicuous and more difficult ously assigned by Nixon (1965) to the car- of Foerster. lo identify than larger host insects, and bonarius-group Apanteles By thus relatively few accurate host-parasite 1985, four Palearctic species (Papp, 1983) records are avaitable for them, except in and three Nearctic species (Whitfield, well-surveyed areas. As an example of 1985, 1995) were known, to which Austin this, Whitfield and Mason (1994) de- and Dangerfield (1992) added one Austra- scribed a new subfamily of braconid lian species. All host records so far assem- wasps that appears to attack concealed bled suggest that Deuterixys spp. always microlepidopterans, and this subfamily attack caterpillars in the genus Bucculatrix now is known to contain at least 10-15 (Bucculatricidae), which in turn feed on a species distributed from South America to wide variety of plant families, most nota- The the southeastern U. S., some of which are bly on Asteraceae (Whitfield, 1985). still being discovered every few years wasps pupate within the elongate ribbed (Valerio and Whitfield, 2000; Yamada and cocoons of their hosts, lining the host co- Penteado-Dias, 2002; Valerio and Whit- coon with some silk of their own. The re- field, 2002). Several genera of the subfam- vision we present below contains no host but ily Microgastrinae are estimated to contain records from the Neotropical Region, at least 100 neotropical species, only a the best current guess is that the known handful of which are currently described pattern of specialization on Bucculatrix (Mason, 1981, Whitfield, 1997). Thus it will be found to be true of this area as comes as no surprise that the revision pre- well. Volume 13, Number 1, 2004 135

Below we describe several new species by the senior author from several other of Denterixys from the Neotropical region collections. The collections and curators from specimens that we have found in responsible for loaning these specimens major collections, and compare these pu- are thanked in the Acknoivledgments. Mu- tative new species to the previously seum abbreviations in the Specimens Ex- known species of the genus. While we can amined sections are as follows: AEI: Amer- make no pretense that the survey we are ican Entomological Institute, Gainesville; presenting is complete, it is intended to CNC: Canadian National Insect Collec- provide a first treatment of this distinctive tion, Ottawa; Humboldt: Humboldt Insti- genus, and should serve to make the in- tute, Villa de Leyva, Colombia; INBio: In- cluded species more recognizable to neo- stituto Nacional de Biodiversidad, Costa tropical scientists. Rica; MCZ: Museum of Comparative Zo- This is a difficult genus in which to de- ology, Harvard University; TAMU: Texas limit species, especially due to the limited A and M University, College Station); material for some species, the small body UKY: University of Kentucky, Lexington; size, and substantial intraspecific variabil- USNM: National Museum of Natural His- ity in body coloration and intensity of tory, Washington, D. C; UWYO: Univer- sculpturing. We recommend using the key sity of Wyoming, Laramie. below first, but strongly recommend fol- Wing drawings were prepared by tem- lowing this by checking the resulting iden- porarily dry slide-mounting wings from tifications by consulting the descriptions ones side of selected specimens, and pro- and the figures. jecting these using a Ken-A-Vision XI 000 The genus appears to be generally un- microprojector onto paper for tracing and common wherever it occurs in the world, recording veins and pigmentation pat- although individual species can some- terns. times be easily collected once their host is Morphological terminology (including located. wing vein nomenclature) follows that of and Wharton and Whit- MATERIALS AND METHODS Sharkey (1997) field (1997). A diagram of this venational Specimens for this study were originally scheme as applied to Deuterixys is pre- sorted and brought together from several sented in Fig. 1. collections by the late W. R. M. Mason, Names of new species are to be attri- and supplemented by specimens sorted buted to both authors.

KEY TO NEW WORLD SPECIES OF DEUTERIXYS MASON

1. Tergite I of metasoma weakly hourglass-shaped, narrowing slightly in anterior half then n. widening again posteriorly (Fig. 9) Deuterixys colombiana, sp. Known so far only from mountains of the northern Andes and Santa Marta (Colombia) Ranges. - Tergite I either narrowing or broadening posteriorly, or relatively parallel-sided, but not 2 narrowest near midlength (Figs. 10, 11, 13, 14) 2. Tergite I of metasoma strongly narrowing posteriorly; tergite III less strongly sculptured Whitfield than tergite II (Fig. 14) Deuterixys padfica North America, ranging well south in Mexico in mountains. - or Tergite I of metasoma either broadening posteriorly (Fig. 16), barrel-shaped (Fig. 15), II 3 relatively parallel-sided; tergite III as strongly sculptured as tergite of 3. Tergite I at posterior end nearly as broad as anterior margin tergite II, and broadening III as as II slightly posteriorly (Fig. 16); tergite nearly long medially tergite Deuterixys quercicola Whitfield 136 Journal of Hymenoptera Research

Western North America, ranging well south in Mexico in mountains. - anterior of and more or less Tergite I posteriorly much narrower than margin tergite II, 4 barrel-shaped or parallel-sided (Figs. 10, 12, 13, 15) bennetti Whitfield 4. Tegulae strongly yellowish Denterixys Gulf coast of U. S., Florida, widespread in Caribbean. - 5 Tegulae deep brown to blackish courses of so 5. Fore wing length > 2.3 mm; mesoscutum slightly depressed along notauli, that there is a suggestion of notauli in dorsal view; Rl (metacarp) only slightly longer and broader than than stigma (Fig. 5); tergite III of metasoma strongly abruptly tergite n. II, and only about 0.83as long as tergite II (Fig. 15) Deuterixys hansom, sp. Known from Costa Rica, tentatively also from Bolivia (both high elevation localities). - Fore wing length < 2 mm; mesoscutum with no hint of notauli; Rl (metacarp) usually approximately 1.53 as long as stigma, sometimes shorter (Figs. 4, 8); tergite III of metasoma slightly broader than tergite II, usually nearly as long as tergite II medially 6 (Figs. 10, 12) 6. Scutellum and posterior 0.2 of mesoscutum polished and nearly impunctate; much or all of head and anterior portions of mesosoma often deep reddish-brown rather than black Deuterixys erythrocephala n. sp. Lower elevations in Antilles and Trinidad; questionably also from Argentina. - Scutellum, and usually also posterior 0.2 of mesoscutum with distinct punctation; head black and thorax mostly black(Veracruz) Deuterixys tehuantepeca n. sp. So far known from mountains of southern Mexico and Central America.

Deuterixys bennetti Whitfield orly; shallowly punctate, becoming less Figs. 2, 13 distinctly and more scarcely punctate posteriorly. Scutellar disc sparsely, shallowly The neotropical specimens of this pre- punctate throughout. dark viously described species are redescribed Propodeum reddish brown with margins and medial for comparison with the other species. carina Female and male. —Body size: Overall longitudinal black, highly polished, for medial length 1.9 mm; fore wing length 1.9 mm. virtually unsculptured except and lateral carinae Head: Antennae approximately same longitudinal strong and weak transverse in in- length as body; scapes and pedicels vivid very ridging mediate of carina yellow, remainder of antennae fulvous- vicinity longitudinal brown; basal 9 flagellomeres with black midbasally. Legs: Legs mainly yellowish, with coxae and trochanters, basal, distal and intermediate patches; api- prothoracic mesothoracic coxae and trochanters and cal 6-7 flagellomeres with only one rank metathoracic trochanters Distal half of placodes; 2nd flagellomere 3x as long pale. as of mesothoracic tibiae and mesothoracic broad; 14th and 15th flagellomeres approximately 1.5X as long as broad. Head tarsi fulvous. Metathoracic legs with distal dark brownish black. Clypeus and genae extreme of femora and tibiae, and tarsi ful- reddish brown; labrum and mandibles, ly brown. Hind apical tibial spurs short, except brown teeth, vivid yellow; glossa whitish, subequal in length. Wings: Tegu- and palpi whitish yellow. Frons nearly lae vivid yellow. Stigma 2.8 X as long as 1.25 X as broad at midheight as midlength; broad. Metacarp (Rl) of fore wing slightly inner margins of eyes weakly converging longer than pterostigma; r and 2Rs sube- towards clypeus. Punctation of head in- qual in length and meeting at rounded, distinct. Mesosoma: Prosternum fulvous approximately 135 degree angle. Venation with yellow margins. Mesoscutum black, of fore wing, including stigma translucent \ lateral dark fulvous patches posteri- pale yellow-brown. Costa and margin of Volume 13, Number 1, 2004 137

Fig. 1. Labelled diagram of wing veins, using the nomenclature adopted in Wharton et al. (1997) and in this paper. pterostigma brown. Metasoma: Tergite I Apical margins of tergite IV and following nearly 1.7X as long as posterior width, tergites brownish yellow. Basal sternites subparallel-sided, virtually parallel-sided, yellowish, distal sternites and hypopy- with fulvous-brown. very slightly divergent anteriorly gium — strong longitudinal groove over anterior Material examined. 16, CUBA: Santa 0.3; coarsely aciculorugose over posterior Clara Prov., San Juan Mrs., Jan.-Feb. 1927 0.7. Tergite II and III subequal in length. C. T. and B. B. Brues (MCZ); 19, DOMIN- Tergite II approximately 1.8X as broad an- ICAN REPUBLIC: Loma el Penon, 676m, teriorly as long and a little more than 1.6X 23-111-1979, L. Masner (CNC); 19, 16% JA- as broad posteriorly as long, with sharp MAICA: Moneague, June, W. S. Brooks anterolateral corners and posterior half (MCZ). — the of virtually parallel sided; surface coarsely Hosts. Along Gulf Coast North aciculorugose, medial raised portion America, Bucculatrix sp. (Lepidoptera: L. sometimes distinctly demarcated by ridg- Bucculatricidae) on Baccharis halimifolia as host es. Tergite III separated from II by crenu- (Asteraceae) serves (Whitfield late furrow, abruptly widening, becoming 1985); in the Neotropical region, the hosts far unknown. broader than tergite II at broadest point; are so — other of surface longitudinally aciculorugose. Hy- Comments. No known species popygium short, blunt, with long hairs New World Deuterixys has the tegulae so the third metaso- subapically. Ovipositor sheaths short, vividly yellowish, and about as long as third metathoracic tarso- mal tergite is also more evenly finely stri- other in mere, decurved, polished, with hairs con- ate in sculpturing than species II III this The centrated apically. Tergites I, and Hemisphere. Jamaican specimens small and less black, rest of tergites brown. Laterotergi- are exceedingly brightly III whitish. colored than to be for the tes and apical margin of tergite appears typical of Hymenoptera Research 138 Journal

D. Figs. 2-8. Fore wings. 2, Deuterixys bennetti Whitfield. 3, D. colombiana n. sp. 4, D. erythrocephala n. sp. 5, hansoni n. sp. 6, D. pacifica Whitfield. 7, D. quercicola Whitfield. 8, D. tehuantepeca n. sp. Volume 13, Number 1, 2004 139 species; it is possible that these represent exemplar. Costa and margin of stigma a distinct species but our best guess at pre- brown. Metasoma: Tergite I 1.8-1 .9 x as sent, based on both morphology and dis- long as posterior width, narrowing in bas- tribution, is that they are dwarfed D. ben- al 0.4, with strong longitudinal groove netti. over anterior 0.5-0.6; coarsely aciculorugose over posterior 0.6. Tergite II and ter- Denterixys colombiana Whitfield and gite III subequal in length. Tergite II su- Oltra, n. sp. bquadrate, slightly broader anteriorly Figs. 3, 9 — than posteriorly, approximately 1.7x as Female and male. Body size: Overall broad as long, surface coarsely aciculoru- length 2.5-3.0 mm; fore wing length 2.5- gose, with parallel dense striations longi- 3.0 mm. Head: Black, with palpi whitish tudinally, medial raised portion distinct. amber. Antennae slightly longer than Tergite III separated from tergite II by body, scapes dark brown yellow infuscate crenulate groove, becoming abruptly wid- and pedicels brown, remainder of anten- er than tergite II; surface longitudinally nae brown. Apical 6 flagellomeres with aciculorugose and medial raised portion only one rank of placodes. Second flagel- as tergite II. Hypopygium short, blunt, lomere 3.5-4.2 X as long as broad, 14th with long hairs mainly concentrated api- and 15th flagellomeres 1.3 X as long as cally. Ovipositor sheaths short, about as broad. Inner margins of eyes weakly con- long as apical metathoracic tarsomere, de- verging towards clypeus. Punctation of curved, polished, with hairs concentrated head indistinct. Mesosoma: Mesoscutum apically. Tergites I, II and III black, tergite black, shallowly punctate anteriorly, be- IV dark brown and rest of tergites brown. coming less distinctly and more sparsely Sternites fulvous-brown. Laterotergites, punctate posteriorly. Scutellar disc sparse- apical margins of tergite III and following ly, shallowly punctate throughout. Pro- tergites, and intersegmental membranes podeum highly polished, virtually un- mainly whitish.— sculptured except for medial and lateral Hosts. Unknown.— longitudinal strong carinae, and weak Material examined. Holotype 9: CO- transverse ridging in inmediate vicinity of LOMBIA: Caldas, 3300-3500m, 5 15'N, longitudinal carina. Propodeum with bas- 76°25'W 3-IV-1973, J. Helava, (CNC). al half brownish black and distal half dark PARATYPES: 59, 66, same data, also 16, reddish brown. Legs: Prothoracic legs Antioquia, 1800m, 7°5'N, 76°30'W, 18-22- brown except whitish trochanters and yel- IV-1973, J. Helava, (CNC), 16, Putumayo, lowish fulvous distal half of femora and 2900m, 1°10'N, 77°15'W, 2-XII-1972, J. He- entire tibiae. Mesothoracic and metatho- lava (CNC). Other material: 29, Id, CO- racic legs mainly brown, except tarsomer- LOMBIA: Magdalena, PNN S. Nevada de es darker and fulvous trochanters, apical Santa Marta, El Ramo, 2500m, 24-30-VI- extreme intermediate femora and basal ex- 2000, J. Castillo, J. Varela (Humboldt, treme tibiae. Apical spurs of hind tibiae UKY). ECUADOR: Id, Napo, above Pa- short, whitish, subequal in length. Wings: pallacta, 3500m, 21-11-1983 (AEI), L. Mas- Tegulae dark brown. Stigma 2.8-3 X as ner; 16, Napo, above Papallacta, 3200m, long as broad. Metacarp (Rl) of fore wing 14-11-1983, Masner and Sharkey (AEI); 16, r sub- Pimo L. slightly longer than stigma; and 2Rs (N. Cahar), 3200m, 10-12-XII-1970, E. 1 San 10 equal in length and meeting at rounded to Pena (AEI). PERU: 9, Jeronimo, E. Feb.-Mar. P. M. Marsh weakly angular, approximately 135 degree km Cusco, 1978, angle. Venation of fore wing light yellow- (USNM). — ish brown, stigma opaque, pale yellowish Comments. Tergite I of this species is to of D. rimulosa white; translucent and light brown in one similar some specimens of Hymenoptera Research 140 Journal

with but it can vex. dark reddish brown (Niez.) from the east of Spain, Propodeum carina from this Palaearc- and medial longitudinal be easily distinguished margins of the combi- black, virtually unsculp- tic species by its possession highly polished, tured for medial and lateral nation of following features: 1) metacarp except longi- I tudinal carinae and transverse longer than stigma, 2) metasomal tergite strong form- in inmediate of three ca- with strong narrowing in basal half, ridging vicinity in females rinae. Prothoracic legs whitish yel- ing an hourglass shape (more Legs: and low fulvous distal tarsomere. Me- parallel-sided in males), 3) coarsely except II and III. sothoracic with trochanters yel- aciculorugose surface of tergites legs light in combina- coxae and femora fulvous, tibiae light The relatively long tergite II, low, is fulvous and tarsi fulvous tion with the very coarse sculpturing, yellowish except in the New distal tarsomere. Metathoracic mostly diagnostic for this species legs dark white tro- World fauna. This species is known cur- fulvous, except yellowish and chanters and basal third to half of tibiae, rently only from the northern Andes, tarsi more from the isolated Santa Marta range in and brown zones of tibiae and northern Colombia. dark. Apical spurs of hind tibiae short, whitish, subequal in length. Wings: Tegu- Whitfield and X Deuterixys erythrocephala lae dark fu i VOus to brown. Stigma 2.75 Oltra, n. of fore sp. as j ong as broad. Metacarp (Rl) 10 Figs. 4, wing up to 1.3X length of stigma; r and in and at Female.—Body size: Overall length 1.7- 2Rs subequal length meeting 135 de- 1.8 mm; fore wing length 1.8-1.9 mm. slightly rounded, approximately includ- Hend: Antennae approximately as long as gree angle. Venation of fore wing, to body; scapes and pedicels fulvous in up- ing stigma, translucent, very pale yellowish brown can per part and vivid yellow in lower part, (stigma appear opaque remainder of antennae brown; basal 8 fla- whitish in some preserved specimens), brown, gellomeres with black basal, distal and in- Costa and margin of pterostigma I from 1.5 X to 2 X termediate patches; apical 8-9 flagello- Metasoma: Tergite up sub- meres with only one rank of placodes; 2nd as long as posterior width, virtually in flagellomere 3-4 X as long as broad; 14th parallel-sided, broadening shallowly and 15th flagellomeres from 1.2X up to distal third and narrowing slightly poste- nearly 1.7 as long as broad. Head dark riorly, with strong longitudinal groove reddish brown in dorsal view; frons, clyp- over anterior 0.3-0.4; coarsely aciculoru- eus deep fulvous to brown; labrum and gose over posterior 0.6-0.8. Tergite II mandibles, except brown teeth, vivid yel- slightly longer than HI. Tergite II parallel- as low; glossa and palpi whitish yellow, sided, from 1.6X up to 2x as broad Frons from 1.3X-1.5X as broad at midh- long; surface coarsely aciculorugose, me- Ter- eight as midlength; inner margins of eyes dial raised portion almost indistinct. fur- virtually parallel to slightly divergent ven- gite III separated from II by crenulate broad- trally towards clypeus. Punctation of head row, abruptly widening, becoming indistinct in dorsal view, punctation of er than tergite II; surface longitudinally face superficial, giving impression of even aciculorugose. Hypopygium short, blunt, rugosity. Mesosoma: Prosternum dark ful- with scattered long hairs subapically. Ovi- vous to reddish brown. Mesoscutum positor sheaths short, about as long as see- mostly dark fulvous to reddish brown, ond /third metathoracic tarsomere, de- with lateral and medial black patches pos- curved, polished, with hairs concentrated occa- teriorly; shallowly punctate, becoming im- apically. Tergites I, II and III black, punctate over posterior 0.2. Scutellar disc sionally shading more reddish brown, rest punctai s virtually throughout, evenly con- of tergites brown. Laterotergites and ster- Volume 13, Number 1, 2004 141

dorsal view: colombiana n. 10, D. n. 11, 12, Figs. 9-12. Metasomal tergites, 9, Deuterixys sp. erythrocephala sp. D. tehuantepeca n. sp. of Hymenoptera Research 142 Journal nites whitish, distal sternites and hypo- num black. Mesoscutum black; punctation and pygium fulvous-brown. shallow dense, becoming impunctate of notauli Material examined.—HOLOTYPE 9: near posterior margin; courses so there is a TRINIDAD: Curepe, 21-VII-1978, malaise somewhat depressed, sugges- PARA- tion of notauli. Scutellar disc black, sparse- trap (no collector given) (CNC). TYPES: 19, same data but XII. 1977 ly and very shallowly punctate, evenly (CNC); 19, DOMINICAN REPUBLIC: La convex. Propodeum black, mostly pol- medial and lateral Cumbre, 400m, 21-111-1978, L. Masner ished, sculptured by carinae and transverse (CNC). More doubtful determination: AR- longitudinal strong GENTINA: 19, Tucuman, Horco Molle, ridging of three carinae giving impression Prothoracic and 10-15-V-1968, no collector listed (MCZ). of even irregularity. Legs: Hosts. —Unknown. mesothoracic coxae dark brown, metatho- black. Prothoracic with Comments—This species can be easily racic coxae legs in the trochanters and femora femora distinguished from the other species yellow, infuscate with tibiae New World by its possession of the fol- dark brown, basally; and tarsi dark distal tarsomere lowing combination of features: 1) tegulae fulvous, Mesothoracic with trochanters dark brown, 2) posterior 0.2 of mesoscu- brown. legs femora dark brown and tum and scutellum virtually impunctate, reddish brown, tibiae and 3) portions of head and mesosoma yellowish distally, gradually chang- reddish brown rather than black. This lat- ing from fulvous basally to brown distally, with distal tarsomere ter feature gives the species its name. The tarsi light brown Metathoracic with tro- specimen from Argentina seems virtually dark brown. legs identical in structural details, but is darker chanters and femora dark brown, femora basal in general coloration. The Trinidad speci- reddish infuscate, approximately rest dark mens differ in some minor details (espe- third tibiae fulvous with the with tarsi dark brown. cially coloration) from the Dominican Re- brown-fulvous, public specimen, but we consider them Hind apical tibial spurs short, whithish, black. conspecific due to their joint possession of subequal in length. Wings: Tegulae the distinctive features listed above. Stigma 2.4X as long as broad. Metacarp (Rl) of fore wing as long as stigma, r and hansoni Whitfield and Oltra, Deuterixys 2Rs subequal in length and meeting at n. sp. about 130 degree angle. Venation of fore 5, 15 Figs. wing mostly brown, discoloured basally; Female.—Body size: Overall length about stigma opaque, whitish yellow; costa and 2.1 mm; fore wing 2.5 mm. Head: Length margins of stigma brown. Metasoma: Ter- bar- of antennae 2.5 mm; antennae dark gite I 2.2X as long as posterior width, brown, pedicels infuscate of dark fulvous; rel-shaped, very weakly widening poste- cor- apical 6 flagellomeres with only one rank riorly, with strongly rounded apical of placodes; 2nd flagellomere 3.3 X as long ners so than greatest width of distal third as broad; 14th flagellomere about 1.5 X as is nearly 1.5X as broad as distal width; long as broad. Head black, clypeus dark with strong longitudinal groove over an- brown, labrum and mandibles light ful- terior 0.4; coarsely rugose over posterior vous infuscate with brown; glossa and 0.8. Tergite II slightly longer than III, with palpi yellowish white. Frons about 1.5X as subparallel sides, shallowly divergent an- broad at midheight as midlength; inner teriorly, with strong anterolateral corners; margins of eyes virtually parallel towards surface coarsely aciculorugose, medial clypeus. Punctation of head superficial, raised portion almost indistinct and nar- dense, giving impression of even rugosity, row. Tergite II about 1.6X as broad as specially in the face. Mesosoma: Prester- long. Tergite III separated from II by cren- Volume 13, Number 1, 2004 143 ulate furrow, abruptly widening, becom- length about 1.9 mm; fore wing approxi- ing broader than tergite II at broadest mately 2 mm. Head: Antennae nearly as point; surface longitudinally aciculorugo- long as body; apical 8 flagellomeres with se. Tergite IV shallowly rugose, remaining only one rank of placodes; 2nd flagello- tergites polished. Hypopygium short, mere 2X as long as broad; 14th and 15th blunt, with long hairs apically. Ovipositor flagellomeres about 1.4X as long as broad. sheaths short, about as long as second Antennae brown. Head black in dorsal metathoracic tarsomere, decurved, pol- view, brown in frontal view, labrum and ished, with hairs concentrated in apical mandibles, except light fulvous teeth, ful- half. Tergites I, II, III and IV black, rest of vous-brown; glossa whitish fulvous; palpi tergites dark reddish brown. Laterotergi- light brown. Frons 1.3X as broad at midh- tes and apical margins of tergites III and eight as midlength; inner margins of eyes IV whitish, basal sternites fulvous-brown. virtually parallel towards clypeus. Punc- Distal sternites and in hypopygium— brown. tation of head virtually indistinct dorsal Material examined. HOLOTYPE 9: view, punctation of face superficial, COSTA RICA: San Jose, 16 km S. Empai- crowded, giving impression of even ru- me, 2600m, II-IV-1989, P. Hanson and I. gosity. Mesosoma: Mesosoma mainly black Gauld (UWYO). More doubtfully associ- dorsally and ventrally; axillae adjacent to ated: BOLIVIA: 19, La Paz, Rio Zongo, scutellum, metanotum and propodeum 3200m, 22-X-1984, L. Pena (AEI). dark reddish brown. Punctation of mesos- Hosts.—Unknown. cutum shallow and fine, rough and more Comments.—This is one of the larger crowded in courses of notauli. Scutellar species of Deuterixys. It differs from the disc shallowly punctate, punctation superficially similar species such as D. crowded giving impression of even rugos- bennetti, D. erythrocephala and D. tehuante- ity laterally and posteriorly. Propodeum peca by its possession of the following dark reddish brown with black margins combination of features: 1) weak sugges- and mid-longitudinal carina, very pol- tion of notauli on mesoscutum, 2) meta- ished, virtually unsculptured except for carp about as long as stigma, 3) tergite I strong medial and lateral longitudinal ca- III transverse weakly widening posteriorly, 4) tergite rinae and very little, superficial, in of abruptly wider than tergite II and about ridging immediate vicinity medial carina. for the most 0.75 length of tergite II. In general the an- Legs: Generally part terolateral corners of tergite II are also brown, more reddish from coxae through more rectangular than in the other species. femora inclusive; prothoracic femora dis- metathoracic tibiae The Bolivian specimen appears to be distally and basally yellowish. Hind tibial tributionally disjunct, but since the speci- apical spurs short, mens are from high elevation and rarely whitish, subequal in length. Wings: Tegu- collected this may not mean much. The lae black. Stigma 2.4 X as long as wide. and fore species is named for Dr. Paul Hanson, of Metacarp (Rl) stigma of wing in 2Rs 1.25X as as r the University of Costa Rica, who has con- subequal length; long at 130 tributed substantially to our knowledge of and meeting about degree angle. Costa Rican Hymenoptera. Venation of fore wing mostly light brown, paler basally; stigma opaque, brown, more Whitfield Deuterixys pacifica light basally; costa brown. Metasoma: Ter- 6, 14 Figs. gite I 3.4X as long as posterior width; sub- and Neotropical specimens of this previous- parallel-sided anteriorly, evenly redescribed over about ly described Nearctic species weakly narrowing posterior with for comparison. 0.7, rounded, convergent posteriorly; Female ami male. —Body size: Overall strong longitudinal groove about over an- of Hymenoptera Research 144 Journal

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Whitfield. Figs. 13-16. Metasomal tergites, dorsal view. 13, Denterixys bennetti Whitfield. 14, D. pacifica 15, D. hansoni n. sp. 16, D. quercicola Whitfield. Volume 13, Number 1, 2004 145

terior 0.5; coarsely rugose over posterior Denterixys quercicola Whitfield 0.6. III 1.4X as Tergite long medially as II. Figs. 7, 16 Tergite II about 2.2 X as broad medially as Neotropical specimens of this previous- long, virtually parallel-sided, narrowing ly described species are redescribed for slightly posteriorly; surface evenly rugose, comparison. medial raised portion little evident. Notch Female and male. —Body size: Overall between tergites II and III evident. Tergite length approximately 2 mm; fore wing III separated from II by narrow crenulate about 2 mm. Head: Antennae as long as furrow, rounded laterally, becoming body, black, 2nd flagellomere 4x as long broader than II at broadest tergite point; as broad, 14th flagellomere approximately surface weakly rugose. Hypopygium 1.4X longer than broad. Head black, clyp- with short, blunt, long hairs subapically. eus black, labrus and mandibles reddish sheaths short, near as as Ovipositor long brown; glossa brownish yellow, palpi yel- third and fourth metathoracic tarsomeres lowish white. Posterior ocelli brownish together, decurved, polished, with hairs red, opaque; anterior ocellus yellow, concentrated II apically. Tergites I, and III translucent. Frons nearly 1.2X as broad at mostly black, except reddish brown midheight as midlength; inner margins of smooth zone basally on tergite I and ful- eyes divergent towards clypeus. Head in dorsal vous zone subdistally on tergite III; rest of view smooth and very polished; of face tergites brownish fulvous. Dorsal portions punctation superficial, crowded, of shallow of laterotergites fulvous. Remaining area giving impression even, rugos- Mesosoma: Mesosoma black of laterotergites and apical margin of ter- ity. dorsally and Punctation of mesoscutum gites III, IV and V whitish. Sternites ventrally. shallow, brown, sterna of basal segments little ligh- becoming impunctate posterior- Scutellar disc smooth and ter, hypopygium fulvous-brown. ly. mostly very with anterolat- Material examined.—19, MEXICO: Mi- polished, light punctation erally. Propodeum black, choacan, 2 mi. S. Carapan, 6-VII-1985, very polished, for Woolley and Zolnerowich (TAMU); 16, shining, virtually unsculptured except medial and lateral longitudinal ca- San Luis Potosi, 7.2 mi. E. San Luis Potosi, strong rinae, and weak, little extensive, trans- 3-VII-1987, G. Zolnerowich (TAMU). verse ridging in inmediate vicinity of ca- Three specimens have also been seen from rinae. Legs: Prothoracic legs mostly the Rocky Mountains in Wyoming (mate- brownish yellow with coxae, basal two rial in UWYO collection), to the adding thirds of femora and last tarsomere black- range from the original description. ish. Mesothoracic legs with black coxae Hosts. —In western North America, this and femora, tibiae brown and tarsi yellow- species attacks Bucculatrix spp. on sage- ish brown. Metathoracic legs mainly brush and a broad of other range compos- black, with two fifth basal of tibiae ful- ites this also (Whitfield 1985); possibly vous. Apical outer spur of hind tibiae for the Mexican material. goes shorter than half length of tibia. Wings: Te- Comments.—This is distinctive species gulae black. Metacarp (Rl) and stigma of the New World in among species having fore wing subequal in length; 2Rs sligthly the first so and tergite strongly narrowed, longer than r and meeting at rounded, ap- in the third much less having tergite proximately 135 degree angle. Stigma strongly sculptured than the second. It re- about 2.7X as long as wide. Venation of D. sembles the Palearctic rimulosa Niez., fore wing brown, stigma translucent yel- but has the first tergite more strongly nar- lowish brown. Costa dark brown. Metaso- rowed posteriorly. ma: Tergite I 1.5X as long as posterior 146 Journal of Hymenoptera Research

1.9 Head: Antennae as as width, subparallel-sided, slightly broad- mately mm. long fulvous in ening in distal 0.4, with strong longitudi- body; scapes and pedicels up- nal groove over anterior 0.3; coarsely aci- per part with brown distal half of pedicels, in remainder of an- culorugose longitudinally over about dis- and yellow lower part, tal 0.8 with a smooth and polished zone tennae brown; apical 8 flagellomeres with mid-longitudinally. Tergite II and III sub- only one rank of placodes; 2nd flagello- as 14th equal in length. Tergite II subparallel-sid- mere approx. 3x as long broad; ed, about 1.7X as broad as long; sculpture and 15th flagellomeres 1.25 X and 1.4X as coarsely aciculorugose, more densely so long as broad respectively. Head black in- la- laterally, medial raised portion almost frontally and dorsally, clypeus brown; distinct. Notch between tergites II and III brum and mandibles, except brown teeth barely evident. Tergite III separated from and infuscate basal part of mandible, yel- II by crenulate furrow, becoming shallow- low; glossa and palpi whitish. Frons slow- ly broader than tergite II; sculpture Ion- ly broader at midheight than midlength; gitudinally aciculorugose, more fine and inner margins of eyes divergent towards dense posteriorly. Of remaining tergites, clypeus. Punctation of head virtually in- only tergites IV and V visible, tergite V distinct in dorsal view, punctation of face short and overlapped by IV. Tergite IV superficial, crowded, giving impression of with fine longitudinal sculpturing over even rugosity. Mesosoma: Mesosoma gen- most of its surface. Hypopygium short, erally black to dark reddish brown dor- blunt, with sparse long hairs. Ovipositor sally and ventrally. Punctation of mesos- sheaths short, near as long as second cutum shallow, becoming impunctate pos- metathoracic tarsomere, decurved, apical- teriorly. Scutellar disc sparsely and very ly hairy. Metasoma mostly black, basal shallowly punctate, evenly convex. Pro- sternites dark fulvous. podeum dark reddish brown with mar- Material examined.—19, MEXICO: Ta- gins and medial-longitudinal carina black, maulipas, Mun. Tula, La Presita, Canon de polished, virtually unsculptured except Coyote, 1900m. 16-111-1987, P. Kovarik, R. for medial and lateral longitudinal strong Jones, R. Trivino (TAMU). 16, MEXICO: carinae and transverse ridging in imme- 6 mi. Durango, W. La Ciudad, 9000', 11- diate vicinity of the three carinae. Legs: VI-1964, W.R.M. Mason (CNC). Prothoracic legs mostly yellowish white, Hosts. —In western North America, this with coxae basally and tarsi distally ful- species parasitizes Bucculatrix spp. primar- vous infuscated. Mesothoracic legs with on oaks ily (Quercus spp.). This is possibly coxae fulvous ventrally, brown dorsally; true also for the Mexican material, but trochanters yellowish fulvous; fulvous there are no rearing records there. femora infuscate basally and dorsally; tib- Comments.— No other New World Deu- iae and tarsi whitish yellow, apical tarso- has the terixys species posterior margin of meres fulvous. Metathoracic legs mainly the first metasomal tergite nearly as broad brown except trochanters and basal two- as the second tergite. This species seems thirds of tibiae yellowish fulvous. Apical to range widely especially in scrubby sa- spurs of hind tibiae short, whitish, sube- vannah-like south vegetation from North qual in length. Wings: Tegulae dark red- America well into southern Mexico. dish brown. Stigma from 2.5 X up to near 2.8 X . as of „ L , , . . , long as wide. Metacarp (Rl) Whitfield and i . « , .. Deutenxys tehnantepeca fore( almost 1.2X„., as as 01fl wing long stigma; r ^ l - anc ^S subec ua m and T7 « 11 i o i length meeting ° ' at rounded, approximately 135 degree an- Female and male.— size: Overall Body gle. Venation of fore wing mostly light about 1.9 fore mm; wing approxi- brown, paler basally; stigma opaque, pale Volume 13, Number 1, 2004 147 to light brown; costa brown. Metasoma: the general region of the Isthmus of Te- I X Tergite about 2-2.2 as long as posterior huantepec, hence the specific epithet. width, to broaden- parallel-sided slightly ACKNOWLEDGMENTS ing posteriorly, posterior corners rounded, with strong longitudinal groove over an- We would like to thank Stefan Cover (Museum of terior 0.5; coarsely aciculorugose over pos- Comparative Zoology, Harvard University), Henri Goulet (Canadian National Collection, Ottawa), Mike terior 0.7. Tergite II slightly longer medi- Sharkey (University of Kentucky in conjunction with ally than III, about 1.75X as broad medi- the Humboldt Institute, Colombia), Scott Shaw (Uni- as with sides; ally long, virtually parallel versity of Wyoming), David Smith (U. S. National surface scrobiculate, medial raised portion Museum), Jesus Ugalde (Instituto Nacional de Bio- almost indistinct, wide. Notch between diversidad, Costa Rica), David Wahl (American En- tomological Institute) and Robert Wharton (Texas A tergites II and III barely evident. Tergite and M University) for loaning specimens used in this III separated from II by crenulate furrow, study, and acknowledge the late William R. M. Ma- this more evident III be- medially; tergite son for doing the same. JBW's work was funded by coming slightly broader than tergite II; U. S. D. A. National Research Initiative Grant no. 95- sculpture longitudinally aciculorugose. 01893. short, blunt, with hairs Hypopygium long LITERATURE CITED subapically. Ovipositor sheaths short, near as long as distal metathoracic tarsomere, Austin, A. D. and P. C. Dangerfield. 1992. Synopsis of Australasian decurved, with hairs concentrated apical- Microgastrinae (Hymenoptera: Braconidae), with a to and II III to key genera descrip- ly. Tergites I, and dark brown tion of new taxa. Invertebrate Taxonomy 6: 1-76. black. IV, V and VI brownish ful- Tergites Mason, W. R. M. 1981. The polyphyletic nature of rest of fulvous. La- vous, polished; tergites Apanteles Foerster (Hymenoptera: Braconidae): a terotergites and apical margin of tergites phylogeny and reclassification of Microgastrinae. Memoirs the Canada 115: IV, V and VI whitish. Basal sternites yel- of Entomological Society of 1-147. lowish, remaining sternites and hypopy- Nixon, G. E. 1965. A reclassification of the tribe fulvous-brown. J. gium Microgasterini (Hymenoptera: Braconidae). Bul- Material examined.—HOLOTYPE 9: letin of the British Museum (Natural History) En- GUATEMALA: Antigua, 1500-1600m, tomology Supplement 2: 1-284. G. E. 1976. A revision of the north-western VII-1980, N. L. H. Krauss (CNC). PARA- Nixon, J. European species of the merula, lacteus, vipio, ul- TYPES: Id, same data; 19, MEXICO: Oa- tor, ater, butalidis, popularis. carbonarius and vali- xaca, 5.7 mi. SE 2100', 21-VII- Quiotepec,

B. Valerio, A. A. and J. Whitfield. 2002. Epsilogaster Marsh, and M. J. Sharkey, eds. Identification Man- faviolae, a new species of Mendesellinae from Co- ual to the Neiv World Genera of the Family Bracon- lombia (Hymenoptera: Braconidae). Zootaxa 41: idae (Hymenoptera). International Society of Hy- 1-7. menopterists Special Publication 1. 439 pp. B. R. Whitfield, J.B. 1985. The nearctic species of Deuterixys Whitfield, J. and W. M. Mason 1994. Mendesel- Mason (Hymenoptera: Braconidae). Pan-Pacific linae, a new subfamily of braconid wasps (Hy- Entomologist 61: 60-67. menoptera, Braconidae) with a review of rela- Whitfield, J.B. 1995. Annotated checklist of the Micro- tionships within the microgastroid assemblage. gastrinae (Hymenoptera: Braconidae) in America Systematic Entomology 19: 61-76. north of Mexico. Journal of the Kansas Entomolog- Yamada, M. V. and A. M. Penteado-Dias. 2002. Men- ical Society 68: 245-262. desella jaraguaiensis, a new species of mendesel- B. 1997. line Whitfield, J. Subfamily Microgastrinae. Chap- wasp (Hymenoptera, Braconidae) from Bra- ter 29, pp. 333-364 in: Wharton, R. A., P. M. zil. Entomological News 113: 236-238. J. HYM. RES. Vol. 13(1), 2004, pp. 149-205

Revision of the Genus Pseudognaptodon Fischer (Hymenoptera: Braconidae: Gnamptodontinae)

Daryl J. Williams

nd Natural Resources Canada, Canadian Forest Service, Northern Forestry Centre, 5320-1 22 street, Edmonton, Alberta, Canada T6H 3S7; email: [email protected]

— Abstract. The New World genus Pseudognaptodon has three previously described species, P. curticauda Fischer, P. tninutus Ashmead, and P. omissus Fischer. Pseudognaptodon attenuatus, P.brevis, P. conjunctns, P. carinatus, P. gibsoni, P. gouleti, P. hemicolor, P labrus, P. langori, P. minimus, P. nitidas, P. ocellatus, P. shaivi, P. striatus, P. whartoni, P. luhitfieldi, and P. xanthus, are described here as new. A key is given to the species of Pseudognaptodon. Problems in the phylogenetic relationship between Gnamptodon and Pseudognaptodon are discussed.

Fischer (1965) described the genus Pseu- documented. The purpose of this study is dognaptodon (Braconidae: Gnamptodonti- to describe these species and discuss char- nae) and included two new species, P. om- acters that may be useful in phylogenetic issus and P. curticauda, each of which was analysis. described from only two specimens from The species of Pseudognaptodon are here the Nearctic. He subsequently (Fischer divided into two species groups, the cur- 1967) included P. minutus Ashmead, based ticauda-group and the omissus-group, which are united venation but on the study of 13 specimens. These three by wing otherwise no more similar to one another species are very similar to species of than either is to Gnamptodon, but differ by the absence of Gnamptodon (Williams, A conservative the forewing r-m vein. Fischer (1977) treat- unpublished data). ap- is taken in of the ed both Gnamptodon (as Gnaptodon) and proach light uncertainty of within the Pseudognaptodon as genera of the Opiinae. phylogenetic relationships so no new They were subsequently transferred to the Gnamptodontinae, genus- names are here. new subfamily Gnaptodontinae by van group proposed Wharton and Achterberg (1983). (1997), METHODS Whitfield and Wagner (1991), have published taxonomic works and keys to the Terms follow Wharton et al (1997), with gnamptodotine genera, including Pseudog- some details clarified as follows: The basal naptodon, but no further species-level work raised area is the anterior portion of the the has been done on the genus since orig- second metasomal tergite (T2), which is inal descriptions from limited material. separated from the posterior portion by a Since that time more material has accu- groove (the defining synapomorphy of the in mulated through biodiversity studies Gnamptodontinae) (Figs. 22, 32, 42). This Central America (S.R. Shaw, R. Wharton, portion is higher posteriorly than anteri- the in lateral and than the pers. comm.) and studies of parasit- orly view, higher oids of Nearctic leaf-mining Lepidoptera posterior portion of the tergite, even in (Whitfield and Wagner 1991), representing species where the groove is somewhat ef- of faced. The anterolateral of the new species and a much wider range grooves third metasomal are those on morphological variation than previously tergite (T3) Research 150 Journal of Hymenoptera

in lateral P. attenuatus: es scrobe. 2, P. Figs. 1-3. Pseudognaptodon species, metasoma view. 1, epicnemial minutus (Ashmead). 3, P. carinatus.

either side of the anterior margin that join Head height: the greatest distance between the suture between the second and third the vertex and ventral margin of the metasomal terga, and separate the antero- gena in lateral view (Fig. 6). lateral corners from the remainder (Figs. Head length: measured from the anterior- 22, 32, 42). The knob of the hind wing R most projection of the middle of the face vein refers to the remnant of the base of to a line intersecting the posteriormost This the RS vein, which may be absent, or vis- curve of the occiput (Fig. 5). meth- ible as a variously developed convexity or od avoids error introduced by variation bump (Figs. 7, 18). in occiput indentation. Most measurements are taken as the Head width: the greatest distance across the greatest length of the body part measur- head in dorsal view, including the eyes able (e.g. femur length, clypeus height, (Fig. 5). etc.). Other measurements used in de- Diameter of median ocellus (MOD): the scriptions are: greatest diameter of the median ocellus Eye length: the greatest width of the eye in (Fig 4). lateral view, along the line of greatest Diameter of lateral (or posterior) ocellus head width (Fig. 6). (POD): the greatest diameter of the left Eye height: the greatest height of the eye in ocellus, usually measured about 45 de- lateral view, measured perpendicular to grees from the long axis of the body the eye length (Fig. 6). (Fig. 4). Gena width: the width of the gena mea- Lateral ocellar length (LOL): The least dis- sured on the same line as for eye length tance between the median ocellus and (Fig. 6). the left lateral ocellus (Fig. 4). Volume 13, Number 2004 1, 151

parastigma

= = Figs. 4-7. Pseudognaptodon attenuatus. 4, Ocelli in dorsal view. LOL lateral ocellar length, MOD diameter = = of median ocellus, POD diameter of lateral ocellus, POL posterior ocellar length. 5, Head in dorsal view. = = = = HL head length, HW head width, OOL ocello-ocular length. 6, Head in lateral view. EH eye height, = = = EW eye width, GW gena width, HH head height. 7, Wings. 152 Journal of Hymenoptera Research

to S.E.M. and Ocello-ocular length (OOL): the shortest dis- mounted directly stubs, tance between the left lateral ocellus gold-coated along with the specimens. and the closest point on the dorsal eye Micrographs are somewhat variable in and of illustrated struc- margin (Fig. 5). quality position Posterior ocellar length (POL): the least dis- tures because of this. For species described tance between the lateral ocelli (Fig. 4). from few specimens it was not desirable of Metasomal Tl length: the distance from the to alter specimens by removal append- of ho- posteriormost point of the tergite to a ages or gold coating. Photographs line drawn through the narrowest point lotypes of these species were taken with a immediately posterior to its' attachment stereomicroscope and digital camera. of insufficient point to the propodeum (Fig. 10). These photographs were Metasomal Tl width: the greatest distance quality for publication, and so were ren- of it's posterior margin (Fig. 10). dered into line drawings by tracing. Metasomal T2 and T3 length: measured on Specimens were obtained from the fol- the midline (Fig. 22). lowing collections, with the name of the Wing vein lengths: vein junctions are grad- originator of the loan: ual thickenings with curved margins. BMNH The Natural Museum, Lengths are measured at the midpoint History Cromwell Road, London, SW7 of the curve between the veins (Fig. 7, 5BD, Suzanne Lew- R and r-m). ENGLAND, is. Descriptions are based on females, and CNCI The Canadian National Collec- are not identical in format between the tion of Insects, ECORC, K.W. species groups, although most characters Neatbly Bldg.-CEF, Ottawa, On- are shared. Characters that are invariable tario, K1A 0C6, CANADA, Hen- in a species group are excluded from spe- ri Goulet. cies treatments. The shape and sculpture DJMW Daryl J. Williams Collection, of the first three metasomal tergites are di- Northern Forestry Centre, Ed- agnostic for most Pseudognaptodon species. monton, Alberta, T6H 3S7, CAN- Males tend to show greater similarity ADA. among species than females for these char- ESUW Department of Plant, Soil, and acters. Males generally have metasomal Insect Science, University of Wy- tergites that are longer, narrower, and less oming, Laramie, 82071, USA, distinctly sculptured than in females, so Scott Shaw. the application of these characters in the FSCA Florida State Collection of Ar- key or diagnoses must be used with cau- thropods, Division of Plant In- tion. Characters of color and other body dustry, Florida Department of parts apply equally to both sexes, and ef- Agriculture, Gainesville, 32602, fort has been made to include these char- USA, Lionel Stange. acters in the key. JWCI James Whitfield Collection, De- Structures were illustrated with scan- partment of Entomology, Uni- electron ning micrographs where possible. versity of Illinois at Urbana- Because of their minute size specimens are Champaign, Urbana, 61801, easily damaged by any attempt to manip- USA. ulate them. So, specimens for S.E.M. were RNHL Rijksmueum von Natuurlijke left on their and points unaltered, except Historie, Raamsteeg 2, Postbus for the removal of wings, which were 9517, 2300 RA Leiden, NETH- mounted i a separate card and later re- ERLANDS, Kees van Achter- v pinned the specimen. Points were berg. Volume 13, Number 1, 2004 153

TAMU of Tex- The Department Entomology, grooves are absent or partially, faintly as A&M University, College Sta- impressed and smooth in the curticauda- tion, 77843-2475, USA, Robert group. This state is found in most other Wharton. Gnamptodon species (van Achterberg 1983, USNM United States National Museum Fig. 63). Another example is the state of of Natural Smithsonian History, the epicnemial scrobe. It is present, and Institution, Washington, DC, joined to the posterior margin of the me- David Smith. 20560-0168, USA, sopleuron by a cuticular fold in species of the ora/'ssi/s-group, and poorly developed Pseudognaptodon Fischer or absent in species of the curticauda- Pseudognaptodon Fischer 1965:182. Fischer 1967: group. I have examined specimens of 973; Fischer 1977:983; Shenefelt 1975:133; Gnamptodon with either of these two Marsh 1979:175; van 1983:26; Achterberg states, and others with the epicnemial Wharton 1997:258; Type species Pseudognap- scrobe present as a depression of varying todon curticauda Fischer by original designa- and (van 1983). A tion. shape depth Achterberg — comprehensive phylogenetic analysis of Relationships. Some questions exist the subfamily is needed in order to dis- about the of validity Pseudognaptodon, cover monophyletic assemblages, identify which is separated from Gnamptodon only synapomorphies, and elucidate phyloge- the loss of the r-m vein. by forewing While netic relationships among species. For this this character has practical utility in iden- reason I have made a conservative choice tification it's phylogenetic reliability has and retained Pseudognaptodon as a separate yet to be assessed. Both genera are specio- taxon. A phylogenetic analysis of Pseudog- se with substantial morphological varia- naptodon is not presented here since the tion that has yet to be phylogenetically an- genus may not be a monophyletic unit, alyzed, and the distribution of states of and outgroup selection would be prob- some of these characters in the two genera lematic at this— level. may suggest relationships among species Diagnosis. Species of Pseudognaptodon and groups of species. Wharton (1997) may be separated from other genera of suggested that generic concepts based pri- Gnamptodontinae by the absence of the marily on the old world species do not ac- forewing vein r-m. count for undescribed variation in new Description. — Head: Oval in anterior world species, and may have to be revis- view; eyes slightly protuberant, with ited. He also suggested that Pseudognap- rounded but parallel inner margins, todon might prove to be one or more spe- straight to slightly concave adjacent to an- cies groups of Gnamptodon. I tend to sup- tennal sockets but margins otherwise con- port that view. The omissus- and curticau- vex; face convex, wider than high; with rfrt-groups each have characters that are curved setae, directed dorsally and dor- more similar to species of Gnamptodon somedially; clypeus arcuate, with straight than to each other. An example of this is ventral and convex dorsal margin, surface the state of the anterolateral grooves of the convex dorsally, concave ventrally, with a third metasomal tergite. These grooves are ventral setose rim. Mesosoma: Length well defined and slightly to moderately about 1.5X height, smooth; notauli deeply crenulate in the omzssws-group. A similar impressed anteriorly, increasingly obso- state is found in Gnamptodon vlugi van lete posteriorly; mesonotum and meta- Achterberg from Europe (van Achterberg pleuron setose, mesosternum and propo- with scattered 1983, Fig. 72), and other undetermined deum sparse, setae. Wings: North American Gnamptodon species 1 Forewing with 2 submarginal cells, r-m of have examined (Wharton 1997, Fig. 3). forewing absent (Fig. 7); first abscissa of 154 Journal of Hymenoptera Research

M present as a short spur, shorter than raised, higher posteriorly than anteriorly, 2RS; RS+Ma and RS+Mb less pigmented smooth, defined posteriorly by a crenu- and thinner than other veins, base of late, striate, or granulate groove; disc of T2 RS + Ma obsolete in some species. Hind posterior to basal raised area granulate, wing with RS and second of M present as granulostriate, or striate, rarely smooth. folds, 2r-m present as fold or absent. Sub- T3 separated from T2 by a well developed, basal cell less than V2X as long as basal smooth to crenulate, evenly curved or lat- cell, first of abscissa M nearly twice as erally decurved groove; T3 smooth to long as M+Cu. Metasoma: Tl about as sculptured on basal V3 with similar sculp- long as or slightly longer than apical ture to T2. Ovipositor barely exserted to width, constricted immediately posterior nearly as long as hind basitarsus. Legs: to junction with propodeum, convex me- Legs stout, hind femur length 3.0-5.0 dially in both lateral and posterior view, greatest width. Femur, tibia, and tarsus with sinuate carinae originating posterior about equally— long. to constriction, the carinae laterally pro- Hosts. One species reared from leaf- x duced on basal h of tergite and conver- mining larvae of the family Nepticulidae gent on remainder. T2 with basal 0.2-0.4 (Lepidoptera).

KEY TO FEMALES OF SPECIES OF PSEUDOGNAPTODON FISCHER

1. Episternal scrobe present as a curved, distinct crease (Fig. 1). T3 of metasoma with anterolateral grooves complete, distinctly impressed, and partially to completely crenulate (complete but smooth in P. attenuatus new species) (Figs. 22, 42). Ovipositor dis- x tinctly exserted, setose portion of sheath 0.5-0.9 as long as hind basitarsus (Fig. 13). oraz'ssMS-group 2 - Episternal scrobe absent (Fig. 2), or present as a shallow impression in P. carinatus new species (Fig. 3). T3 of metasoma usually without anterolateral grooves, rarely with faint grooves near lateral tergal margins or with anterolateral areas partially defined by microsculpture (Figs. 104, 207). Ovipositor barely exserted, setose portion of sheath less than 0.5X as long as hind basitarsus (Fig. 115). curticauda group 9 2. Head and mesosoma yellow to light orange with brown patches 3 - Head and mesosoma brown to black, rarely with lighter patches on head and /or lighter pronotal collar 4 3. smooth Propodeum medially (Fig. 81). Basal cell of hind wing narrow, r-m 0.5-1 .Ox as as R long (Fig. 79). Mesothorax and metasoma unicolorous or with slightly darker areas dorsally p. xanthus new species Propodeum striate medially (Fig. 39). Basal cell of Hind wing wide, r-m 2.5 X as long as R (Fig. 38). Mesothorax with sharply contrasting brown areas dorsally and ventrally, and metasoma with brown spot mid-dorsally P. langori new species 4. Appendages elongate, basal flagellomere 4.8-5.0 x as long as wide and hind femur 4.5- 5.5 X as as wide of long (Fig. 13). T1-T3 metasoma smooth (Fig. 12) P. attenuatus new species Appendages relatively shorter, basal flagellomere 3.0-4.5 X as long as wide and hind femur 3.0-4.0 x as long as wide. Tl of metasoma usually striate medioapically, and T2 and T3 striate usually partially or granulate (Figs. 22, 52) 5 5. T3 of metasoma with anterolateral corners joined by a thin carina, the groove between T2 and T3 appearing double (Fig. 22) P. conjunctus new species T3 of metasoma with anterolateral corners separate, the groove between T2 and T3 single (Figs. 42, 52) 6 of metasoma posterior to basal raised area and base of T3 between anterolateral corners Volume 13, Number 1, 2004 155

striate Hind coarsely (Fig. 73). wing with angle of M and r-m with a faintly pigmented, directed posteriorly spur (Fig. 69) P. striatus new species - T2 of metasoma granulate or finely striate and T3 smooth or with faint microsculpture (Figs. 42, 52). Hind wing with angle of M and r-m without spur or with faint, unpigmented crease (Fig. 47) 7 7. Head bicolored, face orange and remainder of head brown. Tl of metasoma with length than greater apical width (Fig. 30) Pseudogttaptodon gouleti new species Head brown, some specimens with lighter markings around dorsal margin of eye. Tl of with metasoma length and apical width about equal (Fig. 50) 8 8. Anterior tergites of metasoma uniform brown, concolorous with rest of body. POD shorter than to as as long LOL and shorter than POL (Fig. 44). T3 of metasoma with grooves defining anterolateral corners mostly crenulate (Fig. 52.) P. ocellatus new species Anterior 2 or 3 tergites of metasoma lighter than remainder of body. POD about as long as LOL and POL (Fig. 55). T3 of metasoma with grooves defining anterolateral corners mostly smooth (Fig. 63) P. omissus Fischer 9. Propodeum with apical cell triangular, median carina complete (Fig. 101) P. carinatus new species - Propodeum with apical cell arcuate or incomplete, median carina absent (Fig. Ill) .... 10 10. Body length about 1 mm. Tl of metasoma triangular, posterior margin wider than length, nearly flat, with basal carinae and striae obsolete (Figs. 152, 153) P. minimus new species - Body length usually greater than 1.5 mm. Tl of metasoma with shape various, convex medially with distinct basal carinae, and usually striate or granulate on apical half or more (Figs. 112, 113, 163, 164) 11 11. Labrum as large as clypeus, apically truncate, brown (Fig. 138). Forewing vein 2M markedly longer than RS+Ma (Fig. 140) Pseudogttaptodon labrus new species - Labrum smaller than clypeus, semicircular or traingular, concolorous with other mouthparts. Forewing vein 2M shorter than RS+Ma (Fig. 120) 12 12. Forewing vein R shorter than stigma along anterior wing margin, RS evenly sharply curved (Fig. 90) Pseudogttaptodon brevis new species - Forewing vein R as long as or longer than stigma along antrior wing margin, RS variable, in most specimens straighter apically than basally or decurved (Fig. 120) 13 13. Lateral margins of Tl to entire metasoma yellow to light honey-brown, contrasting with head and mesosoma 14 - to concolorous with and Metasomal tergites unicolorous, medium red-brown black, head 16 mesosoma or slightly lighter 14. Tergites of metasoma uniformly yellow, or with slightly darker shade posterior P. gibsoni new species - brown Tergites of metasoma with lighter color on at most T1-T3, remainder or red-brown 12

15. Junction of 1M and RS + Ma at or basal to midlength of discal cell, RS + Ma about as long to black with anterior metasomal as 1M (Fig. 130). Body dark brown yellow tergites P. hemicolor new species - shorter Junction of 1M and RS+Ma apical to midlength of discal cell, RS+Ma clearly than with to anterior meta- 1M (Fig. 192). Body reddish-brown yellow light honey-brown P- whartoui new somal tergites species 16. Metasoma with disc of T2 with granulate microscuplture posterior to basdl raised area, most o\ tor- from a small area immediately posterior to basal raised area to covering P- curticauda Fischer gum (Fig. 114) - of Metasoma with disc of T2 striate or granulostriate, or smooth with posterior margin 17 basal raised area crenulate (Figs. 175, 186) 17. Metasoma with most of T2 and base of T3 coarsely striate, the anterolateral corners of T3 18 defined by sculpture (Figs. 175, 207) 156 Journal of Hymenoptera Research

- Metasoma with T2 partly to mostly striate, T3 smooth or with some striae mid-basally, the anterolateral corners not defined (Figs. 165, 186) 19 18. Vertex with granulate microsculpture obsolete, appearing smooth (Figs. 167, 168). Ocellar Tl basal triangle large, both LOL and POL3 as long as POD (Fig. 167). Metasomal raised area with median projection (Figs. 173, 175) P. nitidus new species Vertex with well developed granular microsculpture (Figs. 199, 200). Ocellar triangle small, LOL shoter than to as long as POD (Fig. 199). Metasomal Tl basal raised area evenly curved or irregular at midline (Fig. 207) P. ivhitfieldi new species 19. T2 of metasoma finely striate or granulostriate, rarely smooth (Fig. 165); groove between T2 and T3 evenly crenulate throughout, rarely slightly widened medially, T3 smooth P. (Fig. 635). POD about as long as LOL (Fig. 156) minutus (Ashmead) T2 of metasoma coarsely striate (Fig. 186); groove between T2 and T3 wider and more strongly crenulate medially than laterally, with striae extending onto disc of T3 (Fig. 186). POD3 as long as LOL (Fig. 177) P. shawi new species

OMISSUS GROUP 10); T2 + T3 of metasoma, including — grooves, smooth and shining (Fig. 12); legs Included species. This species group in- elongate, femur 4.5-5.53 as long as great- cludes Pseudognaptodon omissus Fischer est width (Fig. 13). and the following new species: P. attenu- Female. —Color: Body uniformly medium atus, P. conjunctus, P. gouleti, P. langori, P. to dark brown as follows: Ventral ocellatus, P. striatus, and P. xanthus. except — or ventral and interior surfaces of scape Remarks. Species of this group are sep- brown; collar to me- arated from the curticauda group by the light pronotal light dium brown; with a narrow following combination of characters: mesopleuron brown band near ventral surface in Frons smooth or with areas of faint gran- light some with tro- ulate sculpture between ocelli and anten- specimens; legs forecoxa, nal chanter, ventral surface of femur, and bas- sockets (Fig. 5). Episternal scrobe pre- al Vs to % of tibia to brown. sent as a sharp, curved groove linked to yellow light Head: of antennal 1. 67-2.00 X posterior margin of mesopleuron (Fig. 1). Length scape Propodeum with a small depression and/ width; flagellum with 18-20 flagellomeres: or area of fine of first three wrinkles mediobasally (Fig. L/W flagellomeres 4.8-5.0, 19). T3 of metasoma with anterolateral 3.67-4.00, 3.00-3.67; L/W of apical flagel- grooves complete and partially to com- lomere 3.67-3.75; MOD 0.86-1 .07X as long pletely crenulate (complete but smooth in as POD; POD 0.75-1. 03 X as long as LOL, P. attenuatus) (Fig. 42). Ovipositor conspic- and 0.60-0.80 X as long as POL (Fig. 4); with setose of uously exserted, portion OOL 2.00-2.25 X as long as POL (Fig. 5); sheath at least half as long as hind basi- head length 0.64-0.67 X width in dorsal tarsus (Fig. 13). view; occiput moderately indented (Fig. 51); head L/H 0.90-0.94 in lateral view; Pseudognaptodon attenuatus Williams, eye L/H 0.65-0.71, eyeH/headH 0.67- new species 0.69, eye width /gena width 1.44-1.88; (Figs. 4-13) wider — gena ventrally than dorsally (Fig. 6); Diagnosis. This species is separated face granulate on lateral Vs and smooth on from other om/ssi/s-group species by the median Vz, most setae as long as clypeus following combination of characters: an- height, clypeus W/H 2.00-2.44, clypeus tenna elongate, with basal flagellomere width 1.11-1.38X as long as malar space. 4.8-5.03 as long as wide; Tl of metasoma Wings: Forewing with RS vein evenly with smooth, protuberant spiracles (Fig. curved, less at apex than at base; RS + Ma Volume Number 2004 13, 1, 157 ^1

Figs. 8-13. Pseudognaptodon attenuatus. 8, Mesosoma in dorsal view. 9, Propodeum in dorsal view. 10, II oi = = metasoma in dorsal view. TIL Tl length, T1W Tl width. 11, Tl of metasoma in Literal view. 12, Anterior of end metasoma in dorsal view. 13, Hind leg.

unpigmented on basal %, tubular through- knob faint or absent (Fig. 7). Mesosoma: out length, distance between point of at- Notauli wide but shallow, merged with to tachment of RS+Ma M and parastigma posterior median depression (Fig. 8); pro- 0.63-0.80 X as long as RS+Mb; 2M spur podeum smooth, medioapical carinae 3.0-5.03 as long as RS + Mb; 2-A1 tubular short, straight, slightly convergent, me- on basal quarter, remainder obsolete and dioapical cell open (Fig. 9). Metasoma: Tl faintly pigmented, first subdiscal cell length 1.04-1. 15x as long as apical width, closed by obsolete veins (Fig. 7). Hind lateral margins concave posterior to spi- wing with r-m 1. 4-2.0 X as long as R; angle racle, the spiracle moderately protuberant, of M and r-m without a spur or thickened lateral carinae about 0.3 tergum length, area; apex of R evenly narrowed apically, smooth and shining throughout, (Figs. 10, 158 Journal of Hymenoptera Research

Figs. 19-23. Pseudognaptodon conjunctus. 19, Propo- in dorsal view. Tl of metasoma in dorsal 0.5 mm J> deum 20, view. 21, Tl of metasoma in lateral view. 22, Anterior = Figs. 14-18. Pseudognaptodon conjunctus. 14, Head in end of metasoma in dorsal view. T2L T2 length. view. in lateral view. in = dorsal 15, Head 16, Head T3W T3 length. 23, Hind leg. anterior view. 17, Mesosoma in dorsal view. 18, Wings. Muerte, 20 km S. Empalme, 2800m, III-IV- 1989, (19), VII-VIII-1989, (2 9), P. Hanson, 11); basal raised area of T2 0.23-0.24 of to- (UWYO). tal T2 length, the posterior margin of basal Etymology. —The name attenuatus refers raised area evenly curved, rarely some- to the elongate appendages, particularly what obsolete; T2 smooth and shining; T3 the legs, of this species. 0.66-0.86 X as long as T2, smooth, with an- Williams, terior and lateral grooves smooth and Pseudognaptodon conjunctus new species somewhat effaced laterally (Fig. 12). Legs: 14-23) Hind femur length 4.5-5.53 maximum (Figs. width, it's ventral hairs as long as or lon- Diagnosis.—This species is separated ger than width of femur at point of attach- from other omissus-group species by the ment (Fig. 13). following combination of characters: Head Material examined.—5 9 16. of , HOLO- bicolored orange and brown, remainder TYPE 9 (UWYO), labelled as follows: body uniform light brown; metasomal Tl "Costa Rica: San Jose, Cerro Muerte 20 km length 1.21-1.363 apical width (Fig. 20); S Empalme 2800m iii-vi 1990 P. Hanson". groove between metasomal T2 and T3 ap- Also red label "HOLOTYPE" and bor- pearing double, separated by a partial to dered label "Pseudognaptodon attenuatus complete fine carina (Fig. 22). Williams Holotype det. D. Williams 2003". Female. —Color: Body and appendages PARATYPES: Costa Rica: San Jose: 16 km medium brown except as follows: Scape S. Empalme, 2500m, III-IV-1989, P. Han- yellow to light honey-brown, face and son I & Gauld, 19, 16, (UYWO). Cerro lower part of gena light orange-brown; Volume 13, Number 1, 2004 159

pronotal collar usually light brown; legs 20, 21); basal raised area of T2 0.31-0.40 of with hind tarsi entirely yellow, slightly total T2 length, the posterior margin of darker. Head: of antennal Length scape basal raised area irregular, slightly con- 1.4-1.6X width; flagellum with 16-17 fla- cave, produced medially; T2 striate near first gellomeres, flagellomere slightly basal raised area margin, rarely with a few curved and narrower than others, L/W of coarse striae extending posteriorly; T3 first three flagellomeres 3.0-3.5, 2.6-3.2, 0.75-0.86 X as long as T2, smooth or with 2.5-2.7; L/W of apical flagellomere 3.2- a few striae mediobasally, with anterior 0.91-1. X 3.5; MOD 00 as long as POD, and lateral grooves smooth, median part POD 1.00-1.33 X as long as LOL, and 1.00- of groove doubled, with fine complete or 1.33X as as long POL (Fig. 14); OOL 2.00- partial ridge joining lateral areas (Fig. 22). X 2.33 as long as POL (Fig. 14); head Legs: Hind femur length 3.22-3.50 X max- length 0.64-0.68 X width in dorsal view; imum width, ventral hairs shorter than fe- and occiput narrowly sharply indented mur width at point of attachment (Fig. 23).

(Fig. 14); head L/H 0.79-0.84 in lateral Material examined.—39 . HOLOTYPE 9 view; eye L/H 0.69-0.71, eyeH/headH (TAMU), labelled as follows: "MEXICO: 0.59-0.64, eye width/gena width 1.69- Jalisco 16 mi. S. Autlan on Hwy. 80, 8-VII- 2.00; gena uniformly wide over most of 1984, J.B. Woolley". Also red label "HO- eye height (Fig. 15); face completely cov- LOTYPE" and bordered label "Pseudog- ered by granular microsculpture, setae as naptodon conjunctus Williams Holotvpe long as clypeus height, clypeus W/H det. D. Williams 2003". PARATYPES: 2.25-2.66, clypeus width 1.00-1.07X as Mexico: Jalisco: 16miS. Autlan on hwy. long as malar space (Fig. 16). Wings: Fore- 80, 8-VII-1984, J.B. Woolley, 1 9 (TAMU). wing with RS vein straighter near apex Mexico: Guerrero: 32miS.E. Petalan, 10- than base, slightly decurved at point of at- VII-1985, Woolley and Zolnerowich, 1 9 tachment to R; RS + Ma unpigmented, bas- (TAMU). — al Vz thinner than remainder, point of at- Etymology. The name conjunctus refers tachment to M almost obsolete, distance to the fine carina that connects the antero- between point of attachment of RS + Ma to lateral areas of T3 of the metasoma. M and parastigma 0.50-0.56 X as long as Pseudognaptodon Williams, new RS+Mb; 2M spur 2.00-2.50X as long as gouleti 2-1 A vein as a crease on species RS+Mb; present ~ s - (Fl S 24 33 ) basal %, first subdiscal cell open (Fig. 18). — Hind wing with r-m 1.0-1.1 X as long as Diagnosis. This species is separated R; angle of M and r-m without spur or from other omfsSMS-group species by the thickened area; apex of R moderately following combination of characters: Head im- with face and knobbed (Fig. 18). Mesosoma: Notauli brown orange, mesosoma pressed, nearly merged posteriorly in a U- metasoma brown; ocellar triangle small, me- with POD to or than inter- shape (Fig. 17); propodeum smooth, equal greater dioapical carinae short, anteriorly conver- ocellar distances and OOL 2.0 time as long as Tl of metasoma gent, medioapical cell open (Fig. 19). Me- POL (Fig. 24); longer hind tasoma: Tl length 1.21-1.36X as long as than apical width (Fig. 30); wing with r-m vien as as R apical width, lateral margins straight to long (Fig. 28). lat- —Color: brown ex- slightly concave posterior to spiracle, Holotype female. Body eral carinae complete on entire tergum cept as follows: face and margin of eyes length or present on basal half but contin- light orange; scape light brown; pronotal with base ued by a raised median area on apical collar light brown; legs yellow of hind dorsal surface of and half, sculpture coarsely striate lateral to coxa, femur, carinae and smooth between carinae (Figs, hind tarsus darker. Head: Length of anten- 160 Journal of Hymenoptera Research

0.5 mm 0.5 mm

0.5 mm

Figs. 29-33. Pseudognaptodon gouleti. 29, Propodeum in dorsal view. 30, Tl of metasoma in dorsal view. Tl of metasoma in lateral view. Anterior end in 31, 32, Figs. 24-28. Pseudognaptodon gonleti. 24, Head of metasoma in dorsal view. 33, Hind dorsal view. 25, Head in lateral view. 26, Head in leg. anterior view. 27, Mesosoma in dorsal view. 28, Wings.

parastigma 0.60 X as long as RS + Mb; 2M nal scape 1.6 width; flagellum with 17 fla- spur 1.77X as long as RS+Mb; 2-1A vein gellomeres; L/W of first three flagello- present as a faintly pigmented crease on meres 3.75, 2.43, and 2.43; L/W of apical basal %, first subdiscal cell open (Fig. 28). flagellomere 3.00; MOD 0.88 X as long as Hind wing with r-m as long as R; angle of POD; POD 1.14X as long as LOL, and as M and r-m without spur or thickened long as POL (Fig. 24); OOL 2.00 X as long area; apex of R moderately knobbed (Fig. as POL (Fig. 24); head length 0.63 X width 28). Mesosoma: Notauli deep, merged with in dorsal view; occiput slightly evenly in- posterior median depression (Fig. 27); pro- dented (Fig. 24); head L/H 0.90 in lateral podeum smooth, medioapical carinae view; eye L/H 0.60, eyeH/headH 0.64, short, straight, medioapical cell open (Fig. eye width /gena width 2.0; gena wider 29). Metasoma: Tl length 1.2X as long as ventrally than dorsally (Fig. 25); face com- apical width, lateral margins straight pos- pletely covered by granular microsculp- terior to spiracle, lateral carinae 0.5 of ter- ture except for narrow median stripe, se- gum length, finely striate lateral to carinae tae as long as clypeus height, clypeus W/ and at apex between carinae (Figs. 30, 31); H 2.50, clypeus width 1.1 X as long as ma- basal raised area of T2 0.26 of total T2 lar space (Fig. 26). Wings: Forewing with length, posterior margin of basal raised RS vein straighter near apex than base; area slightly irregular; T2 striate on basal RS+Ma unpigmented on basal half, basal half posterior to basal raised area margin, Vi thinner than remainder, point of attach- with scattered fine striae reaching to pos- ment to M obsolete; distance between terior margin of tergum; T3 0.80 X as long >oint of attachment of RS+Ma to M and as T2, finely striate medially near anterior Volume 13, Number 2004 1, 161

Figs. 39-43. Pseudognaptodon langori. 39, 0.5 mm Propodeum in dorsal view. 40, Tl of metasoma in dorsal view. 34-38. 41, Tl of metasoma in lateral view. 42, Anterior end Figs. Pseudognaptodon langori. 34, Head in of metasoma in dorsal view. dorsal view. 35, Head in lateral view. 36, Head in 43, hind leg. anterior view. 37, Mesosoma in dorsal view. 38, Wings. Pseudognaptodon langori Williams, new species 34-43) — (Figs. groove, with anterior and lateral grooves Diagnosis. This species is separated smooth to slightly crenulate (Fig. 32). Legs: from other omissus-group species by the Hind femur length 3.6maximum width, following combination of characters: Me- ventral hairs shorter than femur width at sothorax yellow with brown areas dorsal- point of attachment (Fig. 33). ly and ventrally; Tl of metasoma with Material examined.—HOLOTYPE 9 posterior half not raised between lateral (CNCI), labelled as follows: "PANAMA, carinae (Fig. 40); hind wing with r-m vien Panama 8°40'N, 19°50'W 850m, Cerro 2.5 X as long as R (Fig. 38); central disc of Helava 7-14.8.73". Also red striate Campana J. propodeum —(Fig. 39). label "HOLOTYPE", and bordered label Holotype female. Color: Body and ap- "Pseudognaptodon gouleti Williams Holo- pendages yellow to orange-yellow except type det. D. Williams". PARATYPE: Mex- as follows: pedicel and flagellum light ico: Jalisco: Chamela, PT, 4-9-VII-1983, M. brown; ocellai triangle brown; mesoscu- Sharkey 19 (CNCI). tum except for parts of notauli and pos- Remarks. —The above desciption is teromedial depression brown; ventral sur- based on the holotype only. The paratype face of mesothorax brown; propodeum from Mexico agrees closely with these brown; metasoma yellow with brown spot characters, but is not intact and has been on posterior half of T3 and anterior half of poorly mounted. T4; Legs entirely yellow. Head: Length of Etymology. —This species is named for antenna] scape 1.6 width; L/W of first Henri Goulet, who has contributed nu- three flagellomeres 3.71, 2.63, and 2.5; merous specimens to this study. MOD 0.84 X as long as POD; POD 1.25X 162 Journal of Hymenoptera Research

than fe- as long as LOL, and 1.11 X as long as POL mum width, ventral hairs shorter width at of attachment (Fig. 34); OOL 2.22 X as long as POL (Fig. mur point (Fig. 43). 34); head length 0.65 X width in dorsal Material examined.—HOLOTYPE 9 labelled as follows: "Texas: view; occiput moderately indented (Fig. (TAMU), Gon- State I-IV-1984 34); head L/H 0.78 in lateral view; eye L/ zales Co. Palmetto Park J. H 0.79, eyeH/headH 0.60, eye width/ Woolley". Also red label "HOLOTYPE", gena width 2.38; gena uniformly wide and bordered label "Pseudognaptodon lan- det. D. Williams". over most of eye height (Fig. 35); face com- gori Williams Holotype — of this pletely covered by granular microsculp- Remarks. The single specimen in the ture except for narrow median stripe, se- species is the only known specimen tae as long as or slightly longer than clyp- species group with a striate propodeum. eus height, clypeus W/H 2.22, clypeus The color pattern of mixed yellow and width as long as malar space (Fig. 36). brown patches on the thorax is unique. Wings: Forewing with RS vein straighter Both flagella are broken, and have 16 fla- near apex than base, slightly decurved at gellomeres present.— point of attachment to R; RS+Ma almost Etymology. This species is named for x entirely pigmented, basal h slightly thin- David Langor, who has allowed me to ner than remainder but well developed; continue working on the systematics of distance between point of attachment of Braconidae when I should be working on RS + Ma to M and parastigma 0.60 X as weevils. long as RS+Mb; 2M spur 2.6X as long as Pseudognaptodon ocellatus Williams, RS + Mb; 2-1 A vein present as a faintly new species pigmented crease and 2cu-a present as a 44-53) pigmented spur, first subdiscal cell nearly — (Figs. closed (Fig. 38). Hind wing with r-m 2.5 X Diagnosis. This species is separated as long as R; angle of M and r-m without from other om/ssws-group species by the spur or thickened area; apex of R strongly following combination of characters: knobbed (Fig. 38). Mesosoma: Notauli shal- Head, thorax and metasoma dark brown low, narrow, merged with posterior me- to black; ocellar triangle large, LOL and dian depression (Fig. 37); propodeum stri- POL larger than MOD and POD, and OOL ate laterally with slightly raised median 1 .3-1.6 X as long as POL (Fig. 44); occiput area confluent with medioapical cell, me- deeply and evenly indented, head appear- carinae dioapical straight or slightly di- ing C-shaped in dorsal view (Fig. 45). vergent, ending in fine wrinkles, medioap- Female.—Color: Dark brown to black ex- ical cell open (Fig. 39). Metasoma: Tl length cept as follows: Scape light brown, or light as long as apical width, lateral margins brown basally and darker brown apically; slightly convex posterior to spiracle, lat- pronotal collar light brown or with light eral carinae 0.6 of tergum length, not brown patches; legs yellow to orange-yel- raised between carinae, sculpture striate low, rarely with base of hind coxa, dorsal lateral to carinae, weakly striate at apex surface— of femur, and hind tarsus darker. between carinae (Figs. 40, 41); basal raised Head. Length of antennal scape 1.4-1.9X area of T2 0.36 of total T2 length, posterior width; flagellum with 16-18 flagellomeres, of basal margin raised area slightly irreg- first flagellomere slightly curved in some ular but evenly curved; T2 sculpture gran- specimens, L/W of first three flagello- ulostriate on basal half posterior to basal meres 3.3-4.5, 2.7-3.9, 3.0-3.9; L/W of api- raised r area wgin; T3 0.84X as long as cal flagellomere 2.8-3.3; MOD 0.8-1. Ox as T2, smooth, 'ith anterior and lateral long as POD, POD 0.8-1. Ox as long as smoo to grooves slightly crenulate (Fig. LOL, and 0.6-0.8 X as long as POL (Fig. 42). Legs: Him femur length 3.15 maxi- 44); OOL 1.3-1.8X as long as POL (Fig. Volume 13, Number 1, 2004 163

head 0.6-0.7X 45); length width in dorsal 52). Legs: Hind femur length 3.3-3.9 max- view; occiput deeply indented, head C- imum width, ventral hairs shorter than re- in dorsal view head at shaped (Fig. 45); L/H mur width point of attachment (Fig. 53). 0.7-0.8 in lateral view; eye L/H 0.6-0.7, Material examined.—HOLOTYPE 9 eyeH/headH 0.6-0.7, eye width/gena (TAMU), labelled as follows: "Mexico: Oa- width wider than 1.6-2.2; gena ventrally xaca 6miles NE Mitla 20-VII-1985 J. Wool- dorsally (Fig. 46); face granulate laterally ley G. Zolnerowich". Also red label "HO- with smooth to medial stripe smooth on LOTYPE", and bordered label "Pseudog- the medial a Vi, with polished, setae as naptodon ocellatus Williams Holotype det. long as clypeus height, clypeus W/H 2.2- D. Williams 2003". PARATYPES: Mexico: width 1.2-1 .4 X as as 3.0, clypeus long ma- Oaxaca: 6miles N.E. Mitla, 20-VII-1985, J. lar space. Wings: Forewing with RS vein Woolley G. Zolnerowich 3c?, 2 9 (TAMU). or evenly curved, straighter near apex 4miW. Miltepec, 21-VII-1984, J.B. Woolley than base and slightly decurved at point 1 9 (TAMU). 8mi N.E. El Punto, 18-VII- of attachment to R, rarely with RS thinner 1985, Woolley & Zolnerowich, 69, 16* and irregular at some points; RS+Ma un- (TAMU). Mexico: Guerrero: lmi N.E. La- pigmented to pigmented on apical half, guna, elev. approx. 5000', 17-VII-1984, J.B. basal Vi thinner than remainder, point of Woolley, 1 9 (TAMU). 6mi N.E. Tixtla, 16- attachment to M almost obsolete in some VII-1984, J.B. Woolley, 1 9 (TAMU). 6miE. specimens, distance between point of at- Xochilapa, 18-VII-1984 19 (TAMU). 15mi tachment of RS+Ma to M and parastigma W. Chichihualco, elev. approx. 1500', 15- 0.5-0.8X as long as RS+Mb; 2M spur 1.5- VII-1984, J.B. Woolley, 19 (TAMU). Mex- 3.0 X as long as RS + Mb; 2-1 A vein present ico: Veracruz: Los tuxtles Bio. Station as a crease on basal %, first subdiscal cell (Malaise), 15-22-VII-1984, G. Steck, IcT open (Fig. 47). Hind wing with r-m 1.3- (TAMU). Mexico: Puebla: 5mi S.E. Izucar 1.8X as long as R; angle of M and r-m de Matamoras, 20-VII-1984, J.B. Woolley, without spur, rarely slightly thickened; 1 9 (TAMU). Panama: Fortuna: Chiriqui, apex of R moderately knobbed (Fig. 47). 8° 44'N: 82° 15'W, 1050m, 31-VIII-6-IX- H. at 1 Mesosoma: Notauli deep, merged with pos- 1977, Wolda,— light, 9 (RNHL). terior median depression (Fig. 48); pro- Remarks. The specimen from Panama podeum smooth, medioapical carinae is anomalous in having the posterior ocelli short, straight, medioapical cell open (Fig. placed close together (about the length of 49). Metasoma: Tl length 0.9-1.1 X as long the median ocellus apart), but falls within as apical width, lateral margins slightly the range of variation of P. ocellatus in all concave to straight posterior to spiracle, other characters. It is provisionally as- x lateral carinae present on basal h to Vi but signed to this—species, continued by a raised median area on api- Etymology. The name ocellatus refers to cal half or coalesced into a single median the ocellar triangle that is distinctly larger other in this ridge, striate lateral to carinae and smooth than species species group. between carinae, striate across apex in .... omissus Fischer 1965 withoutuu *. raised„„a median™^;™ area~>r.~* Pseuaognaptoaon* specimens re ~ s - 54 T2 0.3- (Fl & M) (Figs. 50, 51); basal raised area of — Fla. 0.4 of total T2 length, posterior margin of Holotype female. Okalosa Co., basal raised area straight but irregular or (USNM). Examined. near basal This is slightly concave; T2 striate Diagnosis.— species separated raised area to striate on most of tergum from other omissus-group species by the with fine wrinkles extending to apex; T3 following combination of characters: 0.7-0.8 X as long as T2, smooth, with an- Head, thorax, and posterior part of meta- face and anterior terior and lateral grooves crenulate (Fig. soma brown, part of me- 164 Journal of Hymenoptera Research

m9 ww

Figs. 44-47. Pseudognaptodon ocellatus new species. 44, Ocelli in dorsal view. 45, Head in dorsal view. 46, Head in lateral view. 47, Wings. Volume 13, Number 1, 2004

in Figs. 48-53. Pseudognaptodon ocellatus new species. 48, Mesosoma dorsal view. 49, Propodeum in dorsal view. 50, Tl of metasoma in dorsal view. 51, Tl of metasoma in lateral view. 52, Anterior end of metasoma in dorsal view. 53, Hind leg.

face with rest of head tasoma yellow to light brown; T2 of me- concolorous except tasoma smooth or granulostriate near bas- near eye margins, ocellai triangle dark al raised area margin and smooth on re- brown; pronotal collar light brown in mainder; Hind wing basal cell narrow, most specimens; metasoma yellow with disc of Tl to r-m 0.7-0.12X as long as R (Fig. 58). brown spot on medium Female. —Color: Body and appendages brown with yellow T2+T3. Legs entirely medium brown except as follows: Scape yellow, rarely with hind tarsi slightly with darker. Head: of antennal yellow to light honey-brown; usually Length scape X face, narrow band around dorsal eye mar- 1. 4-1.6 width; flagellum with 16-18 fla- of first three gin, and lower part of gena light orange- gellomeres, L/W flagello- with meres of yellow to light orange-brown, rarely 3.6-4.7, 3.0-3.9, 2.8-3.4; L/W api- 166 Journal of Hymenoptera Research

X as with an- cal flagellomere 3.0-3.4; MOD 0.61-0.86 0.65-0.75 as long T2, smooth, POD, POD 0.88-1. 20 X as long as LOL, terior and lateral grooves smooth to slight- Hind femur and 0.72-1.14X as long as POL (Fig. 54); ly crenulate (Fig. 63). Legs: ventral OOL 1. 40-2.00 X as long as POL (Fig. 55); length 3.41-3.87 maximum width, head length 0.64-0.67 X width in dorsal hairs shorter than femur width at point of indented attachment view; occiput slightly (Fig. 55); (Fig. 64). — head L/H 0.76-0.86 in lateral view; eye L/ Material examined. Mexico: Guerrero: H 0.65-0.81, eyeH/headH 0.55-0.66, eye 32miS.E. Petalan, 10-VII-1985, Woolley & width/gena width 2.42-3.17; gena uni- Zolnerowich, 26 (TAMU). Mexico: Jalis- formly wide over most of eye height to co: 16miS. Autlan on Hwy. 80, 8-VII-1984, face 1 USA: California: slightly widened ventrally (Fig. 56); J. Woolley, 9 (TAMU). completely covered by granular micro- Contra Costa Co.: Mt. Diablo, ex. Copto- sculpture except for narrow median stripe disca on Q. lobata, 4-XI-1984, D. Wagner, which is slightly raised on dorsal half of 16, 19 (JWCI). Mt. Diablo, ex. Copitodisca face, setae as long as clypeus height, clyp- on Q. douglasii, 4-21-XI-1985, D. Wagner, eus W/H 2.57-2.66, clypeus width 1.14- 1 9 (JWCI). Texas: Brazos Co.: College Sta- 1.33 X as long as malar space (Fig. 57). tion, Lick Creek Park, 22-XI-6XII-1987, Wings: Forewing with RS vein straighter Wharton, Praetorius, 1 9 (TAMU). Brew- near apex than base, slightly decurved at ster Co.: Big Bend National Park, Cotton- point of attachment to R; RS+Ma with wood campsite, 2300', 13-14-VII-1982, G. basal V3-% unpigmented, basal xh thinner Gibson 1 9 (DJMW). Comal Co.: Guada- than remainder, point of attachment to M lupe River State Park, 18-VIII-1988, J. almost obsolete in some specimens, dis- Woolley, 1 9 (TAMU). Hidalgo Co.: Ben- tance between point of attachment of sten Rio grande State Park, 15-XII-1983, J. RS+Ma to M and parastigma 0.45-0.86X Woolley, H. Browning, 19 (TAMU). Jim as long as RS+Mb; 2M spur 1. 40-2.57 X as Wells Co.: 8miW. Ben Bolt, La Copita Re- long as RS+Mb; 2-1 A vein present as a search Sta., 20-V-1987, J. Woolley, 1^59 crease to present but obsolete on basal xh, (TAMU). Arizona: Santa Cruz Co.: first subdiscal cell open (Fig. 58). Hind 4.5miN.E. Patagonia, Hwy. 83, sweeping wing with r-m 0.7-1.2X as long as R; angle Bacharis glutinosa and Chrysothamnus sp., of M and r-m without or thickened G. 1 9 spur Gibson, — (DJMW). area; apex of R slightly knobbed (Fig. 58). Remarks. The holotype female has Mesosoma: Notauli shallow, merged with more developed striae on metasomal T2 posterior median depression (Fig. 59); pro- than other specimens examined, but is podeum smooth, medioapical carinae within the range of variation of other short, convergent, medioapical cell open specimens in the other characters as- (Fig. 60). Metasoma: Tl length 0.83-1.1 X as sessed. Specimens from Big Bend National long as apical width, lateral margins Park and College Station Texas are larger straight to slightly concave posterior to and darker than others, with occiput more spiracle, lateral carinae 0.5-0.6 of tergum deeply indented and head somewhat C- length, convergent basally but parallel shaped in dorsal view, apically, striate lateral to carinae and striatus Williams, smooth to weakly striate at apex between Pseudognaptodon new s ecies carinae 61, basal raised area of P (Figs. 62); " Fi s - 65 74 T2 0.3-0.5 of total T2 length, posterior ( g ) of margin basal raised area irregular, Diagnosis.—This species is separated slightly concave medially to evenly from other om/ssws-group species by the cur ed; T2 sculpture smooth to granulos- following combination of characters: Head near basal raised area margin; T3 capsule bicolored, with face light brown Volume 13, Number 1, 2004 167

Ocelli in dorsal view. I lead in dorsal view. Head in Figs. 54-58. Pseudogrfavtodon omissus Fischer. 54, 55, 56, lateral view. 57, Head in anterior view. 58, Wings.

and remainder dark brown; metasomal Tl grooves (Fig. 73); Hind wing with spur on covered with fine striae (Fig. 71); most of angle of M and r-m (Fig. 69). —Color: dark T2 posterior to basal raised area and base Holotype female. Body of T3 covered with very coarse, heavy stri- brown except as follows: Scape of antenna ae, anterolateral corners of T3 defined by and face light orange-brown; ventral collar of Hymenoptera Research 168 Journal

in dorsal view. in dorsal view. Figs. 59-64. Pseudognaptodon omissus Fischer. 59, Mesosoma 60, Propodeum in dorsal 61, Tl of metasoma in dorsal view. 62, Tl of metasoma in lateral view. 63, Anterior end of metasoma

view. 64, Hind leg.

X in dorsal of pronotum with light and medium length 0.65 width view; occiput head brown areas; legs yellow, except apical slightly evenly curved (Fig. 65); L/ tarsomere of fore- and middle leg and en- H 0.81 in lateral view; eye L/H 0.63, width tire tarsus of hind leg which are dark eyeH/headH 0.67, eye width/gena brown. Head: Length of antennal scape 1.88; gena wider ventrally than dorsally 1.83X width; flagellum with 19 flagello- (Fig. 66); face completely covered by gran- as meres; L/W of first three flagellomeres ular microsculpture, setae as long clyp- 3.43, 3.14, 3.14; L/W of apical flagellomere eus height, clypeus W/H 2.44, clypeus 4.0; MOD 0.80 X as long as POD; POD as width 1.20X as long as malar space (Fig. as LOL, and 0.80 X as long as POL 67). Wings: Forewing with RS vein 65); OOL 2.00 X as long as POL; head straighter near apex than base; RS+Ma Volume 13, Number 1, 2004 169 with basal Vs unpigmented, thinner than Pseudognaptodon xanthus Williams, remainder, point of attachment to M al- new species most distance obsolete, between point of (Figs. 75-85) attachment of RS + Ma to M and parastig- Diagnosis. —This species is separated ma 0.60 X as long as RS + Mb; 2M spur from other omissus-group species by the 3.6 X as long as RS+Mb; 2-1 A vein present following combination of characters: Body but obsolete and very faintly pigmented to ocelli x pale yellow light honey-brown; on basal h, first subdiscal cell open (Fig. finely pitted on medial side (Fig. 75); Tl 69). Hind wing with r-m 1.8 X as long as and T2 of metasoma with grooves smooth R; angle of M and r-m with posteriorly di- and microsculpture somewhat obsolete rected, obsolete, and faintly pigmented (Figs. 82, 84). spur; apex of R moderately knobbed (Fig. Female.—Color: Body and appendages 69). Mesosoma: Notauli shallow, not con- pale yellow to honey-brown, most darker vergent posteriorly, merged with lateral specimens darker dorsally than laterally edges of posterior median depression (Fig. or ventrally, except as follows: Flagellum 68); propodeum smooth, medioapical ca- of antenna brown in most specimens; ocel- rinae straight, medioapical cell open (Fig. lar triangle brown; propodeum light 70). Metasoma: Tl length 1.04X as long as brown in most specimens; rarely scutum apical width, lateral margins straight pos- of thorax and terga of abdomen with var- terior to spiracle, lateral carinae about half ious light brown patches or deeper of striate tergum length, finely throughout shades. Head: Length of antennal scape except near base between carinae, (Figs. 1.5-1.6 width; flagellum with 17-18 flagel- 71, 72); basal raised area of T2 0.21 of total lomeres; L/W of first three flagellomeres T2 of basal raised length, posterior margin 3.3-3.8, 3.3-3.8, 3.0-3.3; L/W of apical fla- area bisinuate; basal % of T2 irregularly gellomere 2.7-3.7; MOD 0.75-0.90 X as basal raised area beyond coarsely striate; long as POD; POD 0.95-1.14 x as long as T3 0.64 X as as T2, striate belong coarsely LOL, and 0.75-1. 00 X as long as POL (Fig. tween lateral with anterior and grooves, 75); ocelli with fine pits medially (Fig. 75); lateral crenulate grooves (Fig. 73). Legs: OOL 2.00-2.33 X as long as POL (Fig. 76); Hind femur length 3.46 maximum width, head length 0.61-0.66 X width in dorsal ventral hairs shorter than femur width at view; occiput slightly evenly curved (Fig. point of attachment (Fig. 74). 76); head L/H 0.79-0.89 in lateral view; Material examined.—HOLOTYPE $ eye L/H 0.71-0.82, eyeH/headH 0.61- (TAMU), labelled as follows: "Venezuela: 0.67, eye width/gena width 2.0-2.17; gena Aragua 22 km. south Colonia Tovar 900 uniformly wide over most of eye height to R. meters December 23, 1985 P. Kovarik, slightly widened ventrally (Fig. 77); face Jones". Also red label "HOLOTYPE" and completely covered by granular micro- bordered label "Pseudognaptodon striatus sculpture, setae as long as clypeus height, Williams Holotype det. D. Williams 2003". clypeus W/H 1.8-3.00, clypeus width Remarks. —This species is known only 1.13-1 .28X as long as malar space (Fig. with vein from the holotype. The metasomal sculp- 78). Wings: Forewing RS ture and hind wing spur on the M/r-m straighter near apex than base, slightly de- attachment angle are distinctive characters that sup- curved at point of to R; with basal xh to entire vein port the separation of this specimen from RS+Ma unpig- basal XA thinner than other Pseudognaptodon species. mented, remainder, of attachment to almost obsolete Etymology. —The name striatus refers to point M the metaso- in some distance between at- the sculpture of T2 and T3 of specimens, ma. tachment of RS + Ma to M and parastigma 170 Journal of Hymenoptera Research

0.2 mm

0.2 mm 0.5 mm

Figs. 65-69. Pseudognaptodon striatus. 65, Head in dorsal view. 66, Head in lateral view. 67, Head in anterior view. 68, Mesosoma in dorsal view. 69, Wings. Volume 13, Number 2004 1, 171

long as T2, smooth, with anterior and lateral grooves smooth to slightly crenulate (Fig. 84). Legs: Hind femur length 3.13- 3.88 maximum width, ventral hairs shorter than femur width at point of attachment (Fig. 85). Material examined.—76 , 229. HOLO- TYPE FEMALE (CNCI), labelled as follows: "U.S.A Georgia 15 km. W. Fargo Okefenokee Swamp Nov. 1979 D. Wil- liams coll.". Also red label "HOLOTYPE" and bordered label "Pseudognaptodon xanthus Williams Holotype det. D. Williams 2003". PARATYPES: U.S.A. Georgia: 15 km W. Fargo, Okefenokee Swamp, XI- 1979, D. Williams, Id, 19 (DJMW), 9 9 (CNCI). 15 km W. Fargo, Okefenokee Swamp, 82° 20'W: 40° 30'N, slash pine stand, 13-XI-1979, D. Williams 3 9 (RNHL). Florida: Sarasota Co.: Myakka, Figs. 70-74. Pseudognaptodon striates. 70, Head in dor- 16-18-1-1984, R. Wharton 16, 2 9 sal view. 71, Head in lateral view. 72, Head in anterior (TAMU). Florida: Lake Dorr Rec. Ocala Net. view. 73, Mesosoma in dorsal view. 74, Wings. Area, Forest, 20-XI-1979, D. Williams 1 9 (gold coated for S.E.M.) (DJMW). Texas: San Ja- 0.50-0.75 X as long as RS+Mb; 2M spur cinto Co.: Big Creek Scenic Area, 7-1 II- 1.60-2.25 X as as RS + 2-1 vein R. long Mb; A 1987, Wharton J. Heraty, \6 (TAMU). present as a crease to present but obsolete Tyler Co.: Kirby State Forest, 3miS. War- and on basal 2 very faintly pigmented A, ren, 22-V-1984, J. Woolley 26, 49 first subdiscal cell open (Fig. 79). Hind (TAMU). Montgomery Co.: Jones State wing with r-m 0.5-1.0 X as long as R; angle Forest, 8miS. Conroe, 4-18-X-1987, R. of M and r-m without spur or thickened Wharton, 1 9 (TAMU). Costa Rica: Guan- area; apex of R slightly to moderately acaste: Santa Rosa Nat. Park, 300m, ex. knobbed (Fig. 79). Mesosoma: Notauli shal- Malaise trap, Site# H-2-C, (H) open regen- low, indistinct on posterior half (Fig. 80); erating woodland <10 years old, (C) more propodeum smooth, medioapical carinae or less fully shaded as possible, 24-V-14VI- slightly arcuate and convergent, medioap- 1986, I. Gauld D. Janzen 26 (UWYO). ical — cell open (Fig. 81). Metasoma: Tl length Remarks. Specimens from Florida are 0.85-0.96 X as long as apical width, lateral generally pale, and those from Texas gen- margins straight to slightly concave pos- erally darker. One of the two male speci- terior to spiracle, lateral carinae 0.5-0.8 of mens from Costa Rica has distinctiwk tergum length, weakly striate lateral to ca- medium-brown abdominal terga, but re- rinae and smooth to weakly striate at apex sembles other specimens in most other between carinae basal raised characters. (Figs. 82, 83); — area of T2 0.33-0.41 of total T2 length, pos- Etymology. The name xanthus refers to terior margin of basal raised area evenly the almost entirely yellow body color. and some- curved, rarely slightly irregular CURTICAUDA GROUP what obsolete; T2 sculpture obsolete gran- — ulostriate to weakly striate near basal Included species. This species group in- raised area to basal half; T3 0.60-0.73 X as cludes Pseudognaptodon curticauda Fischer, 172 Journal of Hymenoptera Research

Figs. 75-79. Pseudognaptodon xanthus. 75, Ocelli in dorsal view. 76, Head in dorsal view. 77, Head in lateral view. 78, Head in anterior view. 79, Wings.

P. minutus (Ashmead), and the following Remarks.—The species of this group are new species: P. brevis, P. carinatus, P. gibsoni, separated from the omissus-group by the P. hemicolor, P. lab us, P. minimus, P. nitidus, following combination of characters: Most P. P. whart shawi, ni, and P. whitfieldi. species with frons, including ocellar trian- Volume 13, Number 1, 2004 173

Figs. 80-85. Pseudognaptodon xanthus. 80, Mesosoma in dorsal view. 81, Propodeum in dorsal view. 82, II ol metasoma in dorsal view. 83, Tl of metasoma in lateral view. 84, Anterior end of metasoma in dorsal view.

85, Hind leg.

gle, with fine but distinct granulate sculp- 114). Ovipositor barely exserted, setose ture between posterior ocelli and antenna 1 portion oi sheath less than half as long as hind basitarsus sockets (Figs. 86, 87). Episternal scrobe pre- (Fig. 115). sent as a faint depression or absent (Figs. Pseudognaptodon brevis Williams, new 2, 3). Propodeum without small depression species and /or area of fine wrinkles mediobasally T3 of metasoma with an- (Figs. 86-95) (Figs. Ill, 132). — terolateral grooves lacking, or with grooves Diagnosis. This species is separated other smooth and faintly impressed near lateral from curticauda-group species by the anterolateral ar- combination of characters: margins of tergite, or with following of frons obsolete in eas defined by microsculpture (Figs. 104, Sculpture except ocel- 174 Journal of Hymenoptera Research

Vi, with granular microsculpture laterally, setae as long as clypeus height, clypeus W/H 1.8, clypeus width 1.5X as long as malar space (Fig. 88). Wings: Forewing with RS vein evenly sharply curved, with R vein shorter than length of stigma on anterior margin; RS + Ma slightly pigmented and evenly sclerotized throughout length; 2M spur 3.0 X as long as RS + Mb; 2-A1 present as a sclerotized vein on basal 2 /3 and a pigmented crease on remainder; spur of m-cu present, subdiscal cell closed (Fig. 90). Hind wing with r-m 1.5X as long as R; Rs 5.4 X as long as R; apex of R with moderately developed knob (Fig. 90). Me- sosoma: Mesoscutum with median groove distinct, merged with notauli near posterior margin of mesoscutum, posteromedial depression small (Fig. 89); propodeum with medioapical carinae straight, conver- 0.5 mm gent, basal cell open (Fig. 91). Metasoma: Tl 0.9 X as as width, lateral Figs. 86-90. Pseudognaptodon brevis. 86, Head in dor- long apical sal view. 87, Head in lateral view. 88, Head in ante- margins convex posterior to spiracle, spi- rior view. Mesosoma in dorsal view. 89, 90, Wings. racle moderately protuberant, lateral carinae indistinct, merged with other striae; Tl evenly convex and finely striate lar triangle; MOD and POD half as long throughout, except for basal area which is as interocellar distances or vein R of less; separated from remaider by a transverse shorter than of on forewing length stigma ridge (Figs. 92, 93); basal raised area of T2 anterior wing margin (Fig. 90). 0.25 of total T2 length, posterior margin of —Color: dark Holotype female. Body basal raised area irregular; basal raised brown for the Pedical except following: area margin obscured by coarse striae, T2 and first two flagellomeres orange-brown; otherwise smooth; T3 0.75 X as long as T2, legs light orange-brown with brown hind smooth, with anterior groove crenulate coxa. Head: Length of antennal scape 1.5X (Fig. 94). Legs: Hind femur length 4.3 X width; flagellum with 16 flagellomeres: L/ maximum width (Fig. 95). W of first three flagellomeres 4.3, 3.9, 3.2; Material examined.—HOLOTYPE 9 L/W of apical flagellomere 3.0; MOD 0.9X (CNCI), labelled as follows: "Chile: Mal- as long as POD; POD 0.5 X as long as LOL, X leco: Princessa 20 km W Curacautin, and 0.4 as long as POL (Fig. 86); OOL as 12.XII.1984-16.II.1985, S&J Peck, 300m, long as POL (Fig. 86); vertex without gran- Notho. for.". Also red label "HOLO- ular microsculpture except in ocellar tri- TYPE" and bordered label angle; head length 0.6 X width in dorsal "Pseudognapto- don brevis Williams det. D. Wil- view; occiput narrowly indented (Fig. 86); Holotype head liams 2003". L/H 0.7 in lateral view; eye L/H 0.7, — eyeH/headH 0.6, eye width/gena width Etymology. The name brevis refers to 1.8 (Fig. 87); gena wider ventrally than the very short R vein of the forewing, dorsally (Fig. 87); face smooth on medial unique to this species within the genus. Volume Number 2004 13, 1, 175

microsculpture and fine wrinkles between lateral ocelli and dorsal margin of eye; head length 0.6 X width in dorsal view; oc-

ciput narrowly indented (Fig. 97); head L/ H 0.7 in lateral view; eye L/H 0.7, eyeH/ headH 0.6, eye width/gena width 1.8 (Fig. 98); gena wider ventrally than dorsally (Fig. 98); face granulate with a polished area dorsal to clypeus and a polished stripe that is narrow dorsally, setae as long as clypeus height, clypeus W/H 2.2, clypeus width as long as malar space. Wings: Forewing with RS vein apically decurved; RS+Ma pigmented and tubular, slightly thinner basally than apically; 2M spur 3. OX as long as RS+Mb; 2-A1 absent; spur of m-cu absent, first subdiscal cell open (Fig. 99). Hind wing with r-m 1.6X as long as R; Rs 6.0X as long as R and thickened basally; apex of R with poorly to moderately developed knob (Fig. 99). Mesosoma: Mesoscutum with median Figs. 91-95. Pseudognaptodon brevis. 91, Propodeum groove present near posteromedial de- in dorsal view. Tl of metasoma in dorsal view. 92, pression, notauli merged with one another Tl of metasoma in lateral view. Anterior end 93, 94, on anterior of de- of in margin posteromedial metasoma dorsal view. 95, Hind leg. pression (Fig. 100); propodeum with basal cell complete, triangular, median carina complete to anterior propodeal margin Pseudognaptodon carinatus Williams, (Fig. 101). Metasoma: Tl length as long as new species apical width, lateral margins slightly con- (Figs. 96-105) — cave posterior to spiracle, spiracle mod- Diagnosis. This species is separated erately protuberant, lateral carinae present from other cnrticauda-group species by the on basal half and present but merged with following combination of characters: Pro- coarse striae on apical half; Tl striate on podeum with triangular basal cell and apical half and lateral to carinae (Figs. 102, complete median carina (Fig. 101). 103); basal raised area of T2 0.3X of total Female. —Color: Body dark brown except T2 length, posterior margin of basal raised for the following: Scape light orange- area evenly curved but slightly sinuate brown; Pronotal collar and part of meso- medially; basal raised area margin erenu- pleuron with light orange-brown area; late, with a few obsolete striae near midlegs yellow, except hind tarsus light line; T3 as long as T2, smooth, with ante- brown. Head: Length of antennal scape rior groove smooth, sinuate, and some- 1.5-1.6X width; flagellum with 20-21 fla- what obsolete Laterally (Fig. 104). Legs: gellomeres: L/W of first three flagello- Hind femur length 2.9X maximum width meres 2.2, 2, 2.2-2.4; L/W of apical flagel- (Fig. 105). as as Material examined.— lomere 2.3; MOD as long POD; POD 39 , Id. HOLO- long as LOL, and 0.8-0.9 X as long as POL TYPE 9 (TAMU), labelled as follows: 6.2 (Fig. 96); OOL 2.0 X as long as POL (Fig. "Mexico: Guerrero mi SW Xochipala 97); vertex with well developed granular VII-6-1987 5670 ft R. Wharton". Also red 176 Journal of Hymenoptera Research

96-99. Figs. Pseudognaptodon carinatus. 96, Ocelli in dorsal view. 97, Head in dorsal view. 98, Head in lateral view. 99, Wings. Volume Number 2004 13, 1, 177

—.

Figs. 100-105. Pseudognaptodon carinatus. 100, Mesosoma in dorsal view. 101, Propodeum in dorsal \ lew. 102, Tl of metasoma in dorsal view. 103, Tl of metasoma in lateral view. 104, Anterior end ot metasoma in dorsal view. 105, Hind leg.

label "HOLOTYPE" and bordered label to the complete median carina of the pro- "Pseudognaptodon carinatus Williams Ho- podeum. lotype det. D. Williams 2003". PARA- curticauda Fischer TYPES: Chiapas: Tapachula, 22-24-IX- Pseudognaptodon 1967 1987, R. Wharton, 16* (TAMU). Costa 106-115) Rica: Guanacaste: Santa Rosa Nat. Park, (Figs.— ex. Malaise, 300m, site 27.IX, (H) open re- Holotype female. Mexico, on Mesquite generating woodland <10 years old, (O) leaf, Brownsville no. 34069, 12-X-1943 isolated of 18- Examined. in clearing, fully part day, (USNM). — IX-1986, I. Gauld D. Janzen, 2 9 (UWYO). Diagnosis. This species is separated Etymology. —The name carinatus refers from other curticauda-group species by the 178 Journal of Hymenoptera Research following combination of characters: half; 2M spur 1.6-3.3 X as long as RS+Mb; Head, mesosoma, and most or all of me- 2-A1 absent to present as an unpigmented tasoma uniformly dark brown or black in crease on basal half; spur of m-cu absent a first sub- color, rarely with lighter clypeus or pro- or present as very small lobe, notal collar; metasoma with Tl about as discal cell open (Fig. 109). Hind wing with r-m 0.8-1.3 X as as Rs 3.6-5.6 X as wide as long (Fig. 112), and T2 finely gran- long R; ulate near apex of basal raised area to long as R; apex of R moderately to well most of center of tergum in a semicircular developed knob (Fig. 109). Mesosoma: Me- with median no- pattern (Fig. 114). soscutum groove absent, Female. —Color: Body dark red-brown to tauli merged with one another on anterior black except as follows: Scape and pedicel margin of posteromedial depression (Fig. with cari- yellow to light brown; rarely with clypeus 110); propodeum medioapical light brown; pronotal collar light orange- nae straight, parallel, and widely separat- brown in some specimens; legs yellow to ed to cell closed by complete carina, rarely light orange-brown, rarely with light with irregular very fine wrinkles, or with brown apical tarsomeres and hind tarsus, other irregular small carinae (Fig. 111), or hind tibia darker apically, or femora Metasoma: Tl length 0.9-1. 2 X as long as with darker area on dorsal surface. Head: apical width, lateral margins slightly con- Length of antennal scape 1 .4-1.8 X width; cave to straight posterior to spiracle, spi- flagellum with 13-17 flagellomeres: L/W racle not to strongly protuberant, lateral of first three flagellomeres 2.5-3.9, 2.2-3.3, carinae present on basal half to complete 2.3-3.2, rarely with flagellomeres shorter to apex or nearly so, rarely merged with apically; L/W of apical flagellomere 2.3- coarse striae or continuous with lateral 3.3; MOD 0.6-1. OX as long as POD; POD margins of a medial raised area; Tl striate 0.8-1. OX as long as LOL, and 0.6-1.0 X as throughout except for small area at base long as POL (Fig. 106); OOL 1.4-2.1 X as of tergum to striate on apical half, (Figs, long as POL (Fig. 107); vertex with well 112, 113); basal raised area of T2 0.2-0.4 of developed granular sculpture anterior to total T2 length, posterior margin of basal ocelli to entire vertex granular, with very raised area broadly concave with lateral fine, incomplete suture or fine wrinkles lunules to evenly curved, irregular, slight- present between lateral ocelli and dorsal ly medially produced, or rarely toothed; eye margin; head length 0.6-0.7 X width in basal raised area margin smooth or weak- dorsal view; occiput shallowly, narrowly ly crenulate, smooth to granulate on most to widely indented, head somewhat C- of tergum posterior to basal raised area in in shaped dorsal view (Fig. 107); head L/ a semicircular pattern; T3 0.8-1. OX as long H 0.7-0.8 in lateral view; eye L/H 0.6-0.7, as T2, smooth to granulate on basal V3 of eyeH/headH 0.6-0.7, eye width/gena disc, with anterior groove smooth to width 1.2-1.9 (Fig. 108); gena parallel sid- weakly crenulate, rarely with groove wid- ed face (Fig. 108); granulate, rarely with a ened or decurved medially (Fig. 114). Legs: small polished area dorsal of clypeus and Hind femur length 2.7-3.6 X maximum a narrow median polished strip, setae width (Fig. 115). shorter than or as long as clypeus height, Material examined.—Costa Rica: Guana- clypeus W/H 1.8-3.0, clypeus width 0.8- caste: P.N. Santa Rosa, 200m, 1-1991, P. 1.3X as long as malar space. Wings: Fore- Hanson, 2 9 (UWYO). Guanacaste Conser- wing with RS vein curved but straighter vation Area, Santa Rosa hdq., 200m, light apically than basally, decurved, or de- trap, 7-VII-1997, L. van der Ent, 1? curved and sinuate at midlength; RS+Ma (UWYO). Santa Rosa Natl. Park, 300m, ex. unpigmented on basal Vz and tubular Malaise trap, (H) open regenerating throughout length, rarely thinned on basal woodland <10 years old, (C) more or less Volume 13, Number 1, 2004 179

0.5 mm

curticauda Fischer. 106, Ocelli in dorsal view. 107, Head in dorsal view. 108, Figs. 106-109. Pseudognaptodon Head in lateral view. 109, Wings. 180 Journal of Hymenoptera Research

in dorsal Figs. 110-115. Pseudognaptodon curticauda Fischer. 110, Mesosoma in dorsal view. Ill, Propodeum view. 112, Tl of metasoma in dorsal view. 113, Tl of metasoma in lateral view. 114, Anterior end of metasoma

in dorsal view. 115, Hind leg.

fully shaded as possible, 14-VIII-6-IX-1986 Cerro el Hacha, N.W. Volcan Orosi, 300m (16*), 8-II-2-III-1986 (19), 23-III-13-IV-1986 1988, 29, IS (UWYO). Puntarenas: R.F. (1$), 21-II-14-III-1986 (19), 26-II-14-III- Golfo Dulce, 3 km S.W. Rincon, 10m, XII-

1987 (39,16), 23-111-1986 (1 6 ), I. Gauld D. 1992, P. Hanson, 19 (UWYO). San Vito, Janzen, (UWYO). Santa Rosa Natl. Park, Estac. Biol. Las Alturas, 1500m, III-1992, P. 300m, ex. Malaise trap, (H) open regener- Hanson, 1 9 (UWYO). Mexico: Guerrero: ating woodland <10 years old, (O) in 6.2miS.W. Xochipala, 13-VII-1985, Wool- clearing fully isolated part of the day, 28- ley & Zolnerowich, 3 9 (TAMU). 6miE. XII-1985-18-I-1986, I. Gauld D. Janzen, 1 9 Xochipala, 13-VII-1985, Woolley & Zolne- (UWYO). Estac. Pitilla, 9 km S. Santa Ce- rowich, 3J, 2 9 (TAMU). 6miE. Xochipala, T cillia, 700m, V-1989, Gauld, 1 9 (UWYO). 18-VII-1985, Woolley & Zolnerowich, 29, Volume 13, Number 1, 2004 181

Id (TAMU). 15miW. Chichihualco, elev. Stumbergs Patio Ranch, 5.6miW. Hunt, approx. 1500', 15-VII-1984, J. Woolley, 1 9 2000', 1-2-VII-1982, G. Gibson, 26 2miN. (TAMU). Cacahuamilpa, 19-VII- (DJMW). Presidio Co: Big Bend Ranch 1984, J. Woolley, Id (TAMU). 17miE. Tix- SNA, Agua Adentro, 29° 40' N, 104° 06"W, tla, G. ll-VII-1985, J. Woolley Zolnerow- 14-V-1990, R. Wharton, 1 6 (TAMU). 16 7miW. — ich, 19, (TAMU). Chilapa, 16- Remarks. This species shows the most VII-1984, Id J. Woolley, 39, (TAMU). character variation of any Pseudognaptodon 32miS.E. Petalan, 10-VII-1985, Woolley & in this study. There is much more varia- Zolnerowich, 2 9 (TAMU). Michoacan: tion in P. curticauda than P. minutus (Ash- 49miS.E. Aguila, 13-VII-1984, J. Woolley, mead) and P. whartoni new species for ex- 19 (TAMU). 2miS. Carapan, 6-VII-1985, ample, which are described here from Woolley & Zolnerowich, 1 6 (TAMU). Oa- similar numbers of specimens. Specimens xaca: 8miN.E. El Punto, 18-VII-1985, assigned to this species are united primar- & 19 Woolley Zolnerowich, (TAMU). ily by overall uniform dark body color 10.8miS. El Punto, 6100', 9-VII-1987, R. and granulate sculpture of T2+T3 of the Wharton, 1 9 (TAMU). Puerto Escondido, metasoma, which are common characters 1 in 15-VII-1985, Woolley & Zolnerowich, 9 the genus and may represent the ple- 6miN.E. (TAMU). Mitla, 20-VII-1985, J. siomorphic states. These specimens may Woolley G. Zolnerowich, 59, 4 6 (TAMU). represent several very similar species with 19miS. San Miguel Suchixtepec, 17-VII- overlapping ranges of characters. The ma- 1985, Woolley & Zolnerowich 2 9 jority of specimens are from the adjacent (TAMU). 17miN.W. Tehuantepec, 15-VII- Mexican states Guerrero and Oaxaca. 1987, R. Wharton, 19 (TAMU). lOmiS.E. These specimens are the most uniform in Totolapam, 4000', 20-VII-1987, R. Whar- appearance, and the most similar to the ton, 19 (TAMU). 3.2miS.W. La Cumbre, holotype. Short series or single specimens 18-VII-1985, Wooley & Zolnerowich, Id from Costa Rica and the United States (TAMU). Puebla: 5miS.E. Izucar de Ma- vary in color, wing venation, shape and tamoras, 20-VII-1984, J. Woolley, 3 9, 16 sculpture of metasomal Tl, and other (TAMU). U.S.A.: Arizona: Santa Cruz characters that are usually important in Co.: l.OmiS. Pena Blanca Lk., 4100', 6-VIII- species diagnosis. These differences are 1982, G. Gibson, 1 9 (DJMW). California: consistent, but less in magnitude than dif- S.L.O. Co.: S.L.O. reservoir, 21-V-1975, R. ference among other species in the genus. Wharton, 19 (TAMU). New Mexico: It cannot be determined from the speci- Otero Co.: Cloudcroft, 8600', 24-VII-1982, mens available if this represents geo- G. Gibson, 26 (DJMW). Texas: Brewster graphic variation or separate species. Co.: Big Bend National Park, Oak Cyn, Williams, new Window Trail, 5400', 24-27-VI-1982, G. Pseudognaptodon gibsotti species Gibson, 29 (DJMW). Big Bend National (Figs. 116-125) Park, Cottonwood Campsite, 2300', 13-14- VII-1982, G. Gibson, 26 (DJMW). Culber- Diagnosis. —This species is separated son Co.: 3.6miS. Pine Springs, old Gua- from other curticauda-group species by the dalupe Pass Road nr. Guadalupe Springs, following combination oi characters: Me- 5200', sweeping flowering Acacia constric- tasoma yellow, with slight brown tint on ta, 20-22-VII-1982, G. Gibson, 19, 16 middle of dorsal surface of terga from T3 (DJMW). Guadalupe Nat'l Park, Choza to end of metasoma; Tl of metasoma than Springs, 5100', 22-VII-1982, 19, 26 slightly wider apically long (Fig. (DJMW). Hidalgo Co.: Bensten Rio 122); T2 of metasoma finely granulate, basal raised area to of Grande State Park, 15-XII-1983, J. Woolley near apex of most H. Browning, 19, Id (TAMU). Kerr Co.: center of tergum (Fig. 124). 182 Journal of Hymenoptera Research

Figs. 121-125. Pseudognaptodon gibsoni. 121, Propo- deum in dorsal view. 122, Tl of metasoma in dorsal Figs. 116-120. Pseudognaptodon gibsoni. 116, Head in view. Tl of metasoma in lateral view. An- dorsal view. 117, Head in lateral view. 118, Head in 123, 124, terior end of metasoma in dorsal view. Hind anterior view. 119, Mesosoma in dorsal view. 120, 125, leg. Wings.

strip, setae much shorter than clypeus Female. —Color: Head and mesosoma height, clypeus W/H 2.4-2.6, clypeus dark brown except for the following: width 1.4X as long as malar space (Fig. Scape yellow; clypeus light orange-brown; 118). Wings: Forewing with RS vein pronotal collar light to medium orange- straight on apical half; RS + Ma unpig- brown; metasoma yellow, with light mented, thinned on basal half to obsolete brown tint on dorsal surface of T3 to end near base; 2M spur 1. 8-2.2 X as long as of metasoma. Head: Length of antennal RS+Mb; 2-A1 present as an unpigmented scape 1.6-1.8X width; flagellum with 18 crease on basal half; spur of m-cu absent, flagellomeres: L 9 W of first three flagello- first subdiscal cell open (Fig. 120). Hind meres 2.7-2.8, 2.3, 2.2-2.3; L9W of apical wing with r-m 1.3-1.5 X as long as R; Rs flagellomere 2.3; MOD 0.8-0.9 X as long as 4.4-5.8X as long as R; apex of R with well POD; POD 0.8-0.9 X as long as LOL, and developed knob (Fig. 120). Mesosoma: Me- 0.7X as long as POL; OOL 1.2-1.4 X as soscutum with median groove absent, no- long as POL (Fig. 116); vertex with well tauli merged with one another on anterior developed granular sculpture anterior to margin of posteromedial depression (Fig. ocelli; head length 0.6-0.6 X width in dor- 119); propodeum with medioapical cari- sal view; occiput narrowly indented (Fig. nae slightly converging, widely separated, 116); head L/H 0.6-0.7 in lateral view; eye irregular very fine wrinkles inside basal L/H 0.5-0.6, eyeH/headH 0.6, eye width/ cell nearly closing cell apex (Fig. 121). Me- gena width 0.9-1.2 (Fig. 117), gena wider tasoma: Tl length 1.1-1.3 X as long as api- ventrally than dorsally (Fig. 117); face cal width, lateral margins straight poste- granulate with a narrow median polished rior to spiracle, spiracle slightly protuber- Volume Number 2004 13, 1, lg3

ant, lateral carinae on basal present half; clypeus yellow to light orange-brown in Tl striate on half and lateral to ca- apical some specimens, face lighter brown near rinae 122, 123); basal raised area of than (Figs. eye margins medially; light orange- T2 0.3 of total T2 length, posterior margin brown laterally on first two or three me- of basal raised area basal tasomal evenly curved; terga; legs yellow with light raised area smooth to margin slightly brown first hind tarsomere. Head: Length crenulate, on most of of antennal granulate tergum scape 1.5-1.8X width; flagel- to basal raised area in a posterior semicir- lum with 16-19 flagellomeres: L/W of first cular T3 0.8-0.9X pattern; as long as T2, three flagellomeres 2.3-3.0, 2.1-2.7, 2.1- with anterior smooth, groove weakly cren- 2.5; L/W of apical flagellomere 2.7-2.9; ulate and curved evenly (Fig. 124). Legs: MOD 0.9-1 .Ox as long as POD; POD 0.8- Hind femur 3.5-3.7X length maximum 1.0X as long as LOL, and 0.6-0.8 X as long width as (Fig. 125). POL (Fig. 126); OOL 1. 5-2.0 X as long Material examined.— HOLOTYPE 9 as POL (Fig. 127); vertex with well devel- laballed as follows: (CNCI), "USA Texas oped granular sculpture anterior to ocelli, Ward Co.: St. Monahans Pk., 6.0 mi N.E. very fine suture or fine wrinkles present Monahans 3000' 22.VI.82 G.A.P. Gibson, between lateral ocelli and dorsal eye mar- sweeping Quercus havardi" . Also red label gin; head length 0.6 X width in dorsal "HOLOTYPE" and bordered label "Pseu- view; occiput narrowly, medially indented dognaptodon gibsoni Williams Holotype (Fig. 127); head L/H 0.7-0.8 in lateral det. D. Williams 2003". PARATYPE: 19 view; eye L/H 0.6-0.7, eyeH/headH 0.6, with same data as holotype (CNCI). eye width/gena width 1.2-1.5 (Fig. 128);

. — Ltymology This species is named for gena parallel sided (Fig. 128); face corn- Gary Gibson, whose collection of speci- pletely granulate to a small polished area mens of several undescribed species (in- dorsal of clypeus and a narrow median eluding the above) started me on this pro- polished strip, setae shorter than or as ject. long as clypeus height, clypeus W/H 2.0- 2.3, clypeus width 1.1 X as long as malar Pseudognaptodon hemicolor Williams, . space (Rg U9) wings Forewing with RS new species vein Recurved apically; RS+Ma unpig- (Hgs. 126-135) — mented, tubular throughout length, rank Diagnosis. This species is separated thinned on basal half and obsolete at ex- from other curticauda-growp species by the treme base, about as long as 1M; 2M spur following combination of characters: Ocel- 2.0-3.3X as long as RS+Mb; 2-A1 absent; lar triangle darker in color than vertex, spur of m-cu absent, first subdiscal cell face lighter laterally than medially; pro- open (Fig. 130). Hind wing with r-m 1.0- podeum with basal cell closed (Fig. 132); 1.3X as long as R; Rs 4.2-5. 2x as long a^ metasoma medium to dark brown apical- R; apex of R with well developed knob lateral with ly, yellow to light orange-brown on (Fig. 130). Mesosoma: Mesoscutum margins or most of first two or three terga; median groove absent to faint, notauli junction of RS + Ma and 1M apical of mid- merged with lateral margins of postero- length of discal cell (Fig. 130); Tl of me- medial depression (Fig. 131); propodeum

a bcT-.i I tasoma slightly wider apically than long with medioapical carinae enclosing cell Metasoma: Tl 0.9- (Fig. 133); T2 of metasoma finely granu- (Fig. 132). length X late, near apex of basal raised area to most 1.1 as long as apical width, lateral mar- of tergum disc in a semicircular pattern gins straight posterior to spiracle, spiracle lateral carinae (Fig. 135). slightly protuberant, pre- 2 Female. — Color: Body medium red- sent on basal V3 to A, poorly developed in brown except as follows: Scape yellow; some specimens; Tl weakly granulostriate 184 Journal of Hymenoptera Research

tarsus Head: on apical half and lateral to carinae, rarely tibia and hind light brown. raised of antennal 1.7X fla- nearly smooth (Figs. 133, 134); basal Length scape width; area of T2 0.2-0.4 of total T2 length, pos- gellum with 19 flagellomeres: L/W of first terior margin of basal raised area evenly three flagellomeres 3.3, 2.8, 2.8; L/W of as as curved, rarely obsolete; basal raised area apical flagellomere 2.5; MOD long as and 0.67 X as margin smooth to slightly crenulate, gran- POD; POD long LOL, ulate on most of tergum posterior to basal long as POL (Fig. 136); OOL 1.8X as long raised area in a semicircular pattern; T3 as POL (Fig. 136); head length 0.64 X 0.9-1.1 X as long as T2, smooth, with an- width in dorsal view; occiput evenly terior groove crenulate (Fig. 135). Legs: deeply indented, head somewhat 'c- Hind femur length 2.9-3.7X maximum shaped' in dorsal view (Fig. 136); head L/ width. H 0.7 in lateral view; eye L/H 0.58, eyeH/ Material examined.—HOLOTYPE 9 headH 0.58, eye width/gena width 1.1 (FSCA), labelled as follows: upper label (Fig. 137); gena wider ventrally than dor- "Florida: Alachua Co., Gainesville Doyle sally (Fig. 137); face granulate with medi- Connor Building", and lower label "H.V. an polished stripe, setae as long as clypeus Weems Jr. and C.R. Artaud 12-XI-1971 height, clypeus W/H 1.7, clypeus width Malaise trap". Also red label "HOLO- 0.8 X as long as malar space (Fig. 138); la- " TYPE" and bordered label Pseudognapto- brum as large as clypeus, with truncate don hemicolor Williams Holotype det. D. apex (Fig. 138). Wings: Forewing with RS Williams 2003". PARATYPES: U.S.A.: vein straighter apically than basally; Florida: Alachua Co.: Gainesville, Doyle RS+Ma pigmented and tubular; 2M spur Connor Building, Malaise trap, 12-XI-1971, 6.4 X as long as RS+Mb and markedly lon- H Weems C. Artaud, 1 9 (FSCA). S9-T10S- ger than RS+Ma; 2-A1 present as a taper- R18E, Pierce's Homestead, Malaise trap, 1- ing tubular vein over most of length and XI-1973 (19), 19-XI-1973 (19), 10-V-1974 pigmented crease over remainder; spur of {IS), W. Pierce (FSCA). Wakulla Co.: F.W. m-cu present, subdiscal cell closed (Fig. Mead Sta. 20-IV-1955, C. Muesebeck 19 140). Hind wing, r-m as long as R; Rs 3.6X (FSCA). North Carolina: Wake Co.: Ra- as long as R; angle of M and r-m with pos- leigh, 9-VIII-1983, J. Whitfield 1 9 (JWCI). teriorly directed faintly pigmented spur; Etymology. —The name hemicolor refers apex of R with moderately developed to coloration of the metasoma. knob (Fig. 140). Mesosoma: Mesoscutum with median groove absent, notauli deep- Pseudognaptodon labrns Williams, new ly impressed, joining in a U posteriorly, species posteromedial depression small (Fig. 139); (Figs. 136-145) propodeum with basal carinae short, par- —This is Diagnosis. species separated alell (Fig. 141). Metasoma: Tl length 1.4X from other curticauda-group species by the as long as apical width, lateral margins following combination of characters: Body concave posterior to spiracle, spiracle dark brown; labrum apically truncate, as slightly protuberant, lateral carinae pre- as large clypeus, dark brown (Fig. 138); sent on basal half, convergent, merged Forewing vein 2M markedly longer than with striae on apical half; Tl finely striate 2 RS+Ma (Fig. 140); Forewing vein 2-A1 on apical A (Figs. 142, 143); basal raised pigmented for most of length, first subdis- area of T2 0.2 of total T2 length, posterior cal cell closed (Fig. 140). margin of basal raised area evenly curved; —Color: Holotype female. Body dark T2 finely granulostriate on most of tergum brown to black except for the following: posterior to basal raised area, striae longer with legs orange-brown, apical tarsomer- medially than laterally; T3 0.7X as long as base of es, hind co a, apical half of hind T2, smooth, with anterior groove slightly Volume Number 2004 13, 1, 185

0.5 mm

Figs. 126-130. Pseudognaptodon hemicolor. 126, ocelli in dorsal view. 127, Head in dorsal view. 128, Head in lateral view. 129, Head in anterior view. 130, Wings. crenulate, deeply impressed, slightly pro- Material examined.— HOLOTYPE 9 curved medially (Fig. 144). Legs: Hind fe- (CNCI), labelled as follows: "COLOMBIA, mur length 4.8 X maximum width (Fig. 2900 m. Putumayo 2. XII. 72 1°10'N, 145). 77°15'W. J. Helava". Also red label "HO- 186 Journal of Hymenoptera Research

V.. -avVCA

Figs. 131-135. Pseudognaptodon hemicolor. 131, Mesosoma in dorsal view. 132, Propodeum in dorsal view. 133, Tl of metasoma in dorsal view. 134, Tl of metasoma in lateral view. 135, Anterior end of metasoma in dorsal view.

LOTYPE" and bordered label "Pseudog- Pseudognaptodon minimus Williams, naptodon labrus Williams Holotype det. D. new species Williams 2003". Colombia: PARATYPES: (Figs. 146-155) Quindio, 11 km E Calarca, 7000', 5-III- —This is 1974, S&J Peck Id (CNCI). Peru: Amazon- Diagnosis. species separated as: from other the 6°50'S, 77°38'W, 3200', 13-11-1973, J. curticauda-group species by combination of characters: Helava, 26 (CNCI). Peru: Amazonas: following Body dark brown with brown metasomal 6°48'S, 77°38'W, 2800', 13-11-1973, J. Hela- light va, IS (CNCI). Tl and basal raised area and lateral mar- Etymology.—The name labrus refers to gins of T2; metasomal Tl distinctly wider the labrum, whose size shape, and color is at apex than long, with basal carinae and unique within the genus. striae obsolete (Fig. 152), somewhat flat in Volume Number 2004 13, 1, 187

Figs. 141-145. Pseudognaptodon labrus. 141, Propo- 0.5 mm deum in dorsal view. 142, Tl of metasoma in dorsal view. 143, Tl of metasoma in lateral view. 144, An- 136-140. labrus. Head in terior end of metasoma in dorsal Figs. Pseudognaptodon 136, view. 145, Hind leg. dorsal view. 137, Head in lateral view. 138, Head in anterior view. 139, Mesosoma in dorsal view. 140, Wings. smooth medially, most setae shorter than clypeus height, clypeus W/H 2.0, clypeus 1.2X width as long as malar space (Fig. lateral view (Fig. 153); very minute, total 148). Wings: Forewing with RS vein sinu- less than 1.5 mm. ate on RS+Ma body length— apical half; unpigmented, Female. Color: Body dark brown except tubular but slender, obsolete at base; 2M for the following: Scape, metasomal Tl, spur 2.0 X as long as RS + Mb; 2-A1 present and T2 basal raised area and lateral mar- as a faint crease on basal Vr, spur of m-cu to first gins light brown; legs yellow light absent, subdiscal cell open (Fig. 150). brown, apex of hind tibia and hind tarsus Hind wing, r-m 0.9X as long as R; Rs 3.6X light brown. Head: Length of antenna 1 as long as R; apex oi R with poorly de- scape 1.6X width; flagellum with 13 fla- veloped knob (Fig. 150). Mesosoma: Some- gellomeres: L/W of first three flagello- what dorsoventrally compressed; mesos- meres 3.6, 2.8, 2.6-2.7; L/W of apical fla- cutum with median groove absent, notauli 0.9 X as as gellomere 2.5; MOD long POD; obsolete on posterior half (Fig. L49); pro- POD 0.9 X as long as LOL, and 0.8 X as podeum with basal carinae curved, con- long as POL (Fig. 146); OOL 1.5 X as long vergent, basal cell narrowly open at apex as POL (Fig. 146); head length 0.7X width (Fig. 151). Metasoma: Tl length 0.8X as in dorsal view; occiput evenly shallowly long as apical width, nearly flat dorsoven- indented (Fig. 146); head L/H 0.8 in lat- trally, lateral margins straight posterior to eral view; eye L/H 0.6, eyeH/headH 0.7, spiracle, spiracle not protuberant, lateral eye width/gena width 1.2 (Fig. 147); gena carinae present onlv on base; Tl with a wider ventrally than dorsally (Fig. 147); few poorly developed striae on apical and l face granulate on lateral margins and lateral h (Figs. 152, 153); basal raised area 188 Journal of Hymenoptera Research

co 0J2 mm .---< 151 I

0.2 mm

0.- mm

0.25 mm ~)\ 154

Figs. 146-150. Pseudognaptodon minimus. 146, Head Pro- Figs. 151-155. Pseudognaptodon minimus. 151, in dorsal view. 147, Head in lateral view. 148, Head podeum in dorsal view. 152, Tl of metasoma in dor- in anterior view. 149, Mesosoma in dorsal view. 150, sal view. 153, Tl of metasoma in lateral view. 154, Wings. Anterior end of metasoma in dorsal view. 155, Hind

leg.

of T2 0.4 of total T2 length, posterior margin of basal raised area evenly curved, the small size of poorly impressed; T2 granulate on basal very (<1.5 mm) specimens of this It is the smallest half of tergum posterior to basal raised species. species examined in this area; T3 0.9 X as long as T2, smooth, with study. anterior groove crenulate, poorly devel- Pseudognaptodon minutus (Ashmead) oped grooves defining anterolateral cor- 1894 ners present near lateral margins (Fig. (Figs. 156-166) 154). Legs: Hind femur length 3.8X maxi- — Indies mum width (Fig. 155). Holoytpie male. St. Vincent: W. Material examined.—HOLOTYPE 9 93-331, Leiophron minutus Type Ash., B.M. (CNCI), labelled as follows: "USA Texas TYPE 3?659, Type H.T., H.H. Smith Brewster Co. Big Bend National Park Oak (BMNH). Examined.— Cyn.-Window Trail 5400' 24-27.VI.82 Diagnosis. This species is separated G.A.P. Gibson". Also red label "HOLO- from other curticauda-group species by the TYPE" and bordered label "Pseudognapto- following combination of characters: Body don minimus Williams Holotype det. D. uniformly light to dark brown in color; Tl Williams 2003". PARATYPES: Pima Co.: of metasoma length markedly greater than Brawley Wash, Mile Wide Rd., 1.5 mi E. apical width (Fig. 163); T2 of metasoma Sandaro Blvd., 2500', sweeping Bacharis finely striate or granulostriate near basal glutinosa, 3-VIII-1982, G. Gibson, 2$ raised area margin, rarely with striae ex- (CNCI). tending to posterior margin of tergum Etymology. —The name minimus refers to (Fig. 165); T3 of metasoma smooth, with Volume 13, Number 1, 2004 189

crenulate anterior slightly groove (Fig. to strongly protuberant, lateral carinae 165). present on basal half and merged in some Female.—Color: Body light honey-brown specimens with striae or ridges originat- to dark red-brown as follows: except ing at spiracles, to complete to tergum Scape yellow to light brown; legs yellow apex, Tl raised between carinae and with darker hind with tarsus, rarely light somewhat distinct from flatter posterolat- brown area on dorsal surface of hind fe- eral corners, striate laterally and apically, mur. Head: of Length antennal scape 1.3- (Figs. 163, 164); basal raised area of T2 0.2- 1.7X with 14-16 0.4 width; flagellum flagel- of total T2 length, posterior margin of lomeres: of first three basal L/W flagellomeres raised area irregular, rarely slightly 2.7-3.3, 2.3-3.0, 2.1-2.8; L/W of apical fla- medially produced; T2 with coarse striae gellomere 2.3-3.6; MOD 0.8-1. Ox as long at basal raised area margin to striate or as .2 X POD; POD 0.9-1 as long as LOL, granulostriate on middle half of tergum and 0.7-1. 2 X as as long POL (Fig. 156); about 0.8 of distance to posterior margin, 1. 7-2.5 X as as OOL long POL (Fig. 157); rarely smooth with weakly crenulate basal vertex with obsolete granular sculpture raised area margin, striae longer medially anterior to head ocelli; length 0.6-0.7X than laterally; T3 0.9-1.1 X as long as T2, width in dorsal view; occiput evenly mod- smooth, with anterior groove partially or indented erately (Fig. 157); head L/H 0.8- entirely crenulate (Fig. 165). Legs: Hind fe- 0.9 in lateral view; eye L/H 0.7-0.8, eyeH/ mur length 3.1-3.5 x maximum width headH 0.6-0.7, eye width/gena width 1.8- (Fig. 166). 2.6 — (Fig. 158); gena as wide ventrally as Material examined. Bolivia: Yungas, 50 face setae dorsally (Fig. 158); granulate, as km N. La Paz, 27-1-1973, 2200m, J. Helava, long as clypeus height, clypeus W/H 1.8- 16 (CNCI). Brazil: Caruaru: Pernambuco, 2.3, clypeus width 0.8-1. Ox as long as ma- VII-1972, M Alvarenga, 19 (CNCI). Per- lar space (Fig. 159). Wings: Forewing with nambuco, IV-1972, M Alvarenga, 19, 1 6 middle of RS vein slightly decurved be- (CNCI). Bahia: Encruzilhada, XI-1974, M. yond midlength, vein slightly sinuate, Alvarenga, 29, 16* (CNCI). Represa Rio rarely nearly straight; RS + Ma unpigment- Grande: Guanabara, VII-1972, F.H. Olivi- ed on basal half and tubular throughout era, 19 (CNCI). Vila Vera: M. Grosso, 12° length to unpigmented and thinned on en- 46' S, 55° 30'W, X-1973, 500M, M. Alvar- tire length with basal section obsolete; 2M enga, 19 (CNCI). Para Jacareacanga, XII- spur 1.8-3.2X as long as RS+Mb; 2-A1 ab- 1968, M. Alvarenga, 19 (CNCI). Costa sent, to present as an unpigmented crease Rica: Guanacaste: Santa Rosa Natl. Park, on basal half; spur of m-cu absent, first regenerating woodland <10 years old, di- 6- subdiscal cell open (Fig. 160). Hind wing rect sun daily, wet, 300m, Malaise trap, with r-m 0.8-1.1 X as long as R; Rs 3.1- 27-IX-1986, I. Gauld, 79, 10(5 (UWYO). 5.0 X as long as R; apex of R with moder- Santa Rosa Natl. Park, (H) open regener- in ately to slightly developed knob (Fig. 160). ating woodland <10 years old, (O) Mesosoma: Mesoscutum with median clearing, fully isolated part of day, 300m, groove faint, notauli merged with postero- Malaise trap/27-IX-18-X-1986 (39,^66"), 26- x 14-VIII-6-IX-1986 medial depression on posterior h (Fig. X-16-XI-1986 {26), (26), 161); propodeum with medioapical cari- 8-29-XI-1986 (16), 29-XI-20-IX-1986 (16), nae enclosing a cell, or carinae curved to- 26-X-16-XI-1986 (19), I. Gauld D. Janzen, ward each other but with small gap at the (UWYO). Santa Rosa National Park, re- woodland clear- apex of the cell (Fig. 162). Metasoma: Tl generating <10 years old, length 1.1-1.6X as long as apical width, ing, 300m, Malaise trap, 5-26-VII-1986, I. Santa Rosa Natl. lateral margins straight or slightly concave Gauld, 19, 16 (UWYO). woodland posterior to spiracle, spiracle moderately Park, (H) open regenerating 190 Journal of Hymenoptera Research

0.5 mm

Figs. 156-160. Pseudognaptodon minutus (Ashmead). 156, Ocelli in dorsal view. 157, Head in dorsal view. 158, Head in lateral view. 159, Head in anterior view. 160, Wings.

<10 years old, (C) more or less fully shad- D. Janzen, (UWYO). Santa Rosa Natl, ed as possible, 300m, Malaise trap, 14- Park, (SE) Bosque San Emilio 30yr old de- VIII-6-IX-1986 (3$, 3 6), 8-II-2-III-1986 ciduous forest, (O) in clearing, fully iso- (1$, 26), 2-23-III-1986 (19, 16), 8-18-III- lated part of day, 300m, Malaise trap, 26- 1986 (26), 21-II-14-III-1986 (16), I. Gauld VII-14-VIII-1986, I. Gauld D. Janzen, 16* Volume 13, Number 2004 1, 191

Figs. 161-166. Pseudognaptodon minutus (Ashmead). 161, Mesosoma in dorsal view. 162, Propodeuin in dorsal view. 163, Tl of metasoma in dorsal view. 164, Tl of metasoma in lateral view. 165, interior end of metasoma in dorsal view. 166, Hind leg.

(UWYO). Santa Rosa Natl. Park, (SE) son, 29 (UWYO). 7 km S.W. Bribri, 50m, Bosque San Emilio 30yr old deciduous for- 4-VI-1990, P. Hanson, 1 9 (UWYO). 7 km est, (C) more or less fully shaded as pos- S.W. Bribri, 50m, XI-1989, P. Hanson, 19 sible, 300m, Malaise trap, 13-IV-4VI-1986, (UWYO). 16 km W. Guapiles, 400m, 1-IV- I. Gauld D. Janzen, 16 (UWYO). P.N. San- 1991, P. Hanson, 16 (UWYO). Puntaren- ta Rosa, 200m, 1-1991, P. Hanson, 16 as: Peninsula Osa, Puerto Jimenez, grass) (UWYO). P.N. Guanacaste, below Pitilla, weedy site, 10m, 1 -1 1-1992, P. Hanson, 2 9 500m, 7-8-III-1990, J. Noyes, 29 (UWYO). (UWYO). Peninsula Osa, Puerto Jimemv, Estac. Pitilla, 9 km S. Santa Cecillia, 700m, grassy disturbed site, 10m, 10-XI-1991, P. V-1989, I Gauld, 69,1c? (UWYO). Limon: Hanson, 19, 16 (UWYO). Golfo Dulce, 3 7 km S.W. Bribri, 50m, 1-II-1990, P. Han- km S.W. Rincon, 10m, XII-1989-I-1990, P. 192 Journal of Hymenoptera Research

Hanson 16 (UWYO). R.F. Golfo Duke, 24 E. Concordia, 12-VIII-1964, 3000m, W. Ma- km W. Piedras Blancas, 200m, III.1993, P. son, 16 (CNCI). Peru: Cuzco, 13° 40'S, 70° Hanson 2 9 (UWYO). Monteverde, season- 40' W, 18-1-1973, J.Helava, 1 9 (CNCI). Su- al forest, 11-1980, 1400m, W. Mason, 1 9 riname: Kobo forest Reserve, S.W. of line (CNCI). Alajuela: 5 km W. San Ramon, 391, Malaise trap, 1-5-IX-1978, E. Nee. . . 1200m, X-1997, O. Castro & P. Hanson, (hand written label, collector name illegi- 19, Id (UWYO). 5 km W. San Ramon, ble), 19 (RNHL). 1200m, 11-1997, O. Castro & P. Hanson, 16 Pseudognaptodon mttdus Williams, new (UWYO). San Pedro de la Tigra Cacao, R. 19 species 200m, 1-II-1990, Cespedes, (UWYO). . - Fl s - 167 176 ( § ) San Jose: San Antonio de Escazu, 1300m, IV-1999, W. Eberhard, 19, 3cT (UWYO). Diagnosis.—This species is separated Cuidad Colon, 800m, III-IV-1990, L. Four- from other curticauda-group species by the nier, 16 (UWYO). Cuidad Colon, 800m, following combination of characters: Ver- XII-1989-I-1990, L. Fournier, 1 9 (UWYO). tex of head smooth, or with obsolete gran- Zurqui de Moravia, 1600m, X-1995, P. ulate microsculpture (Figs. 167, 188); ocel- Hanson, 16 (UWYO). Zurqui de Moravia, lar triangle large, interocellar distances 1600m, 24-XII-1988, P. Hanson, 16* longer than POD (Fig. 167); base of Tl of (UWYO). Zurqui de Moravia, 1600m, Mai- metasoma separated from remainder by a aise, 11-1996, P. Hanson, 1 9 (UWYO). He- transverse ridge or sharp convexity, apex redia: 3 km S. Puerto Viejo, OTS-La Selva, irregularly rugose (Fig. 173); T2 of meta- 100m, V-VI-1993, P. Hanson, 19, 36 soma with coarse striae on basal half or (UWYO). 3 km S. Puerto Viejo, OTS-La more posterior to basal raised area; T3 of Selva, 100m, IV-1991, P. Hanson, 18 metasoma medially striate on base, with (UWYO). Carthago: Dulce Nombre, Viv- anterolateral corners defined by bands of ero Linda Vista, 1300m, VII-IX-1994, P. sculpture (Fig. 175). Hanson, 1 9 (UWYO). La Cangreja, 1950m, Female. —Color: Body medium to dark VII-1991, P. Hanson, 1 9 (UWYO). Ecua- brown except for the following: Scape and dor: Napo: Limoncocha, 15-28-VI-1976, pedicel yellow; mesosoma with lighter S&J Peck, 16 (CNCI). 5 km S. Baeza, 13- brown area on pronotal collar and ventral 11-1983, 1700m, Masner & Sharkey, 19 portion of mesopleuron in some speci- (CNCI). Pich.: S. Domingo, 16 km S. Tin- mens; legs yellow, except hind tarsus light alandia, 15-30-VI-1975, 680m, S. Peck, 1 9 brown. Head: Length of antennal scape (CNCI). Rio Palenque R.S., 4-II-1983, 1.6X width; flagellum with 14-16 flagel- 200m, Masner & Sharkey 1 9 (CNCI). lomeres: L/W of first three flagellomeres Mexico: Guerrero: 32 mi S.E. Petalan, 10- 2.9-3.0, 2.8-2.9, 2.7; L/W of apical flagel- VII-1985, Woolley & Zolnerowich, 29,16 lomere 3.3; MOD 0.8X as long as POD; (TAMU). 2 mi N. Cacahuamilpa, 19-VII- POD 0.8-0.9 X as long as LOL, and 0.7X 1984, J. Woolley, 16 (TAMU). 2 mi N. Ca- as long as POL (Fig. 137); OOL 1.6-1 .8 X cahuamilpa, 5300', 4-VII-1987, R. Whar- as long as POL (Fig. 168); vertex smooth ton, 16 (TAMU). Jalisco: 16 mi S. Autlan or with very faint granular microsculp- on in Hwy. 80, 8-VII-1984, J. Woolley, 19 ture; head length 0.7X width dorsal (TAMU). 5.4 mi N. Autlan 7-VII-1984, view; occiput broadly but unevenly in- Schaffner Woolley Carroll Friedlander, 16 dented (Fig. 168); head L/H 0.9 in lateral (TAMU). Oaxaca: 4.4 mi N.E. San Pedro view; eye L/H 0.6-0.7, eyeH/headH 0.7, Mixtepec, 16-VII-1985, Woolley & Zolne- eye width/gena width 1.9 (Fig. 169); gena rowich, 39, 36 (TAMU). 2 mi N. Cande- wider ventrally than dorsally (Fig. 169); laria Loxicha, 17-VII-1985, J. Woolley G. face granulate with a narrow median pol- 'olnerowich, 46 (TAMU). Sinaloa: 20 mi ished ridge on dorsal half, setae shorter Volume 13, Number 1, 2004 193

1 , 5 miiT'\fi* %

0.5 mm

Ocelli in dorsal view. 168, Head in dorsal view. 169, Head m Figs. 167-170. Pseudognnptodon nitidus. 167, lateral view. 170, Wings. 194 Journal of Hymenoptera Research

Figs. 171-176. Pseudognaptodon nitidus. 171, Mesosoma in dorsal view. 172, Propodeum in dorsal view. 173, Tl of metasoma in dorsal view. 174, Tl of metasoma in lateral view. 175, Anterior end of metasoma in dorsal

view. 176, Hind leg.

than clypeus height, clypeus W/H 2.3, (Fig. 170). Mesosoma: Mesoscutum with clypeus width 1.1 X as long as malar median groove absent, notauli merged space. Wings: Forewing with RS vein with one another on anterior margin of slightly sinuate on apical half; RS+Ma un- posteromedial depression (Fig. 171); pro- pigmented, tubular except extreme base, podeum with basal cell complete, some- obsolete; 2M spur 2.0-2.5 X as long as what transverse and slightly raised, cari- RS + Mb; 2-A1 present as a crease on basal nae more developed laterally than at cell 1.1 X Vr, spur of m-cu absent, first subdiscal cell apex (Fig. 172). Metasoma: Tl length open (Fig. 170). Hind wing with r-m 0.9- as long as apical width, lateral margins 1.0X as long as R; Rs 3.5-4.5 X as long as slightly concave posterior to spiracle, spi- R; apex of R with well developed knob racle slightlv protuberant, lateral carinae Volume 13, Number 1, 2004 195

on basal a present Vz, joined medially by deeply impressed, crenulate, laterally de- transverse ridge that defines a semicircu- curved anterior groove, widened mediallv lar basal smooth area; Tl coarsely irregu- with a narrow band of striae extending striate larly throughout and very convex, posteriorly (Fig. 186); forewing 2-A1 vein except for basal area, (Figs. 173, 174); an- pigmented for most of length, first subdi- terior basal raised area of T2 0.3 of length, cal cell closed by veins and pigmented of basal raised area even- creases posterior margin (Fig.— 181). ly curved to slightly irregular, crenulate, Female. Color: Body black except for and medially produced, T2 striate on basal the following: Scape yellow, pedicel light half most of tergum posterior to basal brown; pronotal collar light orange- raised area except for a narrow apical brown; legs yellow, apical tarsomeres, band; T3 0.9 X as long as T2, striate me- base of hind coxa, and entire hind tarsus dially on basal ¥s, with anterior groove light brown, rarely with dorsal surface of crenulate, and anterolateral areas delimit- femur and apex of tibia light brown. Head: ed by narrow bands of sculpture (Fig. Length of antennal scape 1.7-1.9X width; 175). Legs: Hind femur length 2.9 X maxi- flagellum with 17-19 flagellomeres: L/W mum width (Fig. 176). of first three flagellomeres 2.7-2.8, 2.4-2.5, Material examined.—HOLOTYPE 9 2.4-2.5; L/W of apical flagellomere 2.8- (UWYO), labelled as follows: upper label 3.3; MOD 0.8-0.9 X as long as POD and "Costa Rica: Guanacaste Santa Rosa Natl. longer than LOL and POL; POD 1.3-1.7X Park 300m ex. Malaise trap site#: 1 Dates: as long as LOL, and 1.0-1.3X as long as X 18-i.8-ii 1986 I.D. Gauld & D. lanzen" low- POL (Fig. 177); OOL 2.0-2.5 as long as obsolete er label "(H) open regenerating woodland POL (Fig. 178); vertex with gran- <10 years old, (O) in clearing, fully isolat- ular microsculpture and well developed ed part of the day". Also red label "HO- grooves or wrinkles between lateral ocelli LOTYPE" and bordered label "Pseudog- and dorsal margin of eye (Fig 178); head naptodon nitidus Williams Holotype det. D. length 0.6-0. 7x width in dorsal view; oc- Williams 2003". PARATYPES: Costa Rica: ciput narrowly indented (Fig. 178); head Guanacaste: Santa Rosa Natl. Park, 300m, L/H 0.6-0.8 in lateral view; eye L/H 0.6- (H) open regenerating woodland <10 0.7, eyeH/headH 0.6-0.7, eye width/gena wider ven- years old, (C) more or less fully shaded as width 1.5-1.8 (Fig. 179); gena face possible, Malaise trap, 26-VII-14-VIII-1986, trally than dorsally (Fig. 179); granu- setae I. Gauld D. Janzen, 1 9 (UWYO). San Jose: late with median polished stripe, San Antonio de Escazu, 1300m, IV-1999, shorter than clypeus height; clypeus W/H W. Eberhard, 19, IS (UWYO). 2.6, clypeus width 1.2X as long as malar with RS Etymology. —The name nitidus refers to space (Fig 180). Wings: Forewing the smooth appearance of the vertex in vein decurved beyond midlength, slightly and this species, which resembles the state in sinuate; RS+Ma pigmented tubular, thinner at 2M 2. 5-3. Ox omzssws-group species but is rare in the slightly base; spur as 2-A1 as a tu- curt icauda-group. as long RS+ Mb; present bular spur at base and faintly pigmented shawi Williams, new Pseudognaptodon crease over most of length; spur of m-cu species present as a faintly pigmented line, first (Figs. 177-187) subdiscal cell closed (Fig. 181). Hind wing with r-m 1.1-1.3X as as R; Rs 3.5 Diagnosis.—This species is separated long X as of R with well from other curticauda-group species by the 4.2 as long R; apex knob Mcoma: Me- following combination of characters: Ocel- developed (Fig. 181). soscutum with median absent, no- li as long as or longer than inter-ocellar groove with tauli in a U distances (Fig. 177); T3 of metasoma deeply impressed, joining pos- 196 Journal of Hymenoptera Research

Figs. 177-181. Pseudognaptodon shawi. 177, Ocelli in dorsal view. 178, Head in dorsal view. 179, Head in lateral view. 180, Head in anterior view. 181, Wings.

teriorly in some specimens, posteromedial long as apical width, lateral margins depression small (Fig. 852); propodeum slightly concave posterior to spiracle, spi- with basal carinae convergent, ending in racle slightly protuberant, lateral carinae several fine wrinkles, cell broadly open present on basal half, rarely merged with (Fig. 183). Metasoma: Tl length 1.1 X as coarse striae on aniral half: T1 striatp on Volume 13, Number 2004 1, 197 1 5T

Figs. 182-187. Pseudognaptodon shawi. 182, Mesosoma in dorsal view. 183, Propodeum in dorsal view. 184, Tl of metasoma in dorsal view. 185, Tl of metasoma in lateral view. 186, Anterior end of metasoma in dorsal view. 187, Hind leg.

half apical and lateral to carinae, (Figs. curved laterally, and merged with a nar- 184, 185); basal raised area of T2 0.2-0. X row band of coarse striae medially, striae of total T2 length, posterior margin of bas- extending onto disc of T3 (Fig. 186). Legs: al raised area irregular, slightly sinuate or Hind femur length 3.4-3.6 maximum evenly curved; basal raised area margin width (Fig. 187). crenulate, coarsely striae on basal half to Material examined.—HOLOTYPE 9 most of tergum posterior to basal raised (UWYO), labelled as follows: "Costa Rica: area, striae longer medially than laterally; San Jose:Zurqui de Moravia 1600m vii- T3 0.9-1.0 X as long as T2, with anterior 1992 Col. Paul Hanson". Also red label groove crenulate, deeply impressed, de- "HOLOTYPE" and bordered label "Pscu- 198 Journal of Hymenoptera Research

of dognaptodon shawi Williams Holotype det. meres 2.1-3.7, 2.1-2.8, 1.9-2.7; L/W api- D. Williams 2003". PARATYPES: Alajue- cal flagellomere 2.3-2.8; MOD 0.8-0.9 la: 5 km W. San Ramon, 1200m, X-1996 POD; POD 0.8-1 .OX as long as LOL, and X as as (1(5), XII-1996 (16), 11-1997 {16), IV-1997 0.7-0.9 long POL (Fig. 188); OOL (36), X-1997 (19, 36), VII-1997 (36), O. 1.3-2.0X as long as POL (Fig. 189); vertex Castro, P. Hanson, (UWYO). Guanacaste: with well developed granular sculpture su- ACT Bagaces, P.N. Palo Verde Sec. P anterior to ocelli, very fine, incomplete Verde, 200 N.E. Est., Extremo E. de Cameo ture or fine wrinkles present between lat- ocelli dorsal de Atterizaje, Malaise, 0-50m, 8-XI-9-XII- eral and eye margin (Fig 189); in 1999, I. Jiminez, 1 6 (UWYO). Guanacaste head length 0.6-0.7X width dorsal Conservation Area, Santa Rosa Hdq., Mai- view; occiput shallowly, evenly indented, aise, 200m, 3-7-VII-1997, L. van der Ent, head somewhat C-shaped in dorsal view in lateral 16 (UWYO). San Jose: P.N. Braulio Car- (Fig. 189); head L/H 0.6-0.8 0.6- illo, 9.5 km E. Tunel, 1000m, 1-III-1990, P. view; eye L/H 0.6-0.7, eyeH/headH Hanson, 1$ (UWYO). Ecuador: Napo: 0.7, eye width/gena width 1.5-1.9 (Fig. Baeza, 9-12-II-1983, L. Huggert, 19 190); gena parallel sided (Fig. 190); face (RNHL). completely granulate to a small polished Etymology. —This species is named for area dorsal of clypeus and a narrow me- or Scott Shaw, who has been one of the major dian polished strip, setae shorter than contributors of specimens to this study. as long as clypeus height, clypeus W/H 1.8-2.2, clypeus width 0.9-1. 2 X as long as whartoni Williams, Pseudognaptodon malar space (Fig 191) Wings: Forewing new species with RS vein decurved apically; RS+Ma 188-198) —(Figs. unpigmented on basal V3 to entire length Diagnosis. This species is separated and tubular throughout length, rarely from other cnrticaudfl-group species by the thinned on basal V2, longer than 1M; 2M following combination of characters: Me- spur 2.5-5.3X as long as RS+Mb; 2-A1 ab- tasoma medium to dark brown apically, sent to present as an unpigmented crease yellow to light brown on lateral margins on basal half; spur of m-cu absent, first or most of first two or three terga; Tl of subdiscal cell open (Fig. 192). Hind wing metasoma slightly wider apically than with r-m 0.9-1. 3 X as long as R; Rs 3.4- long, granulostriate (Fig. 195); T2 of me- 4.6 X as long as R; apex of R moderately tasoma finely granulate, near apex of basal to well developed knob (Fig. 192). Meso- raised area to most of tergum disc in a soma: Mesoscutum with median groove semicircular pattern (Fig. 197). absent to faint, notauli merged with one Female.—Color: Body dark brown to another on anterior margin of posterome- black except as follows: Scape yellow; dial depression (Fig. 193); propodeum clypeus yellow to light orange-brown in with medioapical carinae straight, paral- some specimens; pronotal collar light or- lei, and widely separated, rarely with ir- ange-brown in some specimens; meso- regular very fine wrinkles or transverse pleuron with lighter bands in some spec- obsolete carina enclosing a compressed Tl imens; and T2 of metasoma light cell (Fig. 194). Metasoma: Tl length 0.9- brown laterally to entirely yellow on first 1.1 X as long as apical width, lateral mar- three metasomal terga; legs yellow with gins slightly concave to straight posterior light brown hind tarsus, apical tarsomeres to spiracle, spiracle not protuberant, lat- and apex of hind tibia darker in some eral carinae present on basal Vi to corn- specimens. Head: Length of antennal scape plete to apex or nearly so, in some speci- 1.5-1.9X width; flagellum with 15-19 fla- mens continuous with lateral margins of a

gellomere ; L/W of first three flaeello- weaklv raised medial area; Tl granulos- Volume Number 2004 13, 1, 199

0.5 mm

Figs. 188-192. Pseudognaptodon whartoni. 188, Ocelli in dorsal view, L89, Head in dorsal view. 190, Head in lateral view. 191, Head in anterior view. 192, Wings.

triate on apical half and lateral to carinae, T2 length, posterior margin of basal raised rarely somewhat more granulate between area evenly curved, slightly medially pro carinae and striate laterally (Figs. 195, duced or toothed in some specimens; bas- 196); basal raised area of T2 0.3-0.4 of total al raised area margin smooth to slightly 200 Journal of Hymenoptera Research

Figs. 193-198. Pseudognaptodon whartoni. 193, Mesosoma in dorsal view. 194, Propodeum in dorsal view. 195, Tl of metasoma in dorsal view. 196, Tl of metasoma in lateral view. 197, Anterior end of metasoma in dorsal view. 198, Hind leg.

crenulate, granulate on most of tergum Material examined.— HOLOTYPE 9 posterior to basal raised area in a semicir- (TAMU), labelled as follows: "Texas: Braz- cular pattern; T3 0.8-1. Ox as long as T2, os Co. College Station Lick Creek Park smooth, with anterior groove smooth to November 1-14, 1987 R. Wharton Mal- weakly crenulate, rarely with groove wid- aise". Also red label "HOLOTYPE" and er medially (Fig. 197). Legs: Hind femur bordered label "Pseudognaptodon whartoni length 3.0-3.8X maximum width (198). Williams Holotype det. D. Williams 2003". Hosts: Reared from Nepticula sp. and PARATYPES: USA: Texas: Brazos Co.: Coptodisca sp. (Lepidoptera: Nepticulidae) College Station, Lick Creek Park, Malaise, on Myrica certifera (Fagales: Mvricaceae). 1-14-XI-1987 (49, 4 6), 20-IX-4-X-1987 (39, Volume 13, Number 1, 2004 201

76), 4-18-X-1987 (39, Id), R. Wharton, Alvarenga, 2 9 (CNCI). Jatai: Goias, XI- (TAMU). College Station, Lick Creek Park, 1972, F.M. Oliviera, 19, Id (CNCI). Can- 16-III-9-IV-1988 (IS), 16-31-V-1988 (29), ada: Ontario: Aylmer West, 23-27-VII- 22-XI-6-XII-1987 (2 9), 2-16-V-1988 (19), 1972, Malaise Trap, Id (CNCI). Domini- 13-18-V-1988 (16*), Wharton, Praetorius, can Republic: El Rio (La Vega), 17-111- (TAMU). College Station, Lick Creek Park, 1978, 1000m, L. Masner, 1 9 (CNCI). USA: 3-9-X-1988 (26), 11-18-IX-1988 (2 9, 16), Arkansas: Mountain Pine: L. Ouachita 11-18-X-1988 (29, 2 6), 18-30-X-1987 (2 9, State Park, V-1972, G. Heinrich, Malaise 3 c?), 23-X-6-XI-1988 (3 9, 1 6), 7-20-XI-1988 Trap, 19 (CNCI). Florida: Alachua: R. Sta- (29), Wharton, (TAMU). College Gainesville, DPI, 1-23-VI-1987, J. Wiley, tion, Lick Creek Park, dissected, 14-IX- Malaise Trap, 29 (CNCI). Georgia: For- 1990, R. Wharton, 19, 16 (TAMU). Lick syth, 1-8-X-1970 (39), 25-VII-1970 (19),

Creek Park, 22-30-VI-1987, J Heraty J. 25-X-1970 (19), 20-XI-1970 (19, Id), VII- Woolley, 16* (TAMU). Harris Co: Bear Cr. 1970 (19), 7-IX-1970 (19, Id), 23-30-VIII- Pk., ex. Nepticula on Myrica certifera, JAP 1970 (39), F.T. Naumann, Malaise trap, No. 82M13, 28-XII-1982, D. Wagner, 19, (CNCI). Louisiana: Evangeline Co.: Bay- 1 6 (JWCI). Bear Cr. Pk., ex. Coptodisca on ou Chicot, 15-V-4-VI-1971, D. Shanek, 2 9 Myrica certifera, JAP No. 82M14, 28-XII- (CNCI). Nachitoches: 20 mi W. Gorum, 3- 1982, D. Wagner, 1 9 (JWCI). Montgomery 18-V-1989, R. Wharton, 19 (TAMU). Co.: Jones State Forest, 8 mi S. Conroe, 8- County?: Lake Bistineau State Park, 15-18- 19-IX-1987, Wharton, Carroll, Praetorius, IV-1972, G. Heinrich, Malaise Trap, 19 19, 26 (TAMU); 18-31-X-1987, Wharton, (CNCI). Lake Bistineau State Park, 22-27- Praetorius, 29, 1 6 (TAMU); 1-6-IX-1987, IV-1972, G. Heinrich, Malaise Trap, Id Wharton, Steck Carroll, 2 9 (TAMU); 20- (CNCI). Maryland: Laurel, ll-V-1965, 26-IV-1987, Wharton, Praetorius, Wang, Malaise Trap, Id (CNCI). Missouri: Wil- 19 (TAMU); 13-19-IV-1987, Wharton, liamsville, 10-IX-5-X-1969, J.T. Becker, Id Praetorius, Wang, 26 (TAMU); 1-7- VI- (CNCI). Williamsville, 15-VII-10IX-1969, 1987, Wharton, Steck, Carroll Wang, 16 J.T. Becker, 19 (CNCI). Williamsville, X- (TAMU); 9-16-VIII-1987, Wharton, Steck, VII7-1969, E.C. Becker, Id (CNCI). North Carroll, 19 (TAMU). Bandera Co.: Lost Carolina: Highlands, 3-VI-1957, 3800', J.R. Maples State Park, 18-VIII-1988, G. Zol- Vockeroth, Id (CNCI). Oklahoma: Lati. nerowich, 29, 16* (TAMU). Lost Maples Co.: 4 mi W. Red Oak, sweep, 1-4- VII- State Park, 22-VIII-1987, R. Wharton, Id 1987, D. Chandler, 1 9 (UWYO). Tennes- (TAMU). Travis Co.: Austin, 8-X-1983, R. see: Lexington: Natchez Trace State Park, Wharton, 16 (TAMU). Bosque Co.: 3 mi 11-15-VI-1972 (19, Id), 15-19-VI-1972 W. Laguna Park, 13-IV-1984, R. Wharton (19), VI-1972 (Id), G. Heinrich, Malaise Id Co.: Mexico: Michoacan: 10 mi J. Woolley, 19, (TAMU). Tyler Trap, (CNCI). 3 mi S. 22-V- S. G. Zol- Kirby State Forest, Warren, Urupan, 7-VII-1985, J. Woolley, 1 9 Walker Co.: Leon: 29 1984, J. Woolley, (TAMU). nerowich, 39 (TAMU). Nuevo Stubblefield Lake, 7-VI-1985, R. Wharton mi W. Linares, S. Rosa Cyn, 730m, 3- VI- 19 (TAMU). Kerr Co.: Center Point, Mal- 1983, M. Kaulbars, 19 (CNCI). Sinaloa: 20 aise trap, 26-VII-2-VIII-1987, Wharton, mi E. Concordia, 3000', 8-VIII-1964, Praetorius, 29 (TAMU). Gonzales Co.: W.R.M. Mason, Id (CNCI). Costa Rica:

'almetto State Pk., l-IV-1984, J. Woolley, San Jose: Cuidad Colon, 800m, VI-VII- 16 (TAMU). Harrison Co.: 9 mi S. 1990, Luis Fournier, 1 9 (UWYO). Guana- rshall, 27-VII-1982, R. Wharton, 19 caste: Guanacaste Conservation Area, San- VMU). OTHER SPECIMENS EXAM- ta Rosa Hdq., Malaise, 200m, 373-VII-1997, 1 ED: Braz L: Ceara: Barbalha, V-1969, 9 L. van der Ent, 1 9 (UWYO). Santa Rosa 4CI). B ia: Encruzilhada, IX-1974, M. Natl. Park, Malaise trap, (H) open regen- 202 Journal of Hymenoptera Research

0.8 X as as erating woodland <10 years old, (O) in flagellomere 3.3; MOD long 14-VI- 1.0-1.3 X as as and clearing, fully isolated part of day, POD; POD long LOL, 5-VII-1986, I. Gauld, D. Janzen, 19 1.0-1.1 X as long as POL (Fig. 199); OOL (UWYO). Santa Rosa Natl. Park, Malaise 2.0-2.3 X as long as POL; vertex with well trap, (H) open regenerating woodland developed granular microsculpture (Fig. 0.7 X width in dorsal <10 years old, (C) more or less fully shad- 200); head length indented ed as possible, 10-31-1-1987, I. Gauld, D. view; occiput broadly evenly (Fig. Janzen, IS (UWYO). P.N. Santa Rosa, 200); head L/H 0.8-0.9 in lateral view; eye 200m, 1-1991, P. Hanson, 1 9 (UWYO). L/H 0.6-0.7, eyeH/headH 0.7, eye width/ wider Remarks.—This species is distributed gena width 1.7-1.8 (Fig. 201); gena from Canada to Brazil. Specimens are re- ventrally than dorsally (Fig. 201); face gran- alativel uniform in appearance, although ulate with a narrow median polished ridge there is variation in color, the size of ocelli, on dorsal half, setae shorter than clypeus and the shape and sculpture of the first height, clypeus W/H 2.3, clypeus width metasomal tergum in specimens from the 1.1 X as long as malar space. Wings: Fore- northern and southern extremes. Because wing with RS vein straight or slightly sin- if this I have selected the type series only uate on apical half; RS+Ma pigmented and from the central part of the range in Texas tubular except extreme base obsolete; 2M and Mexico. — spur 3.0-3.3 X as long as RS+Mb; 2-A1 pre- Etymology. This species is named for sent as a crease on basal Vy, spur of m-cu Robert Wharton, who collected the holo- absent, first subdiscal cell open (Fig. 202). type and most of the specimens of this Hind wing with r-m 1.3X as long as R; Rs species, who has been one of the major 4.5-5.0 X as long as R; apex of R with poor- contributors of specimens to this study, ly developed knob (Fig. 202). Mesosoma: and who has been patient while I retained Mesoscutum with median groove present specimens needed for his work on other but faint near posteromedial depression, Gnamptodontinae. notauli merged with one another on anterior margin of posteromedial depression Pseudognaptodon whitfieldi Williams, . (Fig 203) propodeum with basa i cell corn- new species plete, somewhat transverse, carinae more (Figs. 199-208) developed laterally than at cell apex (Fig. Diagnosis.—This species is separated 204). Metasoma: Tl length 1.1-1.3X as long from other curtica i/da-group species by the as apical width, lateral margins slightly following combination of characters: Ver- concave posterior to spiracle, spiracle tex of head with weak granular micro- slightly protuberant, lateral carinae corn- sculpture; LOL as long as or shorter than plete to apex but merged with coarse striae LOD (Fig. 200); T2 of metasoma almost and ridges originating from spiracles on entirely covered by coarse striae except for very convex medioapical area; Tl coarsely narrow apical band; T3 of metasoma me- striate throughout except for basal area be- striate x dially on basal h, with polished an- tween carinae (Figs. 205, 206); basal raised terolateral corners defined by bands of area of T2 0.3 of total T2 length, posterior sculpture—(Fig. 207). margin of basal raised area evenly curved Female. Color: Body medium to dark or slightly irregular and medially concave; brown except for the following: Scape and basal raised area margin crenulate, striate pedicel yellow; legs yellow, except hind on most of tergum posterior to basal raised tarsus light brown. Head: Length of anten- area except for a narrow apical band; T3 nal scape 1.8X width; flagellum with 14- 0.9-1.1 X as long as T2, striate medially on 16 of first flagellomeres: L/W three flagel- basal Vz, with anterior groove crenulate, lomeres 2.9-3.0, 2.8-2.9, 2.7; L/W of apical and anterolateral smooth areas delimited Volume 13, Number 1, 2004 203

in 1 load in dorsal \\c,\d in 199-202. Pseudognaptodon whitfieldi. 199, Ocelli dorsal view. 200, view. 201, al view. 202, Wings. 204 Journal of Hymenoptera Research

in dorsal view. Figs. 203-208. Pseiidognaptodon whitfieldi. 203, Mesosoma in dorsal view. 204, Propodeum 205, Tl of metasoma in dorsal view. 206, Tl of metasoma in lateral view. 207, Anterior end of metasoma in dorsal

view. 208, Hind leg.

R.F. by narrow bands of sculpture (Fig. 207). PARTYPES: Costa Rica: Puntarenas: Legs: Hind femur length 3.1 X maximum Golfo Dulce, 24 km W. Piedras Blancas, P. width (208). 200m, 11-1993 (1 9 ), III-1993 (1 9 ), Han- Material examined.—HOLOTYPE 9 son, (UWYO). U.S.A.: Texas: Erath Co.: (UWYO), labelled as follows: "Costa Rica: Stephenville, suction trap, 25-31-111-1982, Puntarenas Golfo Dulce, 24 km W. Piedras C. Agnew, 1 9 (TAMU). Blancas, 200m xii-1991. Paul Hanson". Etymology. —This species is named for Also red label "HOLOTYPE" and bor- James Whitfield, who has provided the dered label "Pseiidognaptodon whitfieldi only specimens in this study for which Williams Holotype det. D. Williams 2003". there are confirmed host data. Volume 13, Number 1, 2004 205

REMARKS ON OTHER SPECIMENS provided numerous specimens and waited patiently while this was on a EXAMINED manuscript produced very part- time basis. This study was supported by David Lan- I have examined three problematic gor, who allowed me time that should have been de- voted to other Pseudognaptodon specimens from CNCI projects. that have not been included in the species LITERATURE CITED treatments above. Two are single males that probably each represent new species Achterberg, C. van, 1983. Revisionary notes on the but are not described here because females subfamily Gnaptodontinae, with description of are label is as fol- unknown. Their data eleven new species (Hymenoptera, Braconidae). lows: "Constance Bay Carleton Co., ONT Tijdschrift voor Entomologie 126: 25-57. W.H. 1894. on the VIII-24-1983 M. Sanbourne" (CNCI), and Ashmead, Report parasitic Cyni- pidae, part of the Braconidae, the Ichneumoni- "USA: FL: Alachua Co. Gainesville, DPI dae, the Proctotrypidae, and part of the Chalci- 1-23.VI.1987, MT J (CNCI). Both Wiley" didae. Part II. Braconidae. Journal of the Linnean also labelled "Pseu- specimens have been Society of London, Zoology 25: 108-138. M. 1965. Der der Nearktischen Re- dognaptodon n. sp. det. D.J.M. Williams Fischer, Opiinae II. Polskie Pismo 35:3-212. 2003". The third specimen is a teneral fe- gion Entomologie Fischer, M. 1967. Die Amerikanischen Arten der Gat- male with pale color and incompletely tungen Euopius, Gnaptodon, und Pseudognaptodon. sclerotized and veins, whose legs wing Beitrdge zur Entomologie, Band 17, Nr 5/8: 959- identity cannot be determined with cer- 976. M. 1977. Braconidae tainty. It is labelled as follows: "9 mi. N. Fischer, Hymenoptera (Opiinae Das Tierreich 96:1-1001. Forrest, Man. 29.VIII.1958 R.L. Hurley" II-Amerika). Shenefelt, R.D. 1975. Pars 12, Braconidae 8, Exothe- I have also added a label "Pseu- (CNCI). in: cinae Rogadinae. Pp. 1115-1262. Vecht, J. van omissus teneral det. dognaptodon prob, der and R.D. Shenefelt, (eds.). Hymenoptorum Ca- D.J.M. Williams 2003". I have not includ- talogus (nova edito). Dr. W. Junk, The Hague. ed it in the treatment of P. omissus above Wharton, R.A. 1997. Subfamily Gnamptodontinae, 256-259, in: Wharton, R.A., P.A. Marsh, and because it does not possess all of the di- pp. M.J. Sharkey, (eds.). Manual of the Mew World gen- agnostic characters of that species. Deter- era of the family Braconidae. Special Publication of mination of whether this is a specimen the International Society of Hymenopterists, Num-

P. or . geographical variant of omissus rep- ber 1 resents a new species cannot be made Wharton, R.A., P.A. Marsh, and M.J. Sharkey, (eds.). Manual of the New World the Bra- with this specimen. genera of family conidae. Special Publication of the International So- ACKNOWLEDGEMENTS ciety of Hymenopterists, Number 1. 439 pp. Whitfield, J.M. and D.I.. Wagner 1991. Annotated kej Henri to I thank David Langor, John Huber, and Gou- the genera of Braconidae (Hymenoptera) at- let for providing a critical review of the manuscript. tacking leafmining 1 epidoptera in the Holarctic Scott Shaw, Robert Wharton, and James Whitfield region, journal of Natural History 25: 733-754. J. HYM. RES. Vol. 13(1), 2004, pp. 206

EDITOR'S NOTE

With this issue of the Journal I regret- Monks Wood fully step down as editor. It has been a Abbots Tipton, Huntingdon PE28 2LS UK pleasure (in most cases) to work with the I am extremely grateful to the subject many fine hymenopterists who have con- editors for their noble assistance through- tributed papers over the past seven years. out the editorial process. There was defi- It is time to infuse the journal with new a of duties without much blood and new ideas, and I am confident nitely sharing on their behalf, but re- that your new editor, Gavin Broad, will do recognition they duced efforts so. Dr. Broad may be contacted by email my considerably. They should all receive medals for their consci- at [email protected]; his office phone num- entious and unheralded duties to ber is 01487 772406 and his fax number is largely our 01487 773467. society. Our many reviewers should also be for much time Beginning at this moment, if not before, recognized devoting and attention to their reviews and the all manuscripts should be sent to the fol- health of our lowing address: journal. In bowing out, let me simply say: Keep Dr. Gavin Broad those manuscripts coming—to Gavin's ad- Coordinator of Zoological Data & Re- dress, that is—and may the journal live search long and prosper! Biological Records Centre Centre for Ecology & Hydrology —Eric Grissell

INSTRUCTIONS FOR AUTHORS

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Beginning 1 April 2004, all manuscripts and correspondence should be addressed to: Dr. Gavin Broad Coordinator of Zoological Data & Research A Biological Records Centre Centre for Ecoloev & Hvdroloev CONTENTS (Continued from front cover)

L. TOOKER, J. F, A. R. DEANS, and M. HANKS. Description of the Antistrophus rufus (Hymenoptera: Cynipidae) species complex, including two new species 125

WHITFIELD, J. B. and M. T. OLTRA MOSCARDO. The Neotropical species of Deuterixys Mason (Hymenoptera: Braconidae) 134

WILLIAMS, D. J. Revision of the genus Pseudognaptodon Fischer (Hymenoptera: Braconidae: Gnamptodontinae) 149 EDITOR'S NOTE 206