PKOC. ENTOMOL. SOC. WASH. 104(3). 2002. pp. .'S89-601 DESCRIPTION OF AEGESEVCOELA BUFFINGTON, NEW NAME, WITH NOTES ON THE STATUS OF GRONOTOMA FORSTER (HYMENOPTERA: FIGITIDAE: EUCOILINAE) MATTHIiW L. BUFFINGTON Department of Entomology. Texas A&M University, College Station, TX 77843, U.S.A.; current address: Department of Entomology, University of California, Riverside, CA 92521, U.S.A. (e-mail: [email protected]) — Abstract. Aegeseucoela Buffington, a replacement name for Moneucoela Dalla Torre and Kieffer 1910, is described. Aei>eseucoela contains two previously described species, Aegeseucoela flavotiucta (Kieffer), n. comb., and A. grenadensis (Ashmead), n. comb., both of which are redescribed. The status of Gronotoma Forster is discussed, and the synonymy of Eucoilidea Ashmead with Gronotoma is formally documented. Gronotoma nigricornata Buffington, new name, is proposed to replace Eucoilidea nigricornis Kieffer 1908. Thirty-two new combinations in Gronotoma are given, and a checklist ofthe world species of Gronotoma is provided. Species in both Aegeseucoela and Gronotoma are common parasitoids of Agromyzidae (Diptera). Aegeseucoela is restricted to the New World tropics and subtropics, and Gronotoma is common in the Afrotropics, as well as Asia, Australia, and the Palearctic and Nearctic regions. Key Words: Aegeseucoela, Gronotonui, Eucoilinae, Figitidae. Agromyzidae, Cynipoidea Eucoiline wasps are endoparasitoids of 1982a, 1982b) was the first to treat eucoi- cyclorrhaphous Diptera inhabiting a variety line classification from a phylogenetic point ofhabitats. These wasps are generally shiny of view. Through these works, cleargeneric black to dark reddish brown and range in and species level definitions were provided size from 0.5 mm to 5 mm. The Eucoilinae for the first time for a number of Palearctic contain 82 genera and nearly 1000 species, and cosmopolitan taxa. Nordlander 1982b) ( and are by far the most diverse of all figitid summarized his findings by proposing in- subfamilies (Ronquist 1999). Only two ma- formal genus groups defined by explicit jor bodies of work have attempted to clas- morphological criteria, a first step towards sify all of the eucoiline genera (Dalla Torre a more logical and natural classification and Kieffer 1910, Weld 1952), and for the scheme. most part, eucoiline classification schemes An investigation into the phylogenetics have resulted in a great deal of chaos (dis- and classification of one of these informal cussed in Nordlander 1982b). Presently genus groups, the Gronotoma group (Buf- available identification keys to eucoiline fington, unpublished data), resulted in the genera (Dalla Torre and Kieffer 1910, Weld identification of a clade of eucoiline wasps 1952) are largely useless due to the reliance of questionable taxonomic placement. Two on a few key features, none of which are previously described species were found to very dependable. belong in this clade, described here as Ae- Nordlander (1976, 1978, 1980, 1981, geseucoela, and both species are rede- 590 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON scribed. Both species are restricted to the grenadensis (Ashmead) from type status of New World tropics and subtropics where Moneucoela Dalla Tone and Kieffer, and they have been reared on numerous occa- designated tinctipennis Kieffer (one of two sions from agromyzid flies (O. Lewis, un- species described in Kieffer 1907) as the published data). type species ofMoneucoela Dalla Torre and The first indication of the need for a new Kieffer 1907. Weld (1952) did not include eucoiline genus was uncovered during an grenadensis within the included species list examination of Kieffer's eucoiline types. for Moneucoela, resulting in the placement The type specimen of Rhabdeucoelaflavo- of the species as incertae sedis. tincta Kieffer did not possess any of the In their revision of Moneucoela Dalla diagnostic features of the genus Rhabdeu- Torre and Kieffer 1907, Diaz and Gallardo coela Kieffer. and the species was most (1998) did not mention Moneucoela Dalla likely placed in Rhabdeucoela based on the Torre and Kieffer 1910, nor the two species relatively well-developed mesoscutal keel that were included in this genus when it and large scutellar plate (neither of which was proposed (Dalla Torre and Kieffer are universally diagnostic features ofRhab- 1910). Of the two species originally includ- deucoela). Furthermore, Rhabdeucoela fla- ed in Moneucoela Dalla ToiTe and Kieffer votiticta was later moved to two different 1910, one is congeneric with flavotincta genera simultaneously by Weld (1952), Mo- and is redescribed below as Aegeseucoela. neucoela Dalla Torre and Kieffer 1907 and The second species belongs in Zaeucoila Tropideucoila Ashmead 1903. This species Ashmead (Buffington, unpublished data) is not readily accommodated in any of the and will be treated in a subsequent paper. three genera in which it was previously Furthermore, since grenadensis Ashmead placed, and it was therefore coded separate- was designated as the type species for Mo- ly in a phylogenetic analysis (Buffington, neucoela Dalla Torre and Kieffer 1910 by unpublished data), the results of which in- Rohwer and Fagan (1917), and Moneucoela dicate none of the three genera (i.e., Rhab- Dalla Torre and Kieffer 1910 is preoccupied deucoela, Moneucoela, and Tropideucoila) by Moneucoela Dalla ToiTe and Kieffer will remain monophyletic if this species is 1907, Aegeseucoela is proposed here as a included within them. Therefore, Aegeseu- replacement name for Moneucoela Dalla coela is proposed to accommodate this spe- Torre and Kieffer 1910. cies and a second species (discussed be- The second part ofthis paper is dedicated low). to a discussion on the status ofthe eucoiline The second species, Diranchis grenaden- genus Gronotonui Forster. Gronotoma re- sis Ashmead, also has a confusing taxo- sides within a basal portion ofthe eucoiline nomic history. It was one of two species clade (Fontal et al., in preparation), as is the originally included in Moneucoela Dalla case with Aegeseucoela (though these two Torre and Kieffer 1910, which is itself a genera do not form a monophyletic group) junior homonym of Moneucoela Dalla Tor- (Buffington, unpublished data). Similar to re and Kieffer 1907. Rohwer and Fagan Aegeseucoela, species of Gronotoma have (1917) apparently missed Kieffer's 1907 all been reared from agromyzid flies publication containing the description of (Scheffer unpublished data, Davidson 1963, Moneucoela, and determined that the genus Harding 1965, Viraktamath et al. 1993), was described as new in Dalla Torre and mostly found within the genus Melanagro- Kieffer (1910). Further, Rohwer and Fagan myza Hendel (Abe and Konishi 1995; (1917) designated Diranchis grenadensis Greathead 1969, 1971). The host preference Ashmead as the type species ofMoneucoe- for agromyzid flies appears to be a plesio- la Dalla Torre and Kieffer 1910. Weld morphic feature within the Eucoilinae (1952) considered this a mistake, removed (Fontal et al. in preparation). VOLUME NUMBER 104. 3 591 GroiioToimi presently contains 47 de- to Prefectural University, Kyoto, scribed species (including synonymies pro- Japan. posed below), making this genus the most MRAC Musee Royale de I'Afrique Cen- diverse of all eucoiline genera that special- trale, Tervuren, Belgium. ize on agromyzid hosts. Species of Grono- USNM National Museum of Natural His- toma have a worldwide distribution and the tory, Smithsonian Institution, Afrotropics are particularly speciose (Quin- Washington, DC, USA (D. lan 1986). Species are also very common Smith). in the Oriental (Abe and Konishi 1995), the ZIN Zoological Institute, Russian Palearctic, and the Nearctic regions (Dalla Academy of Sciences, St. Peters- Torre and Kieffer 1910, Weld 1952). Mem- burg, Russia. bers of this genus are rarely recorded from ZMHB Zoologisches Museum, Humboldt the Neotropical Region but are frequently Universitiit, Berlin, Germany (F. collected throughout subtropical Mexico Koch). (Buffington, unpublished data). Because of their potential usefulness in the biological Type specimens (holotypes and para- control of pest Agromyzidae and the rela- types) were generously loaned to me by the tively high global diversity of species with- CAS (containing a large portion of Kieffer's in the genus, the status of Gronotoma is eucoiline types originally housed in Po- reviewed and includes an important syn- mona), the USNM (containing many of onymy. Phylogenetic evidence (Buffington, Ashmead's and Weld's eucoiline types), the BMNH unpublished data) supports the conclusions (containing Quinlan's eucoiline of Hedicke (1930) and Beardsley (1988) types), and the ZMHB (containing many of with respect to the synonymy ofEiicoilidea Forster's eucoiline types). Ashmead with Gronotoma. A world check- While sorting unidentified eucoilines in list of the species of Gronotoma is present- the AEIC in the summer of 2000, I found ed, with nomenclatural notes where appli- an extensive series ofeucoilines conspecific cable. with the type specimen ofAegeseucoelafla- votincta, all collected from Costa Rica. Ad- BMNH The Natural History Museum, ditionally, a long series of Aegeseucoela UK London, (S. Lewis). flavotincta and ofA. grenadensis were sent BPBM Bernice P. Bishop Museum, Hon- to me for identification by O. Lewis (Sil- olulu, HL USA. wood Park, UK); these specimens were of CAS California Academy of Sciences, particular importance since host data ac- San Francisco, CA, USA (W. Pu- companied each specimen. Unidentified laski, R. Zuparko). specimens ofAfrican Gronotoma were gen- CNC Canadian National Collection of erously loaned to me by Dr. Robert Cope- Insects, Ottawa, Canada (J. Hub- land (ICIPE). er, J. Read, L. Masner). Institutions and individuals that donated CUIC Cornell University Insect Collec- ethanol preserved specimens for this study tion, Ithaca, NY, USA (E.R. Hoe- were: CNC; TAMU; and Dr. Owen Lewis, beke). Imperial College at Silwood Park, UK. All IBPR Institute of Biology at the Acad- SEM images utilized in this study were pre- emy of the People's Republic of pared digitally on a JOEL JSM-5600 SEM Romania. (operated by James Ehrman, Digital Micro- ICIPE International Centre of Insect scope Facility, Mt. Allison University, Physiology and Ecology, Nairobi, Sackville, NB, Canada). Kenya (R. Copeland). Terminology follows largely that ofWeld KPU Laboratory of Entomology, Kyo- (1921, 1952), Nordlander (1982b), Ron- 592 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. \. Aegeseucoelaflavotincta. A, Side view, habitus. B, Head, anteriorview; A = orbital furrowcomplete to lateral ocellus. C, Mesosoma, dorsal view; A = parapsidal hair line; B = well developed lateraldorsal pro- jections of scutellum; C = pronotal triangle. quist and Nordlander (1989), and Ronquist groove originating at either the lateral ocel- (1995), with the following modifications: lus or the lateral side of the torulus (de- parapsidal ridges is preferred over parapsi- pending on the taxon) and lining the inner dal furrows, as found in Weld (1952); or- orbit of the eye, terminating at the clypeal bital furrows and pronotal triangle are new margin after fusing with or paralleling the terms and are defined as such—below. malar sulcus. — Orbital furrow (Fig. IB). A distinct Pronotal triangle (Figs. IC, 2C). An — VOLUME 104. NUMBER 3 593 area on the dorsal surface oi' the proiiotum Pronotiiin: Pronotal plate wide, with se- bordered by the lateral pronotal carina, the tae along dorsal margin; slightly crested pronotal plate and the anterior margin ofthe and bifurcate dorsally; pront)tal fovea open. mesoscLitum; found in genera with well de- [*ronolai triangle present (Figs. IC, 2C). veloped pronotal ridges. Pronotal impression absent. Lateral aspect of pronotuni (below pronotal triangle) Aegeseucoela Buffinj^ton, new name smooth and glabrous. Lateral pronotal ca- Moiu'iicoehi Dalla Torre and Kieffer 1910: rina absent. 103. 888. Type species: Diranchis gren- Mesosciitiini: Smooth with some setae. adeiisis Ashmead. designated by Rohwer Mesoscutal keel present, reaching posterior margin of mesoscutum; tapering posterior- and Fagan 1917. Preoccupied by Moneii- coehi Dalla Torre and Kieffer 1907. ly. Parapsidal ridges absent. Parapsidal hair lines present (Figs. IC. 2C). Parascutal im- — Diagnosis. Orbital furrows originating pression incomplete, narrow. Notauli ab- either at lateral ocellus or lateral side of to- sent. lulus. Genal carina well developed, often Mesopectus: Upper part and lower part flanged posterior to compound eye. Meso- of mesopleuron glabrous and smooth. Dor- scutal keel present, at least anteriorly. Par- sal margin of mesopleural triangle well de- apsidal ridges absent. Parapsidal hair lines fined, rounded ventrally. Mesopleural cari- present. Laterodorsal projections of scutel- na simple. Lower part of mesopleuron bor- lum present to absent. Rl never tubular or dered by distinct precoxal carina; anterior pigmented (radial cell open). Most similar surcoxal depression present, reticulate. to Zaeucoila and Agrostocynips, but differs Scufeniini: Scutellar plate ranging from by the presence of the extended orbital fur- medium to large; mid pit placed centrally rows (in some species), the presence ofpar- on plate; plate truncated posteriorly; nearly apsidal hair lines and Rl incomplete. always bearing tubercles and setae on dor- Description. Head: Nearly glabrous, sal surface. Dorsal surface of scutellum re- with a few scattered setae along lower face, ticulate; margined laterally and posteriorly. clypeus. inner orbits of compound eyes, Laterodorsal projections present, present to malar space and gena; orbital hair patches nearly absent (Figs. IC. 2B. 2C); posterior present. Ventral Va of lower face with ad- projections absent. medial clypeal furrows converging towards Metapectal-propodeal complex: Meta- clypeus. Orbital furrows present, originat- pectus nearly glabrous with a few scattered ing at lateral side of torulus or at lateral setae posteriorly. Spiracular groove with a ocelli (species dependent), terminating at well defined dorsal margin and a well de- malar sulcus. Malar sulcus simple. Malar fined to rounded ventral margin. Posterior space smooth with a single prominent con- margin of metapectus ridged. Metapleural ical protuberance. Genal carina present, ex- ridge absent; submetapleural ridge variable tending from malar space to lateral ocelli, from present to absent. Anterior impres- often undulating posterior to compound sions of metepimeron and metepisternum eye. present. Anteroventral cavity semi-circular Antenna: Female. 13 segments, monili- and setose. Propodeum covered in long se- form. semi-clavate; segments 3-13 sube- tae. Lateral propodeal carinae semi-parallel, qual in length; rhinaria present on segments bowed at junction with auxiliary propodeal 3-13. Male. 15 segments, filiform; rhinaria carinae; auxiliary propodeal carinae indis- present on segments 3-15; segments 4-15 tinct. Nucha glabrous, reticulate. subequal in length. Segment 3 slightly lon- Wings: Hyaline, with base of forewing ger than 4, curved outwardly, excavated lat- sometimes darkened; usually setose basally, erally. always setose apically. Rl incomplete, mar- 594 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 2. AegeseiicoeJa grenadensis. A» Side view, habitus. B. Mesosoma, posterolateral view; A = reduced lateral projections ofscutellum. C, Mesosoma, dorsal view; A = parapsidal hair line; B = reduced lateraldorsal projection ofscutellum; C = pronotal triangle. ginal cell as long as deep. Apical fringe pre- coxa with a prominent setal band on hind sent, short. margin. Femora and tibiae sparsely setose; Legs: Fore and mid coxae about the same tarsomeres with dense, appressed setae. size, hind coxa about twice size of either Length of hind tarsomere 1 equal to 0.5X fore or mid coxae. Fore coxa variably cov- combined length of remaining hind tarso- ered in long setae; mid coxa with anterior meres. and posterior dorsoventral setal bands; hind Metasoma: Female: Sub-equal in size to — VOLUME 104. NUMBER 3 595 mesosoma. Base of synlerguni with a hairy 1908:46, hololype in CAS (#10537). Re- ring present, ranging from complete to dor- described below. sally bare, comprised of short, semi-ap- Aegeseucoela grenadensis (Ashmead). n. pressed setae and longer erect setae; re- comb. Diranchis grenadensis Ashmead mainder of metasoma glabrous. Micropunc- 1900: 248. holotype in BMNH. Rede- tures present on posterior V4 of syntergum, scribed below. and on remaining terga. Terga posterior to Aegeseucoelaflavotincta (Kieffer), syntergum directed posteroventrally, result- new combination ing in a 70 degree angle between syntergum and remaining terga. Male: As in female — (Fig. 1) but terga posterior to syntergum abruptly Description. As in description of genus angled ventrally. resulting in a 90 degree except as follows: Head: Orbital furrows angle between syntergum and remaining originating at lateral ocelli (Fig. IB); genal terga. — carina continuing to near lateral ocelli, un- Biology. I have examined specimens dulating posterior to compound eye (Fig. reared by O. Lewis from species of agro- lA). Fronotuni: Pronotal crest prominent, myzid flies in the genera Haplopeodes bifurcate; pronotal ridge well developed. Steyskal (on Solanaceae) and Calycomyzci Scutellum: Scutellar plate large, nearly Hendel (on Fabaceae). round; laterodorsal projections of scutellum — Distribution. Neotropical Region: Cos- sometimes well developed (Fig. IC). Me- ta Rica. Belize. Mexico (Veracruz. Tamau- tapectal-propodeal complex: Submeta- lipas). Panama. Nearctic Region: USA: AZ. pleural ridge usually well developed, con- Previously only known from Guatemala necting ventral margin of spiracular groove (flavotincta) and Grenada {grenadensis). with posterior margin of mesopleuron. Etymology. Aeges; from Greek my- Wings: Base offorewing occasionally dark- thology, the name of Athena's dreaded ened; base of forewing ranging from gla- shield, used here to reference the broad, brous to setose; forewing always setose api- shieldlike pronotal plate present in this ge- cally. Metasoma: Hairy ring at base of syn- nus; eucoela, a suffix frequently used by tergum often highly reduced (narrow), but J.J. Kieffer in his treatments of the Neo- always present. — Material examined. Holotype 9. troCpiocmamleEnutcso.il—inTaeh.ough this genus is close- Champerico, Guatemala. Coll. Baker. CAS ly related to Zaeucoila and Agrostocynips, #10537; the type specimen is in good con- phylogenetic evidence (to be found in a dition, with Kieffer's original determination label (a large red label), the collection data forthcoming publication by the author) sug- labels, depository label and my determina- gests that neither of these genera will re- tion label (slender white label). Additional main monophyletic if the species of Aege- material: BELIZE: Las Cuevas. Chiquibul seucoela are placed within them. The par- Forest. Cayo District. 550 m. (various dates apsidal hair lines, extended lateral protu- between Oct. 1997 and Sept. 1998), O.T. berances on the scutellum and incomplete Lewis (13 9, 5 6). BOLIVIA: Yungas, Rl vein on the forewing are reliable auta- XII.4.84. 2.400 m. Luis Peiia (19); COS- pomorphies for the genus, and reliable syn- TA RICA: S. Rosa Park. Guan., various apomorphies for the species within the ge- dates between 27.V.1976 and I0.VIII.1978, nus. D.H. Janzen. Dry Hill (46 c?. 90 9). MEX- ICO: Veracruz. 2 km SW Fortin, 8°54'N, Included Species 97°00'W, 2.700'. 23.VI.1997, J.B. WooIIey, Aegeseucoela flavotincta (Kieffer), n. screen sweep (1 9); Veracruz, El Crucero comb. Rhabdeucoela flavotincta Kieffer nr Puente Nacional, 19°20'N, 96°26'W, 596 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 13.VI.1997, L.A. Wilson & J.B. Woolley (1 mead's original determination label (in 9); Tamaulipas, 97 km Ciudad Victoria, Ashmead's hand). Below that is the collec- Hwy 70, 3.VII.1986, G. Zolnerowich & R. tion data, and finally below that, my des- Trevino (19). PANAMA: Colon Prov.. 2 ignation label. Additional material: BE- km S Sabanitas, 4-15.VII.1999, 120 m, LIZE: Las Cuevas, Chiquibul Forest, Cayo Gillogly & Woolley, MT 99/033 (1 9). District, 550 m, (various dates between Oct. U.S.A: Arizona, Portal, 19-23.VIII.1987, 1997 and Sept. 1998), O.T Lewis (27 9, H. & M. Towne—s (4 6). 20 (?). MEXICO: Veracruz, 0.7 mi N Jilo- Distribution. Neotropical and southern tepec, 19°36'N, 96°56'W, 3,680', screen & Nearctic regions (see above list of locali- sweep, 14.VI.1997, L.A. Wilson J.B. ties). — Woolley (1 9). PANAMA, Prov. Colon, Biology. I have examined specimens Quebrada Lopez, MT, 2-4.VII.1999, A. reared from the agromyzid flies Haplopeo- Gillogly (1 9). VENEZUELA: Merida, des sp. on Solanum ericmthiim D. Don (So- Merida City, 8°35'54"N, 71°08'42"W, 1860 lanaceae) and Calycomyzci hyptidis Spencer m, sweep veg. along trib. to Chama R., on Hyptis capitata Jacq. and H. urticoides 3.V.1981, L. Ma—sner (1 9). Kunth (Lamiaceae) (data from O. Lewis). Distribution. Neotropical Region (see above list o—f localities). Aegeseucoela grenadensis (Ashmead), Biology. I have examined specimens new combination reared from the agromyzid flies Calycomy- — (Fig. 2) zci verhenivora on Verbena sp. (Verbena- Description. As in description of genus ceae); Calycomyza c.f. cassiae (Frost) on & except as follows: Head: Orbital furrows Senna cohanensis (Britton) H.S. Irwin originating at torulus; genal carina reduced, Barneby (Fabaceae); Haplopeodes sp. on non-undulating, terminating at dorsal mar- Solanum erianthum (Solanaceae) (data gin of compound eye (Fig. 2A). Pronotum: from O. Lewis—). Pronotal crest reduced, sometimes absent; Comments. The type specimen of Dir- pronotal ridge sometimes absent. Scutel- anchis grenadensis Ashmead, as noted luni: Scutellar plate ranging from medium above, is labeled 'paratype.' In the original to small; laterodorsal projections usually re- description of D. grenadensis, Ashmead duced/absent (Figs. 2B, 2C). Metapectal- stated "described from 1 female speci- propodeal complex: Submetapleural ridge men." Since the single specimen, the lo- completely reduced. Wings: Base of fore- cality data, and the description all agree, wings occasionally darkened; usually entire this must be the holotype. forewing surface is setose. Metasoma: Gronotoma Forster Hairy ring at base of syntergum thick and densely pubescent. Gronotoma Forster 1869: 342, 346. Type — Material examined. Holotype 9 Bal- species: Gronotoma sculpturata Forster ; thazar (windward side), Grenada, West In- 1869: 346, by original designation. dies. H.H. Smith (Coll.), BMNH; the type Eucoilidea Ashmead 1887: 150, 154. Type specimen is in poor condition, with the species: Eucoilidea canadensis Ashmead, head plus thorax on one end of a pinned by subsequent designation (Ashmead card, and the metasoma on the other end. 1903); synonymy by Hedicke (1930) and The diagnostic features discussed below for Beardsley (1988). this genus are all visible. Two *type" labels Eucoelidea Dalla Tone 1893: 15; Kieffer are present on the specimen, one with a red 1901: 159. Emendation. circle and the second labeled 'BM Type Afrostilba Benoit 1956: 544. Type species: Hym., y. 50'. Under this lies a label with Afrostilba nirida Benoit, by monotypy. 'paratype' printed on it. Below that is Ash- Synonymy by Quinlan (1986). VOLUME 104. NUMBER 3 597 Ashmead (1887) proposed the genus Eh- hairy ring at base ofsyntergum absent. Spe- coilidea to accommodate E. longiconii.s and cies ofGronotonui are most easily confused E. canadensis, but did not specify a type with Dii^lypliosenui Forster, but easily dis- species. Later, Ashmead (1903) designated tinguished IVom that genus by having the Eucoilidea canadensis as type species. Dal- scuteUum meeting the scutellar plate at an la Torre (1893) emended the original Ash- acute angle (meets at a 90 degree angle in mead spelling o'i Eucoilidea to Encoelidea, Dii^lyphoseina), and the scutellar plate not and cites the former in brackets. Encoelidea as elongate (though some species of Gron- was recognized in Kieffer ( 1901 ), and Kief- otonui tend to have an oval scutellar plate). fer (1907, 1909) described four new species Previous works treating the synonymy of (and two varieties) in 'Encoelidea Ashm.' Eucoilidea with Gronotonui focused pri- None of Kieffer's treatments of Encoelidea marily on the type species; thus, as indicat- mention the name's emendation from Eu- ed in the list below, most of the described coilidea. Based on Article 33.2.1 of the species have not been formally transferred ICZN (1999), Encoelidea is an emendation to Gronotonui. Hence, 32 new combina- of Eucoilidea and is not available. Burks tions are proposed, mainly reflecting the ( 1979) reported two apparent misspellings taxonomic work on this genus by Weld of Eucoilidea, namely Eucoilidia (Kieffer (1952) and Quinlan (1986). Species for 1907) and Eucoelidia (Kieffer 1909); after which type material (holotypes and/or para- consulting the original references, neither types) were examined are indicated by an of these misspellings were found to exist. *. The reader should also note that Kieffer It is apparent from Ashmead's (1887) (1901) did indeed treat four species in original description of Eucoilidea that he Gronotonui (and not Eucoilidea), i.e., made no comparison with Gronotonui be- Gronotonui carinata (Cresson), G. minor fore proposing the new genus, and Hedicke (Provancher), G. nigricornis Kieffer and G. (1930) was the first to propose Eucoilidea oralis (Thomson) (see in list below). as a synonym of Gronotonui. Weld (1952) and Quinlan (1986, 1988), however, sub- Included Species sequently maintained Eucoilidea as distinct from Gronotonui. Beardsley (1988) again adachiae Beardsley 1988: 39, holotype in BPBM. synonymized these two genera. Though the holotype of E. canadensis is in poor con- ''advena (Quinlan), n. comb. Eucoilidea dition, with great portions of the body ob- advena Quinlan 1986: 262, 263, holotype scured by blackish glue, features such as in MRAC, paratypes in BMNH. the notauli, the lateral pronotal carinae and allotriaeforniis (Giraud). Eucoila allotriae- the scutellar plate are all clearly visible. fonnis Giraud 1860: 142. Gronotonui al- Comparing these features with those found lotriaeforniis: Forster 1869: 346. Type in Gronotonui sculpturata clearly indicate depository presently unknown. that indeed, Eucoilidea is ajunior synonym *arcuata (Kieffer), n. comb. Encoelidea of Gronotonui, supporting the decisions of arcuata Kieffer 1909: 65. Type in CAS. Beardsley (1988) and Hedicke (1930). "^bakeri (Kieffer), n. comb. Encoelidea Species of Gronotonui can be recognized bakeri Kieffer 1907: 107. Type in CUIC. by the following diagnostic features: lateral '^bakeri var. cupularis (Kieffer), n. comb. pronotal carina present (a distinct ridge, dis- Encoelidea bakeri var. cupularis Kieffer tal to the lateral margins of the pronotal 1907: 108. Type in CAS. plate, that give species in the this genus the ''^bakeri war.flavipes (Kieffer), n. comb. En- appearance of having a large pronotal coelidea bakeri var. flavipes Kieffer plate); notauli present and well-developed 1907: 108. Type in CAS. in nearly all species; scutellar plate large; *bucca (Quinlan), n. comb. Eucoilidea 598 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON biicca Quinlan 1986: 263, holotype in lacerta (Quinlan), n. comb. Eucoilidea lac- MRAC, paratypes in BMNH. erta Quinlan 1986: 266, 267, holotype '^canadensis (Ashmead). Eucoilidea cana- and paratype in MRAC. densis Ashmead 1887: 154. Gronotoma lana (Quinlan), n. comb. Eucoilidea lana canadensis: Hedicke 1930; Beardsley Quinlan 1986: 267, holotype and para- 1988. Holotype in USNM. type in MRAC. carinata (Cresson). Eucoila ? carinata *'leptis (Quinlan), n. comb. Eucoilidea lep- Cresson 1865: 6. Gronotoma carinata: tis Quinlan 1986: 267, 268, holotype in Kieffer 1901: 159. Type depository pres- BMNH. ently unknown. longicornis (Ashmead), n. comb. Eucoili- *compressa (Quinlan), n. comb. Eucoilidea dea longicornis Ashmead 1887: 154. compressa Quinlan 1986: 263, 264, ho- Type depository presently unknown. lotype in MRAC, paratypes in BMNH. maquilingensis (Kieffer), n. comb. Eucoil- *conversa (Quinlan), n. comb. Eucoilidea idea maquilingensis Kieffer 1914: 184- conversa Quinlan 1986: 264, holotype in 185. Type depository presently unknown. MRAC, paratypes in BMNH and MRAC. '^marcellus (Quinlan), n. comb. Eucoilidea '^crenulata (Kieffer), n. comb. Eucoilidea marcellus Quinlan 1986: 268, holotype in crenulata Kieffer 1908: 47. Type in CAS. BMNH, paratypes in BMNH and MRAC. *dilitata (Kieffer), n. comb. Eucoelidea ^mauri (Quinlan), n. comb. Eucoilidea dilitata Kieffer 1907:108. Type in CAS. mauri Quinlan 1986: 268, 269, holotype domestica Girault 1932: 3. Type depository and paratype in BMNH. presently unknown. melanagromyzae Beardsley 1988: 40, ho- *dubia (Quinlan), n. comb. Eucoilidea du- lotype in BPBM. bia Quinlan, 1986: 264. 265, holotype in micromorpha (Perkins). Eucoilidea micro- BMNH, paratypes in BMNH and MRAC. niorpha Perkins 1910: 676. Gronotoma extraria (Quinlan), n. comb. Eucoilidea ex- micromorpha: Beardsley 1988: 38. Eu- traria Quinlan 1986: 265, holotype and coilidea rufula Yoshimoto 1962b: 845, paratype in MRAC. synonymy by Beardsley (1988), holotype *fetura (Quinlan), n. comb. Eucoilidea fe- and paratypes in BPBM. tura Quinlan 1986: 265, 266, holotype in minor (Provancher). Eucoila minor Pro- MRAC, paratypes in BMNH. vancher 1888: 398. Gronotoma minor: fulvicornis (Hedicke). Ganaspisfulvicornis Kieffer 1901: 159. Type depository pres- Hedicke 1913: 445. Gronotomafulvicor- ently unknown. nis: Hedicke 1934: 704. Holotype and nigra lonescu 1963: 10, holotype in IBPR. four paratypes in ZMHB. '"nigricornata Buffington, new name. Eu- "^'furcula (Quinlan), n. comb. Eucoilidea coilidea nigricornis Kieffer 1908: 48. furcula Quinlan 1986: 266, holotype in Preoccupied by Gronotoma nigricornis BMNH, paratypes in BMNH and MRAC. Kieffer 1901: 159. Paratypes in CUIC. ^'fuscipes (Kieffer), n. comb. Eucoelidea nigricornis Kieffer 1901: 159. Type depos- fuscipes Kieffer 1907: 1 12. Type in CAS. itory presently unknown. gracilicornis Cameron 1889: 15. Type de- '*nitida (Benoit). n. comb. Afrostilba nitida pository presently unknown. Benoit 1956: 544, holotype in MRAC, guamensis (Yoshimoto), n. comb. Eucoili- paratypes in BMNH. Eucoilidea nitida: dea guaniensis Yoshimoto 1962a: 107, Quinlan 1986: 269. holotype and paratypes in BPBM. ovalis (Thomson). Cothonaspis avails hiranoi Abe and Konishi 1995: 309-311, Thomson 1877: 817. Gronotoma ovalis: holotype and paratypes in KPU. Kieffer 1901: 159. Type depository pres- insularis Ashmead 1895: 743, holotype in ently unknown. BMNH. ^pallida (Quinlan), n. comb. Eucoilidea