Journal of Hymenoptera Research Volume 17, Number 2 | October 2008 ISSN #1070-9428 CONTENTS WILLIAMS, K. A. and J. P. PITTS. Three species masquerading as one: updating the taxonomy of Pseudomethoca russeola Mickel and P. donaeanae (Cockerell & Fox) (Hymenoptera: I Ae RP ea Pe EL i Piss asim Ia oa ln ws wii Sie bd hind Va one eb ame bas 427 POLASZEK, A. and S$. MANZARI. A new species of Encarsia (Hymenoptera: Aphelinidae) parasitising Aleuromarginatus tephrosiae (Hemiptera: Aleyrodidae) inIranand Oman... 134 KULA, R. R. Taxonomic status and location of type specimens for species of Coelinidea Viereck and Sarops Nixon (Hymenoptera: Braconidae: Alysiinae) described by Garland T. er sr ce ght wi wie ie eb eek a sai o'e We wide oles > 138 KIMSEY, L. S. The Neotropical chrysidid genus Adelphe Mocsary revisited (Hymenoptera: regres ere SBMS ORC ea yee ects sees ein ee ca Re ek eee ER Eee to¥ HUBER, J. T., G. A. P. GIBSON, L. S. BAUER, H. LIU, and M. GATES. The genus Mymaromella (Hymenoptera: Mymarommatidae) in North America, with a key to described extant Se Oa ee ee eG own ln a ss pela els dle wed a cineca ews eae Wis. FRAMPTON, M.,S. DROEGE, T. CONRAD, S. PRAGER, and M. H. RICHARDS. Evaluation of Beeman preservatives tor WNA analyses of Dees... 2... ee ee 195 BARCENAS, N. M.., N. J. THOMPSON, V. GOMEZ-TOVAR, J. A. MORALES-RAMOS, and J. S. JOHNSTON. Sex determination and genome size in Catolaccus grandis (Burks, voli SOV SMe Ce sh 3 ve Tire 2 2) av ne ae 201 GUPTA, S. K.,S. F GAYUBO, and W. J. PULAWSKI. On two Asian species of the genus Mellinus epeeciie 02) Eivtueneprera: (rabronidae) .. 2.2... 2. be ee eee ee ec eee eee 210 OBITUARY: (OSL LE EDS SSF Le 2S A eee rn 216 INTERNATIONAL SOCIETY OF HYMENOPTERISTS Organized 1982; Incorporated 1991 OFFICERS FOR 2008 Michael E. Schauff, President James Woolley, President-Elect Andrew Deans, Secretary Joseph Fortier, Treasurer Gavin R. Broad, Editor Subject Editors SYMPHYTA AND PARASITICA ACULEATA Biology: Mark Shaw Biology: Jack Neff Systematics: Andrew Deans Systematics: Wojciech Pulawski All correspondence concerning Society business should be mailed to the appropriate officer at the following addresses: President, Plant Sciences Institute, Bldg. 003, Rm. 231 BARC-West, Beltsville, MD 20705, USA; Secretary, Department of Entomology, North Carolina State University, Campus Box 7613, 2301 Gardner Hall, Raleigh, NC 27695-7613, USA; Treasurer, Saint Louis University, 3507 LaClede Ave., St. Louis, MO 63103, USA; Editor, Dept. of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK. Membership. Members shall be persons who have demonstrated interest in the science of entomol- ogy. Annual dues for members are US$45.00 per year (US$40.00 if paid before 1 February), payable to The International Society of Hymenopterists. Requests for membership should be sent to the Trea- surer (address above). Information on membership and other details of the Society may be found on the World Wide Web at http://hymenoptera.tamn.edu/ish/. Journal. The Journal of Hymenoptera Research is published twice a year by the International Society of Hymenopterists, % Department of Entomology, Smithsonian Institution, Washington, D.C. 20560- 0168, U.S.A. Members in good standing receive the Journal. Nonmember subscriptions are $60.00 (U.S. currency) per year. The Society does not exchange its publications for those of other societies. Please see inside back cover of this issue for information regarding preparation of manuscripts. Statement of Ownership 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, 0 Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168, U.S.A. Editor: Gavin R. Broad, Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK. Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 1 October 2008 J. HYM. RES. Vol. 17(2), 2008, pp. 127-133 Three Species Masquerading as One: Updating the Taxonomy of Pseudomethoca russeola Mickel and P. donaeanae (Cockerell & Fox) (Hymenoptera: Mutillidae) KEVIN A. WILLIAMS AND JAMES P. PITTS Utah State University, Department of Biology, Logan, Utah 84322, USA; (KAW) email: kawilliams@biology.usu.edu Abstract.— Pseudomethoca donaeanae (Cockerell & Fox) was described based on females only, while P. russeola Mickel was described based on males only. Manley (1999) synonymized P. russeola with P. donaeanae after associating a male that superficially resembles P. russeola with P. donaeanae. Close examination of male genitalia of specimens currently identified as P. donaeanae, along with additional morphological characters, suggests that three species are actually being misidentified as a single species. Our comparison of the male associated with P. donaeanae with the type specimen of P. russeola (male) suggests that these species are not synonymous. The male of P. donaeanae is described for the first time, and P. russeola new comb., is resurrected from synonymy and redescribed. The third species, P. ajattara sp. nov. also superficially resembles P. russeola and P. donaeanae, but has definitive genitalia with hooked setae located ventrally along the internal margin of the parameres. The females of neither P. russeola or the undescribed species are known. Pseudomethoca Ashmead is one of the largest diurnal mutillid genera in the New World, including almost 50 species in the United States. Pseudomethoca species occur throughout the Americas, from Canada to Argentina (Nonveiller 1990). This range is slightly misleading, however, because Pseudomethoca appears to be an unnatural grouping (pers. obs). Like other mutillid genera, Pseudomethoca species exhibit ex- treme sexual dimorphism. As a result, less than half of the species are known from both sexes (Krombein 1979). Additional problems stem from the relative lack of obvious characters useful for diagnosing species based on males. While many females have unique coloration schemes, males exhibit a limited suite of coloration, with most species having the integument entirely black and the setae mostly silver. In some cases, males with unique colora- tion are immediately recognizable, and additional morphological characters are ignored. Pseudomethoca russeola Mickel (1924), known only from the male, is among the species having unique coloration. The head and mesosoma are black, while the meta- soma is orange, and the entire insect is clothed with silvery setae. The male of P. donaeanae (Cockerell & Fox) was discov- ered by Manley (1999), when he attracted two males to a caged female. He identified these males as P. russeola and synonymized the two species under the name P. donaea- nae presumably based on this “‘unique”’ coloration. A study of male genitalia and other characters in Pseudomethoca led to the discovery of three unique species that currently are identified as P. russeola, with all possessing the unique coloration. The taxonomy and sex associations of these species are addressed in this paper. MATERIALS AND TERMINOLOGY The following acronyms are used for institutions housing the material discussed in the current study: 128 ANSP Department of Entomology, Academy of Natural Sciences, Philadelphia, Pennsylvania, USA. Frank M. Hasbrouck Insect Collection, Department of Zo- ology, Arizona State Universi- ty, Tempe, Arizona, USA. Department of Entomology, Ca- lifornia Academy of Sciences, San Francisco, California, USA. Essig Museum of Entomology, Department of Entomological Sciences, University of Califor- nia, Berkeley, California, USA. C.P. Gillette Arthropod Biodi- versity Museum, Department of Entomology, Colorado State University, Fort Collins, Colo- rado, USA. Personal Collection of Donald G. Manley, Pee Dee Research Center, Florence South Caro- lina, USA. Department of Biology Insect Collection, Utah State Univer- sity, Logan, Utah, USA. Department of Entomology In- sect Collection, Texas A&M University, College Station, Texas, USA. Department of Entomology, Smithsonian Institution, Na- tional Museum of Natural His- tory, Washington, District of Colombia, USA. ASUT CASC GISC CSUC DGM EMUS TAMU NMNH The holotype of P. russeola was exam- ined, but that of P. donaeanae was not available. We have used the acronyms T2, T3, etc., to denote the second, third, etc., metasomal tergites while $2, S3, etc., denote the second, third, etc., metasomal sternites. Lastly, punctures can sometimes be elongate and their posterior edge indistinct. We have used the term “punc- ture width” to indicate the transverse measurement of the width of a puncture. This is the only way to accurately and JOURNAL OF HYMENOPTERA RESEARCH reproducibly measure an elongate punc- ture: Pseudomethoca ajattara, new species (Figs 1, 2, 5, 7, 10-13) Diagnosis.—The male is similar to P. donaeanae and P. russeola in coloration, wherein the integument of the head and mesosoma is black, the integument of the metasoma is orange, and the setae are silvery white (Fig. 1). This species can be separated from other species with this coloration by the following combination of characters: the head is narrower than pro- notum; the clypeus is expanded anteriorly with two medial approximate teeth (Fig. 7); the apical fringes of T2-4 have dense, thick, pale golden setae, while T4-5 have inter- spersed brown and pale golden setae; the paramere has long, elbowed setae along the internal margin (Fig. 10); and the cuspis has an apical finger-like process (Fig. 10). Male holotype description.—Coloration: Head and mesosoma black to dark red- dish-brown, except metapleuron red; me- tasoma orange; legs dark reddish-brown; tibial spurs reddish-brown, lighter than legs. Wings slightly infuscated. Setae of head, mesosoma, and legs silvery white, except mesonotum with erect and ap- pressed dark brown setae. Setae of meta- soma entirely pale golden, except T5-6 and disc of T2 having interspersed brown and pale golden setae. Head: Narrower than pronotum, densely punctate throughout. Mandible oblique, tridentate apically, inner tooth strongly developed (Fig. 7). Clypeus densely punctate, anteriorly expanded, covering inner margin of mandibles, with two approximate median teeth (Fig. 7). Antennal scrobe lacking carina. Ocelli miniscule; ocellocular distance 10 length of lateral ocellus, interocellar distance 3X lateral ocellar length. Flagellomere I 2X pedicel length; flagellomere II 3X pedicel length. Mesosoma: Pronotum moderately punctate; mesonotum and _ scutellum densely punctate; mesopleuron moderately VOLUME 17, NUMBER 2, 2008 129 Figs 1-6: Habitus, Fig. 1: Pseudomethoca ajattara. Metasomal terga, Figs. 2-4; Fig. 2: P. ajattara; Fig. 3: P. donaeanae; Fig. 4: P. russeola. Fore wing, Figs. 5-6: Fig. 5: P. ajattara; Fig. 6: P. donaeanae. punctate with micropunctures anteriorly, metapleuron glabrous; propodeum reticu- late dorsally, horizontally striate laterally. Tegula evenly convex, punctate and pu- bescent throughout. Marginal cell 2.75 length of stigma. Metasoma: Petiole broadly sessile, evenly convex. Apical fringes of T2- 4 forming dense rows of short, evenly _ spaced, slightly curved, pale golden bris- tles; bristles separated by 0.5x bristle width. T1 with ovate punctures; T2 and S52 moderately punctate; T3-6 and S3-6 moderately punctate. S1 with low longitu- dinal carina. Pygidium densely punctate, with micropunctures and fine setae among punctations. Hypopygidium densely punc- tate, apical margin nearly flat. Genitalia (Figs 10-13): Paramere tapering apically, curved ventrally and slightly curved later- ally at apex, with dense ventral brush and long curved bristles along inner margin. Cuspis with apical, finger-like process, 0.4X free length of paramere, setose basal- ly, with apical tuft and short, thick bristles on venter of finger-like process. Basal lobe of cuspis short, glabrous. Penis valve unidentate apically, hooked baso-dorsally. Length.—10 mm. 130 Figs 7 —2ie if Ny ( / i) \\ (A MY i : ci Wet’ \ x ye —. 6 uy rf ‘ a = JOURNAL OF HYMENOPTERA RESEARCH Lh fy fer / //, ig | fi /] NU iY Yi > 4, ” f \N 7 ’ Yi \ \ | Ni Hi i MIS A i Ble s , 16 19 Clypeus, Figs. 7-10; Fig. 7: Pseudomethoca ajattara; Fig. 8: P. donaeanae; Fig. 9: P. russeola. Male genitalia: dorsal view, ventral view, lateral view, and penial valve, Figs 10-21: Figs 10-13: P. ajattara; Figs 14-17; P. donaeanae; Figs 18-21: P. russeola. Female—Unknown. Host.—Unknown. Etymology.—From Finnish mythology, Ajattara is an evil forest spirit. Treat as noun in apposition. Distribution.—USA: southeastern Arizona. Holotype-—USA: ARIZONA: Cochise Co., Portal, 8.1X.1974, H. & M. Townes coll. (EMUS). Remarks.—This new species is closely related to P. nigricula Mickel based on the genitalia, which are virtually identical (Figs 10-13; Fig. 6 in Mickel 1924). These species can be separated by setal and integumental coloration; P. nigricula has the integument and setae entirely black, while P. ajattara sp. nov. has the metasomal integument orange and most of the setae pale golden (Fig. 1). Additionally, P. nigri- cula has coarser punctation, especially on the pronotum and genae, where the punc- tures are deep and contiguous. The curved setae on the internal margin of the para- mere of these two species are unique among United States species. In Mexico, however, at least three undescribed species have been examined with this genitalic feature (pers. obs.), which will be described at a later date. Pseudomethoca donaeanae (Cockerell and Fox) (Figs 3, 6, 8, 14-17) Sphaeropthalma dona-anae Cockerell and Fox, 1897: 136. HOLOTYPE 9, USA, New Mexico (ANSP). Mutilla donae-anae Fox, 1899: 224. 9 Pseudomethoca Donae-Anae André, 1903: 28. 9 Pseudomethoca donaeanae Krombein, 1979: 1302. 9 Pseudomethoca donaeanae Manley, 1999: 32. 9 ¢ VOLUME 17, NUMBER 2, 2008 Female diagnosis.—This species can im- mediately be separated from all other known females of North American Pseudo- methoca by the presence of a prominent rugose tubercle on the dorsum of the propodeum medially, although the females of P. ajattara and P. russeola are unknown. Male diagnosis.—The male is similar to P. russeola and P. ajattara sp. nov. in colora- tion, wherein the integument of the head and mesosoma is black, the integument of the metasoma is orange, and the setae are silvery white. This species can be separated from these species by the following com- bination of characters: head broader than pronotum; clypeus with small, widely separated lateral teeth (Fig. 8); fringes of T2-4 with thick, slightly curved, pale silver setae and T5 with simple intermixed black and silver setae (Fig. 3); paramere covered with simple setae only (Fig. 14); and cuspis rectangular (Fig. 14). Male description.—Coloration: Head and mesosoma black or dark reddish-brown; metasoma orange; legs reddish-brown, lighter than head and mesosoma; tibial spurs white; wings slightly infuscated. Setae of head, mesosoma, and legs silvery white, except mesonotum having ap- pressed black setae interspersed with erect white setae. Setae of metasoma entirely silvery white, except T6-7 and disc of T2 having some black setae. Head: Broader than pronotum. Front and gena densely punctate, vertex moderately punctate. Mandible oblique, tridentate apically, inner tooth strongly developed (Fig. 8). Clypeus weakly punctate, flat anteriorly, with two small, sharp, lateral teeth (Fig. 8). Antennal scrobe lacking carina. Ocelli minuscule; ocellocular distance 10X length of lateral ocellus, interocellar distance 3X lateral ocellar length. Flagellomere I 1.5X pedicel length; flagellomere II 2.5X pedicel length. Mesosoma: Pronotum and scutellum dense- ly punctate; mesonotum and mesopleuron moderately punctate; metapleuron gla- brous; propodeum reticulate dorsally, an- terior margin glabrous laterally. Tegula 131 evenly convex, pubescent anteriorly, gla- brous posteriorly. Marginal cell 1.75 length of stigma (Fig. 6). Metasoma: Petiole broadly sessile, evenly convex. Apical fringes of T2-5 and S2-4 forming rows of short, evenly spaced, slightly curved, sil- very white bristles, those of T2-5 separated by the bristle width, those of 52-4 separat- ed by 2 the bristle width (Fig. 3). T1 with elongate shallow punctures; T2 and S2 moderately punctate; T3-6 and S3-6 densely punctate. S1 with low longitudinal carina. Pygidium densely punctate, with micro- punctures and fine setae among punctations. Hypopygidium densely punctate, apical margin slightly convex. Genitalia (Figs 14— 17): Paramere tapering apically, moderately setose throughout, weakly curved ventrally. Cuspis short, 0.25 free length of paramere, rectangular, setose throughout. Basal lobe of cuspis extending beyond anterior margin of cuspis, dorsally curved, glabrous. Penis valve unidentate with ventral lobe apically, hooked basodorsally. Length.—8-9 mm. Host.—Unknown. Distribution.—USA: southern Arizona and New Mexico, southeastern California; MEXICO: northeastern Baja California. Material examined—USA: ARIZONA: Cochise Co., Portal, 1g, 2.IX.1959, H.E. Evans coll. (DGM); Maricopa Co., Granite Reef Dam, 13, 4.X.1964, J.W. Debolt (ASUT); Pinal Co., Sacaton, 13, Geo. Harrison coll. (NMNH); CALIFORNIA: Imperial Co., El Centro, 1¢ 19, 7.VII.1955, A. Ross coll. (EMUS); NEW MEXICO: Dona Ana Co.: Hatch, 2g, 28-29.VII.1974, H. & M. Townes coll. (EMUS); 2 km E Radium Springs, 13, 2.X.1992, D.G. Manley coll. (DGM); Hidalgo Co.: Rodeo, 13, 28.VIII.1959, H.E. Evans coll. (DGM). MEXICO: BAJA CALIFORNIA: Mexicali, 13, 16.V1I.1956 (CSUC); SONORA: 2.6 mi W La Jollita, 14, 21.1X.1967, G. I. Marsh coll. (CISC). Remarks.—The sex association was dis- covered by Manley (1999), when he attract- ed two males to a caged female specimen in New Mexico. He identified the males as P. russeola Mickel, and synonymized the two species. Although this male keys out to 132 P. russeola using Mickel (1924, 1935), it has numerous morphological differences from the type of that species. Most notably, the head is broader than the pronotum (nar- rower in P. russeola), the clypeus is gla- brous anteromedially (Fig. 8) (punctate throughout in P. russeola) (Fig. 9), and metasomal terga two to four have rows of short silver bristles (Fig. 3) (the terga of P. russeola have simple setae only) (Fig. 4). Manley (1999) also suggests that the record of P. donaeanae from Calexico, CA may be based on a mislabelled specimen, and that it was unlikely that P. donaeanae actually lives that far west. A male and female from El Centro, CA and a male from Mexicali, Baja California have been examined, however, and both of these sites are within 15 miles of Calexico. Thus, we believe that the Calexico locality is legiti- mate. This is a relatively uncommon distribution, but many species of Dasymu- tilla Ashmead that are typically recognized from Arizona and New Mexico, have also been found in the western Sonoran Desert in California (Hurd, 1951). Pseudomethoca russeola Mickel (Figs 4, 9, 18-21) Pseudomethoca russeola Mickel, 1924: 44. NEW COMBINATION. Ho.Lotyre J, USA, Texas, San Diego, 4 May 1901, R.A. Cushman coll. (NMNH). Diagnosis.—The male of this species is similar to P. donaeanae and P. ajattara sp. nov. in coloration, wherein the integument of the head and mesosoma is black, the integument of the metasoma is orange, and the setae are silvery white. This species can be separated from these species by the following combi- nation of characters: head narrower than pronotum; clypeus with moderate, separat- ed lateral teeth (Fig. 9); T2-5 with intermixed sparse, simple, black and silver setae (Fig. 4); paramere covered with simple setae only (Fig. 18); and cuspis rectangular (Fig. 18). Additions to male description—Antennal scrobe lacking carina. Ocelli minuscule; JOURNAL OF HYMENOPTERA RESEARCH ocellocular distance 10x length of lateral ocellus, interocellar distance 3X lateral ocellar length. Flagellomere I 1.5x pedicel length; flagellomere II 2.5x pedicel length. Marginal cell of forewing 1.5x length of stigma. First metasomal sternum with low longitudinal carina. Pygidium densely punctate, with micropunctures and fine setae among punctations. Hypopygidium densely punctate, apical margin slightly convex. Genitalia (Figs 18-21): Paramere tapering apically, moderately setose throughout, weakly curved ventrally. Cus- pis short, 0.25x free length of paramere, rectangular, setose throughout. Basal lobe of cuspis extending beyond anterior mar- gin of cuspis, dorsally curved, glabrous. Penis valve unidentate apically, angulate basodorsally. Length.—8-10 mm. Female.—Unknown. Host.—Unknown. Distribution. USA: southern Texas. Material examined—USA: TEXAS: Bexar Co.: Leon Creek, 13, 19.X.1952, M. Wasbauer coll.; Hidalgo Co.: Bentsen Rio Grande State Park, 1d, 27. 1V.1986, W.J. Pulawski coll. (CASC); 54, 27.V.1979, H. Evans, A. Hook & W. Rubick coll. (CSUC); 4¢, 15.V.1979, H. Evans, A. Hook & W. Rubick coll. (CSUC); 1g, 13.V1.1978, C.C. Porter coll. (DGM); Kleberg Co., Route 2045E, 30 mi. E Kingsville, 13, 3.X1.1990, T. Carlow coll. (TAMU). Remarks.—This species seems to be re- stricted to the humid area of southern Texas, and is likely to extend far south into Mexico as well. We did not find any Mexican P. russeola specimens, most likely because few Pseudomethoca were available from that region. Unlike the other species examined in this paper, this species lacks thickened bristles on the metasomal terga, having only simple setae (Fig. 9). ACKNOWLEDGEMENTS We would first like to thank Donald G. Manley at the Pee Dee Research Center for allowing us to borrow voucher specimens from his previous study. We thank the curators of all the museums who provided specimens for this study. KAW was able to search for specimens in Portal, AZ using funds from the VOLUME 17, NUMBER 2, 2008 Theodore Roosevelt Memorial Fund of the American Museum of Natural History. This research was supported by the Utah Agricultural Experiment Station, and Utah State University, Logan, Utah. Approved as journal paper no. 7921. LITERATURE CITED André, E. 1903. Mutillidae. Genera Insectorum 1: 1-77. Cockerell, T. D. A. and W. J. Fox. 1897. Descriptions of New Hymenoptera from New Mexico. Proceed- ings of the Academy of Natural Sciences, Philadelphia 49: 135-141. Fox, W. J. 1899. The North American Mutillidae. Transactions of the American Entomological Society Boe 19 792. Hurd Jr., P. D. 1951. The California Velvet Ants of the Genus Dasymutilla Ashmead (Hymenoptera: Mu- tillidae). Bulletin of the California Inset Survey Vol. 1, 4: 88-114 + 1 plate. 133 Krombein, K. V. 1979. Chapter 76. Mutillidae. Pp. 1276-1313 in: Krombein, K. V., et al., eds. Catalog of Hymenoptera in America North of Mexico vol. 2. Smithsonian Institution Press, Washington, D. C, xvi + 1199-2209 pp. Manley, D. G. 1999. A synonymy for Pseudomethoca donaeanae (Cockerell & Fox) (Hymenoptera: Mu- tillidae). Pan-Pacific Entomologist 75: 32-34. Mickel, C. E. 1924. A revision of the mutillid wasps of the genera Myrmilloides and Pseudomethoca occur- ring in America north of Mexico. Proceedings of the United States National Museum 64: 1-52. . 1935. Descriptions and records of nearctic mutillid wasps of the genera Myrmilloides and Pseudomethoca (Hymenoptera: Mutillidae). Annals of the Entomological Society of America 29: 29-60. Nonveiller, G. 1990. Catalog of the Mutillidae, Myrmosidae and Bradynobaenidae of the Neo- tropical region including Mexico. Hymenopter- orum Catalogus, Pars 18: 1-350. J. HYM. RES. Vol. 17(2), 2008, pp. 134-137 A New Species of Encarsia (Hymenoptera: Aphelinidae) Parasitising Aleuromarginatus tephrosiae (Hemiptera: Aleyrodidae) in Iran and Oman ANDREW POLASZEK AND SHAHAB MANZARI (AP) Dept of Entomology, the Natural History Museum, London SW7 5BD, U.K. (SM) Insect Taxonomy Research Department, Iranian Research Institute of Plant Protection, P.O. Box 1454, Tehran 19395, Iran; email: manzari@ppdri.ac.ir Abstract—Encarsia indigoferae Polaszek & Manzari, new species, is described and illustrated. It is known from Iran and Oman, and all known specimens were reared from the whitefly Aleuromarginatus tephrosiae Corbett. The purpose of this paper is to describe a new species in the genus Encarsia. E. indigoferae is clearly a member of the Encarsia strenua-group, having scutellar sensilla sep- arated by approximately the maximum diameter of one sensillum (see Fig. 4), a characteristic stigmal vein with an asetose area above it, and a seta present at the junction of the marginal and submarginal veins. It differs from other species of the strenua-group in having a combination of three setae on the submarginal vein and a rugose stemmaticum. The host, Aleuromargi- natus tephrosiae Corbett, was described from Sierra Leone (Corbett 1935) and is wide- spread in Africa and Asia, apparently specific to various Papilionaceae (Bink-Moe- nen 1983). It seems probable that E. indigo- ferae is more widespread than is currently known. It is worth mentioning that within the colony of A. tephrosiae on Indigofera sp. collected in Iran, parasitised pupae of Zaphanera cyanotis Corbett with parasitoid emergence holes were also collected. These two whitefly species, which had heavily infested Indigofera sp. in the collecting areas, were found to be mostly parasitised. It is quite likely that Z. cyanotis is also being parasitised by E. indigoferae but no parasitoid was reared from the former species. To whom correspondence should be addressed: E-mail: a.polaszek@nhm.ac.uk Encarsia species are mostly parasitoids of whiteflies and armoured scale insects (Diaspididae), and are of considerable economic importance. The systematics and biology of the genus are treated in detail by Heraty et al. (2008). Abbreviations.— NHM Natural History Museum, Lon- don, U.K. Hayk Mirzayans Insect Muse- um, Iranian Research Institute of Plant Protection, Tehran, TRAN. HMIM Encarsia indigoferae Polaszek & Manzari new species Figs 14 Description.mFemale Colour. Head and body yellow except the following areas pigmented with brown (Fig. 1): three spots on stemmaticum (Fig. 3), adjacent to ocelli; pronotum and front of mesoscutum, notauli (especially poteriorly); most of axillae, but fading posteriorly; T2 and T3-T6 either just later- ally or more extensively. Antennae and legs uniformly pale brown, or appearing paler, almost yellow. Fore wings hyaline. Morphology. Stemmaticum with densely rugose surface sculpture (Fig. 3). Antennal formula 1,1,3,3 (Fig. 2). Pedicel equal in VOLUME 17, NUMBER 2, 2008 Fig. 1. Encarsia indigoferae holotype female, habitus. length to F1 and F2. Flagellomeres with the following numbers of sensilla: F1: 1-2 (1), P22 (yk: 2-3 (2), F4: 3-4 (3), F5: 3-4 (4), F6: 2-3 (3). Midlobe of mesoscutum (Fig. 4) with 12-15 (14) setae arranged symmetrically, side lobes with 3 setae each. Scutellar sensilla close together, separated by a distance of about the width of a 15 sensillum. Distance between anterior pair of scutellar setae smaller than between posterior pair. Fore wing 2.3-2.5 (2.4) times as long as width of disc. Marginal fringe 0.19-0.20 times as long as width of disc (0.16 times in Oman specimens). Submar- ginal vein with 3 setae (4 on one side in one individual), marginal vein anteriorly with 136 JOURNAL OF HYMENOPTERA RESEARCH Fig. 2. Encarsia indigoferae female, antenna. 6-9 setae. Basal cell with 4-7, leading edge of costal cell with 2-4 distinct setae (less conspicuous in Oman specimens). Tarsal formula 5-5-5. Apical spur of midtibia subequal in length to corresponding basi- tarsus. Tergites laterally with the following numbers of setae: T1: 0, T2: 1, T3: 1, T4: 1, T5: 2-3, T6: 2-3, T7 with 5-8 (6) setae. Ovipositor longer than midtibia, 1.22-1.37 (1.22) and 2.4-2.9 (2.4) times as long as clava. Third valvula 0.20-0.21 (0.21) times as long as ovipositor. Male. Morphology as for female, except for antennal and genitalia characters. F5 and F6 apparently partially fused, as in many males of strenua-group species. Pro- notum, mesoscutum anteriorly and cen- trally, axillae and metasoma entirely dark. Species group placement.—E. strenua group. Distribution.—lran, Oman. Fig. 3. Encarsia indigoferae female, stemmaticum. Host.—Aleuromarginatus tephrosiae Corbett. Material studied.mHolotype 9, IRAN: Sistan-Balouchestan, Chabahar, Noban- dian, 28 m. 18.xii.2006, 25° 28’ 54.8” N, 61° 9’ 21.9" E. (S. Manzari, M. Moghaddam & Durbin), ex Aleuromarginatus tephrosiae on Indigofera sp. (HMIM). Paratypes: 159, 10d, same data as holotype (NHM, HMIM); 49, 13, IRAN: Sistan-Balouchestan, Nikshahr, Geshig, 631 m. 20.xii.2006, 26° 18’ 6.1” N, 60° 20’ 3.7” E. (S. Manzari), ex A. tephrosiae on Indigofera sp. (HMIM). OMAN: 39, 1d, Rumais, 18.1ii.1992, ex A. tephrosiae on weed IIE 22998 (NHM, HMIM). Comments.—E. indigoferae is related to E. dialeurodis Hayat from Pakistan, and to the Australian E. oakeyensis Schmidt & Nau- Fig. 4. Encarsia indigoferae female, mesosoma. VOLUME 17, NUMBER 2, 2008 mann. It differs from both these species in having three setae on the submarginal vein. It can be further distinguished from other species in the strenua-group having 3 or more setae on the submarginal vein by the rugose (rather than reticulate or striate) stemmaticum (Fig. 3). ACKNOWLEDGEMENTS We would like to thank N. Shahbazvar for slide mounting the whitefly specimens. 137 LITERATURE CITED Bink-Moenen, R. M. 1983. Revision of the African whiteflies (Aleyrodidae). Monografieén van de Nederlandse Entomologische Vereniging 10: 1-212. Corbett, G. H. 1935. On new Aleurodidae. Annals and Magazine of Natural History 16: 240-252. Heraty, J. M., A. Polaszek, and M. E. Schauff. 2008. Systematics and Biology of Encarsia. Chapter 4, Pp. 71-87 in: Gould, J., K. Hoelmer, and J. Goolsby eds. Classical Biological Control of Bemisia tabaci in the United States. A review of interagency research and implementation. Springer, 343 pp. J. HYM. RES. Vol. 17(2), 2008, pp. 138-156 Taxonomic Status and Location of Type Specimens for Species of Coelinidea Viereck and Sarops Nixon (Hymenoptera: Braconidae: Alysiinae) Described by Garland T. Riegel ROBERT R. KULA Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, c/o National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, MRC-168, Washington, DC 20013-7012; email: Robert.Kula@ars.usda.gov Abstract—The following species of Coelinidea Viereck and Sarops Nixon described by Garland T. Riegel are transferred to other genera resulting in 28 new combinations: Chorebus pallidus (Riegel), n. comb., Coelinius acicula (Riegel), n. comb., Coelinius acontia (Riegel), n. comb., Coelinius alima, (Riegel), n. comb., Coelinius alrutzae (Riegel), n. comb., Coelinius arizona (Riegel), n. comb., Coelinius arnella (Riegel), n. comb., Coelinius bakeri (Riegel), n. comb., Coelinius baldufi (Riegel), n. comb., Coelinius calcara (Riegel), n. comb., Coelinius columbia (Riegel), n. comb., Coelinius crota (Riegel), n. comb., Coelinius dubius (Riegel), n. comb., Coelinius ellenaae (Riegel), n. comb., Coelinius frisoni (Riegel), n. comb., Coelinius garthi (Riegel), n. comb., Coelinius hayesi (Riegel), n. comb., Coelinius jeanae (Riegel), n. comb., Coelinius marki (Riegel), n. comb., Coelinius marylandicus (Riegel), n. comb., Coelinius minnesota (Riegel), n. comb., Coelinius montana (Riegel), n. comb., Coelinius muesebecki (Riegel), n. comb., Coelinius nellae (Riegel), n. comb., Coelinius niobrara (Riegel), n. comb., Coelinius robinae (Riegel), n. comb., Coelinius ruthae (Riegel), n. comb., and Coelinius sommermanae (Riegel), n. comb. Coelinius ohioensis (Riegel, 1982), and Coelinius wheeleri (Riegel, 1982) are new synonys, and the former is designated the senior synonym because the holotype is a female. The holotypes of Coelinidea antha Riegel, Coelinidea arca Riegel, Coelinidea colora Riegel, and Coelinidea coma Riegel, reportedly deposited at the Academy of Natural Sciences, Philadelphia, Pennsylvania, apparently are lost. Therefore, all four names are considered nomina dubia since each species is known only from the holotype, and information Riegel provided in the original descriptions and key to North American species of Coelinidea is not adequate to apply the names unequivocally. The locations of primary and, where applicable, secondary types are indicated for all other species of Coelinidea and Sarops described by Riegel. Coelinius alima, C. markt, C. ohioensis, and C. robinae are first recorded from Quebec, Wyoming, Wisconsin, and Kansas and Missouri, respectively. Alysiinae is a speciose subfamily of koinobiont endoparastioids of cyclorrha- phous Diptera (Wharton 1997). The sub- family consists of Alysiini and Dacnusini with ~1,245 and ~817 described species, respectively, as of mid-November 2007 for Alysiini and mid-June 2008 for Dacnusini (Yu et al. 2005). Most species of Alysiini with known hosts are parasitoids of sa- prophagous flies; as far as is known, all dacnusines are parasitoids of plant-feeding flies (Wharton 1997). The Palearctic species of Dacnusini have been studied extensive- ly, including comprehensive taxonomic revisions (Griffiths 1964, 1966a, 1966b, 1967, 1968a, 1968b, Nixon 1937, 1943, 1944, 1945, 1946, 1948, 1949, 1954). Few taxonomic treatments have been published for Nearctic dacnusines; notable works include Rohwer (1914), Riegel (1950, 1982), Wharton (1994), and Kula and Zolnerowich (2008). Riegel (1982) revised the Nearctic species of Chaenusa Haliday sensu stricto, Chorebidea Viereck, Chorebidella Riegel, Coelinidea Vier- eck, and Sarops Nixon. The revision, based VOLUME 17, NUMBER 2, 2008 largely on Riegel’s doctoral dissertation from 1947, included descriptions of 44 new species, over half of the ~86 described Nearctic species of Dacnusini (Yu et al. 2005). Several taxonomic changes occurred in the 35 years between completion and publication of the revision. Griffiths (1964) provided hypotheses on character polarity for dacnusines and for the most part only recognized groups he considered mono- phyletic. As a result Griffiths (1964) syn- onymized Chorebidea Viereck, 1913 and Chorebidella Riegel, 1950 with Chaenusa Haliday, 1839 (i.e., Chaenusa sensu lato). Further, Griffiths (1964) treated Coelinidea Viereck, 1913 and Polemochartus Schulz, 1911 as subgenera of Coelinius Nees, 1819 (i.e., Coelinius s.l.) and synonymized Sarops Nixon, 1942 with Synelix Forster, 1862. Riegel (1982), in reference to Griffiths (1964), stated that he was “not convinced that certain genera should have been placed in synonymy” and recognized Chorebidea, Coelinidea, and Sarops as valid genera and described four, 32, and four new species in each genus, respectively. Wharton and Austin (1991) agreed with the synonymies in Griffiths (1964) and deter- mined that Lepton Zetterstedt, 1838 has priority over Coelinidea but considered the division of Coelinius into subgenera “‘pre- mature’ because of undescribed “‘interme- diate forms’” from the Oriental Region. However, Wharton and Austin (1991) did not transfer any of the species described in Riegel (1982) to Chaenusa s.l. or Coelintus s.l. Wharton (1994) considered Coelinius s.l. monophyletic based on three putative apomorphies, the presence of an additional tooth between tooth one and two, a laterally compressed gaster, and the exclu- sive use of chloropids as hosts, and on this basis synonymized Sarops with Coelinius rather than following the synonymy of Sarops and Synelix in Griffiths (1964). Kula and Zolnerowich (2008) transferred the species of Chorebidea described in Riegel (1982) to Chaenusa s.l. and returned Chor- ebidella bergi Riegel to Chaenusa s.l. 169 I agree with Wharton (1994) and consid- er Coelinius s.l. monophyletic based on the apomorphies mentioned above. Kula (2006) recovered Coelinius sensu Wharton (1994) as monophyletic in one of two preliminary cladistic analyses for Dacnu- sini based on morphology, but bootstrap support was low. Clades corresponding to Coelinius s.s., Lepton, and Polemochartus were also recovered, with the included species of Coelinius s.s. and Sarops forming a clade, but only Polemochartus had mod- erate bootstrap support. Therefore, I agree with Wharton and Austin (1991) and Wharton (1994) that Coelinius s.l. should not be split into genera or divided into subgenera at this point in time since only Polemochartus is clearly monophyletic. The species of Sarops described in Riegel (1982) were transferred to Coelinius through the synonymy of Coelinius and Sarops in Wharton (1994), but the taxonom- ic placement of each species has not been verified through the examination of holo- types. The species of Coelinidea described in Riegel (1982) have not been transferred to Coelinius s.l., and holotypes should be examined for these species to verify place- ment before they are transferred. Holo- types of several species described in Riegel (1982) are currently housed in repositories other than those indicated in the original descriptions, and four holotypes apparent- ly are lost. Therefore, the primary objec- tives of this study are to (1) verify and update the taxonomic placement of species of Coelinidea and Sarops described in Riegel (1982) and (2) document the location and condition of holotypes for these species since most are known only from the holotype. Additionally, repositories are indicated for all paratypes of the afore- mentioned species, and new distribution records are provided for four species of Coelinius s.l. MATERIALS AND METHODS Specimens were examined using a Leica Wild M10 stereomicroscope with 25x 140 oculars. Images of holotypes were cap- tured using a Microptics digital camera system, and image clarity was enhanced using Adobe Photoshop 9.0. A color image of each holotype was deposited in Morph- bank (collection number 369162), and if a holotype is damaged, its condition is described. Morphological terminology fol- lows Sharkey and Wharton (1997) except as noted below. All species in this study have three major mandibular teeth; the recogni- tion and numbering of teeth follows Whar- ton (2002) and Wharton (1977), respectively. Tooth one is the dorsal tooth, tooth two is the middle tooth, and tooth three is the ventral tooth. In addition to the three major teeth, a smaller tooth is present between tooth two and three in Chorebus pallidus (Riegel), new combination and is referred to as tooth four. Another smaller tooth is present between tooth one and two in all species and is referred to as tooth five. Thus, the numbering of mandibular teeth in this article differs from Kula and Zolnerowich (2008) in that the latter article referred to a smaller additional tooth as tooth four regardless of its position. The apical rim of the clypeus, metapleural rosette, and tuft of curved setae on the metacoxa are as in Kula and Zolnerowich (2008). The material examined sections are formatted as in Kula and Zolnerowich (2008). Exact label data are reported for holotypes; Riegel (1982) provided more extensive type locality information. The following museum codens (Evenhuis and Samuelson 2007) are used to indicate repositories where type specimens of spe- cies Riegel (1982) described in Coelinidea and Sarops are housed currently: Albert J. Cook Arthropod Research Collection, Mi- chigan State University, East Lansing (MSUC); California Academy of Sciences, San Francisco (CAS); Canadian National Collection of Insects, Ottawa (CNC); Cor- nell University Insect Collection, Ithaca, New York (CUIC); Illinois Natural History Survey, Champaign (INHS); Museum of JOURNAL OF HYMENOPTERA RESEARCH Comparative Zoology, Harvard Universi- ty, Cambridge, Massachusetts (MCZ); Smithsonian Institution National Museum of Natural History, Washington, DC (USNM); Snow Entomological Museum, University of Kansas, Lawrence (SEMC); University of Minnesota Insect Collection, Saint Paul (UMSP); and University of Wyoming Insect Museum, Laramie (ESUW). The holotypes of Coelinidea antha Riegel, Coelinidea arca Riegel, Coelinidea colora Riegel, and Coelinidea coma Riegel, reportedly deposited at the Academy of Natural Sciences, Philadelphia, Pennsylva- nia (ANSP), apparently are lost and could not be examined. All other holotypes of species Riegel (1982) described in Coelinidea and Sarops were examined, as were all paratypes except specimens of Coelinius acontia (Riegel), new combination at ESUW, Coelinius marylandicus (Riegel), new combination at MCZ, and Coelinius niobrara (Riegel), new combination at ESUW. In addition to the repositories listed above, new distribution records and syno- nyms were discovered through examina- tion of specimens borrowed from the American Entomological Institute, Gaines- ville, Florida (AEIC) and the Bohart Muse- um of Entomology, University of Califor- nia, Davis (UCDC). Entries with an asterisk are new distribution records. The specific epithets of C. acontia, Coeli- nius alima (Riegel), new combination, Coe- linius arnella (Riegel), new combination, Coelinius calcara (Riegel), new combination and Coelinius crota (Riegel), new combina- tion, show evidence of being derived from Latin or Greek words. However, they cannot be traced in a standard dictionary and do not follow established rules of Latin grammar. Therefore, in accordance with ICZN Article 31.2.2, “the original spelling is...retained [for the aforementioned spe- cific epithets], with gender ending un- changed” since “the evidence of usage is not decisive’ (ICZN 1999). VOLUME 17, NUMBER 2, 2008 RESULTS AND DISCUSSION Chorebus pallidus (Riegel), new combination (Fig. 1) Coelinidea pallida Riegel 1982: 80, 92 [USNM, examined]. Type material.—Holotype male: Top label (white; partially handwritten, partially type- written) = ““Washgtn [;] 30-6 DC’. Second label (white; typewritten) = “Type’’. Third label (white; handwritten) = ““Chaenon [;] pallidus [] 9 Ashm”. Fourth label (brown; partially handwritten, partially typewritten) = ““HOLO- TYPE ¢ [;] Coelinidea [;] pallida [;] Riegel’ (USNM). Paratypes: 1 [sex unknown] USA, WASHINGTON, DC, 30.vi, Collection Ashmead (USNM). Discussion.—Riegel (1982) indicated that the holotype and paratype of C. pallidus were deposited at USNM. The holotype bears a glass vial with a cork cap between the third and fourth labels. The vial contains the posterior portion of the meta- soma. The antennae are broken, as is the tarsus of one prothroacic leg. One meso- thoracic leg is missing except for the coxa and trochanter; the other is broken at the coxa. One metathoracic leg is broken at the coxa. Hither a meso- or metathoracic leg that has broken off of the specimen at the trochanter is glued to the point. The mandible of C. pallidus has five teeth, with tooth four between tooth two and three and tooth five between tooth one and two. Coelinius s.l. is partially defined by the presence of a small tooth between tooth one and two; Chorebus is partially defined by the presence of a small tooth between tooth two and three. Thus, the mandible is intermediate between Chorebus and Coeli- nius s.1. However, C. pallidus has a complete metapleural rosette, a narrow, smooth sternaulus, and a tuft of curved setae on the metacoxa, features Griffiths (1968b) used to define the Chorebus affinis-group but not found among species of Coelinius s.l. Additionally, the apical rim of the clypeus is present in C. pallidus, and t2 is 141 smooth. The apical rim is present in all species of Chorebus, and t2 is smooth in most species. The apical rim is present in species of Coelinius s.l. that fit Coelinius s.s. (Griffiths 1964, Riegel 1982) and Sarops sensu Riegel (1982) and Maeté (1983), but t2 is striate. Therefore, C. pallidus is transferred to Chorebus because it fits the genus aside from the additional tooth between tooth one and two and otherwise fits the affinis-group. Coelinius acicula (Riegel), new combination (Fig. 2) Coelinidea acicula Riegel 1982: 82, 109 [SEMC, examined]. Type material—Holotype male: Top label (white; typewritten) = ‘Northgate Colo [;] 8- 20-31 [;] L. D. Anderson”. Bottom label (brown; partially handwritten, partially typewritten) = “HOLOTYPE <¢ [;] Coelinidea [;] acicula [;] Riegel” (SEMC). Discussion.—Coelinius acicula is known only from the holotype, which Riegel (1982) indicated was deposited at the “University of Kansas...(KU).’”” The holo- type bears a glass vial with a cork cap between the top and bottom labels. The vial contains the posterior portion of the metasoma. One antenna is broken. Coelinius acontia (Riegel), new combination (Fig. 3) Coelinidea acontia Riegel 1982: 81, 102 [INHS, examined]. Type material.—Holotype female: Top label (white; typewritten) = ““INHS [;] Insect Collec- tion [;] 213,095’’. Second label (white; handwrit- ten) = “Albany Co., Wyo” [;] July 11, 1944 [;] R. E. Pfadt’’. Third label (brown; partially hand- written, partially typewritten) = ““HOLOTYPE 9 [;] Coelinidea [;] acontia [;] Riegel’ (INHS). Paratypes: 1 9 same data as holotype (ESUW); USA, WYOMING: 1 3 Goshen Co., 21.vii.1944, R. E. Pfadt, INHS Insect Collection 213,096 (INHS); 1 3 Platte Co., 13.vii.1944, R. E. Pfadt (ESUW). Discussion.—Riegel (1982) indicated that the holotype and all paratypes of C. acontia JOURNAL OF HYMENOPTERA RESEARCH cf : | =. i Figs 1-6. Holotypes of species of Coelinidea and Sarops described in Riegel (1982) with current taxonomic affiliations. 1, Chorebus pallidus. 2, Coelinius acicula. 3, Coelinius acontia. 4, Coelinius alberta. 5, Coelinius alima. 6, Coelinius alrutzae. VOLUME 17, NUMBER 2, 2008 were deposited at the ‘University of Wyoming...(WYO).” Scott Shaw (in litt.) confirmed that ESUW has two paratypes, but INHS currently has the holotype and a paratype. The holotype bears a glass vial with a cork cap below the third label. The vial contains the posterior portion of the metasoma. The antennae are broken, as is the tarsus of one pro- and mesothoracic leg. One metatarsus is broken; the other is missing. Coelinius alberta (Riegel) (Fig. 4) Sarops alberta Riegel 1982: 56, 57 [CNC, exam- ined]. Coelinius alberta: Wharton 1994: 631 [synonymy of Coelinius and Sarops]. Type material.—Holotype female: Top label (white; partially handwritten, partially type- written) = “Banff, Alta. [;] 16 vi.1922 [;] C. B. D. Garrett”. Second label (yellow; handwritten) = “wing [;] on slide’. Third label (brown; partially handwritten, partially typewritten) = “HOLOTYPE 9 [;] Sarops [;] alberta [;] Riegel’. Fourth label (red; partially handwritten, partial- ly typewritten) = ‘““HOLOTYPE [;] 21202 [;] CNC No.” (CNC). Discussion.—Coelinius alberta is known only from the holotype, which Riegel (1982) indicated was deposited at the “Canadian Department of Agriculture, G. S. Walley (GSW).” The holotype bears a glass vial with a cork cap between the top and second labels. The vial contains some debris but otherwise appears to be empty. The head has been broken off of the specimen and is glued to the pin. The antennae are broken, as are the tarsi except for one mesotarsus. One forewing is mounted on a slide. Coelinius alima (Riegel), new combination (Fig. 5) Coelinidea alima Riegel 1982: 85, 143 [INHS, examined]. Type material_—Holotype female: Top label (white; typewritten) = ‘INHS [;] Insect Collec- 143 tion [;] 201,193’. Second label (white; typewrit- ten) = “Fox Lake, Ill.” [;] June 3, 1943 [;] Ross&Sanderson’’. Third label (brown; partially handwritten, partially typewritten) = ‘““HOLO- TYPE 9 [;] Coelinidea [;] alima [;] Riegel’ (INHS). Paratypes: 1 3 same data as holotype except INHS Insect Collection 201,194 (INHS). Other material examined—*CANADA: QUE- BEC: 1 9 2 § Témiscamingue, Laniel, 8.vii.1944, A. R. Brooks (CNC). Discussion.—Riegel (1982) indicated that the holotype and paratype of C. alima were deposited at INHS. A glass vial with a cork cap was associated with the holotype at some point in time. The cap is still associ- ated, but apparently the vial has been lost. The specimens from CNC expand the range of this species to southeastern Quebec. Coelinius alrutzae (Riegel), new combination (Fig. 6) Coelinidea alrutzae Riegel 1982: 83, 119 [INHS, examined]. Type material.—Holotype male: Top label (white; typewritten) = ““INHS [;] Insect Collec- tion [;] 201,195’. Second label (white; partially handwritten, partially typewritten) = “Algon- quin, Ill. [;] June 15 ’08 [;] Nason 192”. Third label (brown; partially handwritten, partially typewritten) = “HOLOTYPE 4¢ [;] Coelinidea [;] alrutzae [;] Riegel’ (INHS). Discussion.—Coelinius alrutzae is known only from the holotype, which Riegel (1982) indicated was deposited at the ‘University of Illinois...(UILL).”” The University of Illinois insect collection was transferred to INHS in 1979 (P. Tinerella in litt.), and thus, Riegel apparently deposited the holotype in INHS. The holotype bears a glass vial witha cork cap below the third label. The vial contains the posterior portion of the meta- soma. One antenna is broken. One pro- throacic leg is missing except for the coxa; the other has a broken tarsus. Coelinius arizona (Riegel), new combination (Fig. 7) Coelinidea arizona Riegel 1982: 81, 107 [USNM, examined]. 144 Type material—Holotype male: Top label (white; partially handwritten, partially type- written) = ““Huachuca Mts. [;] Ariz., 4-14 1938 [;] R. H. Crandall’’. Second label (white; handwritten) = ‘171’. Third label (white; partially handwritten, partially typewritten) = “Coelinidea [;] sp. [;] det [;] Mues’’. Fourth label (brown; partially handwritten, partially type- written) = “HOLOTYPE ¢ [;] Coelinidea [;] arizona [;] Riegel’ (USNM). Discussion.—Coelinius arizona is known only from the holotype, which Riegel (1982) indicated was deposited at the “University of Arizona...(ARIZ).”” The ho- lotype was transferred to USNM in 1999 (D. Furth in litt.). The holotype bears a glass vial with a cork cap below the third label. The vial contains the posterior portion of the metasoma. The antennae are broken. One prothoracic leg is imbed- ded in glue; the other has a broken tarsus. One mesothoracic leg has a broken tarsus; the other is missing except for the coxa, trochanter, and trochantellus, as is the case with one metathoracic leg. One forewing is missing; the other wings are torn and missing distally. Coelinius arnella (Riegel), new combination (Fig. 8) Coelinidea arnella Riegel 1982: 82, 114 [INHS, examined]. Type material—Holotype male: Top label (white; typewritten) = “INHS [;] Insect Collec- tion [;] 201,099”. Second label (white; partially handwritten, partially typewritten) = ‘Mont. Exp. Sta. [;] Sidney, Mont. [;] June 14, 1913”. Third label (brown; partially handwritten, par- tially typewritten) = “HOLOTYPE ¢ [;] Coeli- nidea [;] arnella [;] Riegel’. Fourth label (white; handwritten) = “[MON]”. Fifth label (white; typewritten) = “INHS [;] TYPE [;] #2030” (INHS). Discussion.—Coelinius arnella is known only from the holotype. Riegel (1982) indicated that the holotype was deposited at ‘‘Montana State College...(MONT),” but INHS currently has the holotype. The holotype bears a glass vial with a cork JOURNAL OF HYMENOPTERA RESEARCH cap below the fifth label. The vial contains the posterior portion of the metasoma. The antennae are broken, and the forewings are torn. Coelinius bakeri (Riegel), new combination (Fig. 9) Coelinidea bakeri Riegel 1982: 83, 123 [USNM, examined]. Type material.—Holotype male: Top label (white; partially handwritten, partially type- written) = “Colo [;] 1563’. Second label (white; typewritten) = ‘Collection [;] CFBaker’’. Third label (brown; partially handwritten, partially typewritten) = “HOLOTYPE @ [;] Coelinidea [;] bakeri [;] Riegel’ (USNM). Other material examined—USA, COLORADO: 3 Q Larimer Co., Estes Park, 5.viii.1947, L. D. Beamer (SEMC). Discussion.—Coelinius bakeri was previ- ously known only from the holotype, which Riegel (1982) indicated was depos- ited at USNM. The holotype bears a glass vial with a cork cap between the second and third labels. The vial contains the posterior portion of the metasoma. The antennae are broken. Riegel (1982) indicat- ed that the holotype was collected in Fort Collins, Larimer County, Colorado. Coelinius baldufi (Riegel), new combination (Fig. 10) Coelinidea baldufi Riegel 1982: 79, 86 [SEMC, examined]. Type material—Holotype male: Top label (white; typewritten) = “Little Beaver Cr. [j] Colo 7 - 11 - 37 [;] C. L. Johnston’’. Second label (white; typewritten) = “Wing on [;] Sl. No.”. Third label (brown; partially handwritten, par- tially typewritten) = ‘““HOLOTYPE ¢ [;] Coeli- nidea [;] baldufi [;] Riegel’’ (SEMC). Discussion.—Coelinius baldufi is known only from the holotype, which Riegel (1982) indicated was deposited at “KU.” The holotype bears a glass vial with a cork cap between the top and second labels. The vial contains the posterior portion of the meta- soma. One forewing is mounted on a slide. VOLUME 17, NUMBER 2, 2008 145 sane nn Na eat et i. os. % % : % & ae z * ¢ % 1 3 a Figs 7-13. Holotypes of species of Coelinidea described in Riegel (1982) with current taxonomic affiliations. 7, Coelinius arizona. 8, Coelinius arnella. 9, Coelinius bakeri. 10, Coelinius baldufi. 11, Coelinius calcara. 12, Coelinius columbia. 13, Coelinius crota. 146 Coelinius calcara (Riegel), new combination (Fig. 11) Coelinidea calcara Riegel 1982: 81, 106 [CAS, examined]. Type material—Holotype female: Top label (white; typewritten) = “Sparks Nev. [;] June 28 1927’. Second label (white; typewritten) = “EPVanDunzee [;] Collector’’. Third label (brown; partially handwritten, partially type- written) = HOLOTYPE 9 [;] Coelinidea [;] calcara [;] Riegel’. Fourth label (white; partially handwritten, partially typewritten) = “Califor- nia Academy [;] of Sciences [;] Type 16687 [;] No.” (CAS). Paratypes: 1 g same data as holotype; USA, CALIFORNIA: 1 9 Inyo Co., Lone Pine, 10.vii.1929, R. L. Usinger; 1 9 San Diego Co., Pine Valley, 24.iv.1920, E. P. VanDu- zee; 1 § same data as previous except W. M. Giffard; 1 3 same data as previous except E. P. VanDuzee, Coelinidea calcara Riegel, 16687 (CAS). Discussion.—Riegel (1982) indicated that the holotype and all paratypes of C. calcara were deposited at the “California Acade- my of Sciences...(CALAS).” The holotype bears a glass vial with a cork cap between the second and third labels. The vial contains the posterior portion of the meta- soma. One antenna is broken. Coelinius columbia (Riegel), new combination (ei Coelinidea columbia Riegel 1982: 85, 144 [CUIC, examined]. Type material.—Holotype female: Top label (white; typewritten) = ‘““Columbia, Mo. [;] May 26-June 8, ’06. [;] C.R. Crosby Coll.”. Second label (brown; partially handwritten, partially typewritten) = “HOLOTYPE 9 [;] Coelinidea [;] columbia [;] Riegel’. Third label (red; partially handwritten, partially typewritten) = ““HOLO- TYPE [;] Cornell U. [;] No. 6491” (CUIC). Paratypes: USA, NEW YORK: 1 9 Tompkins Co., McLean, 2.vii.-3.vii.1904, PARATYPE, Cor- nell U., No. 6491 (CUIC). Discussion.—Riegel (1982) indicated that the holotype and paratype of C. columbia were deposited at “Cornell Universi- ty...(CN).”” The holotype bears a glass vial JOURNAL OF HYMENOPTERA RESEARCH with a cork cap between the top and second labels. The vial contains the poste- rior portion of the metasoma. One antenna is broken. Coelinius crota (Riegel), new combination (Fig. 13) Coelinidea crota Riegel 1982: 79, 90 [INHS, examined]. Type material.—Holotype male: Top label (white; typewritten) = ““INHS [;] Insect Collec- tion [;] 201,196”. Second label (white; typewrit- ten) = “Apple Riv. Can. S.P. [;] l., Aug. 23, 1939 [;] Ross & Riegel’. Third label (brown; partially handwritten, partially typewritten) = HOLOTYPE ¢ [;] Coelinidea [;] crota [;] Riegel” (INHS). Discussion.—Coelinius crota is known only from the holotype, which Riegel (1982) indicated was deposited at INHS. The holotype bears a glass vial with a cork cap below the third label. The vial contains the posterior portion of the metasoma. Coelinius dreisbachi (Riegel) (Fig. 14) Sarops dreisbachi Riegel 1982: 57, 60 [MSUC, examined]. Coelinius dreisbachi: Wharton 1994: 631 [synony- my of Coelinius and Sarops]. Type material_—Holotype male: Top label (white; partially handwritten, partially type- written) = ““Midland Co., Mich. [;] 5-21-42 [;] R. R. Dreisbach”’. Second label (yellow; hand- written) = “wing [;] on slide’. Third label (brown, partially handwritten, partially type- written) = “HOLOTYPE 4¢ [;] Sarops [;] dreis- bachi [;] Riegel’ (MSUC). Discussion.—Coelinius dreisbachi is known only from the holotype. Riegel (1982) indicated that ““R. R. Dreisbach (DREI)” either loaned or donated the holotype but did not specify where the holotype was deposited. The holotype is currently housed in MSUC, presumably donated after Dreis- bach’s death, and bears a glass vial with a cork cap between the top and second label. The vial contains the posterior portion of the VOLUME 17, NUMBER 2, 2008 147 es N > \ \ \18 J 19 Figs 14-19. Holotypes of species of Coelinidea and Sarops described in Riegel (1982) with current taxonomic affiliations. 14, Coelinius dreisbachi. 15, Coelinius dubius. 16, Coelinius ellenaae. 17, Coelinius frisoni. 18, Coelinius garthi. 19, Coelinius hayesi. metasoma. One antenna is broken. One forewing is mounted on a slide, and a forewing and hind wing are torn. Coelinius dubius (Riegel), new combination (Fig. 15) Coelinidea dubia Riegel 1982: 84, 129 [INHS, examined]. Type material.—Holotype male: Top label (white; typewritten) = “INHS [;] Insect Collection [;] 201,197’". Second label (white; typewritten) = “3326”. Third label (white; typewritten) = ‘“Ash- mead [;] Det. ’99’’. Fourth label (brown; partially handwritten, partially typewritten) = HOLO- TYPE ¢ [;] Coelinidea [;] dubia [;] Riegel’ (INHS). Discussion.—Coelinius dubius is known only from the holotype, which Riegel is (1982) indicated was deposited at INHS. The holotype bears a glass vial with a cork cap below the fourth label. The vial contains the posterior portion of the meta- soma. The antennae are broken. Coelinius ellenaae (Riegel), new combination (Fig. 16) Coelimidea ellenage Riegel 1982: 81, 105 [SEMC, examined}. Type material—Holotype male: Top label (white; typewritten) = “Little Beaver Cr.” [j] Colo 7 - 11 - 37 [;] C. L_ Johnston”. Bottom label (brown; partially —— type- written) = “HOLOTYPE 3 [;] Coelinides [] ellenaae [;] Riegel” (SEMO). Discussion—Coelinius ellenaae is known only from the holotype, which Riegel (1982) indicated was deposited at “KU.” The holotype bears a glass vial with a cork cap between the top and bottom labels. The vial contains the posterior portion of the metasoma. One antenna is broken. Coelinius frisoni (Riegel), new combination (Fig. 17) Coelmmidea frisoni Riegel 1982: 83, 117 [INHS, examined]. Type maierial—Holotype female: Top label (white; typewritten) = “INHS [;] Insect Collec- tion [;] 201,198”. Second label (white; typewrit- ten) = “Empire, Colo. [7] July 23, 1938 [;] HH. & J.A. Ross”. Third label (brown; partially hand- writien, partially typewritten) = “HOLOTYPE ° [;] Coelinidea [;] frisoni [;] Riegel” (INHS). Discussion —Coelinius frisoni is known only from the holotype, which Riegel (1982) indicated was deposited at INHS. The holotype bears a glass vial with a cork cap below the third label. The vial contains the posterior portion of the metasoma. One antenna is broken. Coelinius garthi (Riegel), new combination (Fig. 18) Coelmmdea garthi Riegel 1982: 84, 130 [INHS, examined }. JOURNAL OF HYMENOPTERA RESEARCH Type material—Holotype male: Top label (white; t : = “INHS [7] Insect Collec- tion [;] 201,199"". Second label (white; typewzit- ten) = “New Milford, Il [;] July 2, 1936 [;] Ross & Parks”. Teed Beet Gees, pee See ten, partially typewritten) = “HOLOTYPE ¢ [] Coelmidea [;] garthi [;] Riegel”. Fourth label (white; handwnitten) “Freak [;] wmeg” (INHS). Discussion.—Coelinius garthi is known only from the holotype, which Riegel (1982) indicated was deposited at INHS. The holotype bears a glass vial with a cork cap below the fourth label. The vial contains the posterior portion of the meta- soma. The antennae are broken. Coelinius hayesi (Riegel), new combination (Fig. 19) Coelimidea hayesi Riegel 1982: 83, 121 [USNM, examined]. Type material —Holotype male- Top label (white; “Colo [;] 1233”. Second label (white; handwitten) = “31”. Third label (white; typewnitten) = “Collection [;] ee Fourth label (white; rs i partially typewritten) = “HOLOTYPE 3 Ll Coelinidea [;] hayesi [;] Riegel” (USNM). Discussion—Coelinius hayesit is known only from the holotype, which Riegel (1982) indicated was deposited at USNM. The holotype bears a glass vial with a cork cap between the fourth and fifth labels. The vial contains the posterior portion of the metasoma. One antenna is broken at the pedicel; the flagellum of that antenna is stuck to the other antenna. Coelinius jeanae (Riegel), new combination (Fig. 20) Coelimidea jeanae Riegel 1982: 79, 87 [INHS, examined]. Type material—Holotype male: Top label (white; typewritten) = “INHS [;] Insect Collec- tion [;] 201,200". Second label (white; — handwritien, partially typewritten) = “Green Mit Falls, [;] Colo. jn 21, 1938 [;] J-A. Ross”. Third label (brown; partially handwritten, par- VOLUME 17, NUMBER 2, 2008 149 & Figs 20-26. Holotypes of species of Coelinidea described in Riegel (1982) with current taxonomic affiliations. 20, Coelinius jeanae. 21, Coelinius marki. 22, Coelinius marylandicus. 23, Coelinius minnesota. 24, Coelinius montana. 25, Coelinius muesebecki. 26, Coelinius nellae. 150 tially typewritten) = “HOLOTYPE ¢ [;] Coeli- nidea [;] jeanae [;] Riegel’ (INHS). Paratypes: 1 3 same data as holotype except 17.vii.1938, H. H. & J. A. Ross (INHS); 1 § USA, COLORADO: Great Sand Dunes near Botger C.R., 23.vi.1944 (USNM). Discussion.—Riegel (1982) indicated that the holotype and a paratype of C. jeanae were deposited at INHS and that a paratype was deposited at USNM. The holotype bears a glass vial with a cork cap below the third label. The vial contains the posterior portion of the metasoma. The antennae are broken, as is the tarsus of one prothroacic leg. Coelinius marki (Riegel), new combination (Fig. 21) Coelinidea marki Riegel 1982: 80, 97 [INHS, examined]. Type material.—Holotype female: Top label (white; typewritten) = ‘“INHS [;] Insect Collec- tion [;] 221,941”. Second label (white; partially handwritten, partially typewritten) = ‘Green Mt. Falls, [;] Colo., Jly. 17, 1938 [;] J. A. Ross & [;] H. H. Ross”. Third label (brown; partially handwritten, partially typewritten) = ‘““HOLO- TYPE 9Q [;] Coelinidea [;] marki [;] Riegel” (INHS). Other material examined —USA, *WYOMING: 1 Q 1 [sex unknown] Big Horn Co., Cowley, 8.viii.1935, in wheat stem (ESUW). Discussion.—Coelinius marki was _ previ- ously known only from the holotype, which Riegel (1982) indicated was depos- ited at INHS. The holotype bears a glass vial with a cork cap below the third label. The vial contains the posterior portion of the metasoma. The specimens from ESUW expand the range of this species to north- western Wyoming. Coelinius marylandicus (Riegel), new combination (Fig. 22) Coelinidea marylandica Riegel 1982: 83, 125 [USNM, examined]. Type material.—Holotype female: Top label (white; typewritten) = ‘“Md.”. Second label (white; typewritten) = ‘Collection [;] Ash- JOURNAL OF HYMENOPTERA RESEARCH mead”. Third label (brown; partially handwrit- ten, partially typewritten) = “HOLOTYPE 9 [;] Coelinidea [;] marylandica [;] Riegel’”” (USNM). Paratypes: 1 9 same data as holotype; 1 9 same data as holotype except Coelinius marylandicus Ashm, Coelinidea n. sp. Mues.; USA, TENNES- SEE: 1 ¢ Middle Tennessee, Cedar Glade Area, 9.xi, Adelphia Meyer, sweep net; 1 9 same data as previous except 524, Coelinidea n. sp. det. Mues. (USNM); USA, VIRGINIA: 1 9 Falls Church, 20.v, Nathan Banks (MCZ). Discussion.—Riegel (1982) indicated that the holotype and four paratypes of C. marylandicus were deposited at USNM and a paratype was deposited at MCZ. The paratypes at USNM were examined, and Stefan Cover (in litt.) confirmed that the other paratype is at MCZ. The holotype bears a glass vial with a cork cap between the second and third labels. The vial contains the posterior portion of the metasoma. The head is missing, as are the mesothoracic legs except for the coxae. One prothoracic leg is missing except for the coxa; the other has a broken tarsus. One forewing is torn. Coelinius minnesota (Riegel), new combination (Fig. 23) Coelinidea minnesota Riegel 1982: 84, 131 [UMSP, examined]. Type material—Holotype female: Top label (white; typewritten) = “Minneapolis, Minn. [;] Excelsior Blvd. [;] Aug. 13, 1927 [;] A. T. Hertig”. Bottom label (brown; partially handwritten, partially typewritten) = “HOLOTYPE 9 [)] Coelinidea [;] minnesota [;] Riegel’ (UMSP). Paratypes: 1 g same data as holotype (UMSP). Discussion.—Riegel (1982) indicated that the holotype and paratype of C. minnesota were deposited at the ‘University of Minnesota...(MINN).”” The holotype bears a glass vial with a cork cap between the top and bottom labels. The vial contains the posterior portion of the metasoma. Coelinius montana (Riegel), new combination (Fig. 24) Coelinidea montana Riegel 1982: 82, 111 [SEMC, examined]. VOLUME 17, NUMBER 2, 2008 Type material_—Holotype male: Top label (white; typewritten) = ‘Bennett Montana [;] 8- 12-31 [;] J Nottingham’’. Bottom label (brown; partially handwritten, partially typewritten) = “HOLOTYPE ¢ [;] Coelinidea [;] montana [;] Riegel’ (SEMC). Discussion.—Coelinius montana is known only from the holotype, which Riegel (1982) indicated was deposited at “KU.” The holotype bears a glass vial with a cork cap between the top and bottom labels. The vial contains the posterior portion of the metasoma. The head is missing. Coelinius muesebecki (Riegel), new combination (Fig. 25) Coelinidea muesebecki Riegel 1982: 84, 141 [INHS, examined]. Type material.—Holotype female: Top label (white; typewritten) = “INHS [;] Insect Collec- tion [;] 221,942’’. Second label (white; handwrit- ten) = “Ripley, Il. [;] Sept. 1, 1939 [;] Ross & Riegel’. Third label (brown; partially handwrit- ten, partially typewritten) = ““HOLOTYPE 9 [;] Coelinidea [;] muesebecki [;] Riegel’”” (INHS). Discussion.—Coelinius muesebecki is known only from the holotype, which Riegel (1982) indicated was deposited at INHS. The holotype bears a glass vial with a cork cap below the third label. The vial contains the posterior portion of the metasoma. Coelinius nellae (Riegel), new combination (Fig. 26) Coelinidea nellae Riegel 1982: 83, 116 [SEMC, examined]. Type material.—Holotype male: Top label (white; partially handwritten, partially typewrit- ten) = “Summit Co., Ohio [;] 6-9 1937 [;] Louis J. Lipovsky”’. Bottom label (brown; partially hand- written, partially typewritten) = ““HOLOTYPE ¢ [;] Coelinidea [;] nellae [;] Riegel’” (SEMC). Discussion.—Coelinius nellae is known only from the holotype, which Riegel (1982) indicated was deposited at “KU.” The holotype bears a glass vial with a partially broken cork cap between the top and bottom labels. The vial contains the 151 posterior portion of the metasoma. One antenna is broken. Coelinius niobrara (Riegel), new combination (Fig. 27) Coelinidea niobrara Riegel 1982: 83, 122 [INHS, examined]. Type material—Holotype male: Top label (white; typewritten) = “INHS [;] Insect Collec- tion [;] 213,097’. Second label (white; partially handwritten, partially typewritten) = “Nio- brara Co. Wyo. Stop [;] July 1, 1943 [;] Collected by R.E. Pfadt’’. Third label (white; handwritten) = “LWYO”. Fourth label (brown; partially handwritten, partially typewritten) = ‘“HOLO- TYPE ¢ [;] Coelinidea [;] niobrara [;] Riegel” (INHS). Paratypes: 1 3 same data as holotype except INHS Insect Collection 213,098 (INHS); 2 3d same data as holotype (ESUW). Discussion.—Riegel (1982) indicated that the holotype and all paratypes of C. niobrara were deposited at “WYO.” Scott Shaw (in litt.) confirmed that ESUW has two para- types, but the INHS has the holotype and a paratype. The holotype bears a glass vial with a cork cap below the third label. The vial contains the posterior portion of the metasoma. One antenna is broken, as is the tarsus of one metathoracic leg. Coelinius ohioensis (Riegel) (Figs 28, 29) Sarops ohioensis Riegel 1982: 56, 58 [SEMC, examined]. Coelinius ohioensis: Wharton 1994: 631 [synony- my of Coelinius and Sarops]. Sarops wheeleri Riegel 1982: 56, 59 [INHS, examined]. Coelinius wheeleri: Wharton 1994: 631 [synonymy of Coelinius and Sarops]. New synonym. Type material.—Holotype female, Sarops ohtoensis: Top label (white; partially handwritten, partially typewritten) = “Summit Co., Ohio [;] 9-1 1937 [;] Louis J. Lipovsky’”’. Second label (yellow; hand- written) = ‘Wings [;] on slide’. Third label (brown; partially handwritten, partially typewrit- ten) = “HOLOTYPE ¢ [;] Sarops [;] ohioensis [;] Riegel” (SEMC). Holotype male, Sarops wheeleri: Top label (white; typewritten) = “INHS [;] Insect JOURNAL OF HYMENOPTERA RESEARCH Figs 27-32. Holotypes of species of Coelinidea and Sarops described in Riegel (1982) with current taxonomic affiliations. 27, Coelinius niobrara. 28, Coelinius ohioensis. 29, Coelinius wheeleri. 30, Coelinius robinae. 31, Coelinius ruthae. 32, Coelinius sommermanae. Collection [;] 212,943”. Second label (white; partially handwritten, partially typewritten) = “Contoocook [;] N.H. ix-4-’21 [;] E.W. Hall’. Third label (brown; partially handwritten, partially typewritten) = “HOLOTYPE ¢ [;] Sarops [;] wheeleri [;] Riegel’. Fourth label (yellow; hand- written) = “wings [;] on slide” (INHS). Other material examined.—All USA, *WIS- CONSIN; 3 9 Fond du Lac Co., T13N R19E $23, 4.ix.1975, gypsy moth Malaise trap; 1 9 same data as previous except 4.1x.-9.ix.1975; 1 9 1 $ same data as previous except 9.1x.-2.x.1975; 191 ¢ Jackson Co., T2IN R4W 9827, 15.ix.- 22.ix.1975, gypsy moth Malaise trap; 1 9 Oneida VOLUME 17, NUMBER 2, 2008 Co., T35N R11E $17, 12.viii.-25.viii.1975, gypsy moth Malaise trap (AEIC). Discussion.—The holotypes of C. ohioensis (Riegel, 1982) (Fig. 28) and C. wheeleri (Riegel, 1982) (Fig. 29) fit within a mor- phospecies series of seven females and two males from Wisconsin and are simply conspecific female and male specimens, respectively. The species are not referenced in the literature beyond Riegel (1982). Therefore, C. ohioensis is designated the senior synonym because the holotype is a female; in dacnusines females usually have a greater number of diagnostic features compared to males. Coelinius ohioensis and C. wheeleri were described from the holotypes only, which Riegel (1982) indicated were deposited at “KU” and INHS, respectively. The holo- type of C. ohioensis bears a glass vial with a cork cap between the top and second labels. The holotype of C. wheeleri bears a glass vial with a cork cap below the fourth label. The vials contain the posterior portion of the metasoma of each species. One antenna of the C. ohioensis holotype is broken, and one forewing is mounted on a slide. The antennae of the C. wheeleri holotype are broken, as is the tarsus of one meso- and metathoracic leg. The other mesotarsus is missing. One forewing is mounted on a slide. The specimens from AEIC expand the range of this species to northern and central Wisconsin. Coelinius robinae (Riegel), new combination (Fig. 30) Coelinidea robinae Riegel 1982: 83, 118 [USNM, examined]. Type material.—Holotype male: Top label (white; partially handwritten, partially type- written) = “Harry L. Johnson [;] 8-14-1940 [;] So. Meriden Conn.”’. Second label (white; partially handwritten, partially typewritten) = Coelinidea [;] sp. [;] det [;] Mues’’. Third label (brown; partially handwritten, partially type- written) = “HOLOTYPE 4 [;] Coelinidea [;] robinae [;] Riegel’ (USNM). 153 Other material examined.—USA, *KANSAS: 1 § Douglas Co., 23.v.1941, D. E. Hardy;195 3 same data as previous except 20.vi.1945, R. H. Beamer; 2 9 7 3 same data as previous except 21.vi.1945; 2 9 same data as previous except 25.vi.1945 (SEMC); 191 3 Riley Co., Konza Prairie Biological Station, watershed N2B, 39°05.27'N 96°35.09'W, 25.v.-27.v.2001, Zolner- owich, Kula, Brown, Malaise trap; 2 9 1 J same data as previous except 1.vi.-8.vi.2001; 1 9 same data as previous except 8.vi-12.vi.2001; 3 g same data as previous except watershed 4F, 39°04.37'N 96°34.26'W, 1.vi.-8.vi.2001; 5 J same data as previous except 8.vi.-12.vi.2001; 8 3 same data as previous except 15.vi.-19.vi.2001; 1 Q same data as previous except 19.vi.-22.vi.2001; 5 § same data as previous except 22.vi.- 26.vi.2001; 1 g¢ same data as previous except watershed 4B, 39°04.65'N 96°35.75'W, 12.vi.- 15.vi.2001; 1 g same data as previous except 22.v1.-26.vi.2001 (KSUC); *MISSOURI: 2 3 Boone Co., Columbia, 7.vi.1970, Malaise trap (UCDC). Discussion.—Coelinius robinae was previ- ously known only from the holotype, which Riegel (1982) indicated was depos- ited at the “Connecticut Agricultural Ex- periment Station...(CONN).” All types in the Connecticut Agricultural Experiment Station collection were transferred to USNM in 1962 (G. Ridge in litt.), and thus, Riegel apparently deposited the holotype in USNM. The metasoma is intact unlike holotypes of other species described in Riegel (1982). Thus, a glass vial is not associated with the specimen. The meso- thoracic legs are missing, as is one meta- thoracic leg, one forewing, and one hind wing. One prothoracic leg is broken at the trochanter; the rest of the leg, except for the trochantellus, is glued to the card. The specimens from UCDC and KSUC expand the range of this species to central Missouri and northeastern Kansas, respectively. Coelinius ruthae (Riegel), new combination (Fig. 31) Coelinidea ruthae Riegel 1982: 79, 85 [SEMC, examined]. Type material.—Holotype male: Top label (white; typewritten) = ‘Pagosa Springs [;] Colo. 154 7-5-37 [;] C. L. Johnston’’. Second label (white; typewritten) = ‘Wing on [;] Sl. No.”. Third label (brown; partially handwritten, partially typewritten) = “HOLOTYPE 4 [;] Coelinidea [;] ruthae [;] Riegel’ (SEMC). Discussion.—Coelinius ruthae is known only from the holotype, which Riegel (1982) indicated was deposited at “KU.” The holotype bears a glass vial with a cork cap between the top and second labels. The vial contains the posterior portion of the metasoma. The antennae are broken, as is the tarsus of one mesothoracic leg. One forewing is mounted on a slide. Coelinius sommermanae (Riegel), new combination (Fig. 32) Coelinidea sommermanae Riegel 1982: 80, 93 [CUIC, examined]. Type material—Holotype male: Top label (white; typewritten) = ‘‘Downie Creek [;] Selkirk Mts. [;] 14 Aug.’05 Br Col [;] J. Ch. Bradley”. Second label (white with red border; handwritten) = “‘Coelinius [;] Nees’. Third label (brown; partially handwritten, partially typewritten) = “HOLOTYPE 4 [;] Coelinidea [;] sommermanae [;] Riegel’. Fourth label (red; partially handwritten, partially typewritten) = “HOLOTYPE [;] Cornell U. [;] No. 6490’ (CUIC). Discussion.—Coelinius sommermanae is known only from the holotype, which Riegel (1982) indicated was deposited at “CN.” The holotype bears a glass vial with a cork cap between the second and third labels. The vial contains the posterior portion of the metasoma. The antennae are broken. NAMES CONSIDERED NOMINA DUBIA The holotypes of four species of Coelini- dea described in Riegel (1982) apparently are lost: C. antha, C. arca, C. colora, and C. coma. Riegel (1982) indicated that the holotype of each species was deposited at the “‘Philadelphia Academy of Natural Sciences...(PHIL).”” The holotypes do not JOURNAL OF HYMENOPTERA RESEARCH appear in the ANSP primary type data- base. Further, I searched throughout the entire ANSP collection in 2006 but did not find any specimens that could potentially be the holotypes. Museum visits and/or correspondence with curators and collec- tion managers confirmed that the holo- types are not in any of the repositories referenced in Riegel (1982). The holotypes of C. antha, C. colora, and C. coma will be difficult to locate in the absence of an explicit holotype label because the locality label on each of the three specimens is apparently “Col,” (Riegel 1982), an ab- breviation for Colorado that is the only locality information for many specimens in ANSP. Recognition of C. arca in the absence of an explicit holotype label may be possible, as locality data according to Riegel (1982) are ‘‘Cochise Co., Arizona, July 26, 1919, Pinery Canyon, 6000 feet, Chiricahua Mts., Witmer Stone.’”” All four species are known only from the holo- types; the original descriptions and key to North American species of Coelinidea in Riegel (1982) do not provide enough detail to apply the names unequivocally. Therefore, the names C. antha, C. arca, C. colora, and C. coma are considered nomina dubia. ACKNOWLEDGMENTS The following individuals kindly provided speci- mens examined in this study: Philip Clausen (UMSP), Zachary Falin (SEMC), Colin Favret (INHS), Henri Goulet (CNC), Steven Heydon (UCDC), E. Richard Hoebeke (CUIC), Gary Parsons (MSUC), Scott Shaw (ESUW), David Wahl (AEIC), and Robert Zuparko (CAS). I appreciate Stefan Cover (MCZ) and Scott Shaw’s confirmation of paratypes in MCZ and ESUW, respectively. I give special thanks to Zhiwei Liu (Eastern Illinois University [EIU]), Michael Goodrich (EIU), and Jason Weintraub (ANSP) for their efforts in locating misplaced holotypes. I am indebted to Jens Prena (Systematic Entomology Laboratory [SEL] at the USNM) for his advice on Latin grammar. Marie Metz captured and processed all original images. I am grateful to Paul Marsh (Kansas State University), John Brown (SEL), and Thomas Henry (SEL) for their comments on the manuscript. VOLUME 17, NUMBER 2, 2008 LITERATURE CITED Evenhuis, N. L. and G. A. Samuelson. 2007. The Insect and Spider Collections of the World Website Bishop Museum, Honolulu, Hawaii, http://hbs. bishopmuseum.org/codens/codens-r-us.html [accessed 17 November 2007]. Forster, A. 1862. Synopsis der Familien und Gattun- gen der Braconen. Verhandlungen des Naturhistor- ischen Vereines preussischen Rheinlande und West- phalens 19: 225-288. Griffiths, G. C. D. 1964. The Alysiinae (Hym. Braconidae) parasites of the Agromyzidae (Dip- tera). I. General questions of taxonomy, biology and evolution. Beitrige zur Entomologie 14: 823-914. . 1966a. The Alysiinae (Hym. Braconidae) parasites of the Agromyzidae (Diptera). II. The parasites of Agromyza Fallén. Beitriige zur Entomo- logie 16: 551-605. . 1966b. The Alysiinae (Hym. Braconidae) parasites of the Agromyzidae (Diptera). III. The parasites of Paraphytomyza Enderlein, Phytagro- myza Hendel and Phytomyza Fallén. Beitrige zur Entomologie 16: 775-951. . 1967. The Alysiinae (Hym. Braconidae) para- sites of the Agromyzidae (Diptera). IV. The parasites of Hexomyza Enderlein, Melanagromyza Hendel, Ophiomyia Braschnikov and Napomyza Westwood. Beitrige zur Entomologie 17: 653- 698. . 1968a. The Alysiinae (Hym. Braconidae) parasites of the Agromyzidae (Diptera). V. The parasites of Liriomyza Mik and certain small genera of Phytomyzinae. Beitriige zur Entomologie 18: 5-62. . 1968b. The Alysiinae (Hym. Braconidae) parasites of the Agromyzidae (Diptera). VI. The parasites of Cerodontha Rondani s.l. Beitrége zur Entomologie 18: 63-152. Haliday, A. H. 1839. Hymenoptera Britannica: Alysia. Hippolytus Bailliére, London. 28 pp. ICZN. 1999. International Code of Zoological Nomencla- ture. Fourth Edition. The International Trust for Zoological Nomenclature, London. XIX + 306 pp. Kula, R. R. 2006. Systematics of Dacnusini (Hyme- noptera: Braconidae: Alysiinae), Parasitoids of Cyclorrhaphous Diptera. Ph.D. Dissertation Kansas State University, Manhatttan, Kansas. 243 pp. and G. Zolnerowich. 2008. Revision of New World Chaenusa Haliday sensu lato (Hymenop- tera: Braconidae: Alysiinae), with new species, synonymies, hosts, and distribution records. Proceedings of the Entomological Society of Washing- ton 110: 1-60. Maeto, K. 1983. Discovery of two genera, Sarops Nixon and Protodacnusa Griffiths, from Japan (Hyme- noptera, Braconidae). Kontyi# 51: 247-253. 1155 Nees von Esenbeck, C. G. 1819. Appendix ad J. L. C. Gravenhorst conspectum generum et familiarum ichneumonidum. Genera et familias Ichneumoni- dum adscitorum exhibens. Nova Acta Physico- Medica Academiae Ceasareae Leopoldinae-Carolinae 9: 299-310. Nixon, G. E. J. 1937. The British species of Dacnusa (Hym., Fam. Braconidae). Transactions of the Society for British Entomology 4: 1-85. . 1942. A new species of Dacnusa and a new dacnusine genus (Hym., Braconidae). The Ento- mologist’s Monthly Magazine 78: 131-135. . 1943. A revision of the European Dacnusini (Hym., Braconidae, Dacnusinae). The Entomolo- gist’s Monthly Magazine 79: 20-34, 159-168. . 1944. A revision of the European Dacnusini (Hym., Braconidae, Dacnusinae). The Entomolo- gist’s Monthly Magazine 80: 88-108, 140-151, 193-200, 249-255. . 1945. A revision of the European Dacnusini (Hym., Braconidae, Dacnusinae). The Entomolo- gist’s Monthly Magazine 81: 189-204, 217-229. . 1946. A revision of the European Dacnusini (Hym., Braconidae, Dacnusinae). The Entomolo- gist’s Monthly Magazine 82: 279-300. . 1948. A revision of the European Dacnusini (Hym., Braconidae, Dacnusinae). The Entomolo- gist’s Monthly Magazine 84: 207-224. . 1949. A revision of the European Dacnusini (Hym., Braconidae, Dacnusinae). The Entomolo- gist’s Monthly Magazine 85: 289-298. . 1954. A revision of the European Dacnusini (Hym., Braconidae, Dacnusinae). The Entomolo- gist’s Monthly Magazine 90: 257-290. Riegel, G. T. 1947. The American Species of Dacnusinae, Excluding Certain Dacnusini (Hymenoptera: Braco- nidae). Ph.D. Dissertation, University of Illinois, Urbana-Champaign, Illinois. 236 pp. . 1950. A new genus and species of Dacnusini (Hym.: Braconidae). Entomological News 61: 125-129. . 1982. The American species of Dacnusinae, excluding certain Dacnusini (Hymenoptera: Bra- conidae). Novitates Arthropropodae 1: 1-185. Rohwer, S. A. 1914. The Nearctic species of the hymenopterous genus Symphya Foerster. Entomo- logical News 25: 168-172. Schulz, W. A. 1911. Zweihundert alte Hymenopteren. Zoologische Annalen 4: 1-220. Sharkey, M. J. and R. A. Wharton. 1997. Morphology and terminology. Pp. 19-37 in: Wharton, R. A., P. M. Marsh, and M. J. Sharkey, eds. Manual of the New World Genera of the Family Braconidae (Hyme- noptera). The International Society of Hymenop- terists, Washington, DC. 439 pp. Viereck, H. L. 1913. Descriptions of ten new genera and twenty-three new species of ichneumon-flies. Proceedings of the United States National Museum 44: 555-568. 156 Wharton, R. A. 1977. New World Aphaereta species (Hymenoptera: Braconidae: Alysiinae), with a discussion of terminology used in the tribe Alysiini. Annals of the Entomological Society of America 70: 782-803. . 1994. New genera, species, and records of New World Alysiinae (Hymenoptera: Braconi- dae). Proceedings of the Entomological Society of Washington 96: 630-664. . 1997. Subfamily Alysiinae. Pp. 84-117 in: Wharton, R. A., P. M. Marsh, and M. J. Sharkey, eds. Manual of the New World Genera of the Family Braconidae (Hymenoptera). The International Society of Hymenopterists, Washington, DC. 439 pp. JOURNAL OF HYMENOPTERA RESEARCH . 2002. Revision of the Australian Alysiini (Hymenoptera: Braconidae). Invertebrate System- atics 16: 7-105. and A. D. Austin. 1991. Revision of Australian Dacnusini (Hymenoptera: Braconidae: Alysiinae), parasitoids of cyclorrhaphous Diptera. Journal of the Australian Entomological Society 30: 193- 206. Yu, D.S., C. van Achterberg, and K. Horstmann. 2005. World Ichneumonoidea 2004. Taxonomy, Biology, Morphology and Distribution. DVD/CD. Taxapad, Vancouver. Zetterstedt, J. W. 1838. Insecta Lapponica Sectio secunda Hymenoptera. Voss, Lipsiae. 1140 pp. J. HYM. RES: Vol. 17(2), 2008, pp. 157-174 The Neotropical Chrysidid Genus Adelphe Mocsary Revisited (Hymenoptera: Chrysididae: Amiseginae) LYNN S. KIMSEY Bohart Museum of Entomology, Department of Entomology University of California, Davis, CA 95616, USA Abstract—The chrysidid genus Adelphe is reviewed. The species Nesogyne taino Krombein is moved into Adelphe (new combination) and Nesogyne Krombein is synonymized herein as a junior synonym of Adelphe. Thirteen new species are described, namely acuta (Dominican Republic), azurea (Costa Rica, Panama), gibba (Ecuador), glabra (French Guiana), guayanensis (French Guiana, Guyana), hyalophora (Puerto Rico), intermedia (Costa Rica, Panama), lewropos (Ecuador, Peru), lobata (Ecuador, Brazil), lyra (Venezuela), paracubana (Puerto Rico, Dominican Republic), polita (Bolivia) and unidens (Costa Rica). All of these species are based on males, or males and females, with the exception of lyra, which is known only from females. In addition, a key to the species is provided. The genus Adelphe is among the more primitive members of the chrysidid sub- family Amiseginae (Kimsey and Bohart 1991). The subfamily was reviewed by Kimsey and Bohart (1991) who also pro- vided keys to genera. This is the most speciose genus of Amiseginae, with 40 species including those described below. Adelphe was most recently revised by Kimsey (1986), with later additions of species by Kimsey (1993). There are un- doubtedly additional undescribed species, particularly in South America. The genus is primarily Neotropical, with only one spe- cies, anisomorphae Krombein, 1957, occur- ring in North America. This is the only amisegine genus found in the Greater and Lesser Antilles. Adelphe species are characterized by a mixture of derived and primitive features. The most distinctive derived features are the peculiar, flattened male mandible and the well-developed transverse pronotal carina. Configuration of the male mandible is a trait only otherwise seen in the South African genus Anachrysis and South Amer- ican Anadelphe. The transverse pronotal carina extends across the anterior margin of the pronotal disk and wraps laterally, ending at the pronotal lobe. This feature is not found in any other chrysidid genera. Primitive features include the extensively sculptured propodeum and mesopleuron, the presence of a scrobal sulcus and omaulus, presence of ocular setulae, and fully developed wings in both sexes, with one exception. Only the species Nesogyne taino Krombein, which is moved into Adelphe herein (new combination), is known to have brachypterous females. However, females are unknown for some species. With the exception of the reduced wings, Nesogyne shares all of the diagnostic features of Adelphe. Thus Nesogyne is made a junior synonym of Adelphe herein (new synonymy). Brachyptery also occurs in two species of Amisega, the North Ameri- can bella (Krombein) and the Chilean chilensis Kimsey. Among the American amisegine genera only Adelphe has a laterally angulate or dentate propodeum. Other diagnostic characters include the well-developed occipital carina and the presence of an elevated metanotal disk. Sexual dimorphism occurs in Adelphe as it does in most other amisegine genera. 158 Males have broad, foliaceous mandibles and an elongate, slender flagellum. Female mandibles are slender, edentate and are otherwise unmodified, and the flagellum is short and broad; the basal articles may be strikingly pale in females of a few species, such as paradoxa (Kieffer). Fortunately, in some species the sexes share enough features in modifications of the propo- deum, eyes and mesopleuron that they can be associated. The biology of Adelphe is poorly known. Members of the genus are thought to be phasmid egg parasites, but to date only one host, Anisomorpha ferruginea (Beauv.) (Phas- matidae), is known, and that is for the North American species, anisomorphae (Krombein 1957). Males are more common- ly collected than females. This is probably because males spend more time flying and perching on vegetation above the ground. Females may spend the majority of their time on the ground and in leaf litter looking for host eggs. The most effective methods for collecting these wasps are yellow bowls, flight intercept traps (FIT) and Malaise traps. To facilitate identification and the recog- nition of new species, a key to species based on males is given below. Because females are known for only about one- quarter of the species they are not included in the key. MATERIALS AND METHODS This study was made possible by loans of specimens from the following institu- tions and individuals: Canadian National Insect Collection, Agriculture Canada, Ot- tawa, Ontario (CNC) (J. Huber, L. Masner); Natural History Museum of Los Angeles, Los Angeles (LACM) (W. Xie), Museo de JOURNAL OF HYMENOPTERA RESEARCH Insectos, Universidad de Costa Rica, San Jose (MUCR) (P. Hanson) and the Bohart Museum of Entomology, University of California, Davis (BME) (S. L. Heydon). Types are deposited in the museums indicated in the type series by the museum acronym given in parentheses. A number of morphological features are used in the key and descriptions below that require further description. These are shown in Fig. 1. The scapal basin is the area immediately above the antennal sock- ets, which is often concave and cross- ridged. Many measurements are made using the greatest side to side diameter of the midocellus (MOD) (Fig. 1A). The fla- gellomeres are numbered using Roman numerals starting with the flagellomere closest to the pedicel. Flagellomere propor- tions are measured using the greatest breadth and length of each article (Fig. 1A). The postocular distance is measured in dorsal view and is the area between the posterior eye margin and the outer margin of the occipital carina on the side of the head (Fig. 1B). The scrobal sulcus is mea- sured from the anterior margin of the sulcus to the scrobe, or prior to the scrobe if the sulcus abruptly narrows, in which case the length is measured to the point of constriction (Fig. 1D). The greatest width of the sulcus is used in the measurement. Finally, the metanotum has an elevated medial disk, which has different dimen- sions in different species (Fig. 1E). The greatest width is measured relative to the length along the midline. A rough estimate of punctation is given using the puncture diameter (PD) on the structure in question to measure the average distance between punctures. These features need to be measured and not determined by “eye”. KEY TO MALE ADELPHE 1 Mandible with one apical tooth (Figs 4, 7, 13) — Mandible with two apical teeth (as in Fig. 3) VOLUME 17, NUMBER 2, 2008 159 OG -aS midocellus diameter malar space eons subantennal{_ eed distance L ia postocular distance Fig. 1. Adelphe structural features, L = length, W = width. A. Front view of face. B. Dorsal view of head. C. Side view of body, ss = scrobal sulcus. D. Lateral view of mesopleuron. E. Dorsal view of propodeum, a. dorsolateral enclosure, b. dorsomedial enclosure, c. posteromedial enclosure. 2 Scapal basin smooth without cross-ridging (Fig. 7); scrobal sulcus more than 8X as long as broad; eye height 1.9-2.0x as long as malar space length in side view; | ECR (G7 ee oe ae ee ee ee emer hyalophora Kimsey, new species = Scapal basin extensively cross-ridged (as in Figs 4, 13); scrobal sulcus less than 8X as long as broad; eye height less than 1.8X as long as malar space length in side view ...... s. © Face long, narrow, margins below eyes nearly parallel-sided medially (Fig. 4); head in dorsal view breadth 1.3X length (Fig. 25); Ecuador ..... gibba Kimsey, new species = Face broad, margins below eyes strongly converging below (as in Fig. 13); head in pele ry ICCA ty AOC OF TOT CMTE EE cee Sica aah nil ah Pain A) oy far ones Rass = + Subantennal distance 1 MOD long; flagellomere I 2.9-3.0X as long as broad; eye height more than 1.5X malar space length in side view; Brazil, Ecuador . paradoxa (Kieffer) = Subantennal distance more than 1 MOD long; flagellomere I 2.5 as long as broad; eye height 1.5X malar space length in side view; Costa Rica unidens Kimsey, new species 5 Scapal basin cross-ridged or vertically rugose (as in Figs 6, 8,10) ............... 6 - Scapal basin smooth, without cross-ridging (as in Figs 3,5,9, 11,12) ........... 17 6 Scrobal sulcus absent; body with extensive decumbent silvery pubescence; Dominican PSEA ee tS EA: Een ae & ad Wa = Ae ea ee ie argentea Kimsey 160 >a || JOURNAL OF HYMENOPTERA RESEARCH Scrobal sulcus well-developed; body without decumbent silvery setae ........... Z Scrobal sulcus:more than 6.0 as:long. as. brodd \. 2. Be . 3 ieee 8 Scrobal sulcusiless than. 5.5X' as lone as broad...3-.. .a?= hain eee ee. 4 Sa eee 2 Head posteriorly with deep pit on either side of midline between occipital and hypostomal carinae; propodeum without lateral tooth or angle; thorax red; pronotum: without lateraltcarina; Puerto RicO). 7. 3 coat its aie masnert Kimsey Head posteriorly without pit on either side of midline between occipital and hypostomal carinae; propodeum with lateral tooth or angle; thorax black; pronotum with lateral carina; brazil. > en cylindrica Kimsey Flagellomere I 3.0X or less as long as broad; flagellomere II twice as long as broad or TeSSD pooya gt A EE LOE oc ae Re =. Roe ee ee 10 Flagellomere I 3.4X or more as long as broad; flagellomere II more than twice as long as broad Pew. Je Sue Maree ers. VA ae ee) Pie il Malar space 4 MOD long in front view; eye height less than twice as long as malar space length in Jateralhview;,Mexico—s5. oA aens. else mexicana Mocsary Malar space less than 3 MOD long in front view; eye height twice as long as malar space length in lateral view; Canada, USA ............. anisomorphae Krombein Flagellomere I more than 4X as long as broad; scrobal sulcus 3.5X as long as broad; MiexdCO", g:08 tac on as ob Ee ae ce ea areas, se laevis Kimsey Flagellomere I less than 4X as long as broad; scrobal sulcus 4—5X as long as broad ... 12 Pronotal carina obsolescent laterally; Panama .................... confusa Kimsey Pronotalcarina well-developed laterally. 7%. 95. eee: ) ae ere 13 Scrobal sulcus 5X as long as broad; flagellomere I 4X as long as broad; flagellomere II 3x-as; long 7ds broad? Costa (RicatCl. . 2. a). See Siete es limonae Kimsey Scrobal sulcus 4.5X as long as broad or shorter; flagellomere I 3.5X as long as broad or shorter; flagellomere II less than 3X as long as broad ..................... 14 Eye height less than twice malar space length in side view .................-. 15 Eye height twice malar space length in side view ........ Sop ane Mikes 5s here 16 Subantennal distance less than 1.5 MOD long; Panama ............. robusta Kimsey Subantennal distance more than 1.5MOD long; Costa Rica ....... intermedia Kimsey Subantennal distance 1 MOD long; malar space 2.5 MOD long in front view; French Guiana, Guyana oo. fo. er te eee ae guayanensis Kimsey, new species Subantennal distance 2 MOD long; malar space 3 MOD long in front view ECUAGOE” hss oe in eee GS aly RMT te tee lobata Kimsey, new species Scrobal sulcus 3.0-4.5X.as longvas broad. /. 19. ses Jose oe eee 18 serobal sulcus 5x as lone as broad or longer 2 =). 22). 228.525 eee 28 Flagellomere | less than.3.6X as long as broad... 44. .a25.. 2404 eee 19 Flagellomere 1 4X or more’as long as broad]. ) 2... 222 &. -2 ee 528) Malar space 4 MOD long in front view; eye height 1.0-1.5x malar space length in side VIC WE coo sans Tice a veyingeteges inal Padine sg) Sb eae kan en 20 Malar space 3 MOD long in front view; eye height 1.9-2.5x malar space length in side MIGW 2s coe woe ee bce s eetite one Sg ep cus ae erage ete lee ee ee a 21 Flagellomere I 3.5X as long as broad; flagellomere II more than 2.5X as long as broad; Donvintican Republic’ <7. fcr = hee eee ee Se re eee dominicana Kimsey Flagellomere I 2.0-2.2< as long as broad; flagellomere II less than twice as long as broad; Eehader Colombia. oe ee ea ee eee longifacies Kimsey Eye with minute ocular setulae or setulae lacking; scrobal sulcus 3.5 or less as long as broad: Erencm Gdiana. ee ee eee glabra Kimsey, new species Eye with ocular setulae 0.3-0.7 MOD long; scrobal sulcus 3.8-4.0< as long as broad ... 22 Postocular distance short, about 1 MOD wide in dorsal view; eye height more than twice malar space length in side view; Ecuador, Peru _lewropos Kimsey, new species Postocular distance long, 2 MOD wide in dorsal view; eye height less than twice malar space ienetiin side view, Costa Ica 2 ie. wre ee Soe | enor hansoni Kimsey VOLUME 17, NUMBER 2, 2008 161 2S 24 36 oF Scrobalisuleusi30=3.5 xX asilongvas broads s:r: Dies ec) Oe 8s Ow ee ee ee. 24 Serobalisuleus 4.04.5 x as, lone as broad), «Wp. oo 2k on age in oo I OMA... 26 Eye height 1.0-1.5x as long as malar space length in side view; malar space 3.5-3.8 MOD long in front view; postocular distance more than 1.5 MOD wide in dorsal wiewsMenezelais:.. : Gass EI LANE Sie ce etas eae a ait meridae Kimsey Eye height 2-3 malar space length in side view; malar space 2.5-3.0 MOD long in front view; postocular distance 1.0-1.5 MOD wide in dorsal view ........... 7) Subantennal distance 1 MOD long; malar space 2.5 MOD long in front view; eye height twice malar space length in side view; Brazil. ................ metallica (Kieffer) Subantennal distance 1.5 MOD long; malar space 3 MOD long in front view; eye height 2.6X malar space length in side view; Brazil .................. flavipes (Ducke) Eye height twice malar space length in side view; subantennal distance 1 MOD long; malar space 3 MOD long in front view; Costa Rica ............... nitida Kimsey Eye height less than twice malar space length in side view; subantennal distance more than 1.4 MOD long; malar space less than 3 MOD long ................... 27 Flagellomere II 2.5x as long as broad; subantennal distance1.7 MOD long; propodeal medial enclosures smooth; Costa Rica ........... paralaevis Kimsey, new species Flagellomere II more than 3X as long as broad; subantennal distance1.5 MOD long; propodeal medial enclosures coarsely rugoseor cross-ridged; Costa Rica = azurea Kimsey, new species senonalesulcus M0 as lone as broad. 67.5... Gee ee ee ee oe ee ek eg) Seropal sulcusio—ox as lOMe aS brOad= a..e. oe hs one Bete oe 30 Subantennal distance more than 1.5 MOD long; eye height more than 2.6X malar space lemeitnin lateral vaew, Westalmdies 92834 ents ee bee. Coane nesos Kimsey Subantennal distance less than 1.5 MOD long; eye height twice malar space length in laterdleview, Domimican*Republic™ © .....<2r. S46). aki Be yee: minuta Kimsey Flagellomere I 3X or less as long as broad; flagellomere II less than twice as long as [SESE BTV Al es as Bowe es Accor to ee Ae. a or eee om calvata Kimsey Flagellomere I 3-5X as long as broad; flagellomere II more than twice as long as broad .. 31 Flagellomere I 3.0-3.5X as long as broad; flagellomere II 2.8-4.0X as long as broad ... 32 Flagellomere I 4-5X as long as broad; flagellomere II 2.0-2.5x as long as broad ... 36 peropalesuilcus less than 6xXvas long asbrodd 5... ee ee le 33 perooaustiletts! 7—o <1as lome vas, broad)... a .8te ~. es de ee eee ee bee ee 34 Flagellomere IX more than 4.5 as long as broad; subantennal distance less than 1.5 LOW MOmereBCAZU FT. a i. dts ons 2 ahs ae hae Rae ae brasiliensis Kimsey Flagellomere IX 4X as long as broad; subantennal distance 1.5 MOD long or longer; Olina seme. Sen ek RETIN A nue a. w/e... 2g7 polita Kimsey, new species Subantennal distance 2.5 MOD long; postocular distance more than 2 MOD wide; scrobal sulcus 8X as long as broad; Jamaica ..................... ziva Kimsey Subantennal distance 1 MOD long or less; postocular distance less than 2 MOD wide; Seto pal ssulltisn7 “GaswOne AsrbrOAG ts. 6. a... be es ge oe oe ee i ew 35 Eye height less than twice malar space length in side view; postocular distance less than seMOMAwide: Jamaica © 2.0304. Bees le as ek ek bb se bees insula Kimsey Eye height twice malar space length in side view; postocular distance more than 1 MOB wiGe: BatentOuRIcO - jus. oj. - . ieee ties paracubana Kimsey, new species Subantennal distance less than 1 MOD long; postocular distance 1 MOD wide or less; Brazile. sree: 2 teen See. . ean. setae ete Ds be tye antennalis Kimsey Subantennal distance 1 MOD long or longer; postocular distance 1.3-1.6 MOD wide ... 37 Flagellomere II less than 3X as long as broad; eye height twice malar space length or TOT Ca Sie ee aR A ee Bod 2 Sk in Eure ui yea dl Cb oe asl wr awa ec olka aga = ay 38 Nay Oe eee er a Pe eS cic hie gs oe ya AR Be ent gt ase a os ig 39 162 JOURNAL OF HYMENOPTERA RESEARCH 38 Scrobal sulcus 6X as long as broad; malar space less than 2.5 MOD long in front view; Puerto. Rico? 6 9c GRR S3. Mymaromella cyclopterus Gibson et al., 2007: 100 (fig. 166, generic transfer). Material examined—The holotype female (Figs 19, 25, 31) and only known specimen of M. cyclopterus, is on a slide under one large coverslip, labelled: 1. “Galeomymar cyclopterus & A.O. Loreto, Misiones, 29.iv.1933. A.O. Typus!”” 2. Palaeomymar cyclopterus Det. De Santis et Fidalgo Holotypo Museo de la Plata” 3. “3912/1”. Descriptive notes—Female. Body length 409 um (holotype). 185 Head. Ocelli are definitely present (Fig. 31), in contrast to what was stated in Fidalgo and De Santis (1982). The number of ommatidia cannot be counted because the eyes are mostly black (Figs 25, 31). Antenna. Fig. 25. L(W) measurements (holotype) are: scape 56 (15); pedicel 35 G7); fh 157) ntl 19 (7), fl; 21 (6); fl, 21 (6), fl; 24 (8), fl, 23 (8), fl; 27 (11), clava 96 (24). FWL 414, FWW 194, FWL(W) 2.13. Fore wing. Fig. 19. Without a long seta on posterior margin basal to short, spine- like setae of the marginal fringe. Mymaromella mira Girault (igs 22, 23, 28,29) Mymaromella mira Girault, 1931: 4; Dahms, 1984: 823; Gordh et al., 1979: 283 (reprint of original description); Gibson et al., 2007: 101 and figs 13, 35, 65, 66, 93, 94, 97, 98, 162, 168 (revised status from Palaeomymar). Material examined.—The holotype specimen no longer exists, but Fig. 23 is a photograph of it (Gibson et al. 2007). Twenty-one specimens, including 59 and 13 on slides, as follows: AUSTRALIA. ACT: Blundells Creek, 35.22S 148.50E, ii.1987, D.H. Colless, Malaise trap (Figs. 22, 28) (29, ANIC); Canberra, Black Mountain, CSIRO, 1—-15.i1.1999, G. Gibson, YPT (Fig. 29) (49, 13g, CNC); Piccadilly Circus, 1240 m, 35.225 148.48E, xii.1984, J. Lawrence, T. Weir, H.-L. Johnson, light intercept/window/trough trap, figured specimen in the Insects of Australia, 2"? edition (19, ANIC). Victoria: [?Ot]Otway Forest, Ormond, no date given, W.S. Anderson (19, USNM). Descriptive notes—Female. Body length 376 um (n = 1, critical point dried speci- men), 500-543 (n = 3, slide mounted specimens from Blundells Creek and Black Mountain). Mesosoma brown, head, ap- pendages and petiolar segments honey yellow, gaster usually brown but in one specimen yellow. Head. Eye with at least 30 ommatidia (in Black Mountain specimens). Head width 133-142 (n = 2). Sculpture reticulate-striate 186 JOURNAL OF HYMENOPTERA RESEARCH Figs 19, 20.—Mymaromella spp., fore wings. 19, M. cyclopterus, holotype; 20, M. pala, holotype. Scale lines = 50 um. (Gibson et al. 2007, figs 13, 35). Ocelli 2): scape 66-68 (18); pedicel 36-38 (16-22); present. fl, 18-23 (9-11), fl, 20-24 (8-10), fl, 24-26 Antenna. Female antenna (Fig. 28). (8-10), fl, 21-23 (9-10), fl; 26-30 (9-10), fl, L(W) measurements (n = 3 or, for width, 40-44 (9-10), fly 33-35 (13-16), clava 106— VOLUME 17, NUMBER 2, 2008 187 / Figs 21, 22.—Mymaromella spp., fore wings. 21, M. palella; holotype; 22, M. mira. Scale lines = 50 um. 111 (30-32), with slightly pointed apex (Fig. 29; Gibson et al. 2007, fig. 65). Mesosoma. Mesosoma with reticulate sculpture, more distinct dorsally than laterally (Gibson et al. 2007, figs 93, 97, 98). Fore wing. Flat, with broadly rounded apex and with hair-like basal seta (Figs 22, 23; Gibson et al. 2007, fig. 168 — seta not visible in the published image but definite- ly present in original photograph). Fore wing broad: FWL 508-555, FWW 222-243, FWL/W 2.15-2.29, longest marginal setae 184-190, venation length 85-86 (n = 3). Metasoma. Petiolar segment 1 length 72-82, segment 2 length 45-47 (n = 2), both petiolar segments with irregular transverse striations and segment 1 with 2 setae at or just before mid-length (Gibson et al. 2007, figs 93, 94). Ovipositor length (including valves) 86-97 (n = 2). 188 JOURNAL OF HYMENOPTERA RESEARCH jet se scureattia: comet a create ee Fig. 23. Mymaromella mira, holotype photograph. Male. Colour as in female except gaster honey yellow. Body length 376-445 (n = 8, critical point dried specimens). Head. Eye with about 50 ommatidia. Antenna. Fig. 29 and Gibson et al. (2007) fig. 66. Measurements L (W) (n = 1): scape — not accurately measurable, pedicel 34 (15), fl, 14 (8), fl, 17 (9), fl, 19 (9), fl, 20 (9), fls 29 (9), fle 28 (11), fly 28 (13), flg 28 (16), flo 22 (17), flip 22 (15), flay 24 (14). Fore wing. Fig. 23. L/W 2.63 (n = 1). Metasoma. Genitalia (Gibson et al. 2007, fig. 162). Variation.—Girault’s (1931) description mentions six features that can be compared accurately with the slide-mounted speci- VOLUME 17, NUMBER 2, 2008 189 Figs 24-28. Mymaromella spp., female antennae. 24, M. chaoi, paratype; 25, M. cyclopterus, holotype; 26, M. pala (+ head, anterior), holotype; 27, M. palella, holotype; 28, M. mira. Scale lines = 50 um. 190 JOURNAL OF HYMENOPTERA RESEARCH Figs 29-31. cyclopterus, head, mesosoma and first segment of gastral petiole, holotype. Scale lines = 50 um. mens we examined. All but one feature almost exactly matches the specimens from ACT. The one feature that does not match is that fl, is not nearly twice as long as fly but is only 1.2-1.3 times as long on the three females we measured. We do not know how Girault measured the funicular lengths but differences in method of Mymaromella spp. 29, M. mira, male antenna + head, lateral; 30, M. pala, male antenna; 31, M. measurement may partially account for the discrepancy. We consider that there is a close enough match between our speci- mens and the original description and type photograph to be certain that the females are M. mira. By association, we also place the males from Black Mountain in this species although the fore wing is narrower, VOLUME 17, NUMBER 2, 2008 with the dark band not so wide or conspicuous. HOSTS AND BIOLOGY The hosts and biology of Mymaromma- tidae are unknown. However, the informa- tion on distribution and habitat obtained from the literature and from specimens in collections provides us with circumstantial evidence for the likely host group. The evidence presented below is based on mymarommatid morphology, collection data, biogeography, habitats, and palaeon- tology, all of which correlate well with one order of potential hosts — the Psocoptera. Morphology. Because Mymarommatoi- dea belong to the parasitic Hymenoptera, probably as the sister group of Chalcidoi- dea (Gibson et al. 2007), it can reasonably be assumed they are parasitoids of other insects. Their small size, rivalling that of small Mymaridae and Trichogrammatidae, and their very short ovipositors, at most about 110 tm long, suggest they parasitize the egg stage, as do members of the latter two families. We also assume that Mymar- ommatidae are solitary, internal parasit- oids that feed on the egg contents before the host cells have begun to differentiate, thus avoiding the problem of overcoming a host immune system, which does not appear before the host larva develops. Minute wasps generally would have a harder time parasitizing the mobile stage of an insect larva or adult compared to an immobile stage (egg or pupa) because a mobile host could defend itself from attack and it would also have an immune system that would have to be countered. A disadvantage of parasitizing eggs is that the body size of an internal parasitoid is limited to that of its host egg. What kind of eggs could be parasitized? We suggest small, thin-walled eggs from which an adult wasp could emerge in one of two ways, assuming that the parasitoid is solitary and completely fills the egg once development is complete. Mymarommati- dae are unique among Hymenoptera be- 191 cause they have the front and back of the head joined by pleated membrane that extends between the base of each mandible across the top of the head. Either an adult mymarommatid could burst open the host ege simply by flattening the pleated membrane, thus enlarging its head (see Gibson et al. 2007, figs 13-15), through hydrostatic pressure or muscle action. Or the expanded head may not itself burst the host egg but instead provides a buttress for the exodont mandibles (another feature of Mymarommatidae — Fig. 2, 26, 29, and Gibson et al. 2007, figs 23, 25, 28, 41, 44, 49, 50) to tear a hole in the chorion through which the wasp emerges. Psocoptera have a thin egg chorion, about 1 pum thick (Seeger 1979). Because of this it may be fairly elastic and easily distorted, hence difficult for an internal parasitoid to bite through without buttressing from an expanded head. Exodont mandibles may also be more efficient than endodont mandibles in pushing an emergence hole through the soil or bark debris, silk threads or fecal material that many Pso- coptera use to cover and protect their eggs (Hinton 1981), but may make it more difficult for an internal parasitoid to bite through the chorion. Consequently, a mechanism to expand the head and firmly appress the exodont mandibles to the chorion may be required. Abundance and phenology. Mymar- ommatidae are usually collected singly or in small numbers. This is partly an artifact of their small size and the consequent difficulty of seeing them. Occasionally, considerable numbers (50 or more) may be collected in a short time by a particular Malaise or yellow pan trap. This suggests a mass emergence, possibly from hosts that lay clusters of eggs. Specimens of both Mymaromella and Mymaromma Girault have been collected in the field during every month from May to September in mid latitudes of the Northern Hemisphere (Canada, USA, var- ious European countries, Japan, Korea) and 192 have emerged in November from logs maintained in the lab in Michigan). In the Southern Hemisphere (Australia, New Zealand) specimens have been collected every month from October to June. In the tropics (Brazil, Céte d’Ivoire, Gabon, Ha- waiian Is., Taiwan, Thailand) specimens have been collected from November to July. Presumably, a given species of my- marommatid has several generations per year and adults may be found throughout the warm season in higher latitudes and most of the year in the humid tropics. Psocoptera lay eggs either singly or in batches, occasionally with up to 80-90 per batch, and are univoltine or multivoltine (New 1987). A given species may have several generations over many months, thus providing a fairly constant source of eggs to be parasitized. If all the eggs in a cluster were parasitized it would account for a mass emergence of a particular species of Mymarommatidae, especially if many egg clusters were so parasitized. Most Psocoptera overwinter as eggs so their eggs would serve as overwintering sites for diapausing mymarommatids. Biogeography. Specimens of Mymar- ommatidae have been collected from all continents except Antarctica, and from remote oceanic islands such as Hawaii (Beardsley et al. 2000) and some subant- arctic islands of New Zealand including Campbell Island, which has one species of Mymarommatidae (Valentine 1971). Psocoptera occur worldwide including many oceanic islands such as Campbell Island, which has three species (Gressitt 1964, Gressitt and Wise 1971) mainly in moss (Gressitt 1964) among the 380 report- ed arthropod species. The species of Mymarommatidae on Campbell Island must be restricted to one or several of the potential arthropod hosts, possibly Psocop- tera. Psocoptera are also relatively easily dispersed, sometimes (by implication) over long distances (New 1987) and evidently occur wherever mymarommatids have been collected. JOURNAL OF HYMENOPTERA RESEARCH Habitats. Data from the literature and from specimens assembled at the CNC for Gibson et al. (2007) indicate that most Mymarommatidae may be collected in a wide diversity of forested habitats from sea level (Bermuda) to 1050 m (Japan). Based on label data, the habitats and countries from which specimens were seen are: Peucedano-pinetum (Poland), garrigue (France), climax flood forest (Czech Re- public), small meadow in old deciduous forest (Japan), secondary forest (Taiwan), mango patch (Australia), sclerophyll forest (Australia), riverine forest (Thailand), cer- rado (Brazil), dense forest (New Caledo- nia), yellow sticky traps hung on roadside trees (Hawaii — Beardsley et al. 2000), ex ash logs from Metropolitan parks (Michi- gan, USA), maple and white pine litter, mixed cedar and alder litter, Berlese extract of soil from maple-hickory forest (Canada), deciduous forest litter (Canada — Clouatre et al. 1989), Nothofagus forest, litter of Stilbocarpa in Olearia forest (New Zealand), and ex bracket fungus (New York, USA). The only records we have seen from outside forested habitats are: litter of Anisotome latifolia at upper margin of supralittoral zone, litter and peat under Stilbocarpa polaris, and ex Poa tannantiana (New Zealand: Snares, Campbell, Auck- land, and Antipodes Is., from label data and from Valentine 1971), Caprobrotus, Munro Beach cottages (Bermuda), and an old field (USA, Maryland). Psocoptera occur in soil and ground litter, low vegetation, on bark of tree trunks and branches, on foliage (New 1987), and in bracket fungi (Matthewman and Pielou 1971). Palaeontology. Mymarommatoidea are known from at least 100 mya as shown by Cretaceous amber fossils from Lebanon, Canada, and Russia (Gibson et al. 2007). Fossils of Psocoptera are known from the Jurassic and various extant families are known from 100 mya Cretaceous amber from Lebanon and India (Kukalova-Peck 1991) so they were present as potential VOLUME 17, NUMBER 2, 2008 hosts when mymarommatids occurred in the fossil record. Discussion.—Psocoptera are proposed as the most likely insect hosts for Mymarom- matidae because their eggs are small and thin-walled, may be laid in clusters, may be present throughout the period that adult mymarommatids have been collected, and in higher latitudes are the over wintering stage. Psocoptera also occur wherever Mymarommatidae have been collected worldwide and may be abundant in a range of different habitats, including the same ones aS mymarommatids. However, these lines of circumstantial evidence could fit several other groups of possible hosts that have the same distribution, habitats, fossil record and egg size as Psocoptera. Such alternative possible hosts include some Coleoptera (such as Curculionidae or Staphylinidae) and Diptera (such as various Nematocera). Other arthropod groups, notably Acari and Collembola emerged in considerable numbers from over wintered ash logs but we consider them unlikely hosts because parasitic Hy- menoptera have rarely been reared from Acari and never, so far, from Collembola. Lists of species reared from bracket fungus (Matthewman and Pielou 1971) and logs of ash trees (often loaded with lichens) over wintered under laboratory conditions (this study) are fairly short. Matthewman and Pielou (1971) list 6 families and 14 species of Psocoptera among 59 families and 133 species of insects from bracket fungus in Quebec. Our ash rearings in Michigan resulted in about 30 genera of predaceous and parasitic Hymenoptera, about five genera of Diptera, about five genera of Coleoptera, and eight genera of Psocoptera including Atropsocus atratus (Aaron), Blaste subquieta (Chapman), Blastopsocus lithinus (Chapman) and B. semistriatus (Walsh), Echmepteryx hageni (Packard), Loensia moesta (Hagen), Liposcelis sp., Psocus leidyi Aaron, and Trichadenotecnum alexanderae Sommerman. Hymenoptera are unlikely as hosts of Mymarommatidae because they 193 themselves are parasitic and most lay their eggs within a host and would be inacces- sible for parasitism. Psocoptera therefore seem to be the most likely host group, particularly as a diversity of genera and species were reared from ash logs. CONCLUSIONS More species of Mymaromella than the five keyed above are known to us. They are numbered in Gibson et al. (2007) but we leave them undescribed until more mate- rial is collected and the respective regional faunas are studied more thoroughly. The biology of Mymaromella and indeed the entire family Mymarommatidae remains unknown, though we hypothesize Psocop- tera as hosts based on the circumstantial evidence presented above. Whereas some other insect groups, such as certain Diptera or Coleoptera, could also be potential hosts of mymarommatids, the taxa reared from bracket fungi and ash logs seem to make these groups less likely candidates. Our hypothesis can be tested by rearing Pso- coptera eggs. We suggest that the best chance of obtaining a definite rearing of any species of Mymarommatidae would be from Psocoptera eggs collected from brack- et fungi, from litter and mosses collected in the subantarctic islands of New Zealand or from trunks of various ash species in north eastern North America. ACKNOWLEDGEMENTS We thank M. Loiacono for the loan of M. cyclopterus holotype, N.-Q. Lin for the loan of M. chaoi types, E. Mockford for identifications of the Psocoptera reared from ash logs, and K. Wu for translating the type locality information. J. Read is gratefully acknowl- edged for preparing the scanning electron micro- graphs and compiling the plates of illustrations. LITERATURE CITED Bauer, L. S., Liu, H-P., Haack, R. A., Petrice, T. R., and Miller, D. L. 2003. Natural enemies of emerald ash borer in southeastern Michigan. Pp. 33-34, in: The Proceedings of the 2003 Emerald Ash Borer Research and Technology Development Meeting. USDA Forest Service FHTET 2004-02. 194 , Liu, H-P., Gould, J., and Reardon, R. 2007. Progress on biological control of the emerald ash borer in North America. Biocontrol News and Information 28: 51N-54N. Beardsley, J. W., Huber, J. T., and Perreira, W. D. 2000. Mymarommatoidea, a superfamily of Hymenop- tera new for the Hawaiian Islands. Proceedings of the Hawatian Entomological Society 34: 61-63. Clouatre, A., Coderre, D., and Gagnon, D. 1989. Habitat of a new Mymarommatidae found in southern Quebec, Canada (Hymenoptera: Tereb- rantes). The Canadian Entomologist 12: 825-826. Dahms, E. C. 1984. A checklist of the types of Australian Hymenoptera described by Alexandre Arsene Girault: III. Chalcidoidea species F-M with advisory notes. Memoirs of the Queensland Museum 21: 579-842. Fidalgo, A. P. and L. de Santis. 1982. Una nueva especie Argentina de mimarido de la subfamilia Mymaromminae (Insecta, Hymenoptera). Revista del Museo de La Plata (Nueva Serie), Zoologia 127: 1-6. Gibson, G. A. P. 1993. Superfamilies Mymarommatoi- dea and Chalcidoidea. Pp. 570-655 in: Goulet, H., and J. Huber eds. Hymenoptera of the world: an identification guide to families. Agriculture Canada Research Branch Monograph No. 1894E, Ottawa. 668 pp. . 1997, Morphology and terminology. Pp. 16-44 in: Gibson, G. A. P., J. T. Huber, and J. B. Woolley eds. Annotated keys to the genera of Nearctic Chalcidoidea. NRC Research Press, Ottawa. 794 pp. , Read, J., and Huber, J. T. 2007. Diversity, classification and higher relationships of Mymar- ommatoidea (Hymenoptera). Journal of Hymenop- tera Research 16: 51-146. Girault, A. A. 1931. A new habit in an old insect, Homo pudicus and new Eurytomidae. Privately printed. 4 pp. JOURNAL OF HYMENOPTERA RESEARCH Gordh, G., Menke, A. S., Dahms, E. C., and Hall, J. C. 1979. The privately printed papers of A. A. Girault. American Entomological Institute 28: 1-400. Gressitt, J. L. 1964. Insects of Campbell Island. Pacific Insects Monograph 7: 1-663. and Wise, K. A. J. 1971. Entomology of the Aucklands and other islands south of New Zealand: Introduction. Pacific Islands Monograph 271-45, Hinton, H. E. 1981. Biology of insect eggs in three volumes. Vol. II. xviii + 475-778. Pergamon Press, Oxford, UK. 1125 pp. Kukalova-Peck, J. 1991. Fossil history and the evolu- tion of hexapod structures. Pp. 141-179 in: The insects of Australia. Second Edition. Vol. I. Mel- bourne University Press, Carleton, Vic. 542 pp. Matthewman, W. G. and Pielou, D. P. 1971. Arthopods inhabiting the sporophores of Fomes fomentarius (Polyporaceae) in Gatineau Park, Quebec. The Canadian Entomologist 103: 775-847. Lin, N-Q. 1994. First discovery of Mymarommatidae (Hymenoptera) from China, with description of a new species. Entomotaxonomia 16: 120-125. New, T. R. 1987. Biology of the Psocoptera. Oriental Insects 21; I-109: Seeger, W. 1979. Specialmerkmale an Eihiillen und Embryonen von Psocoptera im Vegleich zu anderen Paraneoptera (Insecta); Psocoptera als monophyletische Gruppe. Stuttgarter Beitrige zur Naturkunde, Serie A (Biologie) 329: 1-57. Valentine, E. W. 1971. Entomology of the Aucklands and other islands South of New Zealand: Hyme- noptera: Mymaridae. Pacific Insects Monograph 27: 327-333. Yoshimoto, C. M. 1984. The insects and arachnids of Canada, Part 12. The families and subfamilies of Canadian chalcidoid wasps. Publication 1760, Cana- dian Government Publishing Centre, Supply and Services Canada, Ottawa. 149 pp. J. HYM. RES. Vol. 17(2), 2008, pp. 195-200 Evaluation of Specimen Preservatives for DNA Analyses of Bees MARK FRAMPTON*, SAM DROEGE, TRAVIS CONRAD, SEAN PRAGER, AND MIRIAM H. RICHARDS (MF, SP, MHR) Department of Biological Sciences, Brock University, St. Catharines, Ontario, Canada; (SD, TC) USGS Patuxent Wildlife Research Center, Beltsville, Maryland, USA Abstract.— Large-scale insect collecting efforts that are facilitated by the use of pan traps result in large numbers of specimens being collected. Storage of these specimens can be problematic if space and equipment are limited. In this study, we investigated the effects of various preservatives (alcohol solutions and DMSO) on the amount and quality of DNA extracted from bees (specifically Halictidae, Apidae, and Andrenidae). In addition, we examined the amount and quality of DNA obtained from bee specimens killed and stored at —80°C and from specimens stored for up to 24 years in ethanol. DNA quality was measured in terms of how well it could be PCR-amplified using a set of mitochondrial primers that are commonly used in insect molecular systematics. Overall the best methods of preservation were ultra-cold freezing and dimethyl sulfoxide, but these are both expensive and in the case of ultra-cold freezing, somewhat impractical for field entomologists. Additionally, dimethy] sulfoxide was shown to have adverse effects on morphological characters that are typically used for identification to the level of species. We therefore recommend that the best alternative is 95% ethanol, as it preserves bee specimens well for both morphological and molecular studies. Recent advances in insect molecular systematics, made possible by the polymer- ase chain reaction (PCR) and other molec- ular techniques have made it important to properly preserve rare specimens for main- taining museum collections, or for molecu- lar analysis. Since DNA can be damaged by enzymatic breakdown, oxidation and hy- drolysis (Lindahl 1993, Quicke et al. 1999), specimens need to be preserved from the time of collection to the time of analysis in order to minimize DNA degradation. Sev- eral factors have been reported that affect DNA degradation in stored insect speci- mens, including preservative type and concentration, time in preservative, temper- ature, pH, and the age of the specimen (Dillon et al. 1996). It is generally accepted that the highest quality of DNA is extracted from live specimens (Tayutivutikul et al. 2003), live specimens frozen at —80°C (Dillon et al. 1996), or live specimens quick * Author for correspondence frozen in liquid nitrogen (Quicke et al. 1999). However, these methods are not always practical for field biologists, and several alternatives have been reported for preserving arthropod, mammalian or plant specimens for the purpose of genetic anal- ysis. These include storage in preservatives such as methanol, ethanol, and isopropanol (Post et al. 1993), propylene glycol (Rubink et al. 2003), acetone (Fukatsu 1999), Car- noy’s solution (Post et al. 1993), and dimethyl sulfoxide (Kilpatrick 2002). An important motivation for the current study is an increasing research emphasis on large- scale collections of bees, especially using pan traps that generate specimens used for molecular systematics and population ge- netic studies. We therefore needed to assess the relative merits of various preservatives since specimens may have to be preserved for considerable periods of time before being analysed. While few studies of the efficacy of the various preservation methods have fo- 196 cused on collections of bees, other insect groups have been well represented, but many of these studies focused only on DNA preservation, or morphology. King and Porter (2004) determined that 95% ethanol or 95% isopropanol were ideal for preserving ants for card-point mounting and through a literature review concluded that DNA was best preserved in 95% ethanol. Quicke et al. (1998), studying wasps of the superfamilies Ichneumonoi- dea and Chalcidoidea, had an 80% success rate at PCR amplification of 285 rDNA from specimens preserved in 70% ethanol at room temperature. Austin and Dillon (1997) also suggested that chemical drying methods could be used to generate suffi- cient quantities of quality DNA from ichneumonoid and chalcidoid wasps. In this report we present our findings on the relative efficacies of several storage meth- ods of bees including ultra cold freezing, various alcohols, and pure dimethyl sulf- oxide (DMSO) in terms of the quality of DNA preserved for downstream applica- tions such as PCR, as well as the ability to preserve delicate morphological features that are required for taxonomic identifica- tion. We analyzed 121 individuals repre- senting three families, Halictidae, Apidae, and Andrenidae, as part of several larger studies of bee diversity. To determine the quality of the DNA that was extracted using each of our preservation methods, three amplicons within the mitochondrial- ly encoded gene, cytochrome oxidase subunit I (COI), a gene widely used in insect systematics, were compared. While the majority of the specimens collected for this study were part of a pre-planned experi- ment, we supplemented the data using older, preserved specimens that had been previously collected in the course of other projects. MATERIALS AND METHODS Most of the bee specimens used in this study were collected in southern Ontario, Canada and Maryland, USA, using pan JOURNAL OF HYMENOPTERA RESEARCH traps containing a mixture of water and blue Dawn® dish detergent. Pan traps were placed approximately 10m apart along transects and were subject to both sun and shade conditions for between 6 and 24 hours. Bees were identified and then transferred to methanol (50 or 95% in water), ethanol (50, 70 or 95% in water), an ethanol-methanol solution (70:30, 95:5), or pure DMSO. Bees were stored at room temperature for between one and twelve months after which they were pinned and stored at room temperature until DNA was extracted. In addition to these specimens, we also analyzed specimens that were live caught in nets. These included eight Xylocopa virginica that were killed by freezing at —80°C, four Lasioglossum marginatum caught in Greece and stored in ethanol since 1994, and 24 Halictus poeyi caught in Florida and stored in 70% ethanol since 1982. These latter bees, as well as those stored in DMSO, remained in preservative, or frozen, until DNA was extracted. Prior to DNA extraction, bees were dried over- night at room temperature to remove any remaining preservative. Bees preserved in DMSO were washed with 95% ethanol, and then allowed to dry overnight at room temperature. DNA was extracted using either the Qiagen DNeasy Tissue Kit (insect protocol) or the Sigma-Aldrich GenElute Mammali- an Genomic DNA Purification Kit, follow- ing the manufacturers’ instructions. The quality of the extracted DNA was assessed by agarose gel electrophoresis of 5 uL of sample on 1.2% agarose gels in TAE buffer containing 10 mg/ml ethidium bromide. Total DNA extracted from each specimen was quantified using a Beckman DU-530 spectrophotometer. DNA extractions were based on whole specimens, with the exception of X. virginica, a very large bee species, for which a single leg was used. The effect of body size on the amount of DNA extracted from bees of different species was statistically controlled by VOLUME 17, NUMBER 2, 2008 measuring the average head width of a set of pinned specimens (in halictid and xylocopine bees, head width is highly correlated with body mass). Polymerase chain reaction (PCR) was carried out to determine the quality of the preserved DNA. COI amplicons were amplified using the primers Ron (C1-J- 1751), Nancy (C1-N-2191), Jerry (C1-J- 2183), and Pat (L2-N-3014) (Simon et al. 1994). The first PCR reaction was per- formed with the Ron-Pat primer pair generating an amplicon of 1264 bp. Upon successful amplification of the 1264 bp amplicon, specimens were removed from any further attempts at amplification. For those specimens that did not successfully amplify using the Ron-Pat primers, a second PCR reaction was performed on 1:100 dilutions of the Ron-Pat PCR product and used either the Ron-Nancy or Jerry-Pat primers. The quality and quantity of PCR ampli- fication products were analyzed by agarose gel electrophoresis as described above. Successful amplifications were scored us- ing a system based on the brightness of the bands present on the gel. A score of ‘0’ was given for no amplification, a score of ‘1’ was given for weak amplification not containing enough DNA to be sequenced, and a score of ‘2’ was given for a bright band that would contain a sufficient quantity of DNA for sequencing (note that our lab has successfully used this scoring system for DNA sequencing for more than 5 years). Images were photographed on the Bio-Rad Gel Doc 1000 using Multi-Analyst (Bio-Rad) software. RESUETS Qualitative Observations Specimens that were preserved in alco- hol-based solutions were dehydrated, brit- tle, and easily damaged when handled. Bees preserved in DMSO were less brittle than those stored in alcohol, but many of the morphological characters that are typ- 197 ically used for identification were distort- ed; for instance, wings were shrivelled (Fig. 1). Several halictids that were stored in DMSO, notably the augochlorine bees, changed colour from bright green to reddish gold, although their natural green colour was restored after washing them in ethanol. These morphological changes proved problematic for those specimens that had not been identified prior to preservation. DNA Quality All preservation methods tested pro- duced DNA of varying quality and con- centration (Fig. 2). Pure DMSO was most successful at maintaining genomic quality (the brightest genomic band is Lane 8 in Fig. 2), whereas ethanol and methanol- preserved bees produced weak genomic DNA bands. All three methanol treat- ments, as well as the 50% ethanol and ethanol-methanol blends, showed signs of DNA degradation indicated by extensive smearing. Several factors could affect both the amount and quality of the DNA from our specimens including species, collection method, preservative, and specimen size (Table 1). An ANCOVA analysis showed that specimen size was the strongest contributor to the DNA _ quantity (F=30.03, df=1, p<0.0001) and preserva- tive type had only a slight effect (F=2.10, df=7, p=0.0568). Other potential influenc- es (type of DNA extraction kit and time in preservative) had non-significant effects on the amount and quality of DNA obtained from specimens. For systematics projects, the amount of DNA obtained from specimens is less important than how well the DNA can be PCR-amplified. DNA concentration and preservative type had strong effects on PCR amplification scores; the highest am- plification scores were for specimens that were caught live and immediately frozen in a —80°C freezer, despite the low amount of total DNA that was recovered. The best 198 JOURNAL OF HYMENOPTERA RESEARCH Fig. 1. Pictures of Augochlorella striata that were preserved in 100% DMSO (left column) and 100% ethanol (right column). The top row indicates the colour changes that are apparent in the DMSO preserved bees. The bottom row shows how DMSO dehydrates the wings causing them to become misshapen. 3000 500 Fig. 2. Representative samples of extracted DNA from various preserved specimens separated on 1.2% agarose stained with ethidium bromide. Lanes contain the following: (1) 95% methanol, (2) 50% methanol, (3) 50% ethanol, (4) 70% ethanol, (5) 95% ethanol, (6) 70:30 ethanol-methanol, (7) 95:5 ethanol-methanol, (8) 100% DMSO, and (9) —80°C. 2.5 ug of a 100 bp size marker (Fermentas) is loaded in each of the outside lanes. amplification scores from a liquid preser- vative came from those bees preserved in DMSO, followed by those preserved in 95% ethanol (lower DNA concentrations produced higher amplification scores). As a final note, we also attempted to extract and amplify DNA from 28 speci- mens that had been stored in ethanol (probably 70% ethanol) for 10-24 years. These comprised four Lasioglossum margin- atum specimens that were collected in Greece in 1994, as well as 24 Halictus poeyi that were collected in Florida in 1982. For three of the four L. marginatum, we were able to amplify only the smallest of the COI amplicons (using primers Ron and Nancy), with amplification scores of 1. Attempts at recovering DNA from the much older (and VOLUME 17, NUMBER 2, 2008 Table 1. each of the preservative types tested. 199 Mean DNA concentration, proportion of specimens scoring ‘2’, and amplification score ranks for Average DNA No. specimens Collection concentration with amplification method Preservative No. specimens (ug/mL) (SD) scores of 2 Pan traps 50% Methanol 10 138.9 (141.1) 0 95% Ethanol 69.4 (26.6) 1 50% Ethanol 93.8 (52.7) i) 70% Ethanol 23 50.8 (18.9) 10 95% Ethanol 10 56.3 (25.0) 8 Ethanol:methanol blend 70:30 8 104.1 (60.5) 3 Ethanol:methanol blend 95:5 10 89.6 (59.0) 3 DMSO 8 104.3 (44.3) 6 Live caught = 3G 8 17.0 (10.9) 8 more poorly preserved) H. poeyi specimens were completely unsuccessful. DISCUSSION The ideal preservative for field collec- tions of bees and other insect specimens should be easy to use, cost efficient, and easily transportable. Typically, primary alcohols have been used to meet these requirements, but recent studies have ex- amined the use of propylene glycol (Rubink et al. 2003, Vink et al. 2005), acetone (Fukatsu 1999), and other commercial products (Vink et al. 2005). Preserving the quality of both genomic and mitochondrial DNA is of great importance for conducting molecular studies. We found that ultra cold freezing was the best method for killing and preserving specimens, but this is often impractical either because specimens can- not be captured alive or because ultra cold freezing facilities are not available. The use of ultra cold freezing has been suggested by Reiss et al. (1995) and Dillon et al. (1996). In the later study, ultra cold storage of parasitic wasps did not affect the amount of DNA that could be recovered or success- fully amplified. Among the liquid preservatives DMSO serves as an ideal candidate for denaturing DNA damaging enzymes and for preserv- ing the quality of genomic and mitochon- drial DNA. The drawback to using DMSO as a preservative is that it distorts mor- phological characters required for identifi- cation of specimens although this problem can be overcome by identifying specimens prior to storage in DMSO. Furthermore, DMSO is considerably more expensive than ethanol. Despite the finding that primary alcohols caused advanced signs of genomic degradation, the best alterna- tive appears to be 95% ethanol. It is relatively inexpensive, easy to transport, and does not distort morphological char- acters to the same degree as DMSO. Preservative type and concentration, as well as storage time, affect the quality as well as the quantity of DNA that can be extracted from a given specimen. Addi- tionally, some methods of preservation have adverse effects on morphological characters that need to be preserved for specimen identification. In this paper we propose that 95% ethanol is the best chemical preservative for maximizing the quantity and quality of DNA, as well as for maintaining morphological integrity when ultra-cold freezing is not immediately available. ACKNOWLEDGEMENTS Special thanks to T. Chessar and H. Ginsberg for reviewing the early stages of this manuscript, and S. Rehan for bee measurements. Funding was provided by Brock University, NSERC, USGS. 200 LITERATURE CITED Austin, A. and N. Dillon. 1997. Extraction and PCR of DNA from parasitoid wasps that have been chemically dried. Australian Journal of Entomology 36: 241-244. Dillon, N., A. Austin, and E. Bartowsky. 1996. Comparison of preservation techniques for DNA extraction from hymenopterous insects. Insect Molecular Biology 5: 21-24. Fukatsu, T. 1999. Acetone preservation: A practical technique for molecular analysis. Molecular Ecol- ogy 8: 1935-1945. . Kilpatrick, C. W. 2002. Noncryogenic preservation of mammalian tissues for DNA extraction: An assessment of storage methods. Biochemical Ge- netics 40: 53-62. King, J. R. and S. D. Porter. 2004. Recommendations on the use of alcohols for preservation of ant specimens (Hymenoptera, Formicidae). Insectes Sociaux 51: 197-202. Lindahl, T. 1993. Instability and decay of the primary structure of DNA. Nature 362: 709-715. Post, R. J., P. K. Flook, and A. L. Millest. 1993. Methods for the preservation of insects for DNA JOURNAL OF HYMENOPTERA RESEARCH studies. Biochemical Systematics and Ecology 21: Soe. Quicke, D. L. J., C. Lopez-Vaamonde, and R. Belshaw. 1999. Preservation of hymenopteran specimens for subsequent molecular and morphological study. Zoologica Scripta 28: 261-267. Rubink, W. L., K. D. Murray, K. A. Baum, and M. A. Pinto. 2003. Long term preservation of DNA from honey bees (Apis mellifera) collected in aerial pitfall traps. The Texas Journal of Science 55: 159-169. Simon, C., F. Frati, A. Beckenbach, B. Crespi, H. Liu, and P. Flook. 1994. Evolution, weighting, and phylogenetic utility of mitochondrial gene se- quences and a compilation of conserved poly- merase chain reaction primers. Annals of the Entomological Society of America 87: 651-701. Tayutivutikul, J., W. Pongprasert, L. A. Royce, and K. Ruangrit. 2003. Comparison of preservation tech- niques for silkworm (Bombyx mori L.) DNA based on polymerase chain reaction (PCR) products. Chiang Mai University Journal 2: 107-114. Vink, C. J., S. M. Thomas, P. Paquin, C. Y. Hayashi, and M. Hedin. 2005. The effects of preservatives and temperatures on arachnid DNA. Invertebrate Systematics 19: 99-104. J. HYM. RES. Vol. 17(2), 2008, pp. 201-209 Sex Determination and Genome Size in Catolaccus grandis (Burks, 1954) (Hymenoptera: Pteromalidae) N. M. BARCENAS, N. J. THOMPSON, V. GOMEZ-TOVAR, J. A. MORALES-RAMOS AND J. S. JOHNSTON (NMB) Heritage University, 3240 Fort Rd, Toppenish, WA 98948, USA (NJT) Department of Biology, Texas A&M University 3258, College Station, TX 77883, USA (VGT) SENASICA-SAGARPA Insurgentes Sur 498, 06100 México, D.F. (JAMR) USDA-ARS National Biological Control Laboratory, Biological Control of Pests Research Unit 59 Lee Road Stoneville, MS 38776, USA (JSJ) Department of Entomology, Texas A&M University 2475, College Station, TX 77883, USA Abstract—Complementary sex determination (CSD) is a common form of the haplodiploid sex determination system found in all wasps, ants, and bees (Hymenoptera). Exceptions exist to CSD, but too few have been documented to make phylogenetic conclusions. Males that are homozygous at CSD loci are diploid and often sterile. Any effect that increases homozygosity (inbreeding and small population size) should increase the proportion of diploid males. We use flow cytometry to determine the genome size of males and females of the parasitic wasp Catolaccus grandis (Burks) (Hymenoptera: Pteromalidae) (1 C = 455.4 + 3.4 mb). We then score haploid and diploid males (and females) from populations that were 25% and 50% inbred. None of the 314 males scored were diploid. We conclude that the CSD system is very unlikely to exist in this species and discuss the implications for sex determination systems in the Pteromalidae and other chalcidoids. Evolving independently at least eight and possibly as many as 15 times in mites and insects (White 1973, Mable and Otto 1998) in phylogenic lineages that represent nearly 20% of all animal species (Bell 1982, Bull 1983), haplodiploidy is the sole repro- ductive mode in Hymenoptera, including the pteromalid wasp Catolaccus grandis (Burks, 1954) (Hymenoptera: Pteromali- dae) studied here. The most common system of haplodiploidy is arrhenotokous reproduction, where females develop from fertilized eggs and males from unfertilized eges. The most studied form of arrhenot- oky is the complementary sex determina- tion (CSD) mechanism, whereby a sex locus with multiple alleles determines the sexual development of the offspring (Whit- ing 1943, Beye et al. 2003). In the CSD system, zygotes heterozygous at the CSD locus develop into biparental diploid fe- males; males are produced as uniparental haploids that have one CSD allele. Unipa- rental males are typically produced as unfertilized eggs, but in some cases are produced as zygotes in which one parental genome is excluded post fertilization (Kel- ler et al. 2001, Beukeboom et al. 2007). The CSD form of arrhenotoky fails when the female is fertilized by a male whose single allele at the CSD locus is identical to one of the two CSD alleles that she carries. When this happens, half of her diploid fertilized progeny (all of whom should develop as female) are homozygous for a CSD allele and develop into biparental diploid males. Diploid males were first observed in a single locus CSD (sl-CSD) system by Whiting in Bracon hebetor Say, 1836 (Whiting 1943) but have been ob- served in very many species since (Trent et al. 2006). Diploid males are often sterile, yet lacking the division of chromosomes required to produce haploid sperm, may 202 produce diploid sperm and consequently, sterile triploid females (Whiting 1943, Stouthamer et al. 1992, Cowan and Stahl- hut 2004). Whether in natural populations or in laboratory reared colonies, diploid males and triploid sterile females represent a reproductive cost, affecting fertility and sex ratio (Stouthamer et al. 1992, Wu et al. 2005, see exception in Cowan and Stahlhut 2004). Production of diploid males and triploid females is most apparent when the number of CSD alleles is reduced by population bottlenecks and founder events, or when homozygosity increases as a result of consanguinity. The latter is important, since inbreeding by sib-mating occurs often in hymenopteran species that are solitary or are social parasites (Schrempf et al. 2006). The parasitoid pteromalid wasp, C. grandis, is economically important for use in bio-control of the boll weevil, Anthono- mous grandis (Boheman, 1843) (Coleoptera: Curculionidae), and has been released experimentally for crop management in many areas of the southern U.S.A., includ- ing the Rio Grande Valley, Texas, San Angelo, Texas, and Aliceville, Alabama (Morales-Ramos et al. 1994, 1995a, 1998, Summy et al. 1995, 1997). An ectoparasi- toid, C. grandis lays its eggs on boll weevil larvae (most often the third instar or pupa), greatly suppressing the numbers of the boll weevil and thereby reducing the damage caused to cotton (Summy et al. 1995). As an idiobiont, the female stops host develop- ment with a paralytic venom (Morales- Ramos et al. 1995b) and the emerging larva consumes its paralyzed host. These char- acteristics may give an advantage as an efficient biocontrol agent (Morales-Ramos et al. 1995b). However, production efficien- cy in insect mass rearing of C. grandis and related species can be problematic if sl- CSD is present, because that would neces- sitate a large number of sex alleles to minimize diploid male production. Special measures would need to be taken in colony management to avoid founder events, JOURNAL OF HYMENOPTERA RESEARCH population bottlenecks, and inbreeding leading to a loss of the sex alleles. Economically important hymenopterans are among the best studied, and their sex determination is among the best document- ed. The sI-CSD system is well demonstrated in the red imported fire ant (Solenopsis invicta Buren, 1972) and the honey bee Apis mellifera L., 1758 (Hymenoptera: Apidae); 11 to 19 alleles were determined for the complementary sex determiner (csd) locus in honey bees (Beye et al. 2003), with a smaller number of alleles (8 to 13) in the population of red imported fire ants, that went through a bottleneck when introduced into southern United States (Ross et al. 1993). The occurrence of CSD in the Hymenoptera has been demonstrated in symphytan, ichneumonid, and braconid species, but to date not in chalcidoids (Stouthamer et al. 1992). In several of these groups apparent exceptions exist, specifi- cally in instances where inbreeding has not exposed sl-CSD (Wu et al. 2005). The relative phylogenetic position of pteromalids within the tree of life is largely unknown, although studies are underway to remedy this situation (Castro and Dowton 2005, J. Heraty personal commu- nication). Understanding the kind of sex determination system and the ancestral or derived state of the CSD system should be part of this phylogenetic effort. We assume that CSD is the ancestral mode of repro- duction. There is little published research to prove this, however, especially in the more ancestral taxa (Cook and Crozier 1995). Here we test the hypothesis that a form of the CSD system exists in C. grandis. This represents the second test of CSD in Pteromalidae. We employ a new approach in studying CSD by determining genome size first, and then directly scoring males of consanguineous matings, using flow cy- tometry to score ploidy level. If a CSD system exists in this species, diploid male production, to the extent they survive, will be directly proportional to the level of consanguinity. VOLUME 17, NUMBER 2, 2008 METHODS Live Material Catolaccus grandis from wild-caught in- dividuals were used to initiate a reared population. The lab colony used here was founded with: 6 genomes from El Salvador, Central America (host plant of boll weevil: wild and cultivated cotton), 56 genomes from Tabasco, México (host plant of boll weevil: Hampea nutricia Fryxell (Malva- ceae)), and 15 genomes from Oaxaca, México (host plant of boll weevil: Cienfue- gosia rosei Fryxell (Malvaceae)). All C. grandis in this study were from controlled matings among reared individuals. Crosses A. Sib-mating offspring.—Virgin C. grandis females were mated with a single male. Their offspring were confined in individu- al Petri dishes and allowed to mate (sibling matings). Each female was isolated in a Petri dish 3-4 days after emergence for independent offspring evaluation. Host boll weevil larvae were presented to the C. grandis females encapsulated in Para- film® using the method described by Cate (1987). Twelve Parafilm® encapsulated boll weevil larvae per female per day over five days were provided as hosts. Females that produced only male offspring were dis- carded (presumably they were not fertil- ized and had no opportunity to produce diploid males). Male offspring of fertilized females were prepared for flow cytometry. B. Backcross offspring.—Virgin sibling cou- ples were isolated in Petri dishes, allowed to mate, and were removed after offspring emergence. Families with only male off- spring or without father survival were discarded. Daughters were paired with their fathers to induce mating and allowed to reproduce. Again, offspring with only males were not included in the analysis. Flow Cytometry Individual C. grandis males were pre- pared for flow cytometric analysis of 203 genome size and ploidy level following Johnston et al. (2004). The head and thorax of a C. grandis was placed in a 1.5 ml Kontes Dounce tissue grinder into 1 ml of cold Galbraith buffer (per litre: 4.26 ¢ MgCl, 8.84 g sodium citrate, 4.2 g 3-[N- morpholino] propane sulfonic acid, 1 mL Triton X-100, 20 ug/mL boiled Ribonucle- ase. fy pli 7.2; Galbraith et al. 1983); Chicken red blood cells (CRBCs) were added to act as standards (1C = 1212 Mb; Bennett et al. 2003). To release and isolate nuclei, the head and thorax plus the CRBCs in buffer solution were stroked 15 times with an A pestle and filtered through a 20-um nylon filter. Propidium iodide, an intercalating dye that binds stoichiometrically to DNA and fluo- resces in direct proportion to the amount of DNA present, was added to a final concentration of 50 ppm, and the mixture co-stained in the dark at 4°C for 20- 40 minutes. The mean fluorescence of co- stained nuclei in replicate samples of each sex was quantified using a Coulter Epics Elite (Coulter Electronics, Hialeah, FL) with a laser tuned at 514nm and 300 mW. Individual nuclei separate and pass across the exciting light source in the flow cell of the cytometry, where the nuclei are counted and the fluorescent light emitted from each nuclei is collected and quantified after passing a long pass filter to eliminate any reflected laser light. To avoid counting debris or nuclei with associated cytoplasmic tags, counting was activated by PI fluorescence, and only clean, singlet nuclei with low light scatter were included in the analysis. Genome size was estimated for each male and female C. grandis by first calculating the ratio of the fluorescent intensity of C. grandis nuclei to that of the CRBCs standard. The genome size was then determined by multiplying this ratio by the amount of DNA in the CRBCs. Analysis The genome size of C. grandis is given as the average (+/— standard error) of the 204 estimated genomes. A minimal estimate of the likelihood of scoring a diploid male is based on a binomial distribution, where the expectation of a diploid male is equal to the probability of homozygosity, assuming sufficient numbers of alleles at indepen- dent CSD loci that inbreeding is the only source of homozygosity. RESULTS AND DISCUSSION The genome size of C. grandis was estimated using a chicken red blood cell (CRBC) (1 C = 1212 Mb) standard and measured 1 C = 455.4 + 3.4 Mb (Fig. 1 A, B). As expected, the haploid genome of males was half that of the diploid genome of females. The C. grandis genome is 1.37 times larger than that of the only other pteromalid genome scored to date, N. vitripennis (Walker) (1 C = 332.5) and 3 times larger than the genome of the only scored braconid wasp, Habrobracon juglan- dis (Ashmead, 1889) (1 C = 156.5) (Rasch et al.1975). Catolaccus grandis were easily scored for ploidy level. Females revealed only a diploid peak (Figure 1A). Haploid males showed 2 fluorescent peaks with equal numbers of haploid and diploid (endo- reduplicated) nuclei (Fig. 1 B). This endo- reduplicated peak is observed because of the diploid tissue found in the muscle Johnston et al. 2004). Diploid males (if present) would have shown only one strong diploid peak, as observed in the females. All 124 males from the F-2 gave a strong haploid peak as seen in Figure 2; no diploid males were scored. These results indicate that a CSD system, if it exists, is based on a number of loci in a multilocus complementary sex determination system (ml-CSD). A second set of inbred backcross offspring was produced, collected, and scored in order to estimate more precisely the number of genes involved in a potential ml-CSD system. Backcross offspring were selected because the expected 50% homo- zygosity in the backcross to the haploid male parent reduces the sample size JOURNAL OF HYMENOPTERA RESEARCH needed to detect diploid males (Table 1). All males produced from the backcrosses (190 in total) were haploid. If the CSD system exists for C. grandis, inbreeding should increase homozygosity and result in significant diploid male production. Given haplodiploid sex deter- mination, sib mated and backcross zygotes had a probability of homozygosity at a single locus of 0.25 and 0.50 respectively. The failure to observe diploid males among the 314 scored males indicates that a CSD system, if it occurs at all in C. grandis, involves >5 independent loci. The probability that diploid males will be produced in significant numbers due to sib mating in nature, or due to small effective size in mass rearing colonies is low. A possible source of error in these analyses is cannibalism or other forms of reduced survival of diploid males. Selec- tive cannibalism based on semiochemicals is known to occur in honey bees, where the workers will consume any diploid males (Woyke 1963). A parasitoid of the codling moth, Liotryphon caudatus (Ratzeburg, 1848) (Hymenoptera: Ichneumonidae) with known cannibalism produces a significant diploid male population (T. R. Unruh, personal communication). This suggests, that in some haplodiploid species, canni- balism does not necessarily lead to selec- tive killing of diploid males. In culture, C. grandis females lay eggs either on the boll weevil larvae or on the interior surface of the Parafilm® capsule where the host is presented as Parafilm® encapsulated lar- vae. In the rearing conditions here, the female is expected to lay 1, 2, or 3 eggs per host. An emerged C. grandis larva will move around and eat any other eggs or larvae that it encounters (Morales-Ramos and Cate 1992), and rarely are two para- sitoid pupae found in the same host. Two sources of data suggest that diploid male mortality cannot explain the absence of diploid males in our study. Fully 44.4% of hosts receive a single egg, and the VOLUME 17, NUMBER 2, 2008 205 Number of nuclei Fig. 1. Female 2C Female 4C Male 1C CRBC 2C Male 2C Male 4C 200 400 600 800 1000 Relative Fluorescence (channel number) Relative DNA staining in nuclei from the head of A) female Catolaccus grandis plus chicken red blood cells (CRBCs) and B) haploid male C. grandis plus CRBCs. Neural tissue produces the first 1 C peak in haploid males. DNA in the nuclei of muscle is largely endoreduplicated in haploid males (Johnston et al., 2004) resulting in the diagnostic 1 C, 2 C and small 4 C peaks shown here. Relative DNA amount is calculated as the product of the ratio of the mean fluorescence of the leftmost peak (1 C in haploid males, 2 C in females) /mean fluorescence of CRBC standard multiplied by the estimated genome size of the CRBC standard (1140 Mb; Bennett et al. 2003). JOURNAL OF HYMENOPTERA RESEARCH ~ Diploid (1C) Females 117 156 196 235 274 313 352 391 430 469 509 548 206 8 Haploid (1C) Males ae) = @) £ O o To) = = z= 0.253978 Genome Size (Mb) Fig. 2. Relative genome sizes of individuals of haploid males and diploid females. remainder (55.6%) receive two or more eggs (Fig. 3). Even under the assumption that diploid males from single egg hosts are the only ones that survive cannibalism, a significant number of diploid males in a CSD system would be expected. Addition- al evidence that diploid males should have survived cannibalism is supported by the observation that there was no deviation of the sex ratio from what was expected. Under lab conditions, the sex ratio ob- served for the sib-matings was approxi- mately 2.8:1. The previously published sex ratio in natural populations is 3:1 or 4:1 Table 1. Sample size needed to detect a diploid male in F2 and backcross offspring (BC), given a sex determining system with one to five genes involved (ml-CSD) and a 4:1 female: male sex ratio in fertilized females under random mating based on binomial distribution. F2 BC Number of loci P=95% P=99% P=95% P=99% 1 10 13 4 6 2 18 Di, 6 9 3 66 101 10 15 = 258 396 18 Di, 5 1025 1575 15) 76 (Morales-Ramos and Cate 1992). A reduc- tion in sex ratio (more surviving males) (1.2:1) was observed in the backcrosses, but this could be related to the age of the father. Gomez et al. (1997) exposed a virgin female every day during the life span of C. grandis males and observed a reduction over time in the ability of males to inseminate females. Inbreeding, because it increases the chance of homozygosity and thus increases the appearance of diploid males, reduces the fitness of species with sl-CSD. In species with proven CSD, the effect of population bottlenecks and founder events that decrease the number of alleles and increase inbreeding may be reduced by modification of the CSD system, including inbreeding avoidance behaviors, and the development of a non-CSD system (Cook and Crozier 1995, Dobson ard Tanouye 1998). One modification of CSD is to involve multiple loci, although the sex determination mechanism remains the same: females are heterozygous at one or more loci, while full fertility is expected only in haploid males where each CSD allele is present as a single copy (Paxton et VOLUME 17, NUMBER 2, 2008 Number of Parasitoids 1 2 207 (7.6%) (2.9%) (3.6%) 4 5 >5 Number of eggs per host Fig. 3. The number and percentage (in parenthesis) of 721 hosts parasitized by 1, 2, 3, 4 and 5 or more eggs. al. 2000). In ml-CSD, the chance of a homozygous diploid male is greatly re- duced, even in inbreeding populations. Inbreeding effects have not been well tested experimentally for many Hymenop- tera, however, and it can be argued that ml-CSD has not been persuasively demon- strated (Beukeboom et al. 2007), leaving the as best-known mechanism for inbreeding avoidance in the presence of CSD, the premating dispersal observed in Bracon hebetor (Ode et al. 1995). Because we experimentally controlled matings, mecha- nisms to avoid inbreeding were not an issue, and even an ml-CSD system is unlikely. Single locus CSD has been demonstrated for at least 60 species of hymenopterans (Beukeboom et al. 2007), including most social species that have been studied (see exception in Schrempf et al. 2006). How- ever, the CSD system is not the only mechanism for arrhenotokous reproduc- tion. The absence of sl-CSD system has been conclusively demonstrated in Nasonia vitripennis (Walker, 1836) (Hymenoptera: Chalcidoidea) and the parasitoid Hetero- spilus prosopidis Viereck, 1910 (Hymenop- tera: Braconidae) (Wu et al. 2005). It is also absent in the more distantly related ant Cardiocondyla obscurior Wheeler, 1929 (For- micidae), which shows no evidence of diploid males but does show inbreeding depression (Schrempf et al. 2006). Pro- posed alternative sex determination mech- anisms in arrhenotokous insects include genomic imprinting, fertilization sex deter- mination, genic balance sex determination, and maternal effect sex determination, which were all tested in populations of N. vitripennis (Dobson and Tanouye 1998, Beukeboom et al. 2007). The genomic imprinting model fits well for N. vitripennis (Trent et al. 2006) and may also be present in related Chalcidoidea. Flow cytometry allowed us to quickly score for diploid male production, and we showed that the CSD model is unlikely to be the sex determination mechanism for the pteromalid, C. grandis. These results, along with fairly compelling evidence to reject CSD in one Trichogrammatidae (Chalcidoidea) species (Stouthamer and Kazmer 1994), raise the question of wheth- er the CSD model arose independently or in a common ancestor of these species. 208 LITERATURE CITED Bell, G. 1982. The Masterpiece of Nature: The Evolution and Genetics of Sexuality. University of California Press, Los Angeles. Bennett, M. D., I. J. Leitch; Hoyt Briceand J--S: Johnston. 2003. Comparisons with Caenorhabditis (~100 Mb) and Drosophila (~175 Mb) using flow cytometry show genome size in Arabidopsis to be ~157 Mb and thus ~25% larger than the Arabi- dopsis initiative estimate of ~125 Mb. Annals of Botany 91: 547-557. Beukeboom, L. W., A. Kamping, and _L. van de Zande. 2007. 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PULAWSKI (SKG) Northern Regional Station, Zoological Survey of India, 218, Kaulagarh Road, Dehradun, 248 195, India; email: skgupta48@yahoo.co.in (SFG) Area de Zoologia, Facultad de Biologia, Universidad de Salamanca E-37071, Salamanca, Spain; email: gayubo@usal.es (WJP) Department of Entomology, California Academy of Sciences, 55 Music Concourse Drive, San Francisco, California 94118, USA; email: wpulawski@calacademy.org Abstract.— Mellinus obscurus Handlirsch, 1888, currently recognized as a subspecies of arvensis (Linnaeus, 1758), is restored to full species status based on the unusual structure of its antennal socket. Mellinus orientalis is described from India and Nepal; its main diagnostic characters are a finely sculptured propodeal enclosure and a carina separating the propodeal posterior surface from the side. The following are the abbreviations used in the text below: CAS: California Academy of Sciences, San Francisco, California, USA USAL: Area de Zoologia, Facultad de Biologia, Universidad de Sala- manca, Spain ZSI: Northern Regional Station, Zoological Survey of India, Dehradun, India. Mellinus obscurus Handlirsch, species status restored Mellinus obscurus Handlirsch, 1888:289, E. Ho- lotype: E, Korea: no specific locality (KRA- KOW). — Dalla Torre, 1897:561 (in catalog of world Sphecidae); Maidl and Klima, 1939:39 and 43 (in catalog of world Astatinae and Bembicinae); Yasumatsu, 1943:2 (China: He- bei Province, Inner Mongolia, description of I); Tsuneki, 1965:26 (in key to Bembicinae of Japan and Korea): Haneda, 1968:46 (Japan); Tano, 1968:33 (Japan); Tsuneki, 1969a:18 (Japan: Sapporo area: nesting habits, prey,), 1969b:26 (Japan: specimens in Osaka Muse- um); Tsuneki, 1969c:64 (Japan: Mount Hyo- nosen); Nambu, 1973:152 (Japan: Saitama Prefecture); Suda, 1973:123 Japan: Yamana- shi Prefecture); Siri and R. Bohart, 1974:170 (in key to world Mellinus), 174 (in review of world Mellinus); Nambu, 1975:72 (Japan: Saitama Prefecture); R. Bohart and Menke, 1976:449 (listed); Kazenas, 1980:84 (first rec- ord from Russia: Primorskiy Kray and Kuril Islands); Tsuneki, 1982:18 (known from Korea; as obscrus), 1982b:36 (first record from Taiwan: Chiay Prefecture: Mount Ali); Miyatake, 1996:103 (specimens in Hiroshi Aoki collec- tion). - As Mellinus arvensis obscurus: Nem- kov in Nemkov, Kazenas, Budrys, et Antro- pov, 1995:455 (new status, in key to Sphecidae of Russian Far East); Nemkov, 2005:157 (Russia: Sakhalin Island), 2006:169 (Russia: Primorskiy Kray: Kedrovaya Pad’ Nature Reserve); nec Boesi, Polidori, Gayubo, Tormos, Asis, and Andrietti, 2007:184 (= Mellinus orientalis); Nemkov, 2007:74 (Rus- sia: Kuril Islands: Iturup and Kunashir Is- lands), 2008:20 (in key to Mellinus of Russia). Mellinus tristis Pérez, 1905:156, E. Holotype or syntypes: E, Japan: no specific locality but presumably Tokyo area (MNHN). Synony- mized with Mellinus obscurus by Tsuneki, VOLUME 17, NUMBER 2, 2008 1965:26. — Pérez, 1905:26 (listed); Maid] and Klima, 1939:42 (in catalog of world Astatinae and Bembicinae); Tsuneki, 1946:85 (prey records). — As Mellinus obscurus tristis: Yasumatsu, 1943:3 (comparison with Mellinus obscurus); Maruyama, 1948:7 (nesting habits); Tsuneki and Shimoyama, 1963:48 (Japan: Towada Prefecture). We have not seen the type of Mellinus obscurus, but there is little doubt about the interpretation of this species, as it is the only member of the genus that occurs in the Asian Far East. It was described as a full species, but differentiated from arvensis only by chromatic characters (Handlirsch, 1888; Siri and Bohart, 1974; Nemkov et al., 1995, 2008). In arvensis the mesopleuron, scutellum, gastral terga Il, III and V (female) or VI (male) are marked with yellow, and the hindtibia is yellowish brown, whereas these body parts, except tergum III (and occasionally other terga), are black in obscurus. In addition, arvensis occurs in Europe, Turkey, Kazakhstan, and east to the Irkutsk area and Altai Mts. in Siberia, while obscurus is known from the Russian Far East, Korea, Japan, China (Hebei Province and Inner Mongolia), and Taiwan. Because the differences between arvensis and obscurus were in color only, and because they appeared to be vicariant species, Nemkov et al. (1995) downgraded obscurus to a subspecies of arvensis. Melli- nus obscurus, however, strikingly differs from all its congeners in having the antennal socket with an overhanging fron- tal lobe (compare Figs 1a, b and 2a). In our opinion, this difference alone suffices to treat obscurus as a full species. We have examined 7 9, 6 ¢ from Japan and 19 and 1 ¢g from Kuril Islands, Russia. Mellinus orientalis Gupta, Gayubo, and Pulawski, sp. nov. Mellinus sp.: Gupta, 1997:102 (first record of Mellinus from Oriental Region). As Mellinus arvensis obscurus: Boesi, Polidori, Gayubo, Tormos, Asis, and Andrietti, 211 2007:184 (Nepal; nesting habits, adult mor- phology, description of larva). Name derivation.—Orientalis, a Latin mas- culine and feminine adjective meaning Orien- tal; with reference to this species distribution. Taxonomic history—Gupta (1997) first recorded Mellinus from the Oriental Re- gion, but he determined his specimens to genus only. Boesi et al. (2007) examined 17 females from Nepal, comparing them to the European Mellinus arvensis and the Japanese M. arvensis obscurus. They dis- cussed several sculptural, setal, and chro- matic characters, but not the antennal socket nor the pygidial plate, and conclud- ed that the Nepalese specimens were conspecific with arvensis obscurus. We consider all these specimens to represent a distinct new species, Mellinus orientalis. Diagnosis.—As in the Palearctic arvensis (Linnaeus), crabroneus (Thunberg), obscurus Handlirsch, and also the Mesoamerican costaricae (Bohart) recently transferred to Mellinus from Trachogorytes by Pulawski (2007), orientalis has a well-defined carina that separates the propodeal posterior (oblique) surface from the side. It differs from these four species by five characters: 1. its propodeal enclosure is finely rugose on a narrow median zone (rather than conspicuously rugose on a large portion of the enclosure, compare Figs. 1c and 2b), 2. the gaster is all black or tergum III has a pair of lateral pale spots (at least tergum II has pale spots in the other four species), 3. the female pygidial plate is punctate over more than half its length and ridged only apically (rather than punctate basally and ridged over more than half of its length, compare Figs. le and 2d), 4. male flagellomeres VI- IX each has a narrow, almost linear placoid (rather than a broadly elliptical placoid; male unknown in costaricae), and 5. gono- coxite narrowed apically (rather than con- spicuously broad (compare Fig. 1f and 2e). Also, the erect setae on tergum I are shorter in orientalis than in arvensis and obscurus (compare Figs. 1d and 2c). 212 Big, A Description.—Clypeal free margin with three well-defined teeth. Tentorial pit closer to antennal socket than to inner eye orbit (0.7:1.0) in female, equidistant in male. Frontal and scutal punctation slight- ly finer than in arvensis. Mesopleuron punctate. Propodeal enclosure microscopi- cally areolate and with finely rugose JOURNAL OF HYMENOPTERA RESEARCH Mellinus obscurus Handlirsch: a — female head in dorsal view showing antennal sockets; b — male head in dorsal view showing antennal sockets; c — propodeal enclosure of female; d — female tergum I in lateral view; e — pygidial plate of female; f — male genitalia in dorsal view. median area (Fig. 2b) that is slightly narrower than midocellar width (the ru- gose area is no longer than midocellar width in some specimens, and extends to about enclosure midlength in others); propodeal side punctate, unsculptured anteriorly, separated from _ posterior (oblique) surface by longitudinal carina VOLUME 17, NUMBER 2, 2008 0.2mm Fig. 2. Mellinus orientalis Gupta, Gayubo, and Pulawski: a — female head in dorsal view showing antennal sockets; b — propodeal enclosure of female; c — female tergum I in lateral view; d — pygidial plate of female; e — male genitalia in dorsal view. that starts about two midocellar widths behind propodeal spiracle; posterior sur- face punctured. Tergum I narrow, its maximum width 2.0 X basal width (mea- sured just behind gastro-propodeal articu- lation) in female, 1.7 X in male. Setae erect on dorsum of peduncle of tergum I, markedly shorter than those on sternum I (setal length about 0.5 x mid- ocellar width, Fig. 2c). Head, thorax, propodeum, and gaster black except the following are pale yellow: narrow paraorbital stripe (extending to about orbit midheight), scapal venter, ventral half of clypeus in Nepalese speci- mens (clypeus all black in Indian male and 214 with two admedian and two small lateral spots in Indian female), and pair of spots on pronotal collar in some Nepalese specimens. Legs black except inner fore- tibial surface pale yellow in Nepalese specimens, partly yellowish brown in Indian female, and dark brown in Indian male; and apical tarsomeres yellowish brown to brown. Tergum III with pair of lateral pale spots in most Nepalese speci- mens (all black in two). Q. — Pygidial plate punctate over more than half its length, ridged on remaining apical portion (Fig. 2d). Length: 9.5- 12.0 mm. 3. — Flagellomeres VI-IX each with narrow, almost linear placoid. Genitalia: Fig. 2e. Length 7.5 mm. Geographic distribution.Northern India, Nepal. . Records. —HOLOTYPE: 9, INDIA: Uttarak- hand: Dwali in Almora District, 2734 _m, 31 Aug 1990, P.C. Tak & party (ZSI: NRS/ZSI/ A9837). PARATYPES: INDIA: Himachal Pradesh: Dalhousie, 2132 m, 17 Aug 1972, Gulati (1 J, ZSI: NRS/ZSI/A9838); Narkanda [ca 2700 m], 21 June 1972, Mayank (1 3, CAS). Uttarakhand: between Dwali and Phurkia in Almora District, 2,734-3,260 m , 1 Sept 1990, P.C. Tak & party (1 Q, CAS). NEPAL: Eastern Region: Solu Khumbu District: Sagarmatha National Park at 27°45’— 28°07'N 86°28'-87°07'E, Roberto Boesi, 30 May 2003 (1 9, USAL); 26 June 2003 (2 9, CAS; 3 9, USAL), 30 June 2003 (1 9, USAL), 6 July 2003 (7 0, USAL); 10 July 2003 9, CAS; 20, USAL), ACKNOWLEGMENTS We thank Arnold S. Menke and Robert L. Zuparko for their critical reading of the manuscript and many excellent suggestions. April Nobile generated illustra- tions using Auto-Montage. LITERATURE CITED Bohart, R. M. and A. S. Menke. 1976. Sphecid Wasps of the World. A generic revision. University of Cali- fornia Press, BerkeleyLos AngelesLondon, 1 color plate, IX + 695 pp. Boesi, R., C. Polidori, S. F. Gayubo, J. Tormos, J. Asis, and F. Andrietti. 2007. Nesting biology, morpho- logical remarks, and description of the mature larva of Mellinus arvensis obscurus (Hymenoptera: JOURNAL OF HYMENOPTERA RESEARCH Crabronidae) in Nepal. Florida Entomologist 90: 184-190. Dalla Torre, C. G. 1897. 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The Life Study (Fukui) 12, 42-48 (in Japanese). Maidl, F. and A. Klima. 1939. Pars 8: Sphecidae I. (Astatinae — Nyssoninae). In: Hedicke, H. ed. Hymenopterorum Catalogus. Dr. W. Junk, Verlag fiir Naturwissenschaften, ’s-Gravenhage. 150 pp. Maryuama, K. 1948. Habits of a Diptera-hunter, Mellinus obscurus tristis Pérez (Hymenoptera). Mushi 19: 7-11. Miyatake, Y. 1996. A list of the insect collection by Mr. Hiroshi Aoki. Special Publications from the Osaka Museum of Natural History 28: 1-132 (in Japanese). [Sphecidae on pp. 101-103]. Nambu, T. 1973. Saitama-ken d6butsushi kiso chésa. Rikusei kontyt rui chésa hékoku. Makushi moku — Hymenoptera [= Studies on the Saitama Prefecture fauna. A report on the terrestrial entomofauna. Order Hymenoptera]. Saitama-ken Déobutsushi Kari-no Moku Roku [=Preliminary Re- ports on the Saitama Prefecture Fauna] 3: 147-154 (in Japanese). . 1975. Kontyai-ko makushi moku Saitama-no hati [= Wasps of Saitama (Insecta, Hymenop- tera)]. 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Rauna royushchikh os (Hymenoptera: Sphecidae, Crabronidae) Kurilskikh ostrovov — The digger wasp fauna (Hymenoptera: Spheci- dae, Crabronidae) of Kuril Islands. Euroasian Entomological Journal 6: 67—76 (in Russian). . 2008. A key to the digger wasps of the genus Mellinus (Hymenoptera, Crabronidae, Mellininae) of the Russia. Far Eastern Entomologist 185: 19-20. , V. L. Kazenas, E. R. Budrys, and A. V. Antropov. 1995. Nadsem. Sphecoidea. 67. Sem. Sphecidae — Royushchiye osy. Pp. 368-480 in: Lehr, P. A. ed. Opredelitel’ nasekomykh Dal’nego Vostoka Rossii v shesti tomakh. Tom IV. Setchatokry- lyie, skorpionnitsy, pereponchatokrylyie. Chast’ 1. Nauka, Sankt-Peterburg. 604 + 2 unnumbered pp. (in Russian). Pérez, J. 1905. Hyménopteres recueillis dans le Japon central, par M. Harmand, ministre plénipoten- tiaire de France a Tokio. Bulletin du Muséum d’Histoire Naturelle 1905: 23-39, 79-87, 148-158. Pulawski, W. J. 2007. 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Dasch, 1925-2007 ANDREW BENNETT, DAVID WAHL, AND IAN GAULD (AB) Canadian National Collection of Insects, Ottawa, Ontario (DW) American Entomological Institute, 3005 SW 56™ Ave., Gainesville, FL 32608-5047, USA; email: aei@aei.cfcoxmail.com (IG) The Natural History Museum, London, UK Dr. Clement E. Dasch, 82, of New Concord, Ohio, died Nov. 29, 2007. He was a leading authority of Nearctic Ich- neumonidae for over 40 years. Dr. Dasch was born Nov. 28, 1925 in Steubenville, OH and was a veteran of World War II, having served as a mortarman with the U.S. Army in France and Germany. He was a recipient of the Purple Heart. | He obtained his bachelor and doctorate degrees from Cornell and was a professor of biology at Muskingum College, New Concord for 37 years, retiring in 1990. He was a long-time member of the Entomo- logical Society of America and the Interna- tional Society of Hymenopterists, as well as a Director of the American Entomological Institute. He is survived by his wife Betty, four sons, nine grandchildren and his brother Lawrence. In terms of contributions to Hymenop- tera, Clement Dasch published 10 works on Ichneumonidae from 1958 to 1992. Seven of these were revisionary memoirs of more than 300 pages each. A summary of his major studies is presented in Table 1. Dr. Dasch’s work greatly complemented Henry Townes’s higher level studies, and the meticulous and comprehensive nature of his Nearctic revisions contribute to making the Nearctic probably the best known region of the world for Ichneumoni- dae. What is most notable in Table 1 is the proportion of new species that he described, clearly illustrating the dearth of alpha-level knowledge of the family prior to his work. Perhaps most remarkable was his 1988 revision of the 313 Nearctic species of Glypta, of which 269 were new. As this genus is one of the most important for biological control of lepidopterous forest pests, Dr. Dasch thus leaves a legacy not only in taxonomy, but in applied entomology as well. It must be mentioned that this work would not have been possible without the partnership of his wife Betty, who worked tirelessly with him to produce these studies. In addition to the revisionary work, Clement and Betty Dasch were exceptional collectors, amassing over 300,000 parasitic Hymenoptera from 1948-1998. The bulk of the collection (including 236,000 Ichneu- Table 1. Statistics of Clement Dasch’s major ichneumonid studies. Ichneumonid taxon revised by region Year specimens examined species treated species described % new species Neotropical Diplazontinae 1964 n/a Sy 42 81% Nearctic Diplazontinae 1964 16762 101 58 57% Nearctic Mesochorinae 1978 10057 127. 95 75% Neotropical Mesochorinae 1974 2416 277 263 95% Nearctic Anomaloninae 1979 25304 179 111 62% Nearctic Cremastinae 1984 16002 322 257 80% Nearctic Glyptini 1988 21671 SF 271 86% Nearctic Orthocentrinae s.I. 1992 29500 120 88 73% Total 121712 1495 1185 VOLUME 17, NUMBER 2, 2008 PAWS Fig. 1. Clement Dasch visiting the Provancher Collection, Université Laval, Québec in 1981. monidae) was given to the American that he was not a full-time research Entomological Institute in Gainesville, taxonomist, but a professional teacher. Florida and is housed in the Clement and His outstanding contribution towards ich- Betty Dasch room at the facility. One of the neumonid taxonomy was undertaken as an amazing aspects of Dasch’s research was_ extra; a passion that consumed most of his 218 and his wife’s time. They collected prodi- giously, then mounted and labelled and sorted far more specimens than most museum collections processed. Amongst the sciences, taxonomy is perhaps unique in depending on the efforts of a few individ- uals who have a level of dedication that is truly phenomenal. Notable examples are C.P. Alexander in Tipulidae, Walter Roths- child and Karl Jordan in Sphingidae, and of course Henry and Marjorie Townes in Ichneumonidae. Dasch is on a level with these all-time greats: an individual who has made many subfamilies of ichneumonids accessible to North American biologists during an era when taxonomy was a Cinderella subject and funding for system- atics was in global decline. If we are ever going to really know the fauna of planet Earth, society needs to be able to cultivate more individuals like Clement Dasch. His death leaves a gap that is unlikely to be filled in the foreseeable future. CLEMENT E. DASCH PUBLICATIONS Butler, G. D. Jr. and C. E. Dasch. 1958. The Diplazontinae of Arizona (Hymenoptera: Ichneu- JOURNAL OF HYMENOPTERA RESEARCH monidae). Arizona Agricultural Experimental Sta- tion Technical Bulletin 135: 1-10. Dasch, C. E. 1964. The Neotropic Diplazontinae. Contributions of the American Entomological Insti- tute 1: 1-77. . 1964. Ichneumon-flies of America north of Mexico: 5. Subfamily Diplazontinae. Memoirs of the American Entomological Institute 3: 1-304. . 1971. Ichneumon-flies of America north of Mexico: 6. Subfamily Mesochorinae. Memoirs of the American Entomological Institute 16: 1-376. . 1971. Hibernating Ichneumonidae of Ohio (Hymenoptera). Ohio Journal of Science 71: 270-283. . 1974. Neotropic Mesochorinae (Hymenoptera: Ichneumonidae). Memoirs of the American Entomo- logical Institute 22: 1-509. . 1979. Ichneumon-flies of America north of Mexico: 8. Subfamily Cremastinae. Memoirs of the American Entomological Institute 29: 1-702. . 1984. Ichneumon-flies of America north of Mexico: 9. Subfamilies Theriinae and Anomalo- ninae. Memoirs of the American Entomological Institute 36: 1-610. . 1988. Ichneumon-flies of America north of Mexico: 10. Subfamily Banchinae, tribe Glyptini. Memoirs of the American Entomological Institute 43: 1-644. . 1992. Ichneumon-flies of America north of Mexico: 12. Subfamilies Microleptinae, Helictinae, Cylloceriinae and Oxytorinae. Memoirs of the American Entomological Institute 52: 1470. INSTRUCTIONS FOR AUTHORS General Policy. The Journal of Hymenoptera Research invites papers of high scientific quality reporting comprehensive research on all aspects of Hymenoptera, including biology, behavior, ecology, systematics, taxonomy, genetics, and morphology. 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