carrer nny J ated ener fee peter, eiitate Halas Ad igee fa Be 7 . Neco is pen byt es wee te : ‘ : 4 ; : : : Ee Oe ee eee aiadi bed ot as ; : : ; Seana kaih «Se 6 hie ay , : Yad Sorter ; . vata . ; ; ; : arabe ; Valea 3 z cect! oh iia Fee nie tnd oP hw een ad bene ean Ea it a Fete Behe wer. ae Lay n ny “8 Nan ashy Vi AN: ‘y j % \ nN mh A yi ui . ty QL Ho | VOL. 103 EGTX JANUARY 2001 NO. 1 thle (ISSN 0013-8797) at PROCEEDINGS of the ENTOMOLOGICAL SOCIETY of WASHINGTON PUBLISHED CONTENTS ASCHER, JOHN S.—Hylaeus hyalinatus Smith, a European bee new to North America, with notes on other adventive bees (Hymenoptera: Apoidea) ............... 000. cee cce ccc c eee eeeeeen 184 BAKER, GERALD T.—Morphology and distribution of sensilla on the antenna of Omoglymmius americanus (Laporte 1836) (Coleoptera: Rhysodidae) ................0.000 cece cece cece eee eeee 135 BALDRIDGE, CHARLES D. and MATTHEW D. MORAN—Behavioral means of coexisting in old fields by heterospecific arthropod predators (Araneae: Lycosidae, Salticidae; Insecta: Wslcaptcta AC ABADIGAC S|, 25555 cag cas ew SUS Aes Gh Shas Maratea Rh nie URE SE NAS a TORN ad Stich Bee 81 DEWALT, R. EDWARD, DONALD W. WEBB, and THOMAS N. KOMPARE—The Perlesta placida (Hagen) complex (Plecoptera: Perlidae) in Illinois, new state records, distributions, andvanigentification Key esis Soe oe oe coaigaee ok nctocen BSS GER Tee te aU re 207 GAGNE, RAYMOND J., RUBENS A.M. ODA, and RICARDO F. MONTEIRO—The gall midges (Diptera: Cecidomyiidae) of Mikania glomerata (Asteraceae) in southeastern Brazil.... 110 GOEDEN, RICHARD D.—Life history and description of immature stages of Neaspilota achilleae Johnson (Diptera: Tephritidae) on Stephanomeria spp. (Asteraceae) in southern California.... 60 GOEDEN, RICHARD D.—Life history and description of immature stages of Neaspilota footei Freidberg and Mathis (Diptera: Tephritidae) on Aster occidentalis (Nuttall) Torrey and A. Stray \CASteraceae) in southern) California: oo yuc.c0 ah ac te eee Eee NN ae en ae eee he 191 HUANG, YIAU-MIN—A pictorial key for the identification of the subfamilies of Culicidae, gen- era of Culicinae, and subgenera of Aedes mosquitoes of the Afrotropical Region (Diptera: ECL AE Nera ses sya athe cle «Sera bak nie Slag eae ke CaS NC FE) Mea OLE oA We De OT OR te 1 KEIPER, J. B., J. JANNINO, M. SANFORD, and W. E. WALTON—Biology and immature stages of Zypopsilopa nigra (Williston) (Diptera: Ephydridae), a secondary consumer of damaged Stems of wetland monoGotsy is swe nels sso see ta sie eee Oe RIE at Sea eer 89 MARTINEZ M., IMELDA, CUAUHTEMOC DELOYA, and MARCO DELLACASA—Anatomical and functional data on female and male reproductive systems of some dung beetle species of Aphodiinae and Eupariinae of Mexico (Coleoptera: Scarabaeoidea: Aphodiidae) ............ 227 O’NEILL, KEVIN M.—Observations on the behavior of Machimus occidentalis (Hine) and Machimus formosus (Hine) (Diptera: Asilidae) in Montana ................. 000. ee cece eee ees 222 (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON OFFICERS FOR 2001 JOHN W. Brown, President MIcHAEL G. PoGuE, Treasurer GABRIELA CHAVARRIA, President-Elect RONALD A. OcHoA, Program Chair Stuart H. McKamey, Recording Secretary STEVEN W. LINGAFELTER, Membership Chair Ho .uis B. WILLIAMS, Corresponding Secretary Davip G. Furtu, Past President Jon A. Lewis, Custodian Davip R. Situ, Editor Publications Committee RAYMOND J. GAGNE THOMAS J. HENRY WAYNE N. MATHIS Honorary President LoutsE M. RUSSELL Honorary Members KARL V. KROMBEIN RONALD W. HopGEs DoNALD M. ANDERSON WILLIAM E. BICKLEY All correspondence concerning Society business should be mailed to the appropriate officer at the following address: Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Wash- ington, D.C. 20560-0168. MEETINGS.—Regular meetings of the Society are held in the Natural History Building, Smithsonian Institu- tion, on the first Thursday of each month from October to June, inclusive, at 7:00 PM. Minutes of meetings are published regularly in the Proceedings. MEMBERSHIP.—Members shall be persons who have demonstrated interest in the science of entomology. Annual dues for members are $25.00 (U.S. currency). PROCEEDINGS.—The Proceedings of the Entomological Society of Washington (ISSN 0013-8797) are pub- lished quarterly beginning in January by The Entomological Society of Washington. POSTMASTER: Send address changes to the Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Washington, D.C. 20560-0168. Members in good standing receive the Proceedings of the Entomo- logical Society of Washington. Nonmember U.S. subscriptions are $60.00 per year and foreign subscriptions are $70.00 per year, payable (U.S. currency) in advance. Foreign delivery cannot be guaranteed. All remittances should be made payable to The Entomological Society of Washington. The Society does not exchange its publications for those of other societies. PLEASE SEE PP. 771-772 OF THE JULY 2000 ISSUE FOR INFORMATION REGARDING PREPARATION OF MANUSCRIPTS. STATEMENT OF OWNERSHIP Title of Publication: Proceedings of the Entomological Society of Washington. Frequency of Issue: Quarterly (January, April, July, October). Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash- ington, D.C. 20560-0168. Editor: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Books for Review: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 16 January 2001 Periodicals Postage Paid at Washington, D.C. and additional mailing office. PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PROC. ENTOMOL. SOC. WASH. 103(1), 2001, pp. 1-53 A PICTORIAL KEY FOR THE IDENTIFICATION OF THE SUBFAMILIES OF CULICIDAE, GENERA OF CULICINAE, AND SUBGENERA OF AEDES MOSQUITOES OF THE AFROTROPICAL REGION (DIPTERA: CULICIDAE) YIAU-MIN HUANG Department of Entomology, MRC 534, Smithsonian Institution, Washington, DC 20560-0534, U.S.A. (e-mail: huang.yiau-min @nmnh.si.edu) Abstract.—A pictorial key is presented as a field aid for the identification of aduit mosquitoes of 3 subfamilies of Culicidae, 15 genera of Culicinae, and 11 subgenera of Aedes occurring in the Afrotropical Region. Scanning electron micrographs demonstrate the taxonomic characters used in the key. Key Words: Aedes, Afrotropical Region To assist field workers in identifying mosquitoes in Africa, a pictorial key to adults (males and females) has been pre- pared. This pictorial key includes 3 parts: (1) Family Culicidae (key to subfamilies of Culicidae, p. 10), (2) subfamily Culicinae (key to genera of Culicinae, p. 14) and (3) genus Aedes Meigen (key to subgenera of Aedes, p. 34). An attempt was made to make the key precise and as simple as pos- sible. A few additional characters indicated by a double asterisk (**) were added wher- ever necessary to facilitate identification. Figures 1—9 illustrate the adult structures of the head, thorax, abdomen, leg and wing that are used in the key. The terminology follows Harbach and Knight (1980, 1982), with the exception of “‘tarsal claws,” which is retained for “‘ungues.’” The venational terms follow those of Belkin (1962). Knight and Stone (1977) in their world catalog of mosquitoes classified the family Culicidae into 3 subfamilies: Anophelinae, Culicinae and Toxorhynchitinae. However, see Harbach and Kitching (1998) who treat Toxorhynchites as a tribe of Culicinae. All 3 subfamilies are represented in the Afro- pictorial key, identification, subfamilies, genera, mosquitoes, subgenera of tropical Region. The 3 subfamilies with their constituent genera known to occur in the Afrotropical Region are indicated in Ta- bie I: Afrotropical Aedes has not been studied as a group since Edwards (1941). Edwards (1941), in his ‘Mosquitoes of the Ethiopian Region,” divided the genus Aedes into 9 subgenera. Reinert (1987) removed Aedes marshallii (Theobald) from the apicoannu- latus group of the subgenus Aedimorphus Theobald and defined a new subgenus, Al- buginosus for that species and its relatives. Reinert (1999) removed Aedes longipalpis (Grunberg) from the fulgens group of the subgenus Finlaya Theobald and defined a new subgenus, Zavortinkius for that species and its relatives. Thus, the genus Aedes in the Afrotropical Region now consists of the following 11 subgenera: . Aedimorphus Theobald . Albuginosus Reinert . Diceromyia Theobald . Finlaya Theobald . Mucidus Theobald . Neomelaniconion Newstead (as Bank- sinella Theobald) NAnNbWN wee 8. Ochlerotatus Lynch Arribalzaga Pseudarmigeres Stone and Knight (as Dunnius Edwards) Skusea Theobald Stegomyia Theobald Zavortinkius Reinert. 2 10. Hale Edwards (1932) divided the subgenus Stegomyia into 4 groups which he desig- nated A, B, C, and D. Aedes vittatus (Bigot) was assigned by Edwards to his Group D (vittatus group), a monotypic species group. Huang (1977) removed Group D (vittatus group), a monotypic species group (Aedes vittatus (Bigot)), from the subgenus Stego- myia and placed it in the subgenus Aedi- morphus of the genus Aedes. Since the Vit- tatus Group is monotypic, the species name Aedes vittatus is used in the key. Cornet (1974: 175) described Aedes cozi, a new species from Eastern Senegal, and placed it in the subgenus Stegomyia. Using Tablewe the Afrotropical Region. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON the subgeneric key, users will find that Ae- des cozi Cornet is not in Stegomyia. The taxonomic status of this species will be treated in a separate paper. This study is based on specimens accu- mulated by the Medical Entomology Pro- ject (MEP) and the Systematics of Aedes Mosquitoes Project (SAMP), Department of Entomology, National Museum of Natural History, Smithsonian Institution. Pinned adults were prepared for scanning electron microscopy using standard procedures by Mrs. Susann G. Braden, SEM Laboratory, National Museum of Natural History, Smithsonian Institution. Images of diagnos- tic characters directly taken from specimens using scanning electron microscopy are provided. The “cut-off”? date is October 1999, and changes in taxonomic status since then are not included. Geographically, this key includes all of Africa south of Mo- rocco, Algeria, Libya and Egypt. Classification of the family Culicidae in Subfamily Genus 1. Anophelinae Culicinae 2: 3. Toxorhynchitinae . Anopheles Meigen Aedeomyia Theobald Aedes Meigen . Coquillettidia Dyar . Culex Linnaeus . Culiseta Felt . Eretmapodites Theobald . Ficalbia Theobald . Hodgesia Theobald . Malaya Leicester . Mansonia Blanchard . Mimomyia Theobald . Orthopodomyia Theobald . Uranotaenia Lynch Arri- balzaga . Toxorhynchites Theobald VOLUME 103, NUMBER 1 ACKNOWLEDGMENTS I express my sincere appreciation and gratitude to: Mr. Walter R. Brown and Mrs. Susann G. Braden, SEM Laboratory, Na- tional Museum of Natural History, Smith- sonian Institution, for access to and assis- tance with a scanning electron microscope while conducting this study; to LTC Daniel A. Strickman, Chief, Department of Ento- mology, Walter Reed Army Institute of Re- 3} search (WRAIR), for providing funding for the publication; to Dr. Wayne N. Mathis, Department of Entomology, Smithsonian Institution, Drs. Daniel A. Strickman and Richard C. Wilkerson, Walter Reed Biosys- tematics Unit (WRBU) and Dr. Maria An- ice Mureb Sallum, University of Sao Paulo, Sao Paulo, Brazil/Smithsonian Research Fellow, for critically reviewing this manu- script and for their valuable comments. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MORPHOLOGICAL FEATURES USED IN IDENTIFICATION proboscis antenna Abdomen ADULT VOLUME 103, NUMBER 1 laliateticiealels hindtibia tarsomeres ————_»> 5@ alialeltclesers HINDLEG PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON < eee ~DrObOscis fi | t ' | Pe saantenna pedicel oh vertex occiput ENG 2 g HEAD - DORSAL hindtarsomere 5 tarsal claws +hOEA* HINDTARSOMERE 5 WITH TARSAL CLAWS VOLUME 103, NUMBER 1 midtarsomeres #HHHE MIDLEG Tarsal claws Ventral view Pulvillus aA J/// Pulvillus Empodium TARSAL CLAWS - VENTRAL PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON THORAX - DORSAL VOLUME 103, NUMBER 1 THORAX - LATERAL 6 ppt a oe a & 1 Lk > dercemmnsoee So Cf Mad gli ohn TT) ae ae WING - DORSAL 10 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Part 1. Key to Subfamilies of Culicidae ADULTS Abdomen. Completely or largely with Abdomen. Completely or largely dense, uniform covering of scales without scales to Page 11 to Page 13 VOLUME 103, NUMBER 1 1] Part 1. (continued) Page 11 Thorax. Posterior margin of scutellum Thorax. Posterior margin of scutellum trilobed, with setae in 3 groups on all evenly rounded, with setae evenly lobes distributed Hi XN ‘ Nie Subfamily Culicinae ** see Part 2 Wing. Posterior margin of wing emarginated near tip of vein Cu2 Tagua w" to Page 12 12 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Part 1. (continued) Page 12 ** Head. Proboscis with apical half strongly recurved and more slender than basal half Female Subfamily Toxorhynchitinae Genus Toxorhynchites Theobald VOLUME 103, NUMBER 1 Part 1. (continued) Page 13 ** | Thorax. Posterior margin of scutellum evenly rounded or slightly trilobed, with setae more or less evenly distributed ** | Wing. Posterior margin of wing not emarginated near tip of vein Cu2 Subfamily Anophelinae Genus Anopheles Meigen 13 14 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Part 2. Key to Genera of Culicinae ADULTS aqua.“ Wing. Vein 1A ending before, or near Wing. Vein 1A ending well beyond base of fork of vein Cu base of fork of vein Cu | to Page 16b Wing. Alula without scales, or Wing. Alula with narrow fringe with broad decumbent scales scales to Page 15 to Page 16a VOLUME 103, NUMBER 1 Part 2. (continued) Page 15 Thorax. Prespiracular setae Thorax. Prespiracular setae present absent ** ke Wing. Vein R2+3 much longer than Wing. Veins R2+3, R2, R3 with veins R2, R3 outstanding scales emarginated at tip Uranotaenia Hodgesia PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Part 2. (continued) Page 16a ** Head. Proboscis with apical part distinctly swollen, upturned and hairy Malaya Page 16b Wing. Alula without scales, or Wing. Alula with narrow fringe with broad decumbent scales scales to Page 17 to Page 20b VOLUME 103, NUMBER 1 Part 2. (continued) Page 17 ** Thorax. Paratergite w ithout scales Wing. Plume scales all narrow to Page 18a Wing. Plume scales all broad to Page 18b 7 18 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Part 2. (continued) Page 18a Page 18b “te sk | ** Thorax. Postspiracular setae Thorax. Postspiracular setae present absent Eretmapodites Thorax. Scutellum with all broad Thorax. Scutellum with all narrow scales scales | to Page 19a to Page 19b VOLUME 103, NUMBER 1 19 Part 2. (continued) Page 19a Page 19b ** ** Wing. Alula with broad decumbent Wing. Alula without scales scales Mimomyia ( in part ) Thorax. Lower mesepimeral setae Thorax. Lower mesepimeral setae present absent | Mimomyia ( in part ) to Page 20a PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Part 2. (continued) Page 20a ** Legs. Mid- and hindfemora with large tufts of suberect scales at apex Aedeomyia Page 20b Thorax. Prespiracular setae Thorax. Prespiracular setae present absent to Page 21a to Page 21b VOLUME 103, NUMBER 1 21 Part 2. (continued) De yet ** Wing. Base of vein Sc with a patch of setae on ventral side Culiseta Page 21b SS Leg. Pulvillus present, well Leg. Pulvillus absent, or not developed (pad-like) well developed (hair-like) to Page 22 to Page 23 N tO PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Part 2. (continued) Page 22 kk 1 Kk 2 a Head. Vertex with erect forked scales Thorax. Paratergite without numerous, not restricted to occiput scales kk 3 kk 4 Thorax. Scutellum with all narrow Head. Antenna with flagellomere 1 scales about as long as flagellomere 2 Culex VOLUME 103, NUMBER 1 Part 2. (continued) Page 23 —————— Head. Vertex with erect forked scales Head. Vertex with erect forked scales numerous, not restricted to occiput not numerous, restricted or not restricted to occiput | to Page 28b Thorax. Paratergite without scales Thorax. Paratergite with scales to Page 24 to Page 28a 23 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Part 2. (continued) Page 24 eee Thorax. Scutellum with narrow scales Thorax. Scutellum with broad scales Aedes ( in part ) - see Part 3 Thorax. Postspiracular setae Thorax. Postspiracular setae present absent to Page 25 to Page 26b VOLUME 103, NUMBER 1 25 Part 2. (continued) Page 25 ST Wing. Most plume scales very broad = Wing. Plume scales all narrow and asymmetrical Aedes ( in part ) - see Part 3 ** Thorax. Subspiracular area without scales to Page 26a 26 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Page 26a *** Thorax. Lower mesepimeral setae present Mansonia Page 26b Thorax. Lower mesepimeral setae Thorax. Lower mesepimeral setae present absent to Page 27a to Page 27b VOLUME 103, NUMBER | 7 Part 2. (continued) Page 27a Page 27b **K* | ** Thorax. Subspiracular area without Leg. Midtarsomere 1 distinctly longer scales than tarsomeres 2-5 combined Coquillettidia ** Leg. Midtarsomere 4 shorter than midtarsomere 5 Orthopodomyia 28 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Part 2. (continued) Page 28a ** | Thorax. Postspiracular setae present Aedes ( in part ) - see Part 3 Page 28b Thorax. Paratergite without scales Thorax. Paratergite with scales | to Page 29 to Page 31 VOLUME 103, NUMBER 1 29 Part 2. (continued) Page 29 a a a rea aoe Thorax. Scutellum with narrow scales Thorax. Scutellum with broad scales ** ** Wing. Plume scales all broad Wing. Plume scales all narrow to Page 30a to Page 30b 30 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Part 2. (continued) Page 30a Page 30b ** ** | Thorax. Postspiracular setae Thorax. Postspiracular setae absent present Ficalbia ** Thorax. Subspiracular area with scales Aedes ( in part ) - see Part 3 VOLUME 103, NUMBER I Part 2. (continued) Page 31 Head. Vertex with all broad flat Head. Vertex with broad flat decumbent scales decumbent scales and some narrow forked scales 7% aa Thorax. Scutellum with broad Thorax. Lower prealar scale patch scales present | Aedes ( in part ) - see Part 3 to Page 32 32 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Part 2. (continued) Page 32 kk 1 kk 2 Wing. Plume scales all Thorax. Scutum with all, or narrow | mainly narrow curved scales mk 3 Thorax. Postspiracular setae present to Page 33 VOLUME 103, NUMBER 1 33 Part 2. (continued) Page 33 Thorax. Postprocoxal Thorax. Postprocoxal membrane with scales membrane without scales Aedes ( in part ) - see Part 3 ** Thorax. Lower mesepimeral setae absent Aedes ( in part ) - see Part 3 34 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Part 3. Key to Subgenera of Aedes ADULTS -—_—_ YT Wing. Wing membrane clouded on Wing. Wing membrane not clouded crossveins r-m and m-cu on crossveins r-m and m-cu Mucidus Head. Vertex with erect forked scales ead. Vertex with erect forked scales numerous, not restricted to occiput not numerous, restricted or not restricted to occiput | | to Page 35 to Page 48 VOLUME 103, NUMBER 1 35 Part 3. (continued) Page 35 Thorax. Lower prealar scale patch Thorax. Lower prealar scale patch present absent | to Page 40 ** Thorax. Paratergite with scales to Page 36 36 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Part 3. (continued) Page 36 _———_______|________, Head. Vertex with decumbent scales Head. Vertex with decumbent scales largely narrow largely broad | to Page 38 Thorax. Scutellum with all narrow Thorax. Scutellum with all broad scales scales Ochlerotatus to Page 37 VOLUME 103, NUMBER 1 Part 3. (continued) Page 37 ek 1 *k 2 Thorax. Lower mesepimeral setae Thorax. Acrostichal setae present present KK 3 Head. Pedicel with white broad scales on mesal and lateral surfaces Aedimorphus ( in part ) Vittatus Group Aedes vittatus 37 38 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Part 3. (continued) Page 38 ek | Thorax. Acrostichal setae absent #* Head. Vertex with broad scales along eye margins to Page 39 VOLUME 103, NUMBER 1 Part 3. (continued) Page 39 Thorax. Subspiracular area with Thorax. Subspiracular area without scales scales *#* | #* | Thorax. Scutellum with broad scales Thorax. Scutellum with narrow scales on midlobe on midlobe Finlaya ( in part ) Finlaya ( in part ) Wellmani Group Pulchrithorax Group Aedes pulchrithorax 40 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Part 3. (continued) Page 40 Ee __ Thorax. Acrostichal setae present Thorax. Acrostichal setae absent to Page 47b Thorax. Paratergite without scales Thorax. Paratergite with scales | | to Page 41 to Page 42b 41 VOLUME 103, NUMBER 1 Part 3. (continued) Page 41 Thorax. Scutellum with all broad Thorax. Scutellum with all narrow scales scales Neomelaniconion ** Head. Vertex with a pair of wedge-shaped patches of broad, flat scales on eye margins | to Page 42a PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Part 3. (continued) Page 42a ** | Head. Pedicel with a few small scales and short fine setae on mesal surface a \ Aedimorphus ( in part ) Domesticus Group Page 42b ey Head. Vertex with a pair of Head. Vertex without a pair of wedge-shaped patches of broad, flat wedge-shaped patches of broad, flat scales on eye margins scales on eye margins | | to Page 43 to Page 44b VOLUME 103, NUMBER 1 43 Part 3. (continued) Page 43 Thorax. Subspiracular area with Thorax. Subspiracular area without scales scales ** | kk Head. Pedicel with a patch of white Head. Pedicel with a few small scales broad overlapping scales on mesal and short fine setae on mesal surface surface Albuginosus to Page 44a Ad PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Part 3. (continued) Page 44a #* | Thorax. Scutellum with all broad, flat scales Aedimorphus ( in part ) Argenteopunctatus Group Page 44b enn cca Thorax. Subspiracular area with Thorax. Subspiracular area without scales scales | | to Page 45 to Page 46b VOLUME 103, NUMBER 1 45 Part 3. (continued) Page 45 ae Thorax. Postpronotum with all or most Thorax. Postpronotum with all or most scales broad and flat scales narrow and curved Aedimorphus ( in part ) ** Thorax. Lower mesepimeral setae present to Page 46a PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Part 3. (continued) Page 46a Page 46b - | a Head. Pedicel with setae mixed with Head. Vertex with narrow scales many broad, flat scales on mesal along eye margins surface kk k* Head. Pedicel with 2 patches of Thorax. Scutellum with all broad scales on mesal surface scales Diceromyia | to Page 47a VOLUME 103, NUMBER 1 Part 3. (continued) Page 47a Rk | Head. Pedicel with few setae and scales on mesal surface Aedimorphus ( in part ) Apicoannulatus Group Page 4/7b KK | Head. Pedicel with a long patch of white broad, flat scales on mesal surface Aedes cozi 47 48 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Part 3. (continued) Page 48 ee, Head. Vertex with erect forked scales Head. Vertex with erect forked scales not numerous, not restricted to occiput not numerous, restricted to occiput to Page 51 ** Thorax. Scutellum with all broad scales | to Page 49 VOLUME 103, NUMBER 1 Part 3. (continued) Page 49 a Thorax. Paratergite without scales Thorax. Paratergite with scales ** #* Thorax. Lower prealar scale patch Thorax. Lower prealar scale patch absent present | to Page 50a to Page 50b 50 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Part 3. (continued) Page 50a Page 50b ek *#* Thorax. Acrostichal setae present Thorax. Acrostichal setae absent *k* kk Thorax. Subspiracular area with Head. Pedicel with very few (1-3) scales scales and short fine setae on mesal surface Skusea Zavortinkius VOLUME 103, NUMBER 1 51 Part 3. (continued) Page 51 eK 4 Kk 2 Head. Vertex with all broad, flat Thorax. Acrostichal setae absent decumbent scales KK 3 Kk 4 Thorax. Paratergite with scales Thorax. Scutellum with all broad scales | to Page 52 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Part 3. (continued) Page 52 SS SY Thorax. Postprocoxal membrane Thorax. Postprocoxal membrane with scales without scales Pseudarmigeres Stegomyia VOLUME 103, NUMBER | LITERATURE CITED Belkin, J. N. 1962. The mosquitoes of the South Pa- cific (Diptera, Culicidae). University of California Press, Berkeley and Los Angeles. 2 vols., 608 and 412 pp. Cornet, M. 1974. Aedes (Stegomyia) cozi n. sp., une nouvelle espece de Culicidae au Senegal. Cahiers de Office de la Recherche Scientifique et Tech- nique Outre-Mer Serie Entomologie Medicale et Parasitologie 11 (1973): 175-180. Edwards, KE W. 1932. Diptera. Family Culicidae. Jn Wytsman, P. Genera Insectorum, Desmet-Verte- neuil, Brussels, Fasc. 194, 258 pp., 5 pl. . 1941. Mosquitoes of the Ethiopian region. IfI.—Culicine adults and pupae. London, British Museum (Natural History). 499 pp., 4 pl. Harbach, R. E. and K. L. Knight. 1980. Taxonomists’ glossary of mosquito anatomy. Plexus Publishing, Inc., Marlton, New Jersey. 415 pp. . 1982. Corrections and additions to taxono- mists’ glossary of mosquito anatomy. Mosquito Systematics (1981) 13: 201-217. 53 Harbach, R. E. and I. J. Kitching. 1998. Phylogeny and classification of the Culicidae (Diptera). System- atic Entomology 23: 327-370. Huang, Y. M. 1977. Medical entomology studies. VIII. Notes on the taxonomic status of Aedes vittatus (Diptera: Culicidae). Contributions of the Ameri- can Entomological Institute (Ann Arbor). 14(1): 112-132. Knight, K. L. and A. Stone. 1977. A catalog of the mosquitoes of the world (Diptera: Culicidae), Sec- ond edition. Thomas Say Foundation, Entomolog- ical Society of America. Vol. 6, 611 pp. Reinert, J. E 1987. Albuginosus, a new subgenus of Aedes Meigen (Diptera: Culicidae) described from the Afrotropical Region. Mosquito Systematics (1986) 18: 307-326. . 1999. Description of Zavortinkius, a new sub- genus of Aedes, and the eleven included species from the Afrotropical Region (Diptera: Culicidae). Contributions of the American Entomological In- stitute (Gainesville) 31(2): 1—105. PROC. ENTOMOL. SOC. WASH. 103(1), 2001, pp. 54-59 STATISTICAL MEASURES OF ASSOCIATION BETWEEN AMBLYOMMA SABANERAE STOLL (ACARI: [IXODIDA: [IXODIDAE) AND THE FURROWED WOOD TURTLE, RHINOCLEMMYS AREOLATA (DUMERIL AND BIBRON) (TESTUDINES: EMYDIDAE), IN NORTHERN BELIZE RICHARD G. ROBBINS, STEVEN G. PLATT, THOMAS R. RAINWATER, AND WENDY WEISMAN (RGR) Armed Forces Pest Management Board, Walter Reed Army Medical Center, Washington, DC 20307-5001 U.S.A. (e-mail: robbinrg @acq.osd.mil); (SGP, WW) Wildlife Conservation Society, 185th Street and Southern Boulevard, Bronx, NY 10460-1099 U.S.A. (e-mail: Plattwcs@aol.com and wweisman@wcs.org); (TRR) Institute of Environ- mental and Human Health, Texas Tech University, 1207 Gilbert Drive, Lubbock, TX 79416 U.S.A. (e-mail: trainwater@ttu.edu) Abstract.—Between April 1997 and June 1998, 159 tick collections, comprising 411 specimens, were made from 261 adults and juveniles of the furrowed wood turtle, Rhin- oclemmys areolata (Duméril and Bibron), on a tract of pine forest and savanna in northern Belize. All ticks were determined to be Amblyomma sabanerae Stoll, a common parasite of testudines in the lowland tropics of Central America. Male ticks were collected from more turtles (94) than were females (48) or immatures (82). However, despite the likeli- hood that male turtles have greater home ranges, significantly more ticks parasitized fe- male turtles than males. It is thought that overdispersion of both ticks and turtles declines in the equable climate of the Neotropics, in which case female turtles, being larger than males, would be expected to acquire more ticks. Collections of immature A. sabanerae were more frequently made on fleshy areas of R. areolata, but the mean number of nymphs and larvae collected from anterior areas was not significantly different from that for posterior areas. Key Words: Acari, Ixodidae, Amblyomma sabanerae, host relationships, Belize Neotropical turtles and tortoises (Testu- dines) are the principal hosts of only 8 tick species: the argasid Ornithodoros transver- sus (Banks), and the ixodids Amblyomma crassum Robinson, A. humerale Koch, A. macfarlandi Keirans, Hoogstraal and Clif- ford, A. pilosum Neumann, A. sabanerae Stoll, A. testudinis (Conil), and A. usingeri Keirans, Hoogstraal and Clifford (Hoogs- traal and Aeschlimann 1982). Yet, the ecol- ogy, population dynamics and disease re- lationships of these acarines have seldom been addressed, the most comprehensive studies to date being those of Schneider et al. (1971) and Ernst and Ernst (1977). This is lamentable because, in recent years, tes- tudine populations have been declining pre- cipitously worldwide (Ernst and Barbour 1989), with the result that of 247 nominal species listed by King and Burke (1989), 70 (28%) appear in Appendix I (species threatened with extinction that are or may be affected by trade) or Appendix II (spe- cies not necessarily threatened with extinc- tion but that may become so unless trade is strictly regulated) of the Convention on In- VOLUME 103, NUMBER 1 ternational Trade in Endangered Species of Wild Fauna and Flora. Between April 1997 and June 1998, we were able to methodi- cally collect ticks from a Belizean popula- tion of the furrowed wood turtle, Rhino- clemmys areolata (Duméril and Bibron), a semiterrestrial species that is still common over most of its range: south-central Vera- cruz, Tabasco and eastern Chiapas to the Yucatan Peninsula and Cozumel Island in Mexico, south through Belize, eastern Gua- temala, and perhaps eastern Honduras (Ernst and Barbour 1989). Our results con- trast markedly with those from a similar study of tick populations on tortoises in the arid subtropics of southwestern Russia (Robbins et al. 1998). MATERIALS AND METHODS Turtles were collected on a privately owned 405-ha tract (17.22N, 88.34W) of pine forest and savanna adjacent to the Western Highway in northern Belize, ap- proximately 48 km west of Belize City. The flora of this area is characterized by an overstory of Caribbean pine, Pinus cari- baea Morelet, and oak, Quercus oleoides Schlechtendal and Chamisso, with an un- derstory of craboo, Byrsonima crassifolia (L.) Kunth, sandpaper tree, Curatella amer- icana L., wax myrtle, Myrica spp., miconia, Miconia spp., and eugenia, Eugenia spp. Savannas are dominated by grasses (Gra- mineae) and sedges, Rhynchospora spp. and Scleria spp., with scattered pines, craboo, and clusters of paurotis, Acoelorraphe wrightii (Grisebach and Wendland) Wend- land ex Beccari (Furley 1989). The climate of northern Belize is considered tropical be- cause the average temperature each month exceeds 18°C. Warmest temperatures are re- corded during the dry season in April and May, with average maxima of 32.8 and 33.1°C, respectively. At this season, natural and anthropogenic fires occur in both pine forest and savanna, as they did in April and May 1997, when our study site was swept by a series of catastrophic blazes. Precipi- tation at our site averages 150 cm/yr, with 55 a pronounced wet season from June to No- vember, though this may vary annually (Hartshorn et al. 1984). Wood turtles were sought during the morning (0700-1000 hr) and evening (1530-1800 hr), when they are most active (Vogt and Platt, in press). Although com- mon on the study site, turtles proved diffi- cult to locate in the thick ground vegetation, and most (>75%) were found with the aid of a trained dog. Captured turtles were tak- en to a field laboratory, where a variety of morphometric data were collected. Sex was determined on the basis of tail morphology: males have long, thickened tails with the vent well beyond the carapace margin, whereas females have short tails with the vent beneath the carapace margin (Ernst and Barbour 1989). Sex could not be de- termined for juveniles (carapace length < 11 cm). Each turtle was permanently marked by notching a unique series of mar- ginal scutes (Cagle 1939) and was released at its point of capture within 24 hours. Of 261 male, female, or unsexed juvenile R. areolata taken in this manner, 159 (61%) were parasitized by ticks: 52 (32.7%) male turtles, 75 (47.2%) females, and 32 (20.1%) juveniles. Infested turtles were assigned ac- cession numbers (RAI1-RAI59) and ME- DARKS (Medical Archives) numbers (1997-0546 through 1997-0703) by the Wildlife Conservation Society (formerly New York Zoological Society), Bronx, New York. Tick collections corresponding to these numbers were shipped to RGR in vi- als of Formalin, later changed to 70% iso- propyl alcohol, for identification and anal- ysis. All tick collections have been depos- ited at the American Museum of Natural History, New York, New York. RESULTS A total of 411 tick specimens were re- moved from our 159 infested R. areolata during this study. All were A. sabanerae, whose range overlaps that of R. areolata and extends southward as a parasite of var- ious turtles, e.g., R. annulata (Gray), R. fu- 56 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1: Descriptive statistics for all stages of Amblyomma sabanerae from 261 Rhinoclemmys areolata; Belize District, northern Belize, April 1997—June 1998 (collectors S. G. Platt and T. R. Rainwater). Tick Sex/Stage Number (%) Range* Males 126 (31) 0-4 Females S35) (G3) 0-3 Nymphs 172 (42) 0-13 Larvae 58 (14) 0-24 Mean with Standard Error Standard Deviation Coefficient of Variation OSF=10105 0.8 160.0 O2Ze=0103 0.5 250.0 0.6, = 0:09 1.5 250.0 OH S= (0). 110) od 850.0 * Ticks/turtle. nerea (Cope), R. pulcherrima (Gray) and Trachemys scripta (Schoepf), into the low- land tropics of Panama (Fairchild et al. 1966) and perhaps northern South America (records of A. sabanerae from Colombia (Schulze 1937, Osorno-Mesa 1940) and Su- riname (Oudemans 1902) require confir- mation). Adults of A. sabanerae were sep- arated from other turtle-infesting amblyom- mines using the key of Jones et al. (1972). The preimaginal stages (larva, nymph) of A. sabanerae have not been described and were “‘identified’’ by association with adults. Occasional nymphs of the reptile and amphibian parasite A. dissimile Koch were found on R. areolata at other locations in northern Belize, but the nymphal denti- tion of A. dissimile is 3/3 (Keirans and Dur- den 1998), whereas in nymphs that we are calling A. sabanerae it is 2/2. Descriptive statistics for A. sabanerae on R. areolata appear in Table 1. Males of A. sabanerae were recovered from 94 turtles, nymphs from 77 turtles, but females and larvae from just 48 and 12 turtles, respec- tively. Because male ticks were least over- dispersed (clumped) on the turtle popula- tion, they were employed to test whether observed differences in parasitization be- tween male and female turtles were statis- tically significant (juvenile turtles were ex- cluded from this analysis because too few were parasitized). The mean number of male A. sabanerae on male R. areolata was 0.6 + 0.1 (mean + standard error), while that for female turtles was 1.0 + 0.1. This difference is significant at t = 2.765, df = 126, P < 0.01. The higher mean infestation of female wood turtles in the humid tropics of Belize stands in contradistinction to the much lower mean recorded for female spur- thighed tortoises, Testudo graeca nikolskii Chkhikvadze and Tuniev, relative to male tortoises infested with the tick Hyalomma daegyptium (L.) along the xeric Black Sea coast of southern Russia (Robbins et al. 1998). There are numerous possible expla- nations for this striking difference in mean infestation, among them sex- or species- specific differences in host behavior, the de- velopment in testudines of a degree of age- related immunity to tick parasitism (Wikel 1996, Tembo and Kiwanuka 1997), or the release of pheromones by female ticks that could affect the numbers of males (Sonen- shine 1985, Gothe 1987). Unfortunately, virtually nothing is known about the phys- iological ecology of A. sabanerae, H. ae- gyptium, or the hosts of these tick species. Considering only habitat differences, it is possible to speculate that overdispersion of both ticks and testudines declines in the more equable climate of the Neotropics, where reduced saturation deficiency favors tick survival and luxuriant vegetation pro- vides unbroken cover for turtles. By con- trast, environmental extremes in the arid scrublands of southern Russia often confine ticks to “islands” of favorable habitat and restrict tortoises to their burrows. Male spur-thighed tortoises are thought to acquire more ticks chiefly because they have larger home ranges. Where environmental vari- ables do not play a major limiting role, host attributes more likely account for observed differences in tick infestation. In the case at VOLUME 103, NUMBER 1 hand, the mean carapace length (cm) of fe- male turtles was 14.6 + 0.2, while that for males was only 12.6 + 0.2. This difference is highly significant at t = 6.666, df = 126, P < 0.001. Larger female turtles might ac- quire more ticks than males when moving through habitat in which ticks are less con- tagiously dispersed—even if male turtles have larger ranges. Another attribute affecting tick loads is the nature of turtle integument. In Mesoam- erican populations of four species of Rhin- oclemmys, Ernst and Ernst (1977) noted that 78.5% of adult A. sabanerae were found attached to the shell; conversely, 95.5% of immature ticks were attached to exposed turtle skin, mostly at limb sockets or on the gular fold. Schmidt (1946) pro- vided a photograph of the carapace of a Panamanian specimen of R. annulata on which numerous adult ticks, all clearly A. sabanerae, had attached at the seams be- tween carapacial scutes. By a test of asso- ciation (Sokal and Rohlf 1973), Ernst and Ernst (1977) were able to statistically dem- onstrate this segregation of adult and im- mature ticks on the basis of attachment site, reasoning that the shorter mouthparts of lar- vae and nymphs may restrict these stages to feeding at locations where the skin is thinner and that this situation may reduce competition between developmental stages. In our study, immature A. sabanerae were found attached to fleshy areas (head, neck, legs, tail) on R. areolata 113 times but to the carapace or plastron only 8 times. Col- lections of adult ticks were made from the carapace and plastron 116 times but from fleshy areas just 32 times. Again, a test of association reveals that these frequencies are highly dependent (G = 157.818, P< 0.005) upon the stage of tick development. However, despite our anticipation that prei- maginal ticks would more often be encoun- tered on anterior rather than posterior fleshy areas, as is generally the case with small mammals (Thompson 1974, Barré et al. 1991) and with the reptile tick A. marmo- reum Koch on the leopard tortoise, Geo- 57 chelone pardalis (Bell), in South Africa (Fielden and Rechav 1994), the mean num- ber of nymphs and larvae collected from anterior areas of R. areolata (3.7 + 1.1) was not significantly different from that for posterior areas (2.8 + 0.8; t = 0.651, df = Lit, Ps 005): During the 15 months of this study, no seasonal trend in the percentage of parasit- ized turtles could be discerned. On a month-to-month basis, figures ranged from 0.0% (February 1998) to 85.7% (May 1997), but months are arbitrary time units that, coupled with wide variations in the number of turtles examined (99 in July 1997, none in November of that year), can easily give a false impression of the prev- alence of parasitization. On our population of R. areolata, the overall prevalences, sen- su. Margolis et al. (1982), of adults, nymphs, and larvae of A. sabanerae were 45.2%, 29.5%, and 4.6%, respectively. Nothing is known of the life cycle of A. sabanerae or, indeed, of most Neotropical amblyommines. Until these baseline data are established through rearing studies, the temporal dynamics of tick populations in the field will remain unresolved. ACKNOWLEDGMENTS We thank Bill and Eleanor Hasse, Ri1- chard and Carol Foster, and Matt Miller for allowing access to their properties in Be- lize. Mark and Monique Howells, Lamanai Field Research Center, Lamanai, Belize, provided invaluable logistical support. Steven Brewer, Travis Crabtree, and Thom- as Rhott assisted us in the field. Thanks also to Patricia Craig, Carnegie Institution of Washington, Washington, DC, and J. Ralph Lichtenfels, Biosystematics and National Parasite Collection Unit, Agricultural Re- search Service, U.S. Department of Agri- culture, Beltsville, Maryland, for locating and supplying several older acarological references. Statistical guidance was provid- ed by J. Robert Burge, Department of Bio- metrics, Walter Reed Army Institute of Re- search. This project was funded by the 58 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Wildlife Conservation Society (SGP and WW) and the Lamanai Field Research Cen- ter (TRR). The opinions and assertions ad- vanced herein are those of the authors and are not to be construed as official or reflect- ing the views of the U.S. Departments of the Army or Defense. LITERATURE CITED Barré, N., E. Camus, G. Borel, and R. Aprelon. 1991. Sites de fixation de la tique Amblyomma varie- gatum sur ses h6tes en Guadeloupe (Antilles fran- caises). Revue d’Elevage et de Médecine Vétéri- naire des Pays Tropicaux 44: 453-458. Cagle, EF R. 1939. A system of marking turtles for future identification. Copeia 1939: 170-173. Ernst, C. H. and R. W. Barbour. 1989. Turtles of the World. Smithsonian Institution Press, Washington, DESI Sspp: Ernst, C. H. and E. M. Ernst. 1977. Ectoparasites as- sociated with Neotropical turtles of the genus Cal- lopsis (Testudines, Emydidae, Batagurinae). Bio- tropica 9: 139-142. Fairchild, G. B., G. M. Kohls, and V. J. Tipton. 1966. The ticks of Panama (Acarina: Ixodoidea), pp. 167-219. In Wenzel, R. L. and V. J. Tipton, eds., Ectoparasites of Panama. Field Museum of Nat- ural History, Chicago, Illinois. Fielden, L. J. and Y. Rechav. 1994. Attachment sites of the tick Amblyomma marmoreum on its tortoise host, Geochelone pardalis. Experimental & Ap- plied Acarology 18: 339-349. Furley, P. A. 1989. Further observations on the nature of the savanna vegetation and soils in Belize, pp. 9-35. In Munro, D. M., ed., Ecology and Envi- ronment in Belize: An Account of the University of Edinburgh Expedition to Belize, Central Amer- ica, 1986. Occasional Publication No. 12, Depart- ment of Geography, University of Edinburgh, Scotland. Gothe, R. 1987. Tick pheromones. Onderstepoort Jour- nal of Veterinary Research 54: 439-441. Hartshorn, G., L. Nicolait, L. Hartshorn, G. Bevier, R. Brightman, J. Cal, A. Cawich, W. Davidson, R. DuBois, C. Dyer, J. Gibson, W. Hawley, J. Leon- ard, R. Nicolait, D. Weyer, H. White, and C. Wright. 1984. Belize Country Environmental Pro- file: A Field Study. USAID and Robert Nicolait and Associates, Ltd., Belize City, Belize. 151 pp. Hoogstraal, H. and A. Aeschlimann. 1982. Tick-host specificity. Mitteilungen der Schweizerischen En- tomologischen Gesellschaft/Bulletin de la Société Entomologique Suisse 55: 5-32. Jones, E. K., C. M. Clifford, J. E. Keirans, and G. M. Kohls. 1972. The ticks of Venezuela (Acarina: Ix- odoidea) with a key to the species of Amblyomma in the Western Hemisphere. Brigham Young Uni- versity Science Bulletin, Biological Series 17: 1— 40. Keirans, J. E. and L. A. Durden. 1998. Illustrated key to nymphs of the tick genus Amblyomma (Acari: Ixodidae) found in the United States. Journal of Medical Entomology 35: 489-495. King, E W. and R. L. Burke. 1989. Crocodilian, Tua- tara, and Turtle Species of the World: A Taxonom- ic and Geographic Reference. Association of Sys- tematics Collections, Washington, DC. 216 pp. Margolis, L., G. W. Esch, J. C. Holmes, A. M. Kuris, and G. A. Schad. 1982. The use of ecological terms in parasitology (report of an ad hoc com- mittee of the American Society of Parasitologists). Journal of Parasitology 68: 131-133. Osorno-Mesa, E. 1940. Las garrapatas de la Repub- lica de Colombia. Revista de la Academia Col- ombiana de Ciencias Exactas, Fisicas y Naturales 4: 6-24. Oudemans, A. C. 1902. Notes on Acari. Fourth series. Tijdschrift der Nederlandsche Dierkundige Ver- eeniging, 2nd series 7: 276-310. Robbins, R. G., W. B. Karesh, P. P. Calle, O. A. Leon- tyeva, S. L. Pereshkolnik, and S. Rosenberg. 1998. First records of Hyalomma aegyptium (Ac- ari: Ixodida: Ixodidae) from the Russian spur- thighed tortoise, Testudo graeca nikolskii, with an analysis of tick population dynamics. Journal of Parasitology 84: 1303-1305. Schmidt, K. P. 1946. Turtles collected by the Smith- sonian biological survey of the Panama Canal Zone. Smithsonian Miscellaneous Collections 106: 1-9. Schneider, C. C., B. Roth, and H. D. Lehmann. 1971. Untersuchungen zum Parasit-Wirt-Verhaltnis der Zecke Amblyomma testudinis (Conil 1877). Zeit- schrift fiir Tropenmedizin und Parasitologie 22: 1— Nk Schulze, P. 1937. Beitrage zur Kenntnis der Zecken- gattung Amblyomma. Zeitschrift fiir Parasitenkun- de 9: 690-694. Sokal, R. R. and E J. Rohlf. 1973. Introduction to Bio- statistics. W. H. Freeman and Company, San Fran- cisco, California. 368 pp. Sonenshine, D. E. 1985. Pheromones and other semi- ochemicals of the Acari, pp. 1-28. /n Mittler, T. E., E J. Radovsky, and V. H. Resh, eds., Annual Review of Entomology, vol. 30. Annual Reviews, Inc., Palo Alto, California. Tembo, S. D. and A. Kiwanuka. 1997. Acquisition of protective immunity in Geochelone pardalis against Amblyomma marmoreum (Acari: Ixodi- dae) nymphal ticks. Onderstepoort Journal of Vet- erinary Research 64: 1—4. Thompson, P. H. 1974. Attachment site on white lab- oratory mice by larvae of Dermacentor variabilis (Say) (Acarina). Proceedings of the Entomological Society of Washington 76: 282-284. VOLUME 103, NUMBER 1 Vogt, R. C. and S. G. Platt. In press. Rhinoclemmys areolata—furrowed wood turtle. /n Pritchard, P. C. H. and A. G. J. Rhodin, eds., The Conservation Biology of Freshwater Turtles. Vol. 2. New World Turtles (Nearctic and Neotropical Realms). Oc- casional Papers, IUCN Species Survival Commis- 39 sion, International Union for Conservation of Na- ture, Gland, Switzerland. Wikel, S. K. 1996. Host immunity to ticks, pp. 1-22. In Mittler, T. E., E J. Radovsky, and V. H. Resh, eds., Annual Review of Entomology, vol. 41. An- nual Reviews, Inc., Palo Alto, California. PROC. ENTOMOL. SOC. WASH. 103(1), 2001, pp. 60-73 LIFE HISTORY AND DESCRIPTION OF IMMATURE STAGES OF NEASPILOTA ACHILLEAE JOHNSON (DIPTERA: TEPHRITIDAE) ON STEPHANOMERIA SPP. (ASTERACEAE) IN SOUTHERN CALIFORNIA RICHARD D. GOEDEN Department of Entomology, University of California, Riverside, CA 92521, U.S.A. (e- mail: richard.goeden @ucr.edu) Abstract.—Neaspilota achilleae Johnson is a bivoltine fruit fly (Diptera: Tephritidae) developing solely in the flower heads of Stephanomeria spp. (Asteraceae) belonging to the subtribe Stephanomeriinae of the tribe Lactuceae in southern California. This species is distributed along the east coast of the United States and in California and Arizona, but has not been reported from middle America. Moreover, in the East it is reported from 17 other host-plant species in the subtribes Asterinae, Hieraciinae, Lactucinae, Liatrinae, and Solidaginae of the Tribes Astereae, Eupatorieae, and Lactuceae, but not from Stephan- omeriinae, which are mostly western plant species. The second- and third-instar larvae and puparium are described and figured, and these larval instars are compared with those of other Neaspilota. The anterior thoracic spiracle of the second instar has three papillae, but three and four papillae are reported in the third instar. The second and third instars of N. achilleae have an undetermined number of oral ridges with dentate posterior margins in a vertical series lateral to the oral cavity. The appearance and arrangement of these oral ridges is now known to be a distinguishing generic larval character. The numbers of these oral ridges, along with the patterns of minute acanthae circumscribing the body segments, are useful in distinguishing several species of Neaspilota larvae. The larvae feed mainly on the corollas of florets as first and second instars; however, as third instars, they may extend their feeding into and through the ovules or soft achenes, and sometimes into the receptacle, and supplement their diet with sap. At least two annual generations are produced in southern California. The life cycle is of the aggregative type and over- wintering mainly occurs as sexually immature adults. Pteromalus sp. (Hymenoptera: Pter- omalidae) was reared as a solitary, larval-pupal endoparasitoid of N. achilleae; Eurytoma obtusiventrus Gahan or near and E. veronia Bugbee (Hymenoptera: Eurytomidae), as probable, solitary, larval-pupal endoparasitoids. Key Words: Insecta, Neaspilota, Stephanomeria, Asteraceae, nonfrugivorous Tephriti- dae, biology, taxonomy of immature stages, allopatry, flower-head feeding, aggregative life cycle, seed predation, parasitoids Revision of the genus Neaspilota (Dip- tera: Tephritidae) by Freidberg and Mathis (1986) facilitated identification of speci- mens reared from California Asteraceae (Goeden 1989) and stimulated several life- history studies, including those on N. viri- descens Quisenberry (Goeden and Headrick 1992), N. wilsoni Blanc and Foote (Goeden and Headrick 1999), N. signifera (Coquil- lett) (Goeden 2000a), N. aenigma Freidberg and Mathis (Goeden 2000b), N. appendi- culata Freidberg and Mathis (Goeden VOLUME 103, NUMBER | 2000c), and N. pubescens Freidberg and Mathis (Goeden 2000d). This paper de- scribes some immature stages and the life history of a seventh species from Califor- nia, N. achilleae Johnson. MATERIALS AND METHODS The present study was based in large part on dissections of samples of flower heads of Stephanomeria virgata Bentham (Aster- aceae) mainly collected SE of Barett Junc- tion at 400-m elevation, SE San Diego Co., and from S. pauciflora (Nuttall) Nelson mainly collected at the mouth of Big Rock Creek Canyon at 1,400 m., Angeles Nat. Forest, Los Angeles Co., during 1988 to 1997. One-liter samples of excised, imma- ture and mature flower heads containing eggs, larvae, and puparia were transported in cold-chests in an air-conditioned vehicle to the laboratory and stored under refrig- eration for subsequent dissection, photog- raphy, description, and measurement. Four second- and seven third-instar larvae and six puparia dissected from flower heads were preserved in 70% EtOH for scanning electron microscopy (SEM). Additional prepuparia and puparia were placed in sep- arate, glass shell vials stoppered with ab- sorbant cotton and held in humidity cham- bers at room temperature for adult and par- asitoid emergence. Specimens for SEM were hydrated to distilled water in a de- creasing series of acidulated EtOH. They were osmicated for 24 h, dehydrated through an increasing series of acidulated EtOH and two, 1-h immersions in hexa- methyldisilazane (HMDS), mounted on stubs, sputter-coated with a gold-palladium alloy, and studied and photographed with a Philips XL-30 scanning electron micro- scope in the Institute of Geophysics and Planetary Physics, University of California, Riverside. Most adults reared from isolated prepu- paria and puparia were individually caged in 850-ml, clear-plastic, screened-top cages with a cotton wick and basal water reser- voir and provisioned with a strip of paper 61 toweling impregnated with yeast hydroly- zate and sucrose. These cages were used for studies of longevity and sexual maturation in the insectary of the Department of En- tomology, University of California, River- side, at 25 + 1°C, and 14/10 (L/D) photo- period. Five pairs of virgin males and fe- males obtained from emergence cages also were held in separate, clear-plastic, petri dishes provisioned with a flattened, water- moistened pad of absorbant cotton spotted with honey (Headrick and Goeden 1994) for observations of their courtship and cop- ulation behavior. Plant names used in this paper follow Hickman (1993) and Bremer (1994); te- phritid names and adult terminology follow Foote et al. (1993). Terminology and tele- graphic format used to describe the imma- ture stages follow Goeden (2000a, b, c, d), Goeden et al. (1998), Goeden and Headrick (1992, 1999), Goeden and Teerink (1997a, b, 1998, 1999a, b), Teerink and Goeden (1999), and our earlier works cited therein. Means + SE are used throughout this paper. Voucher specimens of N. achilleae imma- ture stages, adults, and parasitoids reside in my research collections. RESULTS AND DISCUSSION TAXONOMY Adult.—WNeaspilota achilleae was de- scribed by Johnson (1900) from several adults collected from an apparent non-host plant, Achillea millefolium L. (Phillips 1946). Phillips (1923) sketched the pat- terned wing. Benjamin (1934) provided drawings of the male antenna, wing, caudal segment in posterior view, and genitalia; fe- male head in frontal view, wing, oviscape and aculeus. Freidberg and Mathis (1986: 28—32) redescribed this species, designated a male lectotype and three paralectotypes, and pictured the uniquely patterned wing (p. 72), along with drawings (p. 29) of the lateral aspect of the head; male right fore- tarsus, epandrium, distiphallus, epandrium and cerci, aculeus and its apex enlarged, 62 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON and spermatheca. Foote et al. (1993) pro- vided a photograph of a right wing and a drawing of the fifth tarsomere of the male foreleg. Immature stages.—Benjamin (1934) pro- vided generalized descriptions and habit sketches of the third-instar larva and pu- parium as well as sketches of the anterior spiracle, caudal segments, and spiracular plate of the third instar. Phillips (1946) de- scribed the third instar and sketched its ce- phalopharyngeal skeleton, anterior spiracle, caudal segment in posterior view, and pos- terior spiracular plate and interspiracular processes. The first-instar larva remains un- described, but the egg, second- and third- instar larvae and puparium are redescribed below with the aid of scanning electron mi- croscopy. Egg: Only three intact eggs were found and measured in situ within separate, im- mature, preblossom flower heads. These eggs were white, opaque, smooth, elongate- ellipsoidal, and averaged 0.61 + 0.02 (range, 0.58—0.64) mm long and 0.16 + 0.00 mm wide, tapered and smoothly rounded at both ends. As no eggs were ex- amined by scanning electron microscopy, the egg of N. achilleae could only be gen- erally compared with the eggs of N. viri- descens, N. wilsoni, and N. appendiculata, which were described in detail by Goeden and Headrick (1992, 1999) and Goeden (2000a). Second instar: White, elongate-cylindri- cal, rounded anteriorly, truncated postero- dorsally (Fig. 1A), body segments well-de- fined, anterior third of meso- and metatho- rax circumscribed anteriorly by minute acanthae, as is anterior half of abdominal segment A-1l, and all of segments A-2 through A-7 (Fig. 1D); dorsal sensory or- gan not well-defined, flattened (damaged in Figs. 1B-1, C-1); anterior sensory lobe (Figs. 1B-2, C) with terminal sensory organ (Figs. 1B-3, C-2), lateral sensory organ (Fig. 1C-3), supralateral sensory organ (Fig. 1C-4), and pit sensory organ (Fig. 1C- 5); stomal sense organ (Figs. 1B-4, C-6) ventrolaterad of anterior sensory lobe; mouthhook bidentate (Fig. 1B-5); median oral lobe not seen; four papilliform, inte- gumental petals in single row dorsal to each mouthhook (Fig. 1C-7); five oral ridges seen of an unknown total, dentate along posterior margins, in vertical series lateral to oral cavity (Fig. 1B-6); prothorax, cir- cumscribed anteriorly by posteriorly-direct- ed, minute acanthae (Fig. 1B-7); anterior thoracic spiracle with three, doliform papil- lae (Fig. 1D); lateral spiracular complexes not seen; caudal segment with two stelex sensilla dorsolaterad (Fig. 1E) and ventro- laterad (Fig. 1F) of posterior spiracular plate (Fig. 1G); posterior spiracular plate bears three ovoid rimae (Fig. 1G-1), ca. 0.012 mm long, and four interspiracular processes (Fig. 1G-2), each with four, sim- ple branches, longest measuring 0.008 mm; intermediate sensory complex (Fig. 1H) with a stelex sensillum (Fig. 1H-1) and a medusoid sensillum (Fig. 1H-2). The habitus of the second instar of N. achilleae (Fig. 1A) is more like N. wilsoni (Goeden and Headrick 1999), N. signifera (Goeden 2000a), N. aenigma (Goeden 2000b), N. appendiculata (Goeden 2000c), and N. pubescens (Goeden 2000d) than the ‘‘barrel-shaped”’ (doliform) second instar of N. viridescens (Goeden and Headrick 1992). The dorsal sensory organ of N. achilleae is not well-defined in the second instar (Figs. 1B-1, C-1), like N. viridescens (Goeden and Headrick 1992), N. wilsoni (Goeden and Headrick 1999), and N. aenig- ma (Goeden 2000b), but unlike N. signifera (Goeden 2000a), N. appendiculata (Goeden 2000c), and N. pubescens (Goeden 2000d), in which this organ is well defined. In Fig. 1B-1 and 1C-1, the cover of the dorsal sen- sory organ is torn away to reveal the large opening that it protects. The integumental petals of the second instars of all seven spe- cies are papilliform, but are only four in number above each mouth hook in N. achil- leae (Fig. 1C-7), like N. signifera (Goeden 2000a), but are six in number in N. virides- cens (Goeden and Headrick 1992) and N. VOLUME 103, NUMBER 1 63 Acc.V Spot Magn 10.0 kV 30 57x kic.V Spot Magn — & Spot Magn ‘Det=W TOOkKV 3.0 1809x mee OKV 3.0 1809x : Fig. 1. Second instar of Neaspilota achilleae: (A) habitus, anterior to left; (B) gnathocephalon, lateral view, 1 — dorsal sensory organ, 2 — anterior sensory lobe, 3 — terminal sensory organ, 4 — stomal sense organ, 5 — mouthhook, 6 — oral ridge, 7 — minute acanthae; (C) anterior sensory lobe, 1 — dorsal sensory organ, 2 — terminal sensory organ, 3 — lateral sensory organ, 4 — supralateral sensory organ, 5 — pit sensory organ, 6 — stomal sense organ, 7 — integumental petal, 8 — oral ridge; (D) anterior thoracic spiracle; (E) dorsolateral stelex sensillum; (F) ventrolateral stelex sensillum; (G) posterior spiracular plate, 1 — rima, 2 — interspiracular process, 3 — ecdysial scar; (H) intermediate sensory complex, 1 — stelex sensillum, 2 — medusoid sensillum. 64 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON pubescens (Goeden 2000d), seven in N. ap- pendiculata (Goeden 2000c) and N. wilsoni (Goeden and Headrick 1999), and eight in N. aenigma (Goeden 2000b). Neaspilota achilleae has three papillae on the anterior spircle of the second instar (Fig. 1D), like the three or four papillae reported in second instars of N. viridescens (Goeden and Head- rick 1992), N. wilsoni (Goeden and Head- rick 1999), N. signifera (Goeden 2000a), and N. aenigma (Goeden 2000b), but dif- ferent from the five papillae in N. pubes- cens (Goeden 2000b) and eight in N. ap- pendiculata (Goeden 2000c). Finally, the interspiracular processes of N. achilleae (Fig. 1G-2) each bear four branches, like those of N. appendiculata (Goeden 2000c), but not one to four branches like N. aenig- ma (Goeden 2000b) and N. pubescens (Goeden 2000d), nor two to four branches like N. signifera (Goeden 2000a), nor five to nine branches like those of N. viridescens (Goeden and Headrick 1992), nor two to six branches like those of N. wilsoni (Goeden and Headrick 1999). Third instar: Pale yellow, ellipsoidal, with posterior spiracular plate dark brown to black, tapering and truncated anteriorly; posterior spiracular plate on caudal segment flattened and upturned dorsally ca. 60° (Fig. 2A), minute acanthae circumscribe anterior fifth of meso- and metathorax, anterior fourth of first abdominal segment, but all of remaining abdominal segments, except for pleura of segments, A6 and A7; gnathoce- phalon conical (Fig. 2B); dorsal sensory or- gan defined by a crescentric fold (Fig. 2B- 1), attached dorsally (Fig. 2C-1) and punc- tured centrally and peripherally by pores (Fig. 2C-2); anterior sensory lobe (Fig. 2C) bears terminal sensory organ (Fig. 2C-3), lateral sensory organ (Fig. 2C-4), suprala- teral sensory organ (Fig. 2C-5), and pit sen- sory organ (Fig. 2C-6); eight, papilliform (upper row) or spatulate (lower row), inte- gumental petals in two rows above each mouthhook (Figs. 2C-7, D-1); at least six oral ridges (Figs. 2B-2, D-2), at least five toothed ventrally, and all lateral to oral cav- ity; stomal sense organ (Figs. 2C-8, D-3) ventrolaterad of anterior sensory lobe; mouthhook (Figs. 2B-3, D-4) tridentate; median oral lobe laterally flattened, (Figs. 2B-4, D-5); prothorax circumscribed by mi- nute acanthae (Fig. 2B-5); verruciform sen- silla circumscribe prothorax anteriorad of minute acanthae (Fig. 2B-6); anterior tho- racic spiracle on posterior margin of pro- thorax bears three or four oblong papillae (Fig. 2E); mesothoracic lateral spiracular complex with a spiracle (not shown) and six verruciform sensilla (Fig. 2F-1), one above and five below the spiracle; metathoracic lateral spiracular complex with a spiracle (Fig. 2F-2) and five verruciform sensilla (Fig. 2F-3), one above and four below the spiracle; lateral spiracular complexes of ab- dominal segments A-1 and A-2 each with a spiracle (Fig. 2G-1) and six verruciform sensilla (Fig. 2G-2), one dorsoposteriorad of the spiracle, another posteriorad of the spiracle, one pair ventroposteriorad of the spiracle aligned longitudinally, and another ventral pair aligned ventrally; a stelex sen- sillum dorsolaterad, laterad, and ventrola- terad (not shown) of posterior spiracular plate (not shown); each posterior spiracular plate bears three ovoid rimae, ca. 0.03 mm in length, and four interspiracular process- es, each with four, simple or forked, pointed branches, longest branch measuring 0.013 mm; intermediate sensory complex (Fig. 2H) with a stelex sensillum (Fig. 2H-1) and a medusoid sensillum (Fig. 2H-2). The habitus of the third instar of N. achil- leae is like that reported for N. viridescens (Goeden and Headrick 1992), N. wilsoni (Goeden and Headrick 1999), N. signifera (Goeden 2000a), and N. aenigma (Goeden 2000b), N. appendiculata (Goeden 2000c) and N. pubescens (Goeden 2000d). The pat- tern of the minute acanthae that circum- scribe the body segments is unique for UN. achilleae in that the acanthae, which uniquely appear to occur in individual de- pressions, increase their coverage posteri- orly from the anterior fifths of the thoracic segments to include nearly all of the ab- VOLUME 103, NUMBER 1 dominal segments beyond the first; where- as, in N. signifera (Goeden 2000a), N. ap- pendiculata (Goeden 2000c) and N. pubes- cens (Goeden 2000d), the anterior part of each body segment is circumscribed, in N. aenigma the anteriors, pleura, and posteri- ors of each segment are circumscribed (Goeden 2000b); in N. wilsoni, all interseg- mental areas and all abdominal segments except the pleura are circumscribed (Goe- den and Headrick 1999); and in N. virides- cens, the intersegmental areas are free of acanthae (Goeden and Headrick 1992). Like N. viridescens (Goeden and Headrick 1992), N. wilsoni (Goeden and Headrick 1999) and N. pubescens (Goeden 2000d), but unlike N. signifera (Goeden 2000a), N. aenigma (Goeden 2000b) and N. appendi- culata (Goeden 2000c), the dorsal sensory organ is not well-defined and flattened, and not dome-shaped, in the third instar of N. achilleae (Fig. 2C-1). Additional similarities involved the inte- gumental petals in the third instars of all six congeners examined to date, all of which are arranged in a double row above each mouthhook and papilliform (upper row) or spatulate (lower row) (Goeden and Head- rick 1992, 1999; Goeden 2000a, b, c, d). The integumental petals differ in number among species and generally increase in number between the last two instars. The stomal sense organ of the third instar of N. achilleae bears one pair each of campani- form and short papilliform (conical) sensilla (Figs. 2C-8, D-3); therefore, it appears slightly less complex than the stomal sense organ of the second instar, which bears at least six sensilla (Figs. 1B-4, C-6). How- ever, the stomal sense organs of the third instars of the five other congeneric species examined appear more complex than earlier instars and each bears different combina- tions of sensory structures, variously de- scribed as several cone-shaped sensilla in N. viridescens (Goeden and Headrick 1992); as papilliform and pit-type in N. wil- soni (Goeden and Headrick 1999); as ver- ruciform or ““compound verruciform”’ in N. 65 signifera (Goeden 2000a), as verruciform and pit-type in N. aenigma (Goeden 2000b), and as verruciform, pit-type, and cone-shaped or short papilliform in N. pu- bescens (Goeden 2000d). The third instars of all seven species of Neaspilota examined to date have oral ridg- es with dentate ventral margins character- istically arranged in vertical series ventro- laterad of the dorsal sensory organ and lat- erad of the oral cavity (Goeden and Head- rick 1992, 1999; Goeden 2000a, b, c, d; Figs. 2B-2, D-2). Though unfortunately partly hidden and not counted in N. achil- leae (Figs. 2B-2, D-2), the oral ridges num- ber eight in N. pubescens (Goeden 2000d), seven or eight in the third instar of N. aenigma (Goeden 2000b), seven in N. ap- pendiculata (Goeden 2000c), but six in the second and third instars of the other three congeners examined to date. The dentate margins and vertical, parallel arrangement of these oral ridges appears to be a generic character; however, Goeden (2000c, d) con- firmed that the oral ridges vary in number among third instars of some Neaspilota spe- cies. Also, the most ventral, eighth oral ridge of N. pubescens is not ventrally toothed (Goeden 2000d). The third instars of Trupanea imperfecta (Coquillett), T. jo- nesi Curran, T. nigricornis (Coquillett), T. pseudovicina (Hering), 7. signata Foote, and 7. wheeleri Curran also bear serrated oral ridges (Goeden and Teerink 1997b, 1998, 1999a; Goeden et al. 1998; Knio et al. 1996; Teerink and Goeden 1999), but these oral ridges appear to be fewer in num- ber, and are not arranged in a more or less regular, vertical row laterad to the mouth hook, as in Neaspilota. The mouthhooks of the third instars of N. appendiculata, N. aenigma, N. signifera, N. viridescens, and N. achilleae, and prob- ably N. pubescens, are tridentate (Goeden and Headrick 1992; Goeden 2000a, b, c; Goeden 2000d; Figs. 2B-3, D-4); whereas, those of the third instar of N. wilsoni are bidentate (Goeden and Headrick 1999). Such interspecific differences in dentation 66 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON pot Magp- Det WD Ex ) 795% SE Fig. 2. Third instar of Neaspilota achilleae: (A) habitus, anterior to left; (B) gnathocephalon, lateral view, | — dorsal sensory organ, 2 — oral ridge, 3 — mouthhook, 4 — median oral lobe, 5 — minute acanthae, 6 — verruciform sensillum; (C) anterior sensory lobe, 1 — dorsal sensory organ, 2 — pores, 3 — terminal sensory organ, 4 — lateral sensory organ, 5 — supralateral sensory organ, 6 — pit sensory organ, 7 — integumental petal, 8 — stomal sense organ; (D) oral cavity, ventrolateral view, 1 — integumental petals, 2 — oral ridge, 3 VOLUME 103, NUMBER 1 are supported by our findings that the mouthhooks of third-instar Trupanea vicina (Wulp) are bidentate; whereas, those of 12 other congeners examined from California are tridentate (Goeden and Teerink 2000b and citations therein). Phillips (1946) described and illustrated the anterior spiracle of what I presume was the third instar as having three papillae; whereas, three and four papillae were re- ported by Benjamin (1934), as also reported for the anterior spiracles of my specimens (Fig. 2E). Moreover, the interspiracular pro- cesses of eastern U.S. specimens were de- scribed and illustrated by Phillips (1946) as being six to 12 in number, lanceolate, and almost never branched; whereas, my spec- imens had interspiracular processes each of which were four-branched (or four in num- ber in Phillips terminology), pointed and simple (lanceolate) or forked. These results indicate morphological differences in east- ern and western U.S. specimens of third in- stars of N. achilleae (see discussion below on distribution). The number and appearance of the stelex sensilla surrounding the posterior spiracular plate differ among the Neaspilota species examined to date. These number only four in the first instars of N. wilsoni (Goeden and Headrick 1999), N. aenigma (Goeden 2000b), and N. appendiculata (Goeden 2000c), but, unfortunately, were not ob- served with N. signifera (Goeden 2000a) or in the present study. This count of stelex sensilla remains at four in the second instars of N. aenigma (Goeden 2000b) N. appen- diculata (Goeden 2000c), N. pubescens (Goeden 2000d), and N. achilleae (Fig. 1E, F), and increases to six in third instars of N. wilsoni (Goeden and Headrick 1999), N. aenigma (Goeden 2000b), N. appendiculata oe 67 (Goeden 2000c), and N. achilleae. These stelex sensilla also show inter-instar (intra- specific) and interspecific differences in the incidence and appearance of the minute acanthae that may ring them basally, but this was not recognized, studied or recorded by my coworkers and me until recently (Goeden 2000b, c, d). The stelex sensilla of the second instar of N. achilleae are sur- rounded by four to seven, conical, minute acanthae (Figs. 1E, F); however, these ste- lex sensilla lack such basal minute acanthae in the third instar. Puparium: Dirty white to light brown, transversely ringed with narrow, brown or black lines, posterior two or three segments brown to black, ellipsoidal, and smoothly rounded at both ends (Fig. 3A); anterior end bears the invagination scar (Fig. 3B-1) and anterior thoracic spiracles (Fig. 3B-2); cau- dal segment circumscribed by minute acan- thae; three stelex sensilla, dorsolaterad, lat- erad, and ventrolaterad of posterior spirac- ular plates; posterior spiracular plate bears three broadly elliptical rimae (Fig. 4C-1), and four interspiracular processes, each with four, simple or forked, apically pointed branches (Fig. 4C-2); intermediate sensory complex with a medusoid sensillum and a stelex sensillum. Thirty-nine puparia aver- aged 2.65 + 0.04 (range, 2.07—2.99) mm in length; 1.14 + 0.015 (range, 0.92—1.30) mm in width. DISTRIBUTION AND HOosTsS Freidberg and Mathis (1986) described the distribution of N. achilleae as, ‘“‘Eastern and Gulf coast of North America from Newfoundland to Alabama and southwest- ern United States, southern California and Arizona.’ They and Foote et al. (1993) both mapped the distribution to include — stomal sense organ, 4 — mouthhook, 5 — median oral lobe; (E) anterior thoracic spiracle; (F) meso- (left) and metathoracic (right) lateral spiracular complexes, 1 — verruciform sensilla on mesothorax, 2 — spiracle and 3 — verruciform sensilla on metathorax; (G) lateral spiracular complexes of first (left) and second (right) abdominal segments, 1 — spiracle, 2 — verruciform sensilla; (H) intermediate sensory complex, | — stelex sensillum, 2 — medusoid sensillum. 68 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Acc. V Spot Magn 5.00kV 3.0 306x Det WD Exp 100 jim Fig. 3. Puparium of Neaspilota achilleae: (A) hab- itus, anterior to left; (B) anterior end, 1 — invagination scar, 2 — anterior thoracic spiracle; (C) caudal seg- ment, 1 — rima, 2 — interspiracular process, 3 — intermediate sensory complex. only a single location in Arizona. Freidberg and Mathis (1986, p. 32) also remarked that: “The known distribution of N. achilleae is peculiar. It is apparently widespread and com- mon and has been reared from several host plants from numerous sites along the east coast. However, several specimens have also been examined from California and Arizona. The latter specimens are smaller on the av- erage than their eastern counterparts, but oth- erwise they seem to be conspecific. Further collecting and rearing will be necessary to de- termine whether the distribution of this spe- cies is disjunct or if the species actually ex- tends across southern United States.’’ Goeden (1989) discussed the host plants of N. achilleae and noted that Freidberg and Mathis (1986) listed four species of Aster, three species of Chrysopsis, three species of Hieraceum, and one species each of Pren- anthes, Sericocarpus, and Trilisa as hosts of N. achilleae in eastern United States. Up- dating Goeden (1989), I note that I have reared other genera and species of Neaspi- lota, but never N. achilleae, from totals of 50 samples of mature flower heads of 10 species of Aster, 17 samples of three spe- cies of Chrysopsis, 43 samples of 10 spe- cies of Erigeron, five samples of two spe- cies of Hieraceum from California. On the other hand, I have not reared N. achilleae from one sample of Stephanomeria cichor- iaea Gray, eight samples of S. exigua Nut- tall, nor two samples of S. parryi Gray. My host record for S$. pauciflora (see above) is new. Eastern host plants belong to the sub- tribes Asterinae, Hieraciinae, Lactucinae, Liatrinae, and Solidaginae of the tribes As- tereae, Eupatorieae, and Lactu~eae, but in- clude no other Stephanomeriinae which pri- marily are western plant species (Bremer 1994). However, eastern and western UN. achilleae do share hosts in the Lactucinae (Bremer 1994). Thus, in southern Califor- nia, N. achilleae appears to be nearly mo- nophagous; whereas, in the eastern U.S., it is a generalist (Headrick and Goeden 1998). BIOLOGY Egg.—In each of seven, closed, preblos- som, immature flower heads of S. pauciflo- ra a single egg of N. achilleae was inserted pedicel-last; six of these eggs were placed VOLUME 103, NUMBER 1 69 Fig. 4. Life stages of Neaspilota achilleae in Stephanomeria pauciflora (Figs. A, B, D) and S. virgata (Fig. C): (A) egg between ovules in closed, preblossom flower head, (B) early-third instar, (C) late-third instar, (D) newly formed puparium, (E) mating adults, lateral view, (F) mating adults, dorsolateral view. Lines = 1 mm. between an inner phyllary and a peripheral floret with their long axes parallel to the long axes of the flower heads (Fig. 4A). The seventh egg had been inserted through the phyllaries and was embedded for half its length in the corolla of a peripheral flo- ret. Larva.—Upon eclosion, the seven first instars found feeding in separate, preblos- som flower heads either tunneled into an ovule, or into a corolla before entering the ovule to which the corolla was basally at- tached. The receptacles averaged 0.87 + 0.08 (range, 0.57—1.12) mm in diameter and 70 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON an average of 1.3 + 0.19 (range, 1-2) flo- rets/ovules was damaged in these seven flower heads. No receptacle was abraded or pitted by larval feeding. Based on 5 + 0.07 (range, 3—6) as the average total number of florets, ovules or achenes respectively counted in 45 preblossom to postblossom flower heads, about 26 + 3.7% (range, 20— 40%) of the ovules in the seven flower heads were damaged by first instars. Second instars continued feeding on co- rollas or ovules in preblossom flower heads. All fed with their bodies more or less per- pendicular to and their mouthparts directed towards or away from the receptacles, but always well above the receptacles. Recep- tacles of three flower heads containing sec- ond instars averaged 0.84 + 0.05 (range, 0.75—0.93) mm in diameter. These flower heads each contained a single larva that had damaged an average of 3 + 1 (range, 2—5) florets/ovules, or about 60% (range, 40- 100%) of the average total of 5 ovules per flower head. Third instars (Fig. 4B) initially continued to feed mainly on ovules and soft achenes in preblossom, blossom, or postblossom flower heads. Seventy-one flower heads that averaged 1.06 + 0.06 (range, 0.56—3.13) mm in diameter each contained a single third instar or puparium. An average of 5.0 + 0.09 (range, 3—6) of the soft achenes therein were damaged, or 100% of the average total of 5 ovules/soft achenes per flower head (Fig. 4C). These percentages of seed preda- tion per larva per flower head, like those re- ported for N. aenigma (Goeden 2000b), N. appendiculata (Goeden 2000c), and N. pu- bescens (Goeden 2000d), are higher than other florivorous tephritids studied by us to date (Headrick and Goeden 1998). For N. achilleae, this higher percentage results from the small-size flower heads of the host plants attacked in southern California and small number of ovules/achenes contained in each flower head (Fig. 4C). Higher percentage seed destruction per flower head also was evidenced by gregarious florivorous species like Trupanea conjuncta (Adams) (Goeden 1987) and T. pseudovicina Hering (Goeden and Teerink 1998) or by species with large larvae that develop in immature or small flower heads like Paracantha cultaris (Co- quillett) (Cavender and Goeden 1982) and Xenochaeta dichromata Snow (Goeden and Teerink 1997a). Also, the rate of flower head infestation per sample for N. achilleae was very low, e.g., a mean of 2.9% (range, 1%— 6%) for seven subsamples of 200 to 400 dis- sected flower heads, similar to most other Neaspilota that we have studied (unpub- lished data). Third instars in flower heads fed with their long axes oriented perpendicular to and mouthparts directed towards the recep- tacles (Fig. 4C). Only seven (10%) of the third instars in the 71 infested heads con- taining thirds instars or puparia scored or pitted the receptacles; however, most larvae presumably supplemented their diet with sap. Goeden and Headrick (1992, 1999) and Goeden (2000c, d) described and discussed this type of feeding by N. viridescens, N. wilsoni, N. appendiculata, and N. pubes- cens, respectively. Upon completing feed- ing, the larvae oriented with their anterior ends away from the receptacles, retracted their mouthparts, and formed prepuparia (Headrick and Goeden 1998). Most individ- uals pupariated (Fig. 4D) and emerged in summer (July—August) and either produced at least one additional generation in the same, long-blooming host-plant species or in its alternate host plants growing at higher elevations. Adult.—Adults are long-lived in nature. Under insectary conditions, 17 unmated males averaged 69 + 10 (range, 11—139) days, and 11 virgin females averaged 30 + 6 (range, 11—75) days. Such lengthy lon- gevities for males compare favorably with average adult longevities reported for adults of N. viridescens (Goeden and Headrick 1992), N. wilsoni (Goeden and Headrick 1999), N. signifera (Goeden 2000a), N. aenigma (Goeden 2000b), N. appendiculata (Goeden 2000c), and N. pubescens (Goeden 2000d). The shorter average longevities re- VOLUME 103, NUMBER 1 corded for females of N. achilleae are un- explained. The premating and mating behaviors of N. achilleae were not studied in the field, but were observed in petri dish arenas found to be so useful with many other non- frugivorous, tephritid species (Headrick and Goeden 1994). Premating behaviors ob- served with paired N. achilleae were ab- dominal pleural distension and side-step- ping by males while tracking females (Headrick and Goeden 1994) and rapid wing hamation, sometimes combined with lofting about 20° by both sexes (Headrick and Goeden 1994). No trophallaxis or nup- tial gift presentation was noted as reported with N. viridescens (Goeden and Headrick 1992). Six matings, two by one pair during a 2-week period (Figs. 4E, F) were ob- served, most of which began during late af- ternoon at dusk and lasted an average of 153 + 54 (range, 80—420) min (or 2 h and 33 min on average). Except for the single pair that began mating at 08:00 h and re- mained in copula for 420 min (7 h), this average was shorter than the average du- rations of 190 min reported for N. aenigma (Goeden 1999b), 235 min reported for N. wilsoni (Goeden and Headrick 1999), 238 min reported for N. signifera (Goeden 2000a), 285 min reported for N. appendi- culata (Goeden 2000c), 318 min reported for N. viridescens (Goeden and Headrick 1992), and 1032 min reported for N. pu- bescens (Goeden 2000d). All matings were observed only after copulation had begun, suggesting that pre- copulatory behavior was perfunctory. The mating position (Fig. 4E, F) was such that the wings of the male were parted from about 5 to 30°, while the wings of the fe- male were parted at about 30 to 80°, with both pairs of wings centered over their re- spective body midlines. The body of the fe- male was held parallel to the substrate, while the body of the male was elevated about 20° anteriorly (Fig. 4E). The hind- and midtarsi of the male usually rested on the substrate (Figs. 4E, F), but sometimes al his midtarsi grasped the base of the aculeus, while his foretarsi hooked over the anterior abdominal terga of the female (Figs. 4E, F). The male’s mouthpars were positioned above syntergite 1 + 2 of the female (Figs. 4E, F). The pair largely remained quiescent with only their mouthparts pumping, or sometimes both formed regurgitation drop- lets (Headrick and Goeden 1994). Besides tightly clinging to the female, the male would sometimes reposition his foretarsi so as to gain better purchase or rub his mid- tarsi alternately along her oviscape to in- duce passivity as a variation on copulatory induction behavior (Headrick and Goeden 1994). These bursts of activity as the fe- male kicked at the male and arched her body in an attempt to dislodge the male al- ways preceded disengagement, but only in- frequently resulted in disengagement; sim- ilar behavior also was reported for N. pu- bescens (Goeden 2000d). A single mating termination and disen- gagement was observed, which involved the male turning in place and walking away from the female while pulling his phallus from within her, however, this process took 34 sec as the male had difficulty freeing himself. Disengagement by N. pubescens (Goeden 2000d) was reported to last less than 10 s. Postcopulatory behavior by N. achilleae mainly consisted of storing of the genitalia by males and cleaning and groom- ing by both sexes (Headrick and Goeden 1994). Seasonal history.—The life cycle of N. achilleae in southern California follows an aggregative pattern (Headrick and Goeden 1994, 1998) in which the adult is the prin- cipal overwintering stage. Come late spring (May), overwintered adults aggregate on preblossom shoots of S. pauciflora to mate and oviposit in the small, newly-formed, closed, preblossom flower heads begining in June. The F, larvae feed until fully grown, then pupariate in flower heads and emerge as adults in summer (July—August) to produce another generation at higher el- evations on S. pauciflora or on S. virgata 72 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON that flower through September—October. There are at least two generations per year, with F, adults emerging in October—No- vember to overwinter as long-lived adults in reproductive diapause. Neaspilota signi- fera (Goeden 2000a) and N. aenigma (Goe- den 2000b) also overwinter principally as adults in southern California. Natural enemies.—A male and a female of Pteromalus sp. (Hymenoptera: Pterom- alidae) were reared from separate puparia of N. achilleae as solitary, larval-pupal en- doparasitoids. One female each of Euryto- ma obtusiventrus Gahan or near and E. ve- ronia Bugbee (Hymenoptera: Eurytomidae) were reared from separate flower heads of S. virgata and S. pauciflora, respectively, as probable, solitary, larval-pupal endoparasi- toids. ACKNOWLEDGMENTS I thank Andrew C. Sanders, Curator of the Herbarium, Department of Botany and Plant Sciences, University of California, Riverside, for identifications of plants men- tioned in this paper. Krassimer Bozhilov in the Institute of Geophysics and Planetary Physics, University of California, River- side, greatly facilitated my scanning elec- tron microscopy. The parasitoids were iden- tified by Harry E. Andersen, now deceased, Huntington Beach, California, and Michael Gates, Department of Entomology, Univer- sity of California, Riverside. I also am grateful to Jeff Teerink and Kristine Gilbert for technical assistance and to Louie Blanc, now deceased, David Headrick, and Jeff Teerink for their helpful comments on ear- lier drafts of this paper. LITERATURE CITED Benjamin, F H. 1934. Descriptions of some native try- petid flies with notes on their habits. United States Department of Agriculture Technical Bulletin 401: 1-95, Bremer, K. 1994. Asteraceae Cladistics & Classifica- tion. Timber Press, Inc. Portland, Oregon. Cavender, G. L. and R. D. Goeden. 1982. Life history of Trupanea bisetosa (Diptera: Tephritidae) on wild sunflower in southern California. Annals of the Entomological Society of America 75: 400— 406. Foote, R. H., E L. Blanc, and A. L. Norrbom. 1993. Handbook of the Fruit Flies (Diptera: Tephritidae) of America North of Mexico. Cornell University Press, Ithaca, New York. Freidberg. A. and W. N. Mathis. 1986. Studies of Ter- elliinae (Diptera: Tephritidae): A revision of the genus Neaspilota Osten Sacken. Smithsonian Contributions to Zoology 439: 1—75. Goeden, R. D. 1987. Life history of Trupanea con- juncta (Adams) on Trixis californica Kellogg in southern California (Diptera: Tephritidae). The Pan-Pacific Entomologist 63: 284-291. . 1989. Host plants of Neaspilota in California (Diptera: Tephritidae). Proceedings of the Ento- mological Society of Washington 91: 164-168. . 2000a. Life history and description of im- mature stages of Neaspilota signifera (Coquillett) (Diptera: Tephritidae) on Hemizonia pungens (Hooker and Arnott) Torrey and A. Gray (Aster- aceae) in southern California. Proceedings of the Entomological Society of Washington 102: 69— 81. . 2000b. Life history and description of im- mature stages of Neaspilota aenigma Freidberg and Mathis (Diptera: Tephritidae) on Erigeron divergens Torrey and Gray (Asteraceae) in south- ern California. Proceedings of the Entomological Society of Washington 102: 384-397. . 2000c. Life history and description of im- mature stages of Neaspilota appendiculata Freid- berg and Mathis (Diptera: Tephritidae) on Ma- chaeranthera canescens (Pursh) A. Gray (Aster- aceae) in southern California. Proceedings of the Entomological Society of Washington 102: 519— 532) . 2000d. Life history and description of im- mature stages of Neaspilota pubescens Freidberg and Mathis (Diptera: Tephritidae) on Lessingia fi- laginifolia (Hooker and Arnott) M. A. Lane (As- teraceae). Proceedings of the Entomological So- ciety of Washington 103: 878-891. Goeden, R. D. and D. H. Headrick. 1992. Life history and descriptions of immature stages of Neaspilota viridescens Quisenberry (Diptera: Tephritidae) on native Asteraceae in southern California. Proceed- ings of the Entomological Society of Washington 94: 59-77, . 1999. Life history and description of imma- ture stages of Neaspilota wilsoni Blane and Foote (Diptera: Tephritidae) on Hazardia squarrosa (Hooker and Arnott) E. Greene (Asteraceae). Pro- ceedings of the Entomological Society of Wash- ington 101; 897-909. Goeden, R. D. and J. A. Teerink 1997a. Life history and description of immature stages of Xenochaeta dichromata Snow (Diptera: Tephritidae) on Hier- VOLUME 103, NUMBER 1 acium albiflorum Hooker in central and southern California. Proceedings of the Entomological So- ciety of Washington 99: 597—607. . 1997b. Life history and description of im- mature stages of Trupanea signata Foote (Diptera: Tephritidae) on Gnaphalium luteo-album L. in southern California. Proceedings of the Entomo- logical Society of Washington 99: 748-755. . 1998. Life history and description of imma- ture stages of Trupanea pseudovicinia Hering (Diptera: Tephritidae) on Porophyllum gracile Bentham (Asteraceae) in southern California. Pro- ceedings of the Entomological Society of Wash- ington 100: 361-372. . 1999a. Life history and description of im- mature stages of Trupanea wheeleri Curran (Dip- tera: Tephritidae) on Asteraceae in southern Cal- ifornia. Proceedings of the Entomological Society of Washington 101: 414—427. . 1999b. Life history and description of im- mature stages of Trupanea vicina (Wulp) (Diptera: Tephritidae) on wild and cultivated Asteraceae in southern California. Proceedings of the Entomo- logical Society of Washington 101: 742-755. Goeden, R. D., J. A. Teerink, and D. H. Headrick. 1998. Life history and description of immature stages of Trupanea jonesi Curran (Diptera: Te- phritidae) on native Asteraceae in southern Cali- fornia. Proceedings of the Entomological Society of Washington 100: 126-140. 73 Headrick, D. H. and R. D. Goeden. 1994. Reproduc- tive behavior of California fruit flies and the clas- sification and evolution of Tephritidae (Diptera) mating systems. Studia Dipterologica 1(2): 194— MSY). . 1998. The biology of nonfrugivous tephritid fruit flies. Annual Review of Entomology 43: 217-241. Hickman, J. C. (ed.) 1993. The Jepson Manual. Uni- versity of California Press. Berkeley and Los An- geles. Johnson, C. W. 1900. Some notes and descriptions of seven new species and one new genus of Diptera. Entomological News 11: 323-328. Knio, K. M., R. D. Goeden, and D. H. Headrick. 1996. Descriptions of immature stages of Trupanea ni- gricornis and T. bisetosa (Diptera: Tephritidae) from southern California. Annals of the Entomo- logical Society of America 89: 1-11. Phillips, V. T. 1923. A revision of the Trypetidae of northeastern America. Journal of the New York Entomological Society 31: 119-155. . 1946. The biology and identification of try- petid larvae. American Entomological Society Memoirs 12: 1-161. Teerink, J. A. and R. D. Goeden. 1999. Description of the immature stages of Trupanea imperfecta (Co- quillett). Proceedings of the Entomological Soci- ety of Washington 101: 75-85. PROC. ENTOMOL. SOC. WASH. 103(1), 2001, pp. 74-80 A NEW SPECIES OF THE GENUS DILAR RAMBUR (NEUROPTERA: DILARIDAE) FROM BORNEO JOHN D. OSWALD AND NATHAN M. SCHIFF (JDO) Department of Entomology, Texas A&M University, College Station, TX 77843- 2475, U.S.A. (e-mail: j-oswald@tamu.edu); (NMS) Southern Research Station, Center for Bottomland Hardwood Research, U.S. Forest Service, U.S. Department of Agriculture, PO. Box 227, Stoneville, MS 38776, U.S.A. (e-mail: nschiff@fs.fed.us) Abstract.—Dilar macleodi is described as a new species from lowland rainforest habitat in the Malaysian state of Sarawak on the island of Borneo. Diagnoses are provided to distinguish D. macleodi from the four other dilarid species that have been reported from the peninsula of Indochina or the Malay Archipelago. Key Words: The small neuropterous family Dilaridae, “pleasing lacewings,’’ presently includes 67 extant species, with a combined distri- bution encompassing parts of North and South America, Europe, Asia and Africa (Oswald 1998). No extant dilarids are known from Australia or New Guinea. En- gel (1999) recently described the only known fossil dilarid, Cascadilar eocenicus, an adult male from Baltic amber. Dilarids are associated with woodland and forest en- vironments, where their larvae live in cor- ticolous or terricolous microhabitats and feed on small arthropod prey. Old World dilarid species are currently placed in four genera—three in the subfamily Dilarinae: Berothella Banks (2 spp.; China, continen- tal Malaysia), Dilar Rambur (45 spp.; wide- spread throughout the Oriental and southern Palearctic regions) and Neonallachius Na- kahara (1 sp.; India), and one genus in the subfamily Nallachiinae: Nallachius Navas (2 spp.; Vietnam, southeastern Africa). The adults of most species are relatively rarely collected and larvae are known for only five species (Oswald 1998). The purpose of the present paper is to Insecta, Dilaridae, pleasing lacewings, Borneo, Sarawak, taxonomy describe Dilar macleodi as a new species from the Malaysian state of Sarawak, on the island of Borneo. Illustrations of male and female terminalia were prepared with the aid of a drawing tube from abdomens mac- erated in KOH and stained with Chlorozol Black. Male terminalic structures are de- scribed with the gonopons, hemigonarcus, 9th gonocoxite terminology of Oswald (1993a, b), rather than the gonarcus, gono- coxite, paramere terminology of some re- cent European authors (e.g., Aspdéck and Aspock 1995, Monserrat 1988). Dilar macleodi Oswald and Schiff, new species (Figs. 1-10) Diagnosis.—Dilar macleodi is readily distinguishable from other dilarids known from Indochina and the Malay Archipelago by the following characters [D. macleodi character states in square brackets]: D. viet- namensis Zakharenko: forewing with dis- tinct transverse bars [not barred], dorsopro- cessus of male terminalia absent [present], apices of male 9th gonocoxites not oppos- able on hemigonarcus [opposable]; D. VOLUME 103, NUMBER 1 grandis (Banks): male forewing 11—16 mm long [much shorter, ca. 6 mm], dorsopro- cessus of male short and membrane-mar- gined to a rounded apex [dorsoprocessus elongate, apex not membrane margined but well sclerotized and truncate], apices of male 9th gonocoxites bicuspate [unicuspa- te]; D. marmoratus (Banks): dorsoproces- sus absent [present and elongate], apices of male 9th gonocoxites lanceolate, not op- posable on hemigonarcus [recurved and op- posable]; dorsodistal angles of hemigonar- cus prolonged as a pair of very slender, elongate, processes [no analogous process- es]; Berothella phantoma Banks (not seen, comparison by Steve Brooks, see acknowl- edgements): gonarcus less massive [more massive, with proximolateral regions broad in dorsal view], dorsoprocessus larger and more prominent relative to 9t + ectoproct, apex quadrate [smaller with apex expand- ed]. Other details of the male terminalia, e.g., the conformation and armature of the supraanal, also differ considerably among these species. Description (from specimens in ethyl] al- cohol).—Head: Head capsule, mouthparts and antenna yellowish white; vertex with three distinct, setose, pulvini, two postero- lateral, one dorsomedial; eye dark brown. Antenna: Scape prominent, simple; ped- icel small, annular, simple; female flagellum filiform, flagellomeres (mean = 18, n = 2 antennae) cylindrical, decreasing in length distally; male flagellum pectinate, flagel- lomeres (mean = 19.5, range = 18-23, n = 8 unbroken antennae) bipectinate (basal flagellomere only [= two fused flagello- meres?]), unipectinate (central flagellomer- es) or simple (apical 5—7 flagellomeres). Thorax.—Coloration: Yellowish white with 13 consistently-placed, contrasting brown maculae as follows: Pronotum, lat- eral margins (2 maculae); mesonotum, an- terior margin (3); mesoscutum, lateral (2); mesoscutum, posterior parasagittal (2); mesanepisterna (2); metanepisterna (2). Metascutum sometimes also with a pair of less-distinct lateral brown maculae. Female Ws) L (2.0 mm) Figasle Dilar macleodi, male, wings. specimen also with a posteromedian pron- otal macula. Legs pale, apices of podo- meres generally narrowly embrowned. Forewing (Fig. 1).—Length: Male 5.0— 7.3 mm (mean = 5.9 mm, n = 10 wings), female 8.2 mm (n = 1 wing). Coloration: Subcostal veinlets and most crossveins brownish; longitudinal veins with irregular- ly alternating brownish and pale segments; membrane hyaline with narrow brownish margining adjacent to some dark vein seg- ments, not distinctly cross-banded as in some Dilar species. Vestiture (dorsal surface): Longitudinal veins and subcostal veinlets densely setose [but severely rubbed from storage in alco- hol]; crossveins asetose; membrane evenly microtrichose; anal angle of forewing with a patch of stout microtrichia on ventral sur- face. Venation: Trichosores prominent along wing margin from pterostigma through cu- bital region, less distinct or absent on more proximal wing margins; subcostal veinlets simple, rarely forked; subcosta free from, or just touching, R1 in pterostigmal region; subcostal crossveins weak and inconspicu- ous, irregular in number and position (often 4 or more); anterior Rs trace with 3—5 pec- 76 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 2-3. accessory glands damaged and lost during preparation, only the bases of these glands are illustrated). Abbre- viations: 7s, seventh sternite; 7t, 8t, 9t, tergites; 9gcx, ninth gonocoxite; bur, bursa; bag, bursal accessory gland; cg, colleterial gland; ect, ectoproct. tinate branches proximal to pterostigma (in- cluding MA); MA arising from radial stem, basal “‘b”’ crossvein generally present only as a short stub from base of MA; MP deep- ly two-branched; two distinct series of gra- date crossveins present. Nygmata: Two present, proximal nygma located between MA and MP near base of MA, distal nygma located between Rs and MA near base of proximalmost Rs branch. Hind wing (Fig. 1).—Length: Male 3.9— 6.1 mm (mean = 4.9 mm, n = 10 wings), female 6.4 mm (n = | wing). Coloration: Generally similar to fore- wing, but paler. Venation: Basal piece of MA present, nearly linear (not markedly sinuous), joined distally to RS. Nygmata: One present, located between MA and Rs near base of proximalmost Rs branch. Female terminalia.—Tergite and sternite 7 (Fig. 2: 7t, 7s): Hemiannular, unmodified. Tergite 8 (Fig. 2: 8t): Membranously divid- ed dorsally into a pair of lateral hemitergi- tes, hemitergites prolonged ventrolaterally and enclosing spiracles of 8th somite. Sternite 8: Apparently absent. Tergite 9 (Fig. 2: 9t): Membranously di- Dilar macleodi, female. 2, Abdominal apex, lateral. 3, Internal genitalia, lateral (apices of bursal vided dorsally into a pair of elongate lateral hemitergites, hemitergites articulated pos- teroventrally to ipsilateral 9th gonocoxites, each hemitergite braced by a strong internal costa that runs anterodorsally from point of articulation. Ectoproct (Fig. 2: ect): A small [vesti- gial] pulvinate prominence, cercal callus and trichobothria absent [lost]. 9th gonocoxites (Fig. 2: 9gcx): Paired, slender and greatly elongate, together form- ing a flexible ovipositor, each gonocoxite bearing proximally a weak costa that runs longitudinally from point of articulation with ipsilateral 9th hemitergite, apical sty- lus absent [lost]. Postgenitalia: Absent. Subgenitale: Absent. Bursa copulatrix (Fig. 3: bur): Membra- nous or weakly sclerotized, without discrete lobes; colleterial gland present (cg), colle- terial accessory gland apparently absent, one pair of bursal accessory glands (bag) present. Insemination/fertilization canal: Appar- ently absent; floor of bursa apparently lack- ing the well-sclerotized, median, “‘fertiliza- tion canal’’ found in many other neuropter- an families [possibly an artifact of the sin- VOLUME 103, NUMBER I gle, somewhat over-cleared, female available for study]. Male Terminalia.—Tergite and sternite 8 (Fig. 4: 8t, 8s): Weakly sclerotized, hem- iannular, unmodified. Pereite: 9. + ectoproct (Figs.-4, 5: 9t + ect): Ectoproct completely fused to 9th hemitergite, suture line lacking, combined sclerite broad and evenly rounded distally, cercal callus and trichobothria absent [lost], contralateral sclerites joined dorsally by a distinct dorsoprocessus (dor). Dorsoprocessus (Figs. 4—6: dor): A well- sclerotized, slender, longitudinal process; slightly broadened distally to a shallowly concave, truncate apex (Fig. 6); joined to dorsal margins of 9th tergite + ectoproct sclerites by a pair of narrow arms (Fig. 5); each arm underlain internally by a stout costa; costae fused distally and together providing primary rigidity of dorsoproces- sus. Sternite 9: Strongly reduced (often not visible in lateral view), transverse, distal margin rounded. Supraanal (Figs. 4, 7: spa): Bipartite, with dorsal and ventral parts divided by a transverse sulcus; dorsal part consisting principally of a transverse subquadrangular plate, proximodorsal margin of plate shal- lowly emarginate medially, lateral margins narrowly flanged, distoventral margin bear- ing a pair of slender, sharply-pointed, para- sagittal processes, plate surface with a pair of shallow lateral depressions separated by a weak sagittal ridge, surface asetose but partially microtrichose; ventral part of su- praanal consisting of a pair of rounded, ventrally papillate, lobes, each of which bears a sharply-pointed process dorsally. Gonarcus (Figs. 4, 8-10: gon): U-shaped in overall from (dorsal view, Fig. 9); gon- opons (gps) short and narrow; extrahemi- gonarcus (ehgs) large, revolute, directed posteriorly and lying adjacent to length of 9th gonocoxites; intrahemigonarcus (ihgs) restricted to a narrow band at anterior end of extrahemigonarcus; revolute hemigonar- cus broad proximally, narrowed distally. a Mediuncus: Absent. 9th gonocoxites (Figs. 4, 8, 10: 9gcx): Paired, each composed of an elongate basal plate and a free distal process; basal plate membrane-margined, its dorsal, subbasal, margin articulated to gonarcus (art), its dor- sal, subapical margin bearing a field of slender apodemal filaments (ap) indicative of a muscle attachment; distal process arched dorsally, its apex opposable on dis- toventral surface of ipsilateral hemigonar- cus (Fig. 8: opp). Hypandrium internum: Present. Distribution.—Currently known only from the Malaysian state of Sarawak on the island of Borneo. Primary type.—Holotype, male (Texas A&M University Insect Collection). MA- LAYSIA: Sarawak: Borneo [island], Gun- ung [= Mt.] Buda, ca. 64 km S of Limbang, 4°13'N 114°56’E. Verbatim label data: “MALAYSIA: Sarawak: / Borneo, Gunung [= Mt.] / Buda, ca. 64 km S of Limbang / 4°13’N 114°56’E Malaise [trap] / 5— 25.x1.1996 N{[athan]. Schiff’? [white rect- angle], “‘Holotype / Dilar / macleodi Os- wald &/ Schiff /J.D. Oswald 1999” [red rectangle]. Condition: excellent, no parts missing, critical point dried from ethyl al- cohol, point mounted, terminalia not dis- sected. Other material examined.—281,000—2,000/ branch [Sutton 1983]) would facilitate the psyllid’s establishment in North America. ACKNOWLEDGMENTS We gratefully acknowledge Gary Miller (USDA, ARS, Systematic Entomology Laboratory, Beltsville, Md.) for identifying Cacopsylla peregrina and providing copies of several papers on psyllids, Raymond J. Gill (Plant Health and Pest Prevention Ser- vices, California Department of Food and Agriculture, Sacramento) for sharing infor- mation on C. peregrina in California and Oregon, Robert Foottit and Eric Maw (Ag- riculture and Agri-Food Canada, Eastern Cereal and Oilseed Research Centre, Otta- wa) for providing collection data for the specimens from British Columbia, Daniel Burckhardt (Naturhistorisches Museum, Basel, Switzerland) and Peter Adler (De- partment of Entomology, Clemson Univer- sity) for helpful comments on an earlier draft of the manuscript, John Townsend (Department of Biological Sciences, Clem- son University) for identifying the Cratae- gus specimen from Poulsbo, Wash., James Phipps (Department of Plant Sciences, Uni- versity of Western Ontario, London) for comments on the naming of cultivated forms of Crataegus, Iain Robertson (De- partment of Landscape Architecture, Uni- versity of Washington, Seattle) for help in locating ornamental plantings of Crataegus, and Brill Academic Publishers (Leiden, the Netherlands) for granting permission to re- produce figures from Ossiannilsson’s (1992) work. LITERATURE CITED Burckhardt, D. 1987. Jumping plant lice (Homoptera: Psylloidea) of the temperate neotropical region. Part 2: Psyllidae (subfamilies Diaphorininae, Ac- izziinae, Ciriacreminae and Psyllinae). Zoological Journal of the Linnean Society 90: 145-205. . 1994. Psylloid pests of temperate and sub- tropical crop and ornamental plants (Hemiptera, Psylloidea): A review. Entomologia (Trends in Agricultural Science) 2: 173-186. Everett, T. H. 1981. The New York Botanical Garden Illustrated Encyclopedia of Horticulture. Vol. 3 Cha-Di. Garland Publishing, New York. pp. 705— 1058. Hodkinson, I. D. 1980. Present-day distribution pat- terns of the holarctic Psylloidea (Homoptera: In- secta) with particular reference to the origin of the nearctic fauna. Journal of Biogeography 7: 127— 146. . 1988. The Nearctic Psylloidea (Insecta: Ho- moptera): An annotated check list. Journal of Nat- ural History 22: 1179-1243. Hodkinson, I. D. and I. M. White. 1979. Homoptera Psylloidea. Handbooks for the Identification of British Insects. Vol. II, Pt. 5(a). Royal Entomo- logical Society of London, London. 98 pp. Ing, B. 1971. The jumping plant-lice (Psyllidae) of Hertfordshire. Transactions of the Hertfordshire Natural History Society and Field Club 27: 110— 116. Klimaszewski, S. M. 1972. Bemerkungen tiber die Systematik der Gattung Psylla Geoffr., s.1. (Ho- VOLUME 103, NUMBER 1 moptera, Psyllodea). Annales Universitatis Mariae Curie-Sktodowska Sectio C (Biologia) 27: 11-15. . 1973. The jumping plant lice or psyllids (Ho- moptera, Psyllodea) of the Palaearctic: An anno- tated check-list. Annales Zoologici (Warsaw) 30: 155-286. Lal, K. B. 1934a. Insect parasites of Psyllidae. Para- sitology 26: 325-334. . 1934b. Psyllia peregrina Forst., the hawthorn race of the apple sucker, P. mali Schmidb. Annals of Applied Biology 21: 641-648. . 1937. On the immature stages of some Scot- tish and other Psyllidae. Proceedings of the Royal Society of Edinburgh 57: 305-331. Lauterer, P. 1976. Psyllids of wetland nature reserves of the German Democratic Republic, with notes on their biology, taxonomy and zoogeography (Homoptera, Psylloidea). Faunistische Abhan- dlungen Staatliches Museum fiir Tierkunde in Dresden 6(10): 111-122. Loginova, M. M. 1964. Suborder Psyllinea, pp. 437- 482. In Bei-Bienko, G. Ya., ed., Keys to the In- sects of the European USSR. Zoological Institute, Akademia Nauk SSSR, Moscow. [Translated from Russian, 1967, pp. 551—608, by J. Salkind, Israel Program for Scientific Translations, Jerusalem]. 1978. Classification of the genus Psylla Geoffr. (Homoptera, Psyllidae). Entomologiches- koe Obozrenie 57: 808-824 [in Russian: English translation in Entomological Review 57: 555— 566]. Maw, H. E. L., R. G. Foottit, K. G. A. Hamilton, and G. G. E. Scudder. 2000. Checklist of the Hemip- tera of Canada and Alaska. National Research Council of Canada, Ottawa. 220 pp. Missonnier, J. 1956. Note sur la biologie du psylle de l'aubépine (Psylla peregrina E). Annales des Epi- phyties (Paris) 7: 253-262. Novak, H. 1994. The influence of ant attendance on larval parasitism in hawthorn psyllids (Homop- tera: Psyllidae). Oecologia (Berlin) 99: 72-78. Novak, H. and R. Achtziger. 1995. Influence of het- eropteran predators (Het., Anthocoridae, Miridae) 109 on larval populations of hawthorn psyllids (Hom., Psyllidae). Journal of Applied Entomology 119: 479-486. Ossiannilsson, EF 1970. Contributions to the knowledge of Swedish psyllids (Hem. Psylloidea) 1—4. En- tomologica Scandinavica 1: 135-144, - 1992. The Psylloidea (Homoptera) of Fenno- scandia and Denmark. Fauna Entomologica Scan- dinavica Vol. 26. E. J. Brill, Leiden, the Nether- lands. 346 pp. Phipps, J. B. 1998. Introduction to the red-fruited haw- thorns (Crataegus, Rosaceae) of western North America. Canadian Journal of Botany 76: 1863— 1899. Sotherton, N. W., S. D. Wratten, S. B. Price, and R. J. White. 1981. Aspects of hedge management and their effects on hedgerow fauna. Zeitschrift fiir Angewandte Entomologie 92: 425-432. Sutton, R. D. 1983. Seasonal colour changes, sexual maturation and oviposition in Psylla peregrina (Homoptera: Psylloidea). Ecological Entomology 8: 195-201. . 1984. The effect of host plant flowering on the distribution and growth of hawthorn psyllids (Homoptera: Psylloidea). Journal of Animal Ecol- ogy 53: 37-SO. Wheeler, A. G., Jr. and E. R. Hoebeke. 1997. Trioza chenopodii Reuter: First North American record of an Old World psyllid (Homoptera: Psylloidea: Triozidae). Proceedings of the Entomological So- ciety of Washington 99: 409—414. White, I. M. and I. D. Hodkinson. 1982. Psylloidea (nymphal stages) Hemiptera, Homoptera. Hand- books for the Identification of British Insects. Vol. II, 5(b). Royal Entomological Society of London, London. 50 pp. . 1985. Nymphal taxonomy and systematics of the Psylloidea (Homoptera). Bulletin of the British Museum (Natural History) Entomology 50(2): 153-301. Wright, S. E. 1983. Recent amenity tree and shrub planting in the rural landscape of England and Wales. Journal of Environmental Management 16: 17-34. PROC. ENTOMOL. SOC. WASH. 103(1), 2001, pp. 110-134 THE GALL MIDGES (DIPTERA: CECIDOMYIIDAE) OF MIKANIA GLOMERATA (ASTERACEAE) IN SOUTHEASTERN BRAZIL RAYMOND J. GAGNE, RUBENS A.M. ODA, AND RICARDO EF MONTEIRO (RJG) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, c/o National Museum of Natural History, Smithsonian Insti- tution, Washington, DC 20560-0168 (e-mail: rgagne @sel.barc.usda.gov); (RAMO, RFM) Laboratorio de Ecologia de Insetos, Departamento de Ecologia-IB-UFRJ, CP 68020, Ilha do Fundao, 21941-590, Rio de Janeiro, RJ, Brazil (e-mail: monteiro @biologia.ufrj.br) Abstract.—A survey of Mikania glomerata Sprengel (Eupatorieae: Asteraceae) in southeastern Brazil resulted in the discovery of eight kinds of galls made by eight species of gall midges. All of the galls and their gall makers are described. Seven of the gall makers are new to science and two new genera, Liodiplosis Gagné and Mikaniadiplosis Gagné, are described to contain four of the seven new species. The eight gall-making species are: Alycaulus globulus Gagné, Asphondylia glomeratae Gagné, Asphondylia moehni Skuhrava, Liodiplosis cylindrica Gagné, Liodiplosis conica Gagné, Liodiplosis spherica Gagné, Mikaniadiplosis annulipes Gagné, and Perasphondylia mikaniae Gag- né. In addition, another gall midge new to science, Contarinia ubiquita Gagné, is de- scribed that lives gregariously in the galls of five of the eight kinds of galls described. Key Words: This work is part of an ongoing study outlined by Monteiro et al. (1994) on the ecology of insect galls of the Brazilian At- lantic Forest Region, including the restinga or coastal shrub zone. One of the first tasks in studying these galls is the description of their inhabitants, most of them new to sci- ence. In this paper we describe the galls and gall midges associated with the eight galls found on Mikania glomerata Sprengel (Eu- patorieae: Asteraceae). Mikania glomerata is a common plant of the Atlantic forest edge and interior that ex- tends from southeast and southern Brazil into Paraguay and Argentina (King and Robinson 1987). It is one of 415 species of the pantropical but chiefly Neotropical ge- nus Mikania (King and Robinson 1987). This plant is much used in popular medi- Neotropical, gall midges, gall makers, Asteraceae cine, especially as an expectorant (Oliveira et al. 1987). Its numerous and conspicuous flowers bloom from August to December and are much visited by the honeybee, Apis mellifera L. (Cortopassi-Laurino and Ra- malho 1988). Honey from M. glomerata is commercially important. During this study, mainly within Rio de Janeiro State, we found nine species of gall midges associated with eight galls. The galls are described here and illustrated in Figs. 1-8. Eight of the nine species of Ce- cidomyiidae are new to science and are de- scribed and named here, and the ninth is redescribed. The species and their galls are as follows, by tribe, species, and gall type: Alycaulini Alycaulus globulus Gagné in slight epidermal leaf swelling (Fig. 1) VOLUME 103, NUMBER 1 Asphondyliini Asphondylia glomeratae Gagné in leaf petiole or vein blister (Fig. 2) Asphondylia moehni Skuhrava in stem swelling (Fig. 3) Perasphondylia mikaniae Gagné in bud gall (Fig. 8) Cecidomyiini Contarinia ubiquita Gagné, an inqui- line in various galls Clinodiplosini Liodiplosis cylindrica Gagné in cylin- drical leaf or branch gall (Fig. 5) Liodiplosis conica Gagné in conical leaf or branch gall (Fig. 6) Liodiplosis spherica Gagné in spheri- cal leaf or branch gall (Fig. 7) Supertribe Cecidomyiidi, unassigned to tribe Mikaniadiplosis annulipes Gagné in leaf vein, petiole, or branch swelling (Fig. 4) As can be seen from the list, these genera and species are a diverse array of gall midg- es. The six genera belong to five separate tribes of Cecidomyiidae. Even assuming that the two species of Asphondylia and three species of Liodiplosis are more close- ly related to one another than to their con- geners, which cannot be determined here, this means that galls on the host plant are caused by at least five distinct lines of gall- makers and that the inquiline comes from yet another line. The inquiline, Contarinia ubiquita, belongs to a large, cosmopolitan genus of about 300 species whose larvae are otherwise mostly gall makers or free living in flower heads. Two genera are described as new to sci- ence: Liodiplosis with three new species and Mikaniadiplosis with one. Neotropical gall midges are extremely poorly known (Gagné 1994) and many more genera will need to be described as more species are uncovered that do not fit satisfactorily into existing genera. With the eventual discov- ery of more species, the relationships among this fauna should become clearer. A Get case in point is the new species, Pera- sphondylia mikaniae, only the second spe- cies described in Perasphondylia. The two species, both forming bud galls on the tribe Eupatorieae (Asteraceae), now form a sat- isfyingly monophyletic group. All except one species are described or redescribed from their adult, pupal, and lar- val stages. The exception is Contarinia ubi- quita, from which we were not successful in obtaining pupae or adults. This species is nonetheless common as larvae, occurring as an inquiline in the galls of five other spe- cies. Because this paper treats all known species of gall midges from M. glomerata and because two of us, Oda and Monteiro, will be referring to C. ubiquita in future ecological work, it is desirable to describe what we know and name it at this time, despite our knowing only larvae. New noteworthy realizations are reported here concerning the morphology of these gall midges. Species of the subtribe Asphondyliina, which includes Asphondylia and Perasphondylia, have lost the pair of trichoid sensilla commonly seen in other gall midges at the anterior edge of the adult abdominal terga and sterna. Two characters are noticed for Liodiplosis that seem to be shared with other Clinodiplosini. These are the bare, non-setulose venter of the male hypoproct and the smooth horizontal band on the abdominal terga of the pupal abdo- men just posteriad of the row of dorsal pa- pillae. It appears that Clinodiplosini which pupate in their galls have large tergal spines that serve to help in escaping from the galls, spines that are not present in species that drop to the soil to pupate. The larva of Mikaniadiplosis annulipes has a pair of large corniform papillae and 3 pairs of small setiform papillae on the terminal seg- ment that are reminiscent of Contarinia spp., but the otherwise great differences be- tween the two genera indicate that the lar- val characteristics they share are separately derived. i PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-8. Cecidomyiid galls of Mikania glomerata, entire and in section, the sectioned galls enlarged in some. 1, Leaf epidermis swelling of Alycaulus globulus. 2, Vein swelling of Asphondylia glomeratae. 3, Stem swelling of Asphondylia moehni. 4, Leaf vein swelling of Mikaniadiplosis annulipes. 5, Cylindrical leaf gall of Liodiplosis cylindrica. 6, Conical leaf gall of Liodiplosis conica. 7, Spherical leaf gall of Liodiplosis spherica. 8, Bud gall of Perasphondylia mikaniae. VOLUME 103, NUMBER 1 MATERIALS AND METHODS Between February, 1996 and October, 1997, monthly collections of galls on Mi- kania glomerata were made in the Reserva Biologica de Pogo das Antas, Rio de Ja- neiro, and between April, 1998 and March, 1999, trimestral collections were made in the following places: Parati, RJ; Parque Na- cional do Itatiaia, RJ; Parque Nacional da Serra dos Orgaos, RJ; and Parque Nacional da Tijuca, RJ; and Parque Estadual da Serra do Mar, Ubatuba, SP. On each occasion, all galls were removed from 20—30 individual plants. In the laboratory, galls were sepa- rated by kinds, samples of larvae were re- moved, and the remaining galls placed by type in plastic bags or boxes covered with fine screening to obtain adults. Larvae and reared adults with their pupal exuviae were preserved in 70% ethanol. A large number of hymenopterous parasitoids were also reared. These will be the subject of a sep- arate paper in preparation by two of us (Oda and Monteiro). For microscopic study, specimens were mounted in Canada bal- sam, using the method outlined in Gagné (1989, 1994). In addition, some pupal ex- uviae were dried and placed on stubs for scanning electron microscope study. In the description of the new species here, ana- tomical terminology of the adult stage fol- lows McAlpine et al. (1981) and that of the larval stage follows Gagné (1989). The ho- lotypes and additional specimens of the new species will be deposited in the Museu Nacional do Rio de Janeiro; the remaining specimens and voucher specimens of galls are deposited in the U.S. National Museum of Natural History, Washington, D.C. (USNM). The field work for this study was done by Oda and Monteiro, and the tax- onomy and descriptions of the new species were the responsibility of the senior author. The descriptions below appear in alphabet- ical order by genus. Alycaulus globulus Gagné, new species (Figs. 9-17) Adult.—Head: Antenna (Fig. 12) with 13-16 flagellomeres in ¢ (n = 5), 21—23 in i} 2 (n = 5). Mouthparts: labellum semicir- cular in frontal view, with 6—8 lateral setae; palpus usually 3 segmented, first segment as wide as long, second longer and wider than first, third, when present, narrower, shorter to longer than second. Thorax: Wing (Fig. 9) length, ¢, 1.6— 2.1 mm (nm = 5); 2, 2.0—2.4 mm (m = 5); R,; about 0.66 length of wing. Tarsal claws with strong tooth; empodia as long as claws. Vestiture as follows: scutum with sparse dorsocentral rows of setae and nu- merous lateral setae, completely covered elsewhere with scales; anepisternum with scales on dorsal half; katepisternum without scales; anepimeron with 21—25 setae and a few scales. Male abdomen: First through 7th tergites rectangular, about 3 times as wide as long, with single row of setae along posterior margin, a pair of trichoid sensilla on ante- rior margin, and covered elsewhere with scales except 7th tergite with scales only on posterior third; 8th tergite about as wide as long, weakly sclerotized, the anterior pair of trichoid sensilla the only vestiture. Sec- ond through seventh sternites quadrate, with double row of posterior setae, lateral setae, and group of mesal setae, and ante- rior pair of trichoid sensilla, the remaining area covered with scales. Eighth sternite as for 7th except setae and scales only on pos- terior half. Genitalia (Figs. 10—11): hypo- proct entire, slightly wider than a cercus, with pair of posterior setae; aedeagus lon- ger than hypoproct; claspettes apically with bare protuberance tipped with 2-3 short se- tae; gonocoxite cylindrical; gonostylus larg- est at base, tapering abruptly to narrowed apex, setulose basally, carinate beyond. Female abdomen (Figs. 13-14): First through 6th tergites rectangular, about twice as wide as long, with vestiture as in male; 7th tergite only slightly wider than long, with 3 rows of posterior setae, anterior pair of trichoid sensilla, and scales covering only posterior half; eighth tergite unsclero- tized posteromesally, resulting posterolat- eral lobes each with 2-3 short setae, the 114 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 9-17. Alycaulus globulus. 9, wing. 10, Male genitalia (dorsal). 11, Claspettes and aedeagus. 12, Scape, pedicel, and first 3 flagellomeres. 13, Female postabdomen, from sixth segment to cerci (dorsal). 14, Female eighth tergite to fused cerci. 15, Pupal head with additional drawing of antennal horn in lateral view. 16, Larval spatula and associated papillae. 17, Eighth and terminal larval segments. VOLUME 103, NUMBER 1 only other vestiture the anterior pair of trichoid sensilla. Second to 7th sternites rectangular, the vestiture as for male. Eighth sternite undefined. Ovipositor: eighth seg- ment at midlength with group of lateral se- tae on each side and scattered ventral setae, with dorsal, longitudinal rows of spicules on posterior third of dorsum enlarged, com- bined, and modified into retrorse, pointed processes; 9th segment covered on anterior half with many-pointed spicules, on poste- rior half with scattered setae laterally and ventrally; fused cerci spheroid, evenly cov- ered with setae; hypoproct slightly less than half length of cerci, slightly longer than wide, narrowed and curved apically, with 2 apical setae; protrusible portion of ovipos- itor about 2.3 times length 7th tergite. Pupa.—Head (Fig. 15): Antennal horns well developed, the anterior margin finely serrate; cervical sclerite with two elongate setae; face without ventral projections, without apparent papillae. Abdominal ter- gites uniformly spiculose. Last larval instar—Integument exten- sively spiculose dorsally and laterally and on most segments, becoming smoother on last 3 abdominal segments, closely set hor- izontal rows of spicules present anterven- trally on all segments. Spatula (Fig. 16) tri- dentate anteriorly, middle tooth largest. Pa- pillae: laterals in one group of 4 on each side of spatula, 2 with setae, 2 without; ter- minal segment (Fig. 17) with 8 setose pa- pillae, the setae of approximately equal length. Holotype.—Male, from epidermal leaf gall on Mikania glomerata, Pogo das Antas, RJ, Brazil, VII-1997, Oda, deposited in Museu Nacional do Rio de Janeiro. Other material examined (all from leaf blister galls on M. glomerata, Rio de Ja- neiro state, Brazil).—2 2, same data as ho- lotype; 6, pupal exuviae, larva, same data as holotype except VIII-1997; 1 2, Silva Jardim, Pogo das Antas Biological Reserve; 3,9 2,5 pupal exuviae, larva, Parque Na- cional do Itatiaia, II-1999; 2 d, 1 2, 2 pu- pal exuviae, Parque Nacional da Tijuca, XI- 1) 1998; 2 6, 1 °, 4 pupal exuviae, 3 larvae, Parati. Etymology.—The name globulus is an adjective referring to the globular, fused fe- male cerci of this species. Gall (Fig. 1).—This gall is a slight, thin- walled, ovoid distention of the leaf epider- mis. It occurs on either of the leaf surfaces and also on veins and petioles, occasionally directly atop blister galls and vein and pet- iole swellings made by other species. The single larva pupates in the gall and the pupa emerges from an opening near either end of the gall. The gall was found in each of the collecting localities but in small numbers. Remarks.—Two other species of Alycau- lus have been described, both from Mikania in the Neotropics. All three have a similarly attenuate gonostylus, a large, three toothed spatula, a long R;, about % the length of the wing, the longest known for the Alycaulini, and all are from leaf or petiole galls of Mi- kania spp. Alycaulus mikaniae Riibsaamen (1916) was associated with blisters on leaf veins of a Mikania sp. in Amazonas, Brazil. The types are in the Museum fiir Naturkun- de der Humboldt Universitat in Berlin but are not in a condition to travel through the mail. Fortunately, some fine illustrations ac- companied the original description. That of the male genitalia of A. mikaniae appear generally similar to the new species except that the gonostylus of A. mikaniae looks much more narrow throughout than in A. globulus. The larval spatula Riibsaamen drew for A. mikaniae is much broader than that of A. globulus and has five anterior teeth instead of the three found in A. glob- ulus. The fused cercal lobe of the female of the new species is very different from the corresponding structure of A. mikaniae. Riibsaamen drew two separate cerci for his species, possibly the main reason he erected Alycaulus. The bulbous, single cercal lobe of A. globulus, if flattened too much on a slide, folds such that it appears to be two separate cerci. Riibsaamen was an excellent draftsman; nevertheless, Riibsaamen’s spec- imen needs to be seen to determine whether 116 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON he drew a crushed fused cercal lobe that only appeared as two cercl. The second of the two previously de- scribed Alycaulus species, Alycaulus trilo- batus Mohn (1964), was described on the basis of larvae taken from petioles of Mi- kania micrantha H.B.K. from El Salvador. Mohn’s drawing shows a shorter spatula than that of A. globulus but both species have three similar anterior teeth. The ratio of the width of the 3 teeth and length of entire spatula is .27 in A. trilobatus but .21 in A. globulus. Wiinsch (1979) recorded A. trilobatus from stem and petiole galls of Mikania cordifolia and Mikania sp. from Colombia. He redescribed a larva similar to that in Mo6hn’s drawing and described also the pupa and female for the first time. Wiinsch drew the cerci as fused, as it is in A. globulus. The pupal antennal horns of A. trilobatus are broader and more serrate than in A. globulus. Asphondylia glomeratae Gagné, new species (Figs. 18-22, 28-29) Adult.—Head: Antenna (Fig. 18) with scape slightly more than twice length ped- icel; pedicel about as wide as long; first fla- gellomere less than twice length of scape; proportions of female segments and flagel- lomeres as in Fig. 18. Eye facets hexagonal, closely approximated. Frons with 18—20 se- tae. Clypeus asetose dorsally, with several short setae laterally, and setulose ventrally. Hypopharynx closely lined with setulae dorsoapically. Labella setose and setulose laterally, with small area of setulae and O— 1 setae mesally. Palpus 3 segmented, first segment about as long as wide, second seg- ment about twice as long as wide, third nar- rower than second, about 3 times as long as wide, all covered with setulae and setae. Thorax: Wing length, male, 2.0—2.3 (n = 4), female, 2.8—2.9 (n = 2). Dorsocentral setal row at midlength with two rows of setae intermixed with scales, the row con- tinuing onto scutellum to posterior margin. Scutellum bare between dorsocentral rows of setae, with setae posteriorly laterad of dorsocentral rows, and otherwise with sparse scales laterally. Anepisternum with setae and scales on dorsal half. Anepimeron covered with setae. Katepimeron bare. Tar- sal claws (Fig. 20) all equal in size and shape, as long as empodia. Abdomen: Tergites without anterior pair of trichoid sensilla, first through seventh tergites rectangular with mostly single row of posterior setae (sparser in male than in female), mostly double row on seventh ter- gite, several lateral setae, elsewhere cov- ered with scales, width of sixth and seventh tergites about 2 % times length; eighth ter- gite short, bandlike, without vestiture. Ster- nites without anterior pair of trichoid sen- silla, width sixth sternite about twice length, male eighth sternite reduced in size, as wide as long. Male genitalia as in Fig. 19. Distal half of ovipositor approximately .55 length seventh sternite. Pupa (Figs. 28—29).—Antennal horns conical, elongate, evenly tapered from base to pointed apex. Upper frontal horn elon- gate-conical with single apex. Lower fron- tal horn 3-toothed, the middle tooth about twice length of lateral teeth. Abdominal ter- gites 2—8 each with, from posterior margin, a regular horizontal row of stout spines, a short bare space, two irregular, sparser rows of stout spines followed by scattered, much smaller spines towards anterior margin of sclerite. Third larval instar.—Spatula (Fig. 21) ro- bust, elongate, 4-toothed anteriorly, the 4 teeth of similar length. Sternal, the 4 pairs of lateral, and the pair of ventral papillae are with setae. One pair of terminal papillae only barely distinguishable from spicules (Fig. 22). Holotype.—Male, from vein blister gall on Mikania glomerata, Rio de Janeiro, RJ, 6-VII-1996, Oda & Monteiro, deposited in Museu Nacional do Rio de Janeiro. Other material examined (all from vein or petiole blister galls on M. glomerata, Rio de Janeiro state, Brazil).—1 ¢, same data as holotype; 1 ¢, 2 2, 3 pupal exuviae, VOLUME 103, NUMBER 1 Wi7/ Figs. 18-25. Foretarsal claw and empodium. 21, Larval spatula and associated papillae. 22, Eighth and terminal larval seg- ments. 23—25, Asphondylia moehni. 23, Larval spatula. 24, Foretarsal claw and empodium. 25, Midtarsal claw and empodium. Silva Jardim, Pogo das Antas Biological Reserve, [X-1997; larva, Silva Jardim, Poco das Antas Biological Reserve, [X- 1999; 3 larvae, Teresdpolis, VIII-1999; 2 d, pupa, Parati. Etymology.—tThe specific name glomer- 25 18-22, Asphondylia glomeratae. 18, Silhouette of female antenna. 19, Male genitalia. 20, atae is the genitive of the host plant specific name. Gall (Fig. 2).—This gall is a single- celled, thin-walled, green, slight swelling of the leaf veins on either leaf surface and on petioles. It can be readily distinguished 118 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON from two externally similar galls, the swol- len vein and simple epidermal galls, by the presence of fungal mycelium lining the in- side of the larval chamber of A. glomeratae. The single larva pupates in the gall and the pupa emerges from an opening near either end of the gall. The gall was uncommon but found in each of the collecting localities. Remarks.—Asphondylia glomeratae is distinct from A. moehni in having identi- cally shaped tarsal claws, three distinct teeth on the pupal lower frontal horn, and a larval spatula with four uniformly long anterior teeth. Asphondylia ulei Riibsaamen (1907) also forms a leaf gall on Mikania. It was de- scribed from a larva and pupa taken from 5 mm long, hairy, spherical galls on the un- derside of leaves of an undetermined spe- cies of Mikania in Palmeiras, Rio de Janei- ro, Brazil. Galls of A. glomeratae are smooth, not hairy, and a redescription of the type series of A. ulei in Mohn (1973) shows a pupa with only one lower frontal horn, not three as in A. glomeratae, and a larval spatula with uneven anterior teeth, not of similar size and shape as in A. glom- eratae. Asphondylia moehni Skuhrava (Figs. 23-27) tavaresi Mohn 1973: 4, Asphondylia. Sec- ondary junior homonym of Asphondylia tavaresi Riibsaamen 1916. moehni Skuhrava 1989: 203, Asphondylia. New name for Asphondylia tavaresi Mohn. Adult (female only).—Head: Antenna (as for A. glomeratae, Fig. 18) with scape slightly more than twice length pedicel; pedicel about as wide as long; first flagel- lomere almost twice length of scape. Eye facets hexagonal, closely approximated. Frons with approximately 36—40 setae. Clypeus asetose dorsally, with several short setae laterally, and setulose ventrally. Hy- popharynx closely lined with setulae dor- sally. Labella setose and setulose laterally and ventrally. Palpus 3 segmented, first seg- ment about as long as wide, second seg- ment about twice as long as wide, third nar- rower than second, about 3 times as long as wide, all covered with setulae and setae. Thorax: Wing length, 3.4—3.6 (n = 5). Dorsocentral setal row at midlength with three rows of setae intermixed with several scales, the row continuing onto scutellum to posterior margin. Scutellum bare be- tween dorsocentral rows of setae, with setae posteriorly laterad of dorsocentral rows, and otherwise with sparse scales laterally. Anepisternum with setae and scales on dor- sal third. Anepimeron covered with setae. Katepimeron bare. Tarsal claws (Figs. 24— 25) larger on mid- and hindleg than on fore- leg, as long as empodia. Abdomen: Tergites without anterior pair of trichoid sensilla, first through seventh tergites rectangular with mostly single row of posterior setae, mostly double row on seventh tergite, many lateral setae, else- where covered with scales, width of sixth and seventh tergites about 2 % times length; eighth tergite short, bandlike, without ves- titure. Sternites without anterior pair of trichoid sensilla, width sixth sternite about twice length. Distal half of ovipositor ap- proximately .55 length seventh sternite. Pupa (Figs. 26—27).—Antennal horns conical, elongate, evenly tapered from base to pointed apex. Upper frontal horn elon- gate-conical with single apex. Lower fron- tal horn 3-toothed, the middle tooth much larger than lateral teeth, which are litle more than slightly convex carinae. Abdom- inal tergites 2—8 each with, from posterior margin, a regular horizontal row of stout spines, a short bare space, two irregular, sparser rows of stout spines followed by scattered, much smaller spines towards an- terior margin of sclerite. Third larval instar (only spatula avail- able, from cast exuviae).—Spatula (Fig. 23) robust, elongate, 4-toothed anterioly, the two middle teeth appreciably shorter than the lateral teeth. Holotype.—Male, from stem gall on Mi- VOLUME 103, NUMBER 1 Figs. 26-31. Pupal exuviae, anterior segments. 26, Asphondylia moehni, ventral view. 27, Same, lateral. 28, Asphondylia glomeratae, ventral. 29, Same, lateral. 30, Perasphondylia mikaniae, ventral. 31, Same, lateral. Line = 100 pm. 120 kania guaco Humb. & Bonpl., Sao Leopol- do, Rio Grande do Sul, Brazil, in the Ta- vares Collection, at present in the Staatlich- es Museum fiir Naturkunde, Stuttgart, Ger- many. Other material examined (all from stem galls on M. glomerata, Rio de Janeiro state, Brazil).—Parati, II-1999, 2 larvae; Parati, VII-1999, 3 larvae; Parque Nacional da Serra dos Orgaos, II-1999, 5 2, 3 pupa, 6 pupal exuviae; Picinguaba, 4 °, 3 pupal ex- uviae. Gall (Fig. 3).—This species causes a ridged, spongy-textured, stem swelling found usually to one side of but sometimes surrounding the leaf petiole or stem. It com- monly is found with a single larval cell, but may occur in aggregate and have several cells. At first the galls are green but even- tually turn a yellow-brown. The interior of the larval cells is lined with a fungal my- celium. The single larva in each chamber pupates in the gall and the pupa emerges from an opening that may occur anywhere on the surface. This gall was found in most of the collection localities but was never common. Remarks.—Tavares (1909) published a photo showing a similar gall to that on M. glomerata but on Mikania guaco Humb. & Bonpl. from Sao Leopoldo, Rio Grande do Sul, Brazil. Specimens that Tavares reared from that gall were later described and named Asphondylia tavaresi by Mohn (1973). The specimens consisted of a male, a partial pupa with head and thorax lacking, and a larva. We were not able to rear a male to compare with Mo6hn’s illustrations and the pupal abdomen is not specific enough for comparison with the species reared from M. glomerata, but the larval spatula as drawn by Mohn fits that of our specimens. For the present we consider our specimens to be the same as Mohn’s species, now known as A. moehni Skuhrava. Asphondylia moehni is distinct from A. glomeratae in having smaller foretarsal than mid- and hindtarsal claws, one distinct PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tooth on the pupal lower frontal horn, and a larval spatula with uneven anterior teeth. Contarinia ubiquita Gagné, new species (Figs. 32-34) Larva (Figs. 32—34), third instar (only stage known).—White. Integument smooth except for anteroventral, horizontal rows of tiny spicules on venter of second and third thoracic and all abdominal segments. Spat- ula (Fig. 33) bidentate anteriorly. Papillar setae short, barely longer than width of pap- illar bases. Terminal pair of corniform pa- pillae (Fig. 34) dark and pigmented exten- sively beyond bases. Holotype.—Larva, from bud gall of Per- asphondylia mikaniae on Mikania glomer- ata, Silva Jardim, Pogo das Antas Biologi- cal Reserve, RJ, XI-1997, Oda, deposited in Museu Nacional do Rio de Janeiro. Other material examined.—All from M. glomerata, Rio de Janeiro state, Brazil: 2 larvae, same data as holotype; 20 larvae from stem gall of Asphondylia glomeratae, Parque Nacional da Serra dos Orgaos, II- 1999; 2 larvae from unspecified Liodiplosis leaf gall, Silva Jardim, Pogo das Antas B1- ological Reserve, VII-1997; 9 larvae from petiole gall of Mikaniadiplosis annulipes, Parati, VII-1999; 6 larvae from epidermal gall of Alycaulus globulus, Parati, VII- 1909: Etymology.—The name ubiquita is an adjective meaning ubiquitous that refers to the fact that this species occurs as an in- quiline in five of the eight known galls of Mikania glomerata. Remarks.—This species is known only from larvae. These drop to the ground when full grown, unlike the Mikania gall makers, which all pupate in their galls. Contarinia ubiquita larvae were discovered in the course of this work from bud galls of Per- asphondylia mikaniae, epidermal galls of Alycaulus globulus, a stem gall of Asphon- dylia glomeratae, petiole galls of Mikani- adiplosis annulipes, and an_ unspecified Liodiplosis leaf gall. The larvae are gregar- ious and, in the case of the petiole gall, fill VOLUME 103, NUMBER | 121 Figs. 32-40. 32-34, Contarinia ubiquita larva. 32, Whole larva, dorsal view. 33, Spatula. 34, Eighth and terminal segments, dorsal. 35—40, Liodiplosis cylindrica. 35, Wing. 36, Tarsal claw and empodium. 37, Male third flagellomere. 38, Female third flagellomere. 39, Female postabdomen, seventh segment to cerci (dorsolat- eral). 40, Female cerci and hypoproct, detail (lateral). 122 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON the gall cavity. Despite having only larvae of this species, it is named here because it appears in many galls, and it is handy to have a name for further work that will be done with Mikania galls. It differs from other known Contarinia species in the pig- mented area that can be seen surrounding the base of the pair of corniform papillae of the terminal segment (Fig. 34). Larvae of all other known Contarinia species have only the papillae themselves pigmented and not the surrounding area. Liodiplosis Gagné, new genus Adult.—Head: Eyes connate, 11—12 fac- ets long at vertex; facets mostly hexago- noid, all closely adjacent. Occiput with dor- sal protuberance with 2 apical setae. Frons with setae. Labella ellipsoid and pointed apically, each with several lateral setae. Pal- pus 4-segmented. Male antennal flagellom- eres (Fig. 37) binodal; one circumfilum on basal node, two on distal node, loops of the three circumfila subequal in length. Female flagellomeres (Fig. 38) cylindrical, ringed by two appressed circumfila connected by two longitudinal bands and with long necks. Thorax: Wing (Fig. 35) unmarked, R, curved apically, joining C posterior to wing apex; Rs incomplete, closer to arculus than to apex of R,. Tarsal claws (Fig. 36) un- toothed, curved near middle; empodia very short, not attaining bend in claws; pulvilli as long as empodia. Male abdomen: First through sixth ter- gites entire, rectangular, with single poste- rior row of setae, several lateral setae, scat- tered scales, and 2 anterior trichoid sensilla; seventh tergite weakly sclerotized posteri- orly, with few to several posterolateral setae on each side, several anteromesal scales, and anterior pair of trichoid sensilla; eighth tergite weakly sclerotized anteriorly, the only vestiture the anterior pair of trichoid sensilla. First through eighth sternites rect- angular, covered with setae and with 2 an- terior trichoid sensilla; eighth sternite sim- ilar to preceding except longer. Genitalia (Figs. 41—44): cerci rectangular with a few posterior and several lateral setae, lateral margin strongly curved ventrad; hypoproct appreciably longer than cerci, widest basal- ly, narrowing near midlength and deeply di- vided apically, resulting lobes tapering to narrow apices and with a few setae apically, dorsal surface setulose, ventral surface bare and horizontally banded; aedeagus shorter than hypoproct, tapering gradually from base to narrowly rounded apex, with lon- gitudinal rows of sensory pits; gonocoxite cylindrical with slight, obtuse, mesobasal lobe; gonostylus broadest near midlength, tapering beyond, with setulae on basal half, covered beyond with minute carinae and scattered short setae. Female abdomen: First through seventh tergites entire, rectangular, with mostly sin- gle row of posterior setae, several lateral setae, extensively covered with scales, and with 2 anterior trichoid sensilla. Eighth ter- gite unsclerotized, with several posterior se- tae and anterior pair of trichoid sensilla. Second through seventh sternites quadrate, extensively covered with setae and scales and with anterior pair of trichoid sensilla. Ovipositor (Figs. 39—40) only slightly pro- trusible, venter of eighth segment with setae on posterior surface, dorsum of ninth and tenth segments without vestiture, venter of ninth and tenth segments with setae, cerci ovoid, with pair of apical sensory setae and scattered setae elsewhere, hypoproct short, broad, with 2 posterior setae. Pupa (Figs. 49—58).—Head: Antennal horns widely separated, each tapering to single apical point, the apex projecting ven- troapically; cervical sclerite with papillae situated on two prominences, each with short seta; face without ventral projections, with usually 3, occasionally fewer papillae, on each side of base of labrum, and with group of three papillae near base of palpus. Prothoracic spiracle elongate. Abdominal second through eighth tergites (Fig. 58) an- teriorly with mostly double row of pig- mented spines, tergal surface covered else- where with uniformly small spicules, ex- VOLUME 103, NUMBER 1 123 F w < ~o c) ° ° 46 47 48 Figs. 41-48. Liodiplosis spp. 41, Liodiplosis cylindrica, male genitalia (dorsal view). 42, Same, aedeagus, hypoproct and a cercus (ventral). 43, Liodiplosis conica, male genitalia (dorsal). 44, Liodiplosis spherica, male genitalia (dorsal). 45, Liodiplosis cylindrica, eighth and terminal larval segments (dorsal). 46, Same, spatula and associated papillae. 47, Liodiplosis conica, spatula and associated papillae. 48, Liodiplosis spherica, spatula and associated papillae. cept for bare horizontal band posteriad of spread apart as larva grows. Antenna about dorsal papillae. twice as long as wide. Spatula (Figs. 46— Third larval instar—White. Integument 48) with 2 widely separated anterior teeth, mostly covered with large spicules that the space between occasionally weakly 124 toothed; shaft reduced posteriorly. Lateral thoracic papillae in 2 groups of usually 3 on each side of central line, 2 papillae in each group with tiny seta. Dorsal and pleu- ral papillae with elongate setae. Terminal segment (Fig. 45) rounded, with only 6 pa- pillae, each with elongate seta, those of one pair slightly longer than remainder. Type species.—Liodiplosis cylindrica Gagné. | Etymology.—The name Liodiplosis is a combination of the Greek word “‘leio” (shortened here to lio) meaning smooth, and “‘diplosis,”’ a suffix commonly used for genera of the supertribe Cecidomyliidi. The prefix refers to the smooth venter of the male hypoproct, although that character is not exclusive to this genus. The suffix means “double,” in reference to the two- noded male flagellomeres. The generic name is feminine. Remarks.—The new genus belongs to the tribe Clinodiplosini and will key to Cli- nodiplosis in couplet 56 of Gagné (1994). The unique characters of the tribe Clinodi- plosini are the obtuse mesobasal lobe of the gonocoxite, the usually quadrate, some- times secondarily bilobed, male cerci, and a particular conformation of the larval ter- minal papillae in which two pairs of the four pairs are corniform, the other two pairs setiform, the more lateral of the two pairs of setae longer than the more mesal pair (cf. Gagné 1989, 1994). The male genitalic characters of Liodiplosis conform to the tribal expectation but the larval characters do not. In the new genus, the larval termi- nal papillae number only six, all of them setiform, the most lateral pair of setae slightly longer than the other two pairs that are subequal in length. A seventh, small, setiform papilla occurs on one specimen of one species, Liodiplosis spherica. The usual presence of only six papillae indicates a loss, of course, but the fact that none is cor- niform indicates either that a pair of cor- niform papillae reverted to setiform or that this new genus shows a character state that PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON is primitive to that found in all other Cli- nodiplosini. Unusual also for clinodiplosines is the modified larval spatula, evidently an adap- tation for life in the particular galls. Cli- nodiplosini usually have a clove shaped spatula, two rounded apical lobes at the end of a long shaft, its length more than 3.5 times as long as widest part just behind the anterior lobes. In Liodiplosis the spatula is more robust, wider than the larval head but only about twice as long as wide. The large spines on the second to eighth abdominal terga of the pupa of Liodiplosis appear only in some, not all, Neotropical Clinodiplosini that pupate in their galls. Other Clinodiplosini with such spines are Clinodiplosis eupatorii Felt, Houardodi- plosis rochae Tavares, Iatrophobia brazil- iensis Ritibsaamen, and Schismatodiplosis lantanae Riibsaamen. The spines are an ad- aptation that allows the pupa to gain pur- chase on the interior surface of the gall while breaking out of the gall. A pupal abdominal character (Fig. 58) brought to light here may be peculiar to the tribe Clinodiplosini because it occurs also in other genera of that tribe for which RJG has seen pupae. It is a horizontal band pos- teriad of the dorsal papillae that, unlike the remainder of the spinulose and spiny terga, is smooth, although somewhat crumpled and obscured in some dry specimens. This character does not occur in other tribes, e.g., Lopesiini, and may possibly be unique to Clinodiplosini. Three new species are described below from three distinct leaf galls on Mikania glomerata (Figs. 5—7). The species are quite similar. They differ in the shape of the male genitalia and characteristics of the pupal head. Liodiplosis cylindrica Gagné, new species (Figs. 35—42, 45—46, 49-50, 55) Adult.—Head: Male third antennal fla- gellomere as in Fig. 37. Female third fla- gellomere as in Fig. 38. VOLUME 103, NUMBER 1 Figs. 49-54. Pupal exuviae, anterior segments. 49, Liodiplosis cylindrica, ventral view. 50, Same, later 51, Liodiplosis conica, ventral. 52, Same, lateral. 53, Liodiplosis spherica, ventral. 54, Same, lateral. Line 100 pm. 126 Thorax: Wing length, male 2.2 mm (n = 3), female 2.5—2.9 mm (n = 5). Tarsal claws as in Fig. 36. Male abdomen: Genitalia (Figs. 41—42): gonocoxite narrow, elongate-cylindrical. Female postabdomen: As in Figs. 39-— 40. Pupa.—Head (Figs. 49-50, 55): Anten- nal horns pointed, the apex projecting ven- troanteriorly. Cervical sclerite smooth, without spicules, cervical papillae on two lateral bumps each with one or two short setae, with no other bumps present. Notum with anterior surface smooth. Third larval instar—Spatula and associ- ated papillae as in Fig. 46. Eighth and ter- minal abdominal segment as in Fig. 45. Holotype.—Male, from cylindrical galls on M. glomerata, Silva Jardim, Pogo das Antas Biological Reserve, RJ, Brazil, from cylindrical gall on M. glomerata, Oda & Monteiro, deposited in Museu Nacional do Rio de Janeiro. Other material examined (all from cylin- drical galls on M. glomerata, Rio de Janeiro state, Brazil)—2 3, 2 2, 6 pupal exuviae, 3 larvae, same data as holotype; 3 ¢, 3 pu- pal exuviae, 2 larvae, Rio de Janeiro, 6-XII- 1996; pupa, Rio de Janeiro, 7-I-1996; 1 3, 3 2, 3 pupal exuviae, Parati. Etymology.—tThe specific name cylindri- ca is an adjective that refers to the shape of the gall made by this species. Gall (Fig. 5).—This is a single-celled, thin-walled but rigid, elongate, unevenly cylindrical gall, generally widest at about midlength and tapered apically. It occurs on either surface of the leaf blade and less commonly on the petioles and young branches. The gall is light green when young, later turning red basally and dark green apically. The single larva pupates in the gall and the pupa emerges from an oper- culum near the gall apex. The gall was fair- ly abundant and the most commonly en- countered in each of the regular collection localities. Remarks.—The female and larva of this species are similar to those of the other two PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON species. The male gonocoxite (Fig. 41) is distinct in being much narrower than in L. conica and L. spherica. The pupa of L. cy- lindrica is generally similar to that of L. conica, but the latter has a slight mesal prominence on the cervical sclerite. Neither of these two species has spiculae on the cer- vical sclerite and notum, nor two large pos- terior bumps on the cervical sclerite as does L. spherica. Liodiplosis conica Gagné, new species (Figs. 43, 47, 51-52, 56) Adult.—Head: Male and female third antennal flagellomeres as for L. cylindrica (Figs. 37-38). Thorax: Wing length, male 2.4—2.6 mm (n = 2), female 2.9 mm (n = 2). Male abdomen: Genitalia (Fig. 43): gon- ocoxite elongate-cylindrical. Female abdomen: As for L. cylindrica (Figs. 39—40). Pupa.—Head (Figs. 51-52, 56): Anten- nal horns pointed, the apex projecting ven- troanteriorly. Cervical sclerite smooth, without spicules, cervical papillae on two lateral bumps each with one or two short setae, and with a slight mesal prominence. Notum with anterior surface smooth. Third larval instar—Spatula as in Fig. 47. Holotype.—Male, from conical gall on Mikania glomerata, Parque Nacional da Ti- juca, RJ, Brazil, Il-1999, deposited in Mu- seu Nacional do Rio de Janeiro. Other material examined (all from coni- cal galls on M. glomerata, Rio de Janeiro state, Brazil)—4 @, 5 pupal exuviae, same data as holotype; 2 larvae, XI-1998, other- wise same data as holotype; 2 6, 2 @, 2 pupal exuviae, 2 larvae, Silva Jardim, Pogo das Antas Biological Reserve, Oda & Mon- teiro. Etymology.—The specific name conica is an adjective that refers to the shape of the gall made by this species. Gall (Fig. 6).—This is a single-celled, thin-walled, conical, light green, rigid gall found on either surface of the leaf blade and VOLUME 103, NUMBER 1 Figs. 55-58. sclerite). 56, Liodiplosis conica, anterior segments. 57, Liodiplosis spherica, anterior segments. 58, Same, third through fifth tergites (“‘a’’ denotes row of spines and ‘‘b” the smooth area, now shrivelled, just ventrad of the dorsal row of papillae. Line = 100 pm. also on the petioles and young branches. The single larva pupates in the gall and the pupa emerges from an operculum near the gall apex. The gall was uncommon, found in only three of the six regular collection sites, and always in heavy shade. Remarks.—See under L. cylindrica for specific differences among the three species of Liodiplosis. Liodiplosis spherica Gagné, new species (Figs. 44, 48, 53-54, 57-58) Adult.—Head: Male and female third antennal flagellomeres as for L. cylindrica (Figs. 37-38). Pupal exuviae. 55, Liodiplosis cylindrica, anterior segments (dorsal; arrow denotes cervical Thorax: Wing length, male 2.8 mm (n = 2), female 3.0—3.7 mm (n = 2). Male abdomen: Genitalia (Fig. 44): gon- ocoxite elongate-cylindrical. Female abdomen: As for L. cylindrica (Figs. 39-40). Pupa.—Head (Figs. 53-54, 57): Anten- nal horns pointed, the apex projecting ven- troanteriorly. Cervical sclerite spiculose, cervical papillae on two lateral bumps each with one or two short setae, and with two large posterior prominences. Notum ante- rior surface spiculose. Third larval instar.—Spatula as in Fig. 47. 128 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Holotype.—Male, from spherical gall on Mikania glomerata, Parati, RJ, Brazil, de- posited in Museu Nacional do Rio de Ja- neiro. Other material examined (all from spher- ical galls on M. glomerata, Rio de Janeiro state, Brazil).—2 2, 2 pupal exuviae, same data as holotype; 1 6,3 2, 2 pupal exuviae, 2 larvae, Silva Jardim, Pogo das Antas Bi- ological Reserve; 2 2, same data as pre- ceding except VII-1997; 3 2, 3 pupal ex- uviae, 3 larvae, Rio de Janeiro, 6-XII-1996. Etymology.—The specific name spherica is an adjective that refers to the shape of the gall made by this species. Gall (Fig. 7).—The gall of L. spherica is spherical, single-celled, and has thick, spongy walls. It protrudes evenly from both sides of the leaves, petioles, and young branches. It is sometimes found in large ag- gregations. Its green color is similar to that of the leaves. The single larva pupates in the gall and the pupa emerges from an oper- culum near either of its two apices. The gall was fairly abundant and found in all six of the regular collection sites. Remarks.—See under L. cylindrica for specific differences among the three species of Liodiplosis. Mikaniadiplosis Gagné, new genus Adult.—Head: Eyes connate, 10—11 fac- ets long at vertex; facets mostly hexago- noid, all closely adjacent. Occiput without dorsal protuberance but with several large setae at extreme vertex. Frons with 10—14 setae. Labella ellipsoid and pointed apical- ly, each with several lateral setae. Palpus 3- segmented. Male antennal flagellomeres (Fig. 61) binodal; one circumfilum present on basal node, two on the distal node, the loops of the three circumfila subequal in length, short, reaching only half way to next circumfilum. Female flagellomeres (Fig. 60) cylindrical, surrounded by two ap- pressed circumfila connected by two lon- gitudinal bands and with necks about as long as wide. Thorax: Wing (Fig. 59) with light and dark pattern; R; curved apically, joining C posterior to wing apex; Rs incomplete, much closer to arculus than to apex of R,. Tarsal claws (Fig. 62) each with 1 large and 2 smaller teeth, curved near basal third; em- podia not attaining bend in claws; pulvilli slightly shorter than empodia. Male abdomen: First through sixth ter- gites entire, rectangular, with mostly single posterior row of setae, several lateral setae, extensive covering of scales, and 2 anterior trichoid sensilla; seventh tergite weakly sclerotized mesoposteriorly, with several setae on posterolateral margin, several lat- eral setae, scattered scales, and anterior pair of trichoid sensilla; eighth tergite weakly sclerotized, the only vestiture the anterior pair of trichoid sensilla. First through eighth sternites rectangular, covered with setae and with 2 anterior trichoid sensilla. Genitalia (Figs. 63—64): cerci tapering gradually from base to narrowly rounded apex, with nu- merous marginal setae especially laterad; hypoproct about as long as cerci, widening gradually from base to apex, the posterior margin slightly concave mesally with many ventral preapical setae; aedeagus about as long as hypoproct, tapering gradually from base to narrowly rounded apex, with lon- gitudinal rows of sensory pits; gonocoxite cylindrical with slight, obtuse, mesobasal lobe; gonostylus in dorsoventral view broadest near midlength, tapering beyond, with setulae on basal fourth, covered be- yond with minute carinae and scattered short setae. Female abdomen: First through seventh tergites entire, rectangular, with mostly sin- gle row of posterior setae, several lateral setae, extensively covered with scales, and with 2 anterior trichoid sensilla. Eighth ter- gite smaller than seventh, less strongly sclerotized, with several widely spaced pos- terior setae and anterior pair of trichoid sen- silla the only vestiture. Second through sev- enth sternites quadrate, extensively covered with setae and scales and with anterior pair of trichoid sensilla, eighth sternite weakly discernable, somewhat longer than corre- VOLUME 103, NUMBER 1 sponding tergite, covered with setae on pos- terior half and with pair of anterior trichoid sensilla. Ovipositor (Figs. 65—66): only slightly protrusible, eighth tergum about 2x as long as its tergite, bare except on tergite; ninth segment bare dorsally, setose laterally and ventrally on posterior half, cerci ovoid, with pair of large, apical sensory setae and scattered setae elsewhere, hypoproct short, broad, with several posterior setae. Pupa.—Head (Fig. 69): Antennal horns elongate, widely separated, each tapering to two unequal points; cervical sclerite with papillae situated on two prominences, each with seta as long as antennal horn; face without ventral projections, with 2 papillae, each with short seta, at base of labrum, and with group of 3 papillae, each with short seta, near base of palpus. Prothoracic spi- racle elongate. Abdominal second through eighth tergites anteriorly with several irreg- ular horizontal rows of pigmented spines, tergal surface covered elsewhere with uni- formly small spicules. Third larval instar—White. Integument roughened. Antenna about twice as long as wide. Spatula (Fig. 67) broadest anteriorly, anterior margin with 2 widely separated teeth near middle, otherwise weakly serrate. Lateral thoracic papillae in 2 groups of 3 on each side of central line, 2 papillae in each group with tiny seta. Dorsal and pleu- ral papillae with short setae, their length barely surpassing width of spiracles. Ter- minal segment (Fig. 68) with pair of papil- lae greatly enlarged into pair of caudal hooks, the 6 remaining papillae each with short seta. Type species.—Mikaniadiplosis annuli- pes Gagné. Etymology.—The name Mikaniadiplosis is a combination of the host plant generic name and “‘diplosis,’”» commonly used as a suffix when naming genera of the supertribe Cecidomyiidi. The suffix refers to the two- noded male flagellomeres. The name is feminine. Remarks.—The new genus is briefly characterized by the following adult char- 129 acters: a dorsal occipital protuberance is lacking, the male flagellomeres have gen- erally short internodes, necks, and circum- fila, the palpi are reduced to three segments, wing vein Rs is much closer to the arculus than to the end of R,, the tarsal claws have multiple teeth, the gonocoxites are splayed apart, and the male hypoproct is unique for being much wider at its apex than base. The female abdomen is unmodified, the seventh segment not reduced and the ovipositor barely protrusible. The larva, with its pair of large, corniform terminal papillae and three pairs of setose papillae, suggests Con- tarinia, except that the corniform papillae are never so large in Contarinia. In the key to genera in Gagné (1994), Mikaniadiplosis will not run past couplet 29, unless one ignores the three-toothed claws, in which case the genus will run to couplet 40. To make a place for the new genus in the key, one can add a new couplet there by changing couplet ‘‘40” to *‘40a,” deleting the last line of the second half of the couplet, adding “‘40b” at the end, and inserting the following new couplet after 40a: 40b. Occipital protuberance present; palpus 4- segmented; Rs equidistant between arculus and end of R,; gonocoxite with large me- sobasal lobe; male hypoproct no broader thankcelcusi- ere Ieee eee Dactylodiplosis — Occipital protuberance absent; palpus 3-seg- mented; Rs much closer to arculus than to end of R,; gonocoxite with only slight me- sobasal lobe; male hypoproct as wide as bOthycerci fr4 a ee en oe Mikaniadiplosis In addition, the second half of couplet 37 would need “‘palpus four-segmented”’ changed to “‘palpus three- or four-segment- ed”’ to accommodate Mikaniadiplosis. Mikaniadiplosis annulipes Gagné, new species (Figs. 59-69) Adult.—Head: Male antennal flagellom- eres (Fig. 61) dark on basal node and distal half of neck, light elsewhere. Female fla- 130 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 59-69. Mikaniadiplosis annulipes. 59, Wing. 60, Female third flagellomere. 61, Male third flagello- mere. 62, Foretarsal claw and empodium. 63, Male genitalia (dorsal view). 64, Aedeagus and hypoproct (ventral). 65, Female postabdomen, seventh tergite to cerci (dorsolateral). 66, Detail of female cerci and hypoproct (lateral). 67, Larval spatula and associated papillae. 68, Larval eighth and terminal segments (dorsal). 69, Head of pupal exuviae (ventral). VOLUME 103, NUMBER 1 74 Figs. 70-75. 131 Perasphondylia mikaniae. 70, Silhouette of female antenna. 71, Female third flagellomere. 72, Foretarsal claw and empodium. 73, Male genitalia (dorsal). 74, Larval spatula and associated papillae. 75, Eighth and terminal larval segments (dorsal). gellomeres (Fig. 60) dark on basal third and distal half of neck, light elsewhere. Thorax: Wing (Fig. 59) with alternating light and dark areas; length, 2.5—2.7 mm (n = 2) in male, 2.9-—3-7 (n = 4) in female. Legs with alternating light and dark color pattern. Tarsal claws as in Fig. 62. Male abdomen: Genitalia as in Figs. 63— 64. Female postabdomen: As in Figs. 65— 66. Pupa.—Head as in Fig. 69. Third larval instar.—Spatula as in Fig. 67. Posterior segments as in Fig. 68. Holotype.—Male, from petiole swelling on Mikania glomerata, Rio de Janeiro, RJ, Brazil, XI-1998, deposited in Museu Na- cional do Rio de Janeiro. Other material examined (all from leaf vein or petiole swellings on M. glomerata, Rio de Janeiro state, Brazil).—1 6,2 2,4 pupal exuviae, same data as holotype; 1 &, 1 pupal exuviae, Silva Jardim, Pogo das Antas Biological Reserve; 1 6, 2 pupal ex- 132 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON uviae, Parati; larva, Parati, XI-1998; 1 °, 1 pupal exuviae, Parati, 6-VII-1996; 2 larvae, Rio de Janeiro, 6-VII-1996; 3 larvae, Ter- esopolis, VIII-1999. Etymology.—The specific name annuli- pes is an adjective meaning “banded leg”’ that refers to the striking light and dark pat- tern of the legs of this species. Gall (Fig. 4).—The gall of M. annulipes is an elongate, swelling of the leaf veins on either surface, and of the petioles and young branches. Mature larval chamber walls are slightly rigid and surrounded with spongy tissue. When young the gall appears to be a simple blister but as it matures de- velops the characteristic spongy texture. The exterior color is green, similar to that of the leaves. One or more larvae may be found in a gall, where they pupate and eventually emerge from an opening near ei- ther end of the gall. The gall was fairly abundant in each of the collecting localities. Perasphondylia mikaniae Gagné, new species (Figs 30-31, 70-75) Adult.—Head: Antenna (Figs. 70-71) with scape slightly more than twice length pedicel; pedicel slightly wider than long; first flagellomere of male about 1.5 times longer than scape, that of female more than twice length of scape. Eye facets hexago- nal, closely approximated. Frons with ap- proximately 35—40 setae. Clypeus asetose dorsally, with several short setae laterally, and setulose ventrally. Hypopharynx close- ly lined with short setae dorsally. Labella setose and setulose laterally and ventrally, with a patch of short setae mesally. Palpus 3 segmented, first segment about as long as wide, second segment about twice as long as wide, third about 3 times as long as wide, all covered with setulae and setae. Thorax: Wing length, male 3.2 mm (n = 3), female 4.1—4.4 (n = 5). Dorsocentral setal row at midlength with three rows of setae intermixed with several scales, the row continuing onto scutellum to posterior margin. Scutellum bare between dorsocen- tral rows of setae, with posterior setae lat- erad of dorsocentral rows, and covered with scales laterally. Anepisternum with setae and scales on dorsal half to two-thirds. Anepimeron covered with setae. Katepi- meron bare. Tarsal claws (Fig. 72) subequal in size and similar in shape, as long as em- podia. Abdomen: Tergites without anterior pair of trichoid sensilla, first through seventh tergites rectangular with mostly single row of posterior setae, many lateral setae, else- where covered with scales, width of sixth and seventh tergites about 2% times length; eighth tergite short, bandlike, without ves- titure. Sternites without anterior pair of trichoid sensilla, width of sixth sternite about twice length, male eighth sternite re- duced in size, as wide as long. Male geni- talia as in Fig. 73. Distal half of ovipositor approximately twice length seventh sternite. Pupa (Figs. 30—31).—Antennal horns dorsoventrally flattened, acute apically, me- sal edge serrate. Frons with elongate, point- ed upper horn and three-pointed lower horn, the mesal much shorter than the lat- eral points. Abdominal tergites 2—8 each with, from posterior margin, a row of stout spines, a short bare space, another row of stout spines followed by two or three irreg- ular horizontal rows of scattered, generally smaller spines, these diminishing in size and growing sparser toward anterior margin of sclerite. Larva.—Spatula (Fig. 74) with two large, widely separated, anterior teeth, the space between sometimes minutely serrate. Ster- nal, the 4 pairs of lateral, and the pair of ventral papillae are with setae. Terminal pa- pillae not distinguishable from spicules (Fig. 75). Holotype.—Male, from bud gall on Mi- kania glomerata, Silva Jardim, Pogo das Antas Biological Reserve, RJ, XI-1997, Oda & Monteiro, deposited in Museu Na- cional do Rio de Janeiro. Other material examined (all from bud galls on M. glomerata, Rio de Janeiro state, Brazil).—1 3, 6 2, 6 pupal exuviae, same VOLUME 103, NUMBER 1 data as holotype; 1 ¢, 3 2, 4 pupal exuviae, same data as holotype except 6-XII-1996; 1 3,5 2, 6 pupal exuviae, same data as ho- lotype except XII-1997; 2 pupa exuviae, Parati, VII-1997; 5 larvae, Parque Nacional do Itatiaia. Etymology.—The specific name mikan- iae is the genitive of the host plant name. Gall (Fig. 8).—Perasphondylia mikaniae forms a bud gall on the terminal and axilary buds of young stems of M. glomerata. It causes an extreme foreshortening of the stem on which the leaves become concen- trated to form a rosette. Stem growth does not proceed beyond the gall. The galls con- tain several to as many as thirty ovoid cells, each inhabited by one larva. Pupae escape from holes near the gall apex. The gall was found in most of the collecting localities but was the least abundant kind of gall found. Remarks.—The only other described congener, Perasphondylia reticulata Mohn, is known from bud galls on Chromolaena odorata L. (K.&R.) collected in Trinidad, El Salvador, and Ecuador. Both species are relatively large and similar in the adult Stage, although available specimens show slightly smaller male hypoproct lobes in the new species. The pupal upper frontal horn is undivided and stronger in the new spe- cies, divided and not so robust in P. retic- ulata. Both species have 3-pointed lower frontal horns, but in the new species the middle point is much shorter than the other two, while in P. reticulata all the points are the same length. The larva of M. mikaniae has four lateral papillae, while that of P. reticulata has five. The terminal larval pa- pillae are not visible in the new species but apparent in P. reticulata. ACKNOWLEDGMENTS We are grateful to: Paulo Ormindo for the drawings of the galls; the staff of Re- serva Biol6gica de Pogo das Antas (IBA- MA) for permission to collect and for the use of facilities; FAPERJ, CNPq and the Fundacao O Boticario de Prote¢ao a Natu- reza for financial support; Nit Malikul for 133 preparing the microscopic slides; Lucrecia H. Rodriguez for computer assistance in preparing the plates; and for their com- ments on drafts of the manuscript: Keith M. Harris, formerly of the International Insti- tute of Entomology, CAB, London, UK; Peter Kolesik, The University of Adelaide, Australia; Valéria Cid Maia, Museu Na- cional, Rio de Janeiro; Douglass R. Miller and Allen L. Norrbom of the Systematic Entomology Laboratory, USDA, Beltsville, MD, and Washington, DC, and an anony- mous reviewer. LITERATURE CITED Cortopassi-Laurino, M. and M. Ramalho. 1988. Pollen harvest by africanized Apis mellifera and Trigona spinipes in Sao Paulo (Brasil) botanical and bio- logical viewes. Apidologie 19: 1—24. Gagné, R. J. 1989. The Plant-Feeding Gall Midges of North America. Cornell University Press, Ithaca, New York. xi and 356 pp. and 4 pls. Gagné, R. J. 1994. The Gall Midges of the Neotropical Region. Cornell University Press, Ithaca, New York. xv and 352 pp. King, R. M. and H. E. Robinson. 1987. The Genera of the Eupatorieae (Asteraceae). Missouri Botan- ical Garden, St. Louis. 581 pp. McAlpine; J: EB: V. Beterson; G: E> Shewell) Ee ar Teskey, J. R. Vockeroth, and D. M. Wood (eds.). 1981. Manual of Nearctic Diptera. Vol. 1. Re- search Branch, Agriculture. Canada Monograph No 27. vi + 674 pp. Mohn, E. 1960. Gallmiicken (Diptera, Itonididae) aus El Salvador. 2. Teil. Senckenbergiana Biologica 41: 197-240. . 1964. Gallmiicken (Diptera, Itonididae) aus El Salvador. 7. Teil: Lasiopteridi. Beitrage zur En- tomologie 14: 553—600. . 1973. Studien tiber neotropische Gallmticken (Diptera, Itonididae). 2. Teil. Stuttgarter Beitrage zur Naturkunde (A) 257: 1-9. Monteiro, R. FE, FE FE E Ferraz, V. C. Maia, and M. A. P. de Azevedo. 1994. Galhas entomogenas em res- tingas: uma abordagem preliminar. Atas do III Simp6sio de Ecossistemas da Costa Brasileira Vol. Ill (ACIESP No. 87): 210-220. Oliveira, E, G. Akisue, M. K. Akisue, B. Mancini, and M. Chumzum. 1987. Morfodiagnose de ax6fito de guaco—Mikania gloverata Spreng. Revista de Ciéncias Farmacéuticas, Araraquara 8/9: 11-24. Riibsaamen, E. H. 1907. Beitrage zur Kenntnis aus- sereuropdischer Zoocecidien. III. Beitrag. Gallen aus Brasilien und Peru. Marcellia 6: 110-173. . 1916. Beitrag zur Kenntnis aussereuropdisch- 134 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON er Gallmiicken. Sitzungsberichte der Gesellschaft cecidologiae Braziliae. Brotéria, Série Zoolégica Naturforschender Freunde zu Berlin 1915: 431— 8: 5—28, pls. I-VIII. 481. Wiinsch, A. 1979. Gallenerzeugende Insekten Nord- Skuhrava, M. 1989. Taxonomic changes and records kolumbiens, speziell Asphondyliidi und Lasiop- in Palaearctic Cecidomyiidae (Diptera). Acta En- teridi (Diptera, Cecidomyiidae) aus dem Kiisten- tomologica Bohemoslovaca 86: 202-233. bereich um Santa Marta. A. Wiinsch, Waiblingen, Tavares, J. S. 1909. Contributio prima ad cognitionem West Germany. 238 pp. PROC. ENTOMOL. SOC. WASH. 103(1), 2001, pp. 135-142 MORPHOLOGY AND DISTRIBUTION OF SENSILLA ON THE ANTENNA OF OMOGLYMMIUS AMERICANUS (LAPORTE 1836) (COLEOPTERA: RHYSODIDAE) GERALD T. BAKER Department of Entomology and Plant Pathology, Mississippi State University, Box 9775, Mississippi State, MS 39762, U.S.A. (e-mail: gbaker@entomology.msstate.edu) Abstract.—Morphology, distribution, and number of sensilla on the female and male antennae of Omoglymmius americanus (Laporte 1836) are described using scanning elec- tron and light microscopy. Both sexes have the same types of sensilla, 3 types of sensilla trichodea (ST), 3 types of sensilla basiconica (SB), 1 type each of sensillum coeloconicum (SCO), and sensillum ampullaceum (SA). ST, and ST, are the only non-porous types, whereas all other types are porous as indicated by their staining with silver nitrate and crystal violet. ST, is found on the scape, pedicel and all flagellomeres; ST, on all flagel- lomeres; ST, on flagellomeres 3—9; SB, and SB, on flagellomeres 3—9; SB, on flagellom- eres 7—9; SCO and SA on the apical flagellomere. The males have significantly more SB, and SCO. Patches of microtrichia and pores are situated on the scape, pedicel and fla- gellomeres 1 and 2. Key Words: Species of the family Rhysodidae are considered to be caraboids that have a mod- ified head and mouthparts, and some tax- onomists have placed these beetles in the Carabidae (Bell 1970, 1998; Bell and Bell 1982). These beetles feed on slime molds (Bell 1994) and have modified mouthparts which are used for wedge-pushing between layers of wood. There are a number of sys- tematic and ecological studies on this group of beetles, but studies are lacking on how these beetles perceive stimuli such as chem- ical cues for host and/or mate location. To date, there are no studies on the sensilla that perceive various mechanical and chemical stimuli which are encountered by these bee- tles. This investigation presents data on the morphology, number and distribution of sensilla on the antennal flagellum of Omo- glymmius americanus (Laporte 1836) (Co- leoptera: Rhysodidae). cuticular sensory receptors, caraboid beetle METHODS For scanning electron microscopy (SEM), specimens of Omoglymmius amer- icanus were placed in half strength Karnov- sky’s fixative in 0.1/M phosphate buffer, pH 7.2 at 4°C overnight and then post-fixed in 2% OsO, in the same buffer for 4h. The specimens were dehydrated in a graded se- ries of ethanol, critical point dried and then mounted on aluminum stubs with carbon sticky tabs (Baker and Monroe 1995). After coating with gold-palladium, the material was examined with a Cambridge (LEO) 360 SEM at 15kV. To determine the porosity of the different types of sensilla, the crystal violet proce- dure of Slifer (1960) and silver nitrate method of Schafer and Sanchez (1976) were used on three female and three male specimens. These same specimens were also used to make counts and measurements 136 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON of the sensilla. The measurements are given as a mean plus the range. Student t-test was used to compare the mean number of sen- sillar types between females and males. RESULTS Antenna The antenna consists of a scape, a pedicel and nine flagellomeres that make-up the fla- gellum (Fig. 1). The scape is 260 jum (256— 263) long and 255 jm (252-258) wide and square whereas the pedicel is 148 pm (145— 151) long and 200 wm (196-206) wide (Fig. 2) and rectangular in shape. The nine flagellomeres are moniliform with a tapered distal end which encloses the ball-like base of the succeeding flagellomere and are globular and 200 wm (196-204) long and 254 wm (252-257) wide (Fig. 1). On the scape, pedicel, and the first two flagellomeres there are patches of microtri- chia (PA, Fig. 2). The patches are scattered over the surface of the scape and pedicel but are found only in association with the distal sensilla trichodea that encircle the fla- gellomeres (A, and A,) (Fig. 2). The mi- crotrichia are 3.0 to 4.5 wm long and a patch may contain from about 50 to 200 microtrichia (MT, Fig. 4). Pores are also sit- uated in these patches of microtrichia (P, Fig. 4). The distal half of the last flagellomere is conical and its surface is covered with sen- silla whereas flagellomeres 3 to 8 have a distinct band of sensilla in a recessed area on the distal end of each flagellomere (Fig. 1, arrowheads pointing out distal bands of sensilla). The pedicel, scape, and flagellom- ers One and two do not have a distinct band of sensilla, but the few sensilla on their sur- face are all concentrated at the distal end of the flagellomere (Fig. 2). Based upon their morphology and porosity, 3 types of sen- silla trichodea (ST), 1 type of sensillum am- pullaceum (SA), | type of sensillum coel- conicum (SC), and 3 types of sensilla bas- iconica (SB) are present on the antennae of the female and male. Their distribution and numbers are given in Table | and Figs. 12, 13. No differences in types, number and di- mensions occur between the female and male except the male have significantly more sensillum basiconicum III and sensil- lum coeloconicum. The distribution pat- terns of each type of sensillum on both sex- es also are very similar (Figs. 12, 13). Sensillum trichodeum (ST).—The first type of sensillum trichodeum (ST),) is lo- cated on the scape, pedicel and flagellom- eres 3 to 9 in the center of each flagellom- ere. They form a ring of sensilla around each flagellomere and project outwards from the surface of the flagellomere (Fig. 1). ST, is distinctly curved with a pointed apex, and longitudinal ridges occur on the surface of the sensillum (Fig. 5). This sen- sillum is 80 wm (76-85) long and 4.4 wm (4.3—4.6) wide at the base. ST, is located on all flagellomeres and occurs in a ring around the apical portion of each flagellom- ere (Figs. 1, 2). It is straight, pointed and with well-defined longitudinal ridges on the cuticular shaft of the sensillum (Fig. 6). This sensillum is 87 wm (85—91) long and 5.1 wm (4.9—5.3) wide at the base. ST, and ST, increase in number from the proximal end to the distal end of the antenna (Fig. 12) and they did not stain with crystal vi- olet or silver nitrate. The third type of sen- sillum trichodeum (ST TI) is found in the distal sensory band on the flagellomeres (Fig. 7). There are 4—6 ST, that encircle a flagellomere which is 44 wm (42-47) long and 3.9 wm (3.7—4.1) wide at the base. The cuticular shaft is straight, with slight lon- gitudinal fluting, and a blunt tip with an apical pore. This sensillum stained with crystal violet and silver nitrate and also in- creases in number from flagellomere 3 to 9 (Fig. 12). Sensillum basiconicum (SB).—Type I sensillum basiconicum (SB,) is 23 wm (19— 25) long and 2.7 wm (2.6—2.9) wide at the base and distinctly curved (Figs. 3, 8). It has a blunt tip and smooth surface. Type II (SB,) is similar morphologically to SB, but is 35 wm long (33-38) and 2.8 wm (2.6— VOLUME 103, NUMBER 1 Figs. 1-6. Antennomeres and sensilla of Omoglymmuis americanus. 1, Distribution pattern of sensilla on the flagellomeres 1-9. 2, Sensilla and patches of microtrichia on the scape, pedicel and flagellomeres 1-2. 3, Three types of sensilla basiconica and sensillum ampullaceum on the terminal flagellomere. 4, Microtrichia and pores on the scape. 5, Long, slender, curved, pointed and grooved sensillum trichodeum, type one. 6, Long, straight and deeply grooved sensillum trichodeum, type two. Abbreviations: MT = microtrichia: P = pores; PA = patches of microtrichia; SA = sensillum ampullaceum; SB = sensillum basiconicum; ST = senillum tricho- deum; double arrow heads in Fig. 1 showing the apical concentration of sensilla on each flagellomere. 138 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Number (mean + SBE) of sensilla on the antennal flagellum. Type Female Male Si Spe) ae 2219)II 56.10 + 2.83 ST, 139.89 + 4.01 142.74 + 3.83 Silk 43.92 + 2.96 45.18 + 3.21 SB, 178.84 + 2.95 181.25 = 3.14 SB, 196.85 + 4.12 DOW) 574.22 SB, 28.94 + 2.02 Nokes as M7 SCO Pr Tfs) 25 (OY IIS) 53,010) 25 O40} SA 5.50 + 0.10 5:66: = 0:20 * Indicates significant difference on mean number of sensilla on the antennal flagellum between female and male (p < 0.05). 2.9) wide at the base (Fig. 9). SB, and SB, form a distinct distal band (which is called the pubescent, subapical band in the taxo- nomic literature) on flagellomeres 3 to 9. The number of SB, and SB, sensilla in- creases from flagellomere 3 to flagellomere 9 (Fig. 13). A third type of sensillum bas- iconicum (SB,) is on flagellomeres 7 to 9 and is 19 wm (17-21) long and 3.0 pm (2.8—3.3) wide at the base (Fig. 10). This sensillum is not distinctly curved and has a stout appearance. All 3 types of sensilla basiconica stained intensely with crystal vi- olet and silver nitrate which indicates their porosity. Sensillum coeloconicum (SCO).—It is located on the last flagellomere of the fe- male and male antennae. Females have 2— 3, whereas the males have 5—6. The pit in which the sensillum is situated measures 1.9 wm (1.7—2.2) in diameter, and the cu- ticular peg portion is 1.7 wm (1.5—2.0) long and 0.9 wm (0.8—1.1) wide at the base (Fig. 11). The apical portion of the sensillum is distinctly pointed, and the cuticular peg also stained with the crystal violet and silver ni- trate indicating its porosity. Sensillum ampullaceum (SA).—This type of sensillum is found only on the last flagellomere of male and female antennae. The opening of the sensillum is in the cen- ter of a slightly raised area that is delineated by an outer raised cuticular rim (Fig. 3) and is 1.4—1.7 ym in diameter. The peg portion can only be seen in specimens that were depigmented after the silver nitrate staining and it appears as a blackened, rod-shaped speck. Males and females have the same number of this sensillum, five or six. DISCUSSION Two types of sensilla trichodea (ST, and ST,) are probably mechanoreceptors. They have a well developed basal socket; they are not porous as indicated by the lack of staining, and every broken ST, and ST, sensillum, when viewed with the SEM, has a solid cuticular shaft. These above-men- tioned characteristics are typical of certain types of mechanoreceptive sensilla. ST, and ST, resemble the trichoid sensilla type I on Carabus fiduciarius saishutoicus Csiki (Carabidae) (Kim and Yamasaki 1996), Ne- bria brevicollis (FE) (Carabidae) (Daly and Ryan 1979), Tenebrio molitor L. (Tene- brionidae) (Harbach and Larsen 1977) and the non-porous hairs of Oryzaephilus suri- namensis L. (Silvanidae) (White and Luke 1986). These two sensilla (ST, and ST,) make up a large proportion of the total number of antennal sensilla, and their po- sition on the antenna also indicates that mechanoreception is an important function of the antenna. Mechanoreception would be an important sensory input as these beetles pushed their way through wood fibers in search of slime molds. The blunt tip with an apical pore on ST, and the staining by the crystal violet and silver nitrate, which indicates porosity, are characteristics of a chemoreceptive sensil- lum. This sensillum on Omoglymmius americanus is similar to the type II sensil- lum trichodeum of Nebria brevicollis (Daly and Ryan 1979), Carabus fiduciaruuis sais- hutoicus (Kim and Yamasaki 1996) and the gently grooved sensillum in Diabrotica vir- gifera LeConte (Chrysomelidae) (Staetz et al. 1976). The ultrastructural data presented by Daly and Ryan (1979) clearly showed that this type of sensillum trichodeum is a contact chemoreceptor. The three types of sensilla basiconica on VOLUME 103, NUMBER 1 Figs. 7-11. Sensilla on the antenna of Omoglymmius americanus. 7, Short, stout, straight, blunt and grooved sensillum trichodeum type 3. 8, Short, slender, curved sensillum baciconicum type 1. 9, Long, slender, curved sensillum basiconicum type 2. 10, Stout, straight sensillum basiconicum type 3. 11, Peg and pit of the sensillum coeloconicum. Abbreviations: SB = sensillum basiconicum; SCO = sensillum coeloconicum; ST = sensillum trichodeum. 140 25 -- - -ST1 Male -- O--ST1 Female — O-— ST2 Male — O-— ST2 Female —t— ST3 Male —O— ST3 Female N fo) sf a a 4 Mean Number of Sensilla Trichodea/Flagellomere rs) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON / pe Wes “A Yi, “A Y 7; / 7 Le Le, ree ae sof — a a Be ee tage ee -7oO----- o pen ae aa =O oO ZO F1 F2 F3 F4 F5 F6 F7 F8 F9 Flagellomere Riss 12% the male and female antennae are stained intensely by the silver nitrate and crystal violet, indicating a porous cuticle. At high magnification with the SEM many pores can be seen which cover the surface of these sensilla. SB, and SB, on Omoglymius americanus are morphologically similar to the basiconic sensilla found on the antenna of Dendroctonus ponderosae Hopkins (Scolytidae) (Whitehead 1981), Oryzaephi- lus surinamensis L. (White and Luke 1986), Agriotes obscurus L. (Elateridae) (Merivel et al. 1997), and Carabus fiduciarius sais- hutoicus Csiki (Kim and Yamasaki 1996) whereas the SB, of O. americanus resemble the type III found or the male and female antennae of Psacothea hilaris (Pascoe) (Cerambycidae) (Dai and Honda 1990), C. fiduciarius saishutoicus (Kim and Yamasaki 1996) and Pterostichus spp. (Carabidae) (Symondson and Williams 1997). It is well documented that sensilla basi- Distribution of the three types of sensilla trichodea (ST) on the female and male antennae. conica respond to various odors such as those from conspecifics and food sources. The sensillum coeloconicum and sensillum ampullaceum of Omoglymmius americanus are morphologically similar to those on the antennae of other beetles, especially cara- bids (Juberthie and Massoud 1977, Nagel 1979, Daly and Ryan 1979, Kim and Ya- masaki 1996, Skilbeck and Anderson 1996, Symondson and Williams 1997). Both types are porous as indicated by the black deposit from the silver nitrate staining, so they are probably involved in some type of chemo- reception like the sensilla basiconica (SB,,,) and sensillum trichodeum, ST;. The type of sensillum ampullaceum on O. amer- icanus, which has an opening in the middle of a large, round, smooth plate-like area, is found only on several species of Carabidae (Nagel 1979, Kim and Yamasaki 1996, Sy- mondson and Williams 1997). Distribution patterns, numbers and types VOLUME 103, NUMBER 1 60 + -- 1 --SB1 Male -- O --SB1 Female — O-— SB2 Male — O-— SB2 Female —t— SB3 Male —O— SB3 Female en 55 + .S a ae 30 + 25 + SB2 20 + SS eS Sa ai a Soaps Cegeetcpoc yee Sg oSaosuO 0- Mean Number of Sensilla Basiconica/Flagellomere 141 / Site fine “Ye Lie BO Ee Goes Bae a. “ Bo nae Zag e Cet ete as Bo Orne a. SB3 SB1 FA F2 F3 F4 FS Flagellomere Fig. 13. of sensilla on Omoglymmius americanus are similar to what is found on the antenna of carabid beetles (Daly and Ryan 1979, Nagel 1979, Kim and Yamasaki 1996, Sy- mondson and Williams 1997). Sexual di- morphism exhibited between the females and males of O. americanus in the number of SB, and SCO is also found in other bee- tles such as Psacothea hilaris (Pascoe) (Dai and Honda 1990) and Semiadalia undecim- notata Schneider (Coccinellidae) (Jourdan et al. 1995). The sensilla on Nebria brevi- collis cover the flagellar surface which dif- fers from the distinct apical bands of sen- silla in a recessed portion of flagellomeres 3 to 8 of O. americanus. This difference may be due to the different habitat and feeding strategies. Nebria brevicollis is an active predator in open spaces and has a more graceful antenna. Omoglymmius americanus is found in confined space pushing through solid wood. The antennal sensilla may need to be protected from me- chanical damage, and the antenna itself is a Distribution of the three types of sensilla basiconica (SB) on the female and male antennae. tough ball and chain structure with a sharp- ened apical end, presumably also used for pushing through wood. The clusters of microtrichia and pores on the flagellomeres of Ommoglymmius amer- icanus also are situated on other body re- gions, especially around the antennal-clean- er located on the protibiae. These structures are also found in other Coleoptera (Faustini and Halsted 1982). They may be involved in secreting material that could be used in cleaning debris off the antennal surface, or they may secrete behavior ellicting chemi- cals (Martin 1975, Barbier et al. 1992). ACKNOWLEDGMENTS I thank R. Brown, P. Ma, and A. Cohen for reviewing the manuscript and A. Ladd for typing it. Partial support from NSF Grants DIR-9001493 and DEB 9200856. I also thank R. Kuklinski for technical assis- tance. This is publication No. J9563 of the Mississippi Agricultural and Forestry Ex- periment Station. 142 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON LITERATURE CITED Baker, G. T. and W. A. Monroe. 1995. Sensory recep- tors on the adult labial and maxillary palpi and galea of Cicindela sexguttata (Coleoptera: Cicin- delidae). Journal of Morphology 226: 25-31. Barbier, R., A. Ferran, J. LeLannic, and M. R. Allo. 1992. Morphology and ultrastructure of integu- mentary glands of Semiodalia undecimnotata Schn. (Coleoptera: Coccinellidae). International Journal of Insect Morphology and Embryology 21: 223-234. Bell, R. T. 1970. The Rhysodini of North America, Central America and the West Indies (Coleoptera: Carabidae or Rhysodidae). Miscellaneous Publi- cations of the Entomological Society of America 6: 289-324. . 1994. Beetles that cannot bite: functional mor- phology of the head of adult rhysodines (Cole- optera: Carabidae or Rhysodidae). Canadian En- tomologist 126: 667—672. . 1998. Where do the Rhysodini (Coleoptera) belong? pp. 261—272. In Ball, G. E., A. Casale, and A. Vigna Taglianti (Eds). Phylogeny and Classification of Caraboidea (Coleoptera: Adepha- ga). Atti del Museo Regionale di Scienze Naturali 5: 1-543. Bell, R. T. and J. R. Bell. 1982. Rhysodini of the world part III. Revision of Omoglymmius Ganglbauer (Coleoptera: Carabidae or Rhysodidae) and sub- stitutions for preoccupied generic names. Quaes- tiones Entomological 8: 127—259. H. G. and H. Honda. 1990. Sensilla on the anten- nal flagellum of the yellow spotted longicorn bee- tle, Psacothea hilaris (Pascoe) (Coleoptera: Cer- ambycidae). Applied Entomology and Zoology 225: 273-282. Daly, P. J. and M. FE Ryan. 1979. Ultrastructure of antennal sensilla of Nebria brevicollis (Fab) (Co- leoptera: Carabidae). International Journal of In- sect Morphology and Embryology 8: 169-181. Faustini, D. L. and D. G. H. Halstead. 1982. Setiferous structures of male Coleoptera. Journal of Mor- phology 173: 43-72. Harbach, R. E. and J. R. Larsen. 1977. Fine structure of antennal sensilla of the adult mealworm beetle Tenebrio molitor L. (Coleoptera: Tenebrionidae). International Journal of Insect Morphology and Embryology 6: 41—60. Jourdan, H., R. Barbier, J. Bernard, and A. Ferran. 1995. Antennal sensilla and sexual dimorphism of the adult lady bird beetle Semiadalia undecimno- tata Schm. (Coleoptera: Coccinellidae). Interna- tional Journal of Insect Morphology and Embry- ology 24: 307-322. Juberthie, C. and Z. Massoud. 1977. L equipement sen- soriel de lantenne d’un coleoptera troglobie Dai, = Aphaenops crypticola Linder (Coleoptera: Trechi- nae). International Journal of Insect Morphology and Embryology 6: 147-160. Kim, J. L. and T. Yamasaki. 1996. Sensilla of Carabus (Usiocarabus) fiduciarius saishutiocus Csiki (Co- leoptera: Carabidae). International Journal of In- sect Morphology and Embryology 25: 153-172. Laporte, E L. 1836. Etudes entomologique, ou descrip- tions d’insectes nouveaux et observations sur la synonymie. Revue Entomologique 4: 5—60. Martin, N. 1975. Ultrastructure des glandes dermiques de l’antenne d’un Coléoptére Cavernicole troglo- phile Choleva spec. (Coleoptera: Silphidae). Zeit- schrift fiir Morphologie die Tiere 80: 261—275. Merivel, E., M. Rahi, and A. Luik. 1997. Distribution of olfactory and some other antennal sensilla in the male click beetle Agriotes obscurus L. (Co- leoptera: Elateridae). International Journal of In- sect Morphology and Embryology 26: 75-83. Nagel, P. 1979. Aspects of the evolution of myrme- cophilous adaptations in Paussina (Coleoptera: Carabidae). Miscellaneous Papers of the Landbou- whogeschool Wageningen 18: 15—34. Schafer, R. and T. V. Sanchez. 1976. The nature and development of sex attractant specificity in cock- roaches of the genus Periplaneta. I. Sexual di- morphism in the distribution of antennal sense or- gans in five species. Journal of Morphology 149: 139-158. Skilbeck, C. A. and M. Anderson. 1996. The ultra- structure, morphology and distribution of sensilla on the antennae of the adult parasitoids Aleochara bilineata Gyll. and Aleochara bipustulata L. (Co- leoptera: Staphylinidae). International Journal of Insect Morphology and Embryology 25: 261—280. Slifer, E. H. 1960. A rapid and sensitive method for identifying permeable areas in the body wall of insects. Entomological News 71: 179-182. Staetz, C. A., H. J. Ball, and S. D. Carlson. 1976. Antennal morphology of Diabrotica virgifera adults. Annals of the Entomological Society of America 69: 695-698. Symondson, W. O. C. and I. B. Williams 1997. Low- vacuum electron microscopy of carabid chemo- receptors: a new tool for the identification of live and valuable museum specimens. Entomologia Experimentalis et Applicata 85: 75-82. White, P R. and B. M. Luke. 1986. Fine structure, function and distribution of antennal sensilla in the saw-toothed grain beetle, Oryzaephilus suri- namensis. Physiological Entomology 11: 227— 238. Whitehead, A. T. 1981. Ultrastructure of sensilla of the female mountain pine beetle, Dendroctonus pon- derosae Hopkins (Coleoptera: Scolytidae). Inter- national Journal of Insect Morphology and Em- bryology 10: 19-28. PROC. ENTOMOL. SOC. WASH. 103(1), 2001, pp. 143-146 A TAXONOMIC STUDY OF THE GENUS ANOMALOSIPHUM TAKAHASHI FROM CHINA (HOMOPTERA: APHIDIDAE: GREENIDEINAE) G. X. QIAO AND G. X. ZHANG Institute of Zoology, The Chinese Academy of Sciences, 19 Zhongguancun Road, Haidian, Beijing 100080, People’s Republic of China Abstract.—The aphid genus Anomalosiphum in China is reviewed. Three species occur in China, including A. scleroticum, n. sp. which is described and illustrated. A key is provided to the alate viviparous females. Key Words: In the classification of the Aphidoidea, as interpreted by Raychaudhuri and Chatterjee (1980), Anomalosiphum is regarded as a member of the small tribe Cervaphidini within the subfamily Greenideinae. Ghosh (1982) provided a generic key to the nine genera within Cervaphidini. The genus Anomalosiphum was de- scribed by Takahashi (1934) from alate vi- viparous females of the type species, An- omalosiphum pithecolobii, collected on Pi- thecolobium lucidum (Fagaceae) in Taiwan, China. Tao (1947) described a second spe- cies, A. takahashii, from Taiwan, China, and Ghosh et al. (1971) described a third species, A. indigoferae, from India. Martin and Agarwala (1994) studied the genus An- omalosiphum and described three new spe- cies, A. murphyi, A. philippinensis and A. tiomanensis. Based on Remaudiére and Re- maudiére’s (1997) catalogue, this genus comprises six species, all in Asia. In China, Anomalosiphum includes only three spe- cies, one of which is new. This paper deals with the species from China, except for the information on the type species from Takahashi (1934). The specimens were collected by the author and are deposited in Zoological Museum, Insti- tute of Zoology, Chinese Academy of Sci- ences. Homoptera, Aphididae, Greenideinae, Anomalosiphum, new species, China The terminology follows Tao (1947) and Martin and Agarwale (1994). Measure- ments are in millimeters (mm). Anomalosiphum Takahashi, 1934 Anomalosiphum Takahashi 1934: 54; Ghosh 1982: 83; Raychaudhuri and Chat- terjee; 1980:.316; Tao. 19902105. type species: Anomalosiphum pithecolobii Takahashi, 1934, by original designation. Diagnosis.—Antenna 4-segmented in ap- terae and 5-segmented in alatae, secondary rhinaria semiannular. Tergites VII and VIII each with a pair of pronounced hair-bearing processes in apterae, these reduced in alatae and usually distinguishable on the tergite VIII only as large hair-bearing tubercles. Cauda transversely rounded triangular, with a pronounced median stylus in apterae which is reduced in alatae. Fore wing with median vein one branched, hind wing with one obliquus. Embryo with dorsal hairs fan-shaped at apex, metanotum to abdominal tergite VI each with 1 pair of spinal and 1 pair of pleural hairs; thorax each with 2 pairs of marginal hairs, pronotum with 1| pair of spi- nal hairs; abdominal tergites I-VII each with 1 pair of marginal hairs; tergites VII and VIII each with 1 pair of spinal hairs. 144 Host plants.—Species of Anomalosiphum have been recorded colonizing members of the plant families Connaraceae, Fagaceae, Leguminosae, Mimosaceae and Polygala- ceae. Distribution.—China (Taiwan, Guangxi, Sichuan), India, Papua New Guinea, Phil- ippines, Singapore, Western Malaysia. KEY TO SPECIES OF ANOMALOSIPHUM (ALATE VIVIPAROUS FEMALES) FROM CHINA (INCLUDING A. TIOMANENSIS MARTIN) 1. Antennal segment III bearing up to 20 second- ary rhinaria, often rather unevenly distributed; segments IV and V without secondary rhinaria mo. cha yl ch diol cin ter ee ee ee Ri takahashii Tao — Antennal segment III bearing more than 30 secondary rhinaria, distributed over whole length of segment (Fig. 2); segment IV, and sometimes also segment V, with secondary rhi- Manta (RNG 2)y 7.2 .taseenn hs soe eet eo Epo eerie ey: 2 . Ultimate rostral segment about 1.85 times as long as second hind tarsal segment...... pithecolobii Takahashi — Ultimate rostral segment at most 1.6 times as NO long as second hind tarsal segment........ 3 3. Antennal segment V without secondary rhinar- ia; ultimate rostral segment 0.47 times siphun- culus length; abdominal segment VII with 1 pair of short marginal processes; spino-pleural band on abdominal tergite VII fused with an irregular brown patch on tergites HI—VI; length of hair on apex of process VIII as long as pro- cess VIII scleroticum, nN. sp. — Antennal segment V with 1—3 secondary rhi- naria; ultimate rostral segment 0.63 times length of siphunculus; abdominal segment VII without short marginal processes; tergite VII with a subtrapezoidal brown sclerite medially; length of hair on apex of process VIII 1.8 times as long as process VIII ..... tiomanensis Martin Anomalosiphum scleroticum Qiao and Zhang, new species (Figs. 1-12) Material examined.—Holotype: Alate vi- viparous female, No. Y1046-1-1-2, April 23, 1974, Guangxi Autonomous Region (Nanning City, 22.8°N, 108.3°E, Alt. 200 m), on Dalbergia hupeana Hance (Legu- minosae), by Wang Zhongfu; paratype 1 alate viviparous female, No. Y1046-1-1-1, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON other data same as holotype; the aphids were colonizing host plant. Description.—Alate viviparous female: Body, long, elliptical. Measurements: Body 1.588 in length, 0.706 in width. Antenna 0.909, length of segments I-V: 0.062, 0.046, 0.427, 0.188, 0.139 + 0.046, respec- tively. Ultimate rostral segment 0.093 in length, 0.036 in basal width. Hind femur 0.319, hind tibia 0.500, 2nd hind tarsal seg- ment 0.088 in length. Siphunculus 0.193 in length. Cauda 0.093 in length. Mounted specimens.—Head and thorax dark brown, eye black. Antenna, apical to rostrum, legs, and siphunculi dark brown. Posterior margin of pterostigma and veins dark brown. Abdominal tergites I and II each with a transverse pigmented band; spi- nal and pleural patches on tergites III—VII fused with each other forming an irregular brown patch, abdominal tergites each with a pair of small sclerotic marginal patches. Venter of abdomen with spinulose stripes, abdominal tergites I, VI-VIII spinulose, an- tennal segments and tarsi with transverse imbrications. Dorsal hairs of body thick and short; ventral hairs long and acute, length of ventral hairs 4 times as long as dorsal hairs. Head with 2 pairs of cephalic hairs, 5 or 6 dorsal hairs; abdominal tergite VII with 12 hairs, tergite VIII with 2 hairs. Ab- dominal segment VII with | pair of short marginal processes 0.016 in length, tergite VIII with 1 pair of long spinal processes 0.057 in length, each process with a long and acute hair at apex. Length of hairs 4.33 times as long as process on abdominal seg- ment VII, length of hairs as long as process on tergite VIII. Length of cephalic hairs 0.018, 0.44 times as widest diameter of an- tennal segment III. Spiracles elliptical, closed, spiracular plates black. Medial front and antennal tubercles slightly developed. Antenna 5-segmented, 0.57 times as long as body, length in proportion of segments I— V: 14, 11, 100, 44, 33 + 11. Processus ter- minalis 1/3 as long as base of segment V. Antennal hairs short and sharp. Antennal segments III-IV with 38—40, and 9 or 10 VOLUME 103, NUMBER 1 Figs. 1-12. NS Se ad oS RO eS SSeS ~ Anomalosiphum scleroticum, alate viviparous female. 1, Dorsal view of head. 2, Antennal seg- ments I-V. 3, Ultimate rostral segment. 4, Dorsal view of abdomen. 5, Marginal process on abdominal segment VII. 6, Medial process on abdominal tergite VIII. 7, Dorsal hair of body. 8, Ventral hair. 9, Siphunculus. 10, Cauda. 11, Anal plate. 12, Embryo. Scale bar: Figs. 1, 2, 5-11 = 0.1 mm; Fig. 3 = 0.05 mm; Figs. 4, 12 = 0.2 mm. semiannular secondary rhinaria, respective- ly, distributed over whole length of each segment; segment V without secondary rhi- naria. Rostrum reaching mid-coxae, length of ultimate rostral segment 2.57 times as long as its basal width, 1.06 times as long as 2nd hind tarsal segment. Legs normal. Hind femur 0.75 times as long as antennal segment III, hind tibia 0.31 times as long as body. Hairs on legs sharp, on hind tibia 0.033, 1.48 times as long as mid-width of segment. First tarsal segment chaetotaxy: 3, 3, 3. Siphunculus 0.196 in length, 0.12 times as long as body length. Cauda broad and round-triangular, bearing 6 hairs. Anal plate with 8—10 hairs. Genital plate with 8 hairs. Embryo: Oval, 0.556 in length, 0.247 in width. Dorsal hairs of body thick and short, mushroom-shaped at apex. Head with 2 pairs of cephalic hairs, 2 pairs of spinal hairs, | pair of pleural and 1 pair of mar- ginal hairs. Pronotum with 1 pair of spinal hairs and 2 pairs of marginal hairs. Meson- otum and metanotum each with | pair of spinal, | pair of pleural and 2 pairs of mar- ginal hairs. Abdominal tergites I-V each with 1 pair of spinal, 1 pair of pleural and 1 pair of marginal hairs. Tergite VI with 1 pair of spinal and 1 pair of pleural hairs. Tergite VII with 1 pair of spinal and 1 pair of marginal hairs. Tergite VIII with | pair of spinal hairs. Eye 3-faceted. Rostrum reaching abdominal segment I. Tarsi nor- mal, thick and short. First tarsal segment chaetotaxy: 2, 2, 2. Second hind tarsal seg- ment 0.052 in length. Siphunculus pore- shaped. Cauda small, circular at apex, with 2 short and sharp hairs. Anal plate with 4 hairs. 146 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Discussion.—The new species is closely related to Anomalosiphum tiomanensis Martin 1994, but differs from it by: anten- nal segment V without secondary rhinaria (the latter: with 1—3 secondary rhinaria), ul- timate rostral segment 0.47 times siphun- culus length (the latter: 0.63 times), abdom- inal segment VII with 1 pair of short mar- ginal processes (the latter: absent), spino- pleural band on abdominal tergite VII fused with an irregular brown patch on tergites II-VI (the latter with tergite VII with a subtrapezoidal brown sclerite medially), length of hair on apex of process VIII as long as process VIII (the latter: 1.80 times). Anomalosiphum pithecolobii Takahashi 1934 Anomalosiphum pithecolobii Takahashi 1934: 54; Martin and Agarwala 1994: 422; Tao 1990: 105. Host plant.—Pithecolobium lucidum Benth (Mimosaceae). Distribution.—China: Guangdong, Hu- bei, and Taiwan (Takahashi 1934). Anomalosiphum takahashii Tao 1947 Anomalosiphum takahashii Tao 1947: 149— L553: Host plant.—unknown. Distribution.—China: Guangxi Auto. Reg. (Beiliu County, 22.7°N, 110.3°E, Alt. 200 m, No. Y6745; Nanning City, Alt. 200 m, No. Y995), Sichuan (Xichang) (the ma- terial from Tao 1947 and Martin and Agar- wala 1994). ACKNOWLEDGMENTS The authors are grateful to Mr. Wang Zhongfu for collecting the specimens. The kindness of Prof. J. H. Martin (Department of Entomology, The National History Mu- seum, Cromwell Road, London SW7 5BD) and Prof. B. K. Agarwala (Department of Life Sciences, Tripura University, Agartala 799004, India), in sending original material of A. murphyi, A. philippinensis, and A. tiomanensis for study, is also gratefully ac- knowledged. This work was supported by the National Natural Sciences Foundation of China (Grant: 39700015), a biological and technical innovation grant from the Chinese Academy of Sciences (Grant: C2999084), and a major project from the National Natural Science Foundation of China (Grant: 39899400). LITERATURE CITED Blackman, R. L. and V. FE Eastop. 1984. Aphids on the World’s Crops. Chichester, John Wiley & Sons, 466 pp. Ghosh, A. K. 1982. Ceraphidini (Homoptera: Aphi- doidea) of the world. Oriental Insects 16(1): 77— 97. Ghosh, A. K., M. R. Ghosh, and D. N. Raychaudhuri. 1971. Studies on the aphids (Homoptera: Aphi- didae) from eastern India. VII. New species and new records from west Bengal. Oriental Insects 5(2): 209-222. Martin, J. H. and B. K. Agarwala. 1994. A taxonomic study of the genus Anomalosiphum Takahashi (In- secta, Aphidoidea). Zoological Journal of the Lin- nean Society 111: 417—429. Raychaudhuri, D. N. and M. Chatterjee. 1980. Sub- family Greenideinae, pp. 314-358. Jn Raychau- dhuri, D. N., ed., Aphids of Northeast India and Bhutan. The Zoological Society, Calcutta. Remaudiére, G. and M. Remaudiére. 1997. Catalogue of the World’s Aphididae. Homoptera: Aphidoi- dea. Institut National de la Recherche Agrono- mique, 147, rue de |’ Université, 75338 Paris Ced- ex 07. 473 pp. Takahashi, R. 1934. Two new genera of Aphididae (Hemiptera). Stylops 3(3): 54-58. Tao, C. C. 1947. Descriptions of three new aphids from west China. Notes d’Entomologie Chinoise 11(5): 149-155. . 1990. Aphid-fauna of Taiwan Province, Chi- na. Taipei, Taiwan Provincial Museum, 328 pp. PROC. ENTOMOL. SOC. WASH. 103(1), 2001, pp. 147-156 A REVISION OF THE PATAGONIAN PREDACEOUS MIDGE GENUS BORKENTHELEA SPINELLI AND GROGAN (DIPTERA: CERATOPOGONIDAE) GUSTAVO R. SPINELLI AND WILLIAM L. GROGAN, JR. (GRS) Instituto de Limnologia “Dr. Raul A. Ringuelet’’, casilla de correo 712, 1900 La Plata, Argentina, and Departamento de Entomologia, Museo de La Plata, Argentina (e-mail: spinelli@museo.fcnym.unlp.edu.ar); (WLG) Department of Biological Sciences, Salisbury State University, Salisbury, Maryland 21801, U.S.A. (e-mail: wlgrogan@ssu. edu) Abstract.—A revision of the Patagonian predaceous midge genus Borkenthelea Spinelli and Grogan revealed three undescribed species that we describe and illustrate as new species: Borkenthelea quatei, from Argentina; B. harii, from Argentina and Chile; and B. nerudai, from Chile. A key is provided for the recognition of males and females of the four species in this genus. Resumen.—La revision del género Borkenthelea Spinelli y Grogan de la Patagonia, revelo la presencia de tres especies nuevas, las que son aqui descriptas e ilustradas: Bor- kenthelea quatei, de la Argentina; B. harii, de la Argentina y Chile; y B. nerudai, de Chile. Ademas, se presenta una clave para el reconocimiento de machos y hembras de las cuatro especies de este género. Key Words: Patagonia The predaceous midge genus Borkenthe- lea Spinelli and Grogan was proposed for the previously undescribed species B. nothofagus Spinelli and Grogan (1993). The type series of B. nothofagus included the holotype female and allotype male from Rio Negro Province, Argentina and two female paratypes from nearby Chile. A subsequent collecting trip to Patagonia by GRS and Larry Quate during 1994 yielded additional material that has al- lowed for a better understanding of the species composition of this apparently very rare genus. In this article, we present changes to the generic diagnosis, describe and illustrate three undescribed species, and provide a Diptera, Ceratopogonidae, Borkenthelea, predaceous midges, new species, key for the recognition of males and fe- males of all four species. For general ter- minology of Ceratopogonidae, see Downes and Wirth (1981); for terminology of pre- daceous midges in the tribe Ceratopogonini, see Wirth and Grogan (1988). Our previous paper on this genus (Spi- nelli and Grogan 1993) contains a detailed generic diagnosis and comparison of this genus with other similar appearing genera. Recently acquired specimens were all mounted on microscope slides in the man- ner of Wirth and Marston (1968). Types are deposited in the Museo de La Plata, Argen- tina (MLPA) or the Canadian National Col- lection in Ottawa, Canada (CNCI), as in- dicated for each new species. 148 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Borkenthelea Spinelli and Grogan Borkenthelea Spinelli and Grogan, 1993: 321. Type-species, Borkenthelea notho- fagus Spinelli and Grogan, by original designation. Diagnosis.—The original generic diag- nosis by Spinelli and Grogan (1993) is ad- equate for the recognition of all included species with the following emendations: Male antenna with flagellomeres 1—2 sepa- rated, 3-10 fused, 11—13 distinctly separat- ed. Hind claws of female of similar diam- eter to claws of other legs or more massive. Costal ratio of female 0.54—0.63; of male 0.51—0.60. Male tergite 9 very short and tri- angular to moderately long and semiquad- rate; apicolateral processes short and bul- bous to elongated and slender. Aedeagus very short, or of moderate length. Paramer- es separate or fused basally. Comments.—The condition of the male flagellum in which flagellomeres 3—10 are fused, apparently represents an apomorphic character state that is unique to the family (A. Borkent, personal communication) and is further evidence of the monophyly of this genus. KEY TO SPECIES OF BORKENTHELEA IR emialey yee A sens Graeme ota ieee eee rons oy a) Male vis fae eco sen ether eee marie ado & 5) . Hind claw not massive (Fig. 2); halter brown- ish; sternite 9 represented by two reduced, slender plates (Fig. 4); spermathecae with very korny mee (ENE D) ccogasccace B. harii, n. sp. — Hind claw massive; halter pale; sternite 9 rep- resented by two conspicuous plates; sperma- NEEAS Wwitiln Soar MEI socosvssoosccc0€ 3 3. Cell cua, without macrotrichia (Fig. 12) .... N | ORS £8 be! GA Ol DLa cae DER eek eae B. nerudai, n. sp. — Cell cua, with several macrotrichia........ 4 4. Large’ species (wing length 1.28-1.35 mm); mandible with 9-10 teeth; clypeus with 2-3 pairs of setae; antennal scape with 4 setae; an- tennal ratio 0.88—0.90; lumen of 2nd radial cell shontitriangulante sapere elon clo Cir B. nothofagus Spinelli and Grogan — Smaller species (wing length 1.12 mm); man- dible with 7 small teeth; clypeus with one pair of setae; antennal scape with 2 setae; antennal ratio 0.82; lumen of 2nd radial cell elongated (Fig. 19) «i ae Gu Payaen se aaa B. quatei, n. sp. 5. Sternite 9 with one or more caudal excavations — Sternite 9 without caudal excavations (Fig. 17) 6. Sternite 9 with a single shallow caudomedian excavation (Fig. 8); gonocoxite with a mesal truncated protuberance (Fig. 8); gonostylus dis- tinctly shorter than gonocoxite; aedeagus with distal portion bearing 2 pairs of strong lateral (KSeAilos tahoylovvaral (Ene, 110) 5 occ od oc B. harii, n. sp. — Sternite 9 with 3 deep caudal excavations, one medial and 2 lateral (Fig. 24); gonocoxite with- out mesal protuberance; gonostylus clearly lon- ger than gonocoxite; distal portion of aedeagus without lateral teeth, tip pointed ........ EH A ea es trees ara nD UNA SY AMS B. quateéi, n. sp. 7. Gonocoxite with mesal protuberance directed caudomesally (Fig. 17); gonostylus deeply curved; parameres separated (Fig. 18); aedea- gus with distal portion bearing 2 pointed scler- ites that cross each other at their midlength (ar tea WD ace eires Grohe ieewr che sok ce B. nerudai, n. sp. — Gonocoxite with mesal protuberance directed mesally; gonostylus slightly curved; parameres fused basally; aedeagus without pointed scler- itesvonudistalg portion) ee eee B. nothofagus Spinelli and Grogan Borkenthelea harii Spinelli and Grogan, new species (Figs. 1-11) Diagnosis.—Distinguished from other species of Borkenthelea by the following combination of characters: female, hind claws non-massive, sternite 9 represented by 2 reduced slender plates, spermathecae with very long necks, and wing with sub- equal radial cells lacking macrotrichia in central portion of cell cua,; male, tergite 9 broad caudally; sternite 9 with a single cau- domedian excavation; gonocoxite with a mesal truncated protuberance; gonostylus shorter than gonocoxite; aedeagus with 2 pairs of pointed lateral teeth on distal por- tion, tip bifid; and parameres separate. Female.—Head: Dark brown. Eyes pu- bescent, narrowly contiguous (V-shaped where contiguous). Antennal scape with 2 setae; flagellum (Fig. 1) brown; flagellom- ere 1 with 3—6 subapical sensilla coelocon- ica; lengths of flagellomeres (of allotype) in um 41-31-33-37-38-38-38-36-36-33-38-43- 61; antennal ratio 0.78 (0.75—0.81, n = 2). VOLUME 103, NUMBER 1 149 Dy; Ses LI] 7p 7 CCI Ve VELOPED PP TT 1’) Figs. 1-11. Borkenthelea harii. 1, Flagellomeres of female flagellum. 2, 5th tarsomeres and claws of female, from top to bottom of fore, mid and hind legs. 3, Female wing. 4, Sternites 8 and 9 of female. 5, Spermathecae. 6, Male flagellum. 7, Male wing. 8, Male genitalia. 9, Detail of apicolateral process. 10, Aedeagus. 11, Parameres. Scales = 0.05 mm. 150 Clypeus with 2 pairs of setae. Palpus brown; lengths of segments (of allotype) in um 15-28-28-18-28; segment 4 with 2—3 se- tae. Mandible with 8 teeth. Thorax: Dark brown; 4 prealar setae, | postalar seta; scutellum with one medial and two lateral setae. Wing (Fig. 3) mem- brane infuscated, veins brown; macrotrichia present in cells r,, m, and m,, only margin- ally in cell cua, and anal cell; radial cells equal sized, well developed, Ist with nar- row lumen (slightly broader in paratype), 2nd with broader lumen; radial veins not unusually thickened; media with long peti- ole, branching slightly distad to level of R,,3, M, obsolete at extreme base; M,,, and Cu, branching at level of midlength of Ist radial cell; wing length 1.00 (n = 2) mm, breadth 0.44 (n = 2) mm, costal ratio 0.62 (0.61—0.63, n = 2). Halter brownish. Legs brown; hind tibial comb with 7 setae; hind tarsal ratio 2.00 (n = 2); tarsomeres 5 (Fig. 2) slender with moderately large, unequal sized claws without basal teeth, those of hind leg not massive, claw length propor- tions of fore, mid, hind legs 12:7, 9:6, 10:6: Abdomen: Dark brown, segments 7—9 more heavily sclerotized than others. Gen- italia as in Fig. 4. Sternite 8 with pair of bluntly pointed posterior extensions. Ster- nite 9 represented by two reduced slender plates. Two ovoid spermathecae with long, slender necks plus rudimentary 3rd (Fig. 5), largest measuring 0.072 by 0.058 mm, neck 0.026 mm, and 0.062 by 0.046 mm, neck 0.026 mm. Male.—Similar to female with the fol- lowing notable sexual differences: flagel- lum (Fig. 6) with flagellomeres 1—2 sepa- rated, 3-10 fused, 11—13 distinctly separat- ed with lengths (of holotype) in um 92-77- 94; wing (Fig. 7) length 1.03 (1.00—1.04, n = 3) mm, breadth 0.36 (0.35-0-37, n = 3) mm, costal ratio 0.60 (n = 3). Genitalia as in Figs. 8—11. Sternite 9 a third as long as greatest breadth with deep caudomedian ex- cavation; tergite 9 with caudal margin near- ly straight, apicolateral processes (Fig. 9) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON with a single subapical seta, cercus short, with 2 apical setae. Gonocoxite stout, 2.7 times longer than greatest breadth, with stout, strongly sclerotized, truncated mesal protuberance; gonostylus slender with nu- merous small setae, clearly shorter than gonocoxite, distal half greatly curved and gradually tapering to slender, pointed tip. Aedeagus (Fig. 10) triangular, basal arch 0.5 of total length, distal portion with 2 pairs of strong lateral teeth, tip bifid. Para- meres (Fig. 11) separated, nearly contigu- ous anteriorly; basal arms very stout basal- ly; distal portions divergent with slender pointed tips. Distribution.—Argentina (Andean forests of Chubut province); Chile (Llanquihue, Osorno). Types.—Holotype 6, Argentina, Chubut, P.N. Los Alerces, 9/12-XII-1994, L. Quate, Malaise trap; Allotype 2, Chile, Llanqui- hue, Las Cascadas, 13 km N. Ensenada, %- XII-1994, L. Quate, Malaise trap. Paraty- pes, 3 6, 1 2, Chile, Osorno, Puyehue, 6/ 8-XII-1994, L. Quate, Malaise trap (depos- ited in MLPA). Etymology.—tThe specific name is a pa- tronym in honor of Dr. Hari Bhat, of the National Institute of Virology, Pune, In- dia, for his friendship during the 1994 trip to Patagonia with Dr. Larry Quate and GRS. Discussion.—The females of the three other species in this genus differ from this new species in having massive hind claws, a pale halter, sternite 9 composed of two conspicuous plates, and spermathecae with short necks. Males of B. nothofagus and B. nerudai differ from this new species in lacking caudomedial excavations on ster- nite 9. Males of B. quatei differ from this new species in having sternite 9 with three deep caudomedian excavations, a gonocox- ite with mesal protuberance, a gonostylus clearly longer than gonocoxite, and an ae- deagus with a pointed tip that lacks lateral teeth on its distal portion. VOLUME 103, NUMBER 1 Borkenthelea nerudai Spinelli and Grogan, new species (Figs. 12-18) Borkenthelea nothofagus Spinelli and Gro- gan 1993: 323, in part (female paratype from Cautin, Chile). Diagnosis.—Distinguished from other species of Borkenthelea by the following combination of characters: female, with massive hind claws, sternite 9 halves broad, spermathecae with short necks, and a wing with subequal sized radial cells that lacks macrotrichia in cell cua,; male, with tergite 9 tapering distally, sternite 9 without cau- domedian excavation, gonocoxite with a slender pointed caudomesally directed pro- tuberance, gonostylus shorter than gono- coxite, aedeagus with distal portion bearing 2 heavily sclerotized pointed sclerites that cross mesally, and parameres separate. Allotype female.—Head: Dark brown. Eyes pubescent, narrowly contiguous (V- shaped where contiguous). Antennal scape with 5 setae; flagellum brown; flagellomere 1 with 3 subapical sensilla coeloconica; lengths of flagellomeres in um 41-23-26- 26-26-28-28-28-38-34-34-38-62; antennal ratio 0.92. Clypeus with 1 pair of setae. Pal- pus brown; lengths of segments in um 15- 20-38-23-36; segment 4 with 2 setae. Man- dible with 10 teeth. Thorax: Dark brown; 5 prealar setae, 1 postalar seta; scutellum with 2 medial, 2 lat- eral setae. Wing (Fig. 12) membrane infus- cated, veins brown; macrotrichia sparse in cells r;, m,, m,, absent in cell cua, and anal cell; radial cells well developed, subequal sized, Ist with elongated narrow lumen, 2nd with broader triangular lumen, radial veins thickened especially distal of 2nd ra- dial cell; media with long petiole, branching slightly distal to level of R,,;, M, obsolete at extreme base; M,,, and Cu, branching at level of midlength of Ist radial cell; wing length 1.00 mm, breadth 0.46 mm, costal ratio 0.54. Halter pale. Legs brown; hind tarsal ratio 2.00 (n = 2); hind tibial comb with 7 setae; tarsomeres 5 slender with 151 moderately large, unequal sized claws with- out basal teeth, those of hind leg most mas- sive, claw length proportions of fore, mid, hind legs 9:7, 8:5, 6:4. Abdomen: Dark brown, segments 7—9 heavily sclerotized. Genitalia as in Fig. 13. Distal portion of sternite 8 and most of ster- nite 9 covered with spiculate membrane; sternite 9 halves represented by two stout plates. Two ovoid spermathecae with mod- erately short necks, plus rudimentary 3rd (Fig. 14), measuring 0.056 by 0.038 mm, neck 0.010 mm, and 0.052 by 0.036 mm, neck 0.010 mm. Holotype male.—Similar to female with the following notable sexual differences: Flagellum (Fig. 15) with flagellomeres 1—2 separated, 3-10 fused, 11—13 distinctly sep- arated, lengths of flagellomeres 11—13 in um 66-71-82; wing (Fig. 16) length 0.98 mm, breadth 0.38 mm, costal ratio 0.51. Genitalia as in Figs. 17—18. Sternite 9 twice as broad as long, caudal margin without ex- cavations; tergite 9 short, rounded, apico- lateral processes closely approximated near midline, with single subterminal seta; cer- cus short, with 3 apical setae. Gonocoxite elongate, 3 times longer than broad, meso- basal lobe very slender, sharply pointed, di- rected caudomesally; gonostylus slender with numerous small setae, distal half great- ly curved (nearly 90°), tip sharply pointed. Aedeagus with stout, low basal arch ex- tending % of total length; distal portion rep- resented by 2 heavily sclerotized pointed sclerites that cross mesally. Parameres (Fig. 18) separate, closely approximated distally; basal arms well developed, bilobate; distal portion stout, outer margin heavily sclero- tized, tip bluntly rounded. Distribution.—Southern Chile; known only from the type locality. Types.—Holotype d, allotype 2, Chile, Cautin, 1150 m, Conguillo Nat. Park. 4/5- II-1988, L. Masner (deposited in CNCI). Etymology.—The specific name is a pa- tronym in honor of the Chilean poet Pablo Neruda, who, in addition to his other mon- umental literary works, admirably de- 152 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 12-18. Borkenthelea nerudai. 12, Female wing. 13, Sternites 8 and 9 of female. 14, Spermathecae. 15, Male flagellum. 16, Male wing. 17, Genitalia, parameres removed. 18, Parameres. Scales = 0.05 mm. VOLUME 103, NUMBER 1 scribed the temperate forests of southern Chile in many of his books. Discussion.—The female of this new species differs from all other species in the genus in lacking macrotrichia in cell cua). The male of B. harii and B. quatei differ from this new species in having one or more caudomedian excavations on sternite 9. The male of B. nothofagus differs from this new species in having a gonocoxite with mesally directed mesal protuberance, a slightly curved gonostylus, parameres that are fused basally, and the distal portion of the aedeagus lacks pointed sclerites. Borkenthelea nothofagus Spinelli and Grogan Borkenthelea nothofagus Spinelli and Gro- gan 1993: 323 (2, 6; Argentina, Chile); Spinelli and Wirth 1993: 39 (Argentina; description; figs. d and ¢); Borkent and Wirth 1997: 91 (in world catalog). Diagnosis.—Distinguished from other species of Borkenthelea by the following combination of characters: female, large size (wing length 1.28—1.35 mm), clypeus with 2-3 pairs of setae, mandible with 9— 10 teeth, hind claws massive, sternite 9 halves in the form of 2 stout plates, sper- mathecae necks short, wing with lumen of 2nd radial cell short and central portion of cell cua, with macrotrichia; males, tergite 9 tapering distally, sternite 9 without caudo- median excavations, gonostylus longer than gonocoxite, aedeagus triangular, strongly sclerotized with a low basal arch and blunt tip, and parameres fused, stout. Distribution.—Argentina (west of Rio Negro province); Chile (Valdivia). Remarks.—Spinelli and Grogan (1993) gave the range of female wing lengths for this species as 1.00—1.35 mm. However, this lowest value was for one paratype from Cautin, Chile. Therefore, because this spec- imen is herein recognized as belonging to B. nerudai, the actual range of female wing lengths for B. nothofagus based upon avail- able specimens is 1.28—1.35 mm. 153 Borkenthelea quatei Spinelli and Grogan, new species (Figs. 19-25) Diagnosis.—Distinguished from other species of Borkenthelea by the following combination of characters: female, hind claws massive, mandible with 7 teeth, ster- nite 9 halves represented as 2 stout plates, spermathecae necks short, wing with lumen of 2nd radial cell elongated and central por- tion of cell cua, with macrotrichia; male, tergite 9 tapering distally, sternite 9 with 3 caudal excavations, gonostylus sickle- shaped; aedeagus heavily sclerotized with long basal arms and high basal arch, and parameres separate, stout. Allotype female-—Head: Dark brown. Eyes pubescent, contiguous for a distance equal to diameter of 2 ommatidia. Antennal scape with 2 setae; flagellum brown; fla- gellomere | with 3 subapical sensilla coe- loconica; lengths of flagellomeres in um 41- 28-3 1-28-31-31-31-33-36-36-36-41-66; an- tennal ratio 0.82. Clypeus with one pair of setae. Palpus brown; lengths of segments in um 18-28-31-23-28; segment 4 with 2 se- tae. Mandible with 7 small teeth. Thorax: Dark brown; prealar setae not visible; scutellum with 2 medial, 2 lateral setae. Wing (Fig. 19) membrane infuscated, veins brown; macrotrichia present in cells r;, M,, m5, cua,, only marginally in anal cell; radial cells well developed with broad lumen, Ist slightly longer than 2nd; radial veins thickened, especially posteriorly; me- dia with long petiole, branching slightly distad to level of R,,,, M, nearly contacting media, obsolete on extreme base; M,,, and Cu, branching at level of midlength of Ist radial cell; wing length 1.12 mm, breadth 0.52 mm, costal ratio 0.59. Halter pale. Legs brown; hind tibial comb with 8 setae; hind tarsal ratio 2.00; tarsomeres 5 slender with moderately large, unequal sized claws without basal teeth, those of hind leg most massive, claw length proportions of fore, mid, hind legs 8:4, 7:5, 10:7. Abdomen: Dark brown, segments 7—9 154 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 19-25. Borkenthelea quatei. 19, Female wing. 20, Sternites 8 and 9 of female. 21, Spermathecae. 22, Male flagellum. 23, Male wing. 24, Male genitalia. 25, Parameres. Scales = 0.05 mm. VOLUME 103, NUMBER 1 heavily sclerotized. Genitalia as in Fig. 20. Distal portion of sternite 8 and most of ster- nite 9 covered with spiculate membrane; sternite 9 halves represented by two stout plates. Two subequal sized ovoid sperma- thecae with slender offset necks plus rudi- mentary 3rd (Fig. 21), measuring 0.038 by 0.034 mm, neck 0.008 mm. Holotype male.—Similar to female with the following notable sexual differences: Flagellum (Fig. 22) with flagellomeres 1—2 separate, 3-10 fused, 11—13 distinctly sep- arated, lengths of flagellomeres 11—13 in um 105-71-112; wing (Fig. 23) length 1.11 mm, breadth 0.48 mm, costal ratio 0.53. Genitalia as in Figs. 24—25. Sternite 9 with 3 deep caudomedian excavations; tergite 9 short, tapering distally, apicolateral process- es blunt, each with a single, small seta; cer- cus elongated with 3 apical setae. Gono- coxite stout, twice as long as broad, without mesobasal lobe; gonostylus slender, greatly curved, sickle-shaped, surface with small setae especially near tip. Aedeagus heavily sclerotized; main portion small, triangular, basal arch extending % of total length, tip pointed; basal arms very long, straight. Par- ameres (Fig. 25) separate, heavily sclero- tized; each half with main portion stout, tips slightly curved outwards; basal apodemes slender on proximal %, broadening distally, tip obliquely truncated. Distribution.—Southwestern Argentina; known only from the type locality. Types.—Holotype <4, allotype °, Argen- tina, Chubut, P. N. Los Alerces, 9/12-XII- 1994, L. Quate, Malaise trap (deposited in MLPA). Etymology.—The specific name is a pa- tronym in honor of Dr. Larry Quate, Re- search Associate at the Natural History Mu- seum of Los Angeles County, who collect- ed most of the specimens described in this paper during our 1994 trip to Patagonia. Discussion.—Females of all other spe- cies in the genus differ from the female of this new species in having a mandible with 8—10 teeth. The female of B. harii is further distinguished from the female of this new 155 species in having non-massive hind claws, a pale halter, sternite 9 represented by two conspicuous plates, and spermathecae with short necks. The female of B. nerudai is further distinguished from the female of this new species in lacking macrotrichia in cell cua,. The female of B. nothofagus is further distinguished from the female of this new species in being larger (wing length 1.28-135 mm), having a clypeus with 2-3 pairs of setae, a larger antennal ratio (0.88—0.90), and the lumen of the 2nd radial cell is short and triangular. Males of B. nothofagus and B. nerudai differ from the male of this new species in lacking caudomedian excavations on ster- nite 9. The male of B. harii differ from the male of this new species in having sternite 9 with only a single caudomedian excava- tion, a gonocoxite with a mesal truncated protuberance, a gonostylus clearly shorter than gonocoxite, and the distal portion of the aedeagus has two pairs of lateral teeth and a bifid tip. ACKNOWLEDGMENTS We gratefully acknowledge grants to GRS from the National Geographic Society (NGS #5265-94) and the Eppley Founda- tion. GRS extends special thanks to Hari Bhat and Larry Quate for their companion- ship and assistance during the 1994 field trip to Patagonia. We also thank Art Bor- kent and Jean-Claude Delecolle for their helpful reviews of an earlier draft of the manuscript. LITERATURE CITED Borkent, A. and W. W. Wirth. 1997. World species of biting midges (Diptera: Ceratopogonidae). Bulle- tin of the American Museum of Natural History, No. 233, 257 pp. Downes, A. and W. W. Wirth. 1981. Chapter 28. Cer- atopogonidae, pp. 393—421. In McAlpine, J. F, B. iV. Beterson, Gaibs shewellSsbn ws Leskey. er Re Vockeroth, and D. M. Wood, eds., Manual of Ne- arctic Diptera, Vol. 1. Agriculture Canada Mono- graph, 27, 674 pp. Spinelli, G. R. and W. L. Grogan, Jr. 1993. Borkenthe- lea, a new predaceous midge genus from subant- arctic Argentina (Diptera: Ceratopogonidae). Pro- 156 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ceedings of the Entomological Society of Wash- ceous midges of the world (Diptera: Ceratopogon- ington 95: 321-326. idae; Tribe Ceratopogonini). Flora and Fauna Spinelli, G. R. and W. W. Wirth. 1993. Los Cerato- Handbook No. 4. E. J. Brill, New York, xv + 160 pogonidae de La Argentina. Vol. 38. Diptera. pp. Fasc. 3. Ceratopogonidae. Fauna de Agua Dulce Wirth, W. W. and N. Marston. 1968. A method for de La Republica Argentina. Museo de La Plata, mounting small insects on microscope slides in La Plata, Argentina, 124 pp. Canada balsam. Annals of the Entomological So- Wirth, W. W. and W. L. Grogan, Jr. 1988. The preda- ciety of America 61: 783-784. PROC. ENTOMOL. SOC. WASH. 103(1), 2001, pp. 157-183 THE SPECIES OF XANTHONIA BALY 1863 (COLEOPTERA: CHRYSOMELIDAE: EUMOLPINAE) IN NORTH AMERICA EAST OF THE MISSISSIPPI RIVER C. L. STAINES AND D. M. WEISMAN (CLS) 3302 Decker Place, Edgewater, Maryland 21037, U.S.A. (e-mail: staines. charles@nmnh.si.edu); (DMW) Systematic Entomology Laboratory, PSI, Agricultural Re- search Service, U.S. Department of Agriculture, c/o National Museum of Natural History, Washington DC 20560-0168, U.S.A. (deceased) Abstract.—The genus Xanthonia Baly is revised for eastern North America, with nine species. The species X. decemnotata (Say) and X. villosula (Melsheimer) are redescribed; X. stevensi Baly is removed from synonymy with X. villosula. Neotypes are designated for X. decemnotata, X. stevensi, and X. villosula. The new species X. angulata from Maryland to Kansas, X. furcata from Illinois and Missouri, X. intermedia from Alabama and Iowa, X. monticola from the mountains of New York and North Carolina, X. serrata from Massachusetts to Louisiana, and X. striata from Massachusetts to Texas are described and a key to the species is presented. Key Words: The genus Xanthonia was erected by Baly (1863) for the new species stevensi from Canada. Baly (1874) described placi- da from Japan. Horn (1892) transferred Co- laspis decemnotata Say, Eumolpus villosu- lus Melsheimer, and Tricotheca vagans LeConte to Xanthonia and synonymized X. stevensi and Myochrous plagiata Melshei- mer with X. villosula. Jacoby (1882) de- scribed guatemalensis, nigrofasciata, and plagiata from Guatemala, marmorata from Mexico and Panama, and tuberosa from Mexico. Clavareau (1914) renamed plagia- ta Jacoby jacobyi because the name was preoccupied by plagiata Melsheimer. Schaeffer (1933) described pinicola from Arizona. Chen (1935) described signata and sinica from China. Chen (1940) de- scribed collaris and varipennis from China and transferred minuta Pic from Aoria. Blake (1954) described flavoannulata from Mexico. Bechyné (1955) described umbili- Coleoptera, Chrysomelidae, Eumolpinae, Xanthonia, new species cata from Mexico. Kimoto and Gressitt (1982) described morimotoi from Thailand and synonymized Microlypesthes Pic with Xanthonia which moved coomani Pic into Xanthonia. Takizawa (1987) described ful- va and nepalensis from Nepal. Takizawa and Basu (1987) described oblonga from India. The entire genus has never been revised. Horn (1892) presented a key to three North American species. Gressitt and Kimoto (1961) presented a key to five species from China. Kimoto and Gressitt (1982) present- ed a key to three species from southeast Asia. This project was begun as a Master of Science thesis by Weisman (1960). The ma- terial was brought to the Systematic Ento- mology Laboratory, USDA, when Weisman accepted a position there. Due to a number of factors Weisman never completed the re- vision and the material remained in the col- 158 lection until I became interested in the group. This work is based on Weisman’s unpublished thesis with additional material identified by myself. Also the nomenclature is updated to fulfill the requirements of the current Code (ICZN 1999). There is little published biological infor- mation on the genus. Adults of Nearctic species have been collected in association with a wide variety of plants but the actual larval host plant has not been determined for any species. Due to the number of new species, the reported plant associations are of questionable value. Packard (1890) re- ported X. stevensi from. Carya (Juglanda- ceae). Hamilton (1895) and Felt (1906) re- ported X. decemnotata on Quercus (Faga- ceae); Chagnon (1937) reported it from Quercus and Fagus; Blatchley (1910), Craighead (1950), and Wilcox (1954) re- ported it from Quercus, Fagus, and Ulmus (Ulmaceae); Phipps (1926) reported it from Malus, Rubus (Roseaceae), and Tilia (Ti- laceae); Wilcox (1979) reported it from Fa- gus grandifolia Ehrh., Hamamelis sp. (Ha- mamelidaceae), and Ulmus sp.; Dearborn and Donahue (1993) reported it from Abies sp. (Pinaceae), Betula sp. catkins (Betula- ceae), Picea sp., and Tsuga sp. (Pinaceae). Stauffer (1865) reported X. villosula dam- aging Vitis (Vitaceae); Douglass (1929), Chagnon (1937), and Wilcox (1954) re- ported it from Quercus; Hamilton (1895) and Blatchley (1910) reported it from Quer- cus and Corylus (Betulaceae); Wellhouse (1922) reported it from Crataegus (Rosea- ceae); Blatchley (1924) reported it from Myrica (Myrtaceae); Felt (1906) reported it from Quercus, Carya, Corylus, and Popu- lus (Salicaceae); Wellhouse (1919) reported feeding on Carya ovata (Mill.) K. Koch, Crataegus punctata Jacq., Tilia americana L., Quercus alba L, Vitis riparia Michx., Rubus sp., Rosa sp., Carpinus caroliniana Walt., Ostrya virginiana (Mill.) K. Koch (Betulaceae), Acer saccharum Marsh (Ac- eraceae), Hamamelis virginiana L., Wald- stenia fragarioides (Michx.) Trattinick, and Fragaria virginiana Duch. (Roseaceae). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Schaeffer (1933) reported X. pinicola beat- en from Pinus. Blake (1954) noted that X. flavoannulata was associated with various Orchidaceae in Mexico. Moldenke (1971) reported X. tuberosa associated with Quer- cus purulbana Trel. Gressitt and Kimoto (1961) mentioned that X. signata was as- sociated with Viburnum spp. (Caprifoli- aceae). Wellhouse (1919) described the eggs of X. villosula and found that they were laid in masses in moist earth just be- low the surface of the soil. Phipps (1926) found that adult X. decemnotata fed active- ly at night but hid in rolled leaves during the day. Riley and Enns (1979) reported taking X. decemnotata and X. villosula at light. Lee (1993) obtained larvae of X. pla- cida by rearing adults on leaves of Morus bombycis Koidz. (Moraceae). For this study, measurements were taken with an ocular micrometer. Total length was measured from the base of the antennae to the apex of the elytra. In recording label data from type specimens, a slash (/) di- vides data on different labels. Specimen de- pository is noted in parentheses, and acro- nyms are given in Acknowledgments. Brackets ([{]) contain explanatory notes. Xanthonia Baly Xanthonia Baly 1863: 151 [Type species: Xanthonia stevensi Baly (by monotypy)]. Crotch 1873a: 96; Chapuis 1874: 273; Ja- coby 1882: 164; LeConte and Horn 1883: 343; Lefévre 1885: 74; Horn 1892: 199; Wickham 1896: 172; Blatchley 1910: 1142; Clavareau 1914: 75; Chen 1935: 359; Chagnon 1937: 250; Chen 1940: 491; Bechyné 1950: 288; Monroés 1952: 189; Bechyné 1953: 248; Wilcox 1954: 401; Chaj6 1956: 108; Seeno and Wilcox 1982: 61; Flowers’ 1996252: Microlypesthes Pic 1936: 15. Type species: M. coomani Pic (by monotypy). [Syn- onymized by Kimoto and Gressitt 1982]. Body oblong, subcylindrical; covered with fine pubescence or scales. Head: Exserted, rounded, perpendicular; VOLUME 103, NUMBER 1 covered with coarse punctures; eyes round- ed, entire, convex; front usually with im- pressed medial line; mentum with anterior margin concave; labrum slightly emargin- ate; last segment of maxillary palpus ovate- acuminate. Antenna: Half body length; slender, fili- form; segment I incrassate; II thickened, oval, shorter than I; III and VI subequal in length, slender, each longer than I; VII to XI slightly shorter than III, slightly thick- ened; XI oval, short. Pronotum: Subcylindrical, transverse, widest point behind middle; slightly flat- tened; lateral margins obsolete; sides broad- ly rounded; longitudinal sulcus present be- hind anterior margin; large impression pres- ent at middle of each side; surface covered with coarse punctures with a seta arising from each puncture; pubescence varying from long silky to short recumbent. Scutellum: Trapezoidal. Elytra: Wider than base of pronotum; hu- meri prominent and with an obscure to prominent callosity at middle of base and an interhumeral sulcus present between the callosity and humerus; surface slightly de- pressed adjacent to scutellum; lateral mar- gins parallel; surface densely punctured, ar- rangement varying from completely con- fused to regularly seriate on disc, punctures always confused near scutellum; intervals completely flat or tending to become cos- tate especially at sides and apex; pubes- cence with setae arising both in punctures and on intervals, setae always appressed and varying in length from short and in- conspicuous to the same length as the in- terval setae; interval setae usually uniseriate on each interval but denser in some species, varying from erect to strongly inclined; api- ces broadly rounded. Venter: Prosternum oblong-elongate, narrow or straight, convex between coxae, depressed before and behind, anterior mar- gin continuous with episternum; mesoster- num oblong; metasternum with parapleura linear, obtuse at apex, with coarse punctures at sides; abdomen with coarse punctures on 159 first and fifth segments, each segment with shallow impression at each side near mar- gin; lateral margins usually entire but sterna 3 to 5 may be irregularly notched or serrate in some species; last sternite of male oc- casionally with tubercle on each side of middle. Legs: Femora slightly thickened, profe- mur usually with ventral tooth, meso- and metafemora with ventral tooth present; tib- iae simple; tarsi with segment 1 shorter than 2 or 3; claws bifid, inner tooth incurved, shorter. Notes.—In North America, Xanthonia can be distinguished from all other Eumol- pinae by the following combination of char- acters: anterior margin of pronotum straight beneath, not forming ocular lobes; prono- tum without distinct lateral margins; body dorsally without metallic colors; head with- out supra-orbital groove; eyes rounded (not emarginate); antennal segment three not longer than two; pronotum transverse; and body covered with scales or dense pubes- cence. The genus most similar to Xanthonia is Fidia Walsh 1867. Fidia differs by having a cylindrical pronotum and antennal seg- ment three longer than two. KEY TO THE XANTHONIA OF NORTH AMERICA EAST OF THE MISSISSIPPI RIVER 1. Elytral punctures confused on disc, tending to form rows only at suture and along lateral mar- UTS) |W eyiagerie eats eaeaere or omen lc etey Oke UR eee 2 — Elytral punctures in rows on disc, punctures are confused near scutellum, behind sulcus and be- LOW INUINERUS sete esac i cgclckn San ae aa ae 3 i) . Abdomen with lateral margins of segments 4 and 5 serrate (Fig. 8); male genitalia as in Fig. 13c; Massachusetts to Louisiana ....... serrata, new species — Abdomen with lateral margins of all segments entire (Fig. 3); male genitalia as in Fig. 12b; Nova Scotia to South Carolina, west to Loui- siana and Montana decemnotata (Say) 3. Abdomen with lateral margins of some seg- ASUS KOA! OP SEAS “Seg odougccacan 4 — Abdomen with lateral margins of all segments SNUIMe: De. war ditste eer ae eeeeah oe ie 5 4. Pronotum widest at middle; elytral punctures smaller than width of an interval; male geni- 160 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON talia as in Fig. 15b; Ontario to South Carolina, west to Texas and Yukon — villosula (Melsheimer) — Pronotum widest behind middle; elytral punc- tures as wide or wider than the width of an interval; male genitalia as in Fig. 13a; Ala- bama, Iowa intermedia, new species 5. Pronotum widest at middle, rounded....... 6 — Pronotum widest behind middle, angulate ... 7 6. Punctures on head and pronotum very coarse (Fig. 6); male genitalia as in Fig. 13b; moun- tains from New York to North Carolina .... Ree Pee Cera Groner ae tosr ae monticola, new species — Punctures on head and pronotum moderately coarse (Fig. 9); male genitalia as in Fig. 14; Quebec to North Carolina and Nebraska .... SE ee ce ioe tk ah ie ee stevensi Baly 7. Elytra with long setae in single lines on every other interval and in each puncture; male gen- italia as in Fig. 12c; Illinois and Missouri . . . furcata, new species — Elytra with long setae on each interval, setae in punctures inconspicuous 8. Pronotum with lateral margins slightly rounded and converging apically; male genitalia as in Fig. 15a; Massachusetts to Texas Se el iy wae, Am to os ea aan striata, new species — Pronotum with lateral margins almost straight and converging apically; male genitalia as in Fig. 12a; Maryland to Kansas angulata, new species Xanthonia angulata Staines and Weisman, new species (Figs. 1, 12a) Holotype.—d, Glen Echo, VI-28-—30, Md/J. C. Bridwell coll./Holotype Xanthonia angulata Staines and Weisman [red label] (USNM). Allotype.—?, Glen Echo Md/JCBrid- well coll. VII-14-30/Allotype Xanthonia angulata Staines and Weiseman [red label] (USNM). Paratypes.—19, each with Paratype Xan- thonia angulata Staines and Weisman [red label]: 1 3: Glen Echo, 23-6-29 Md/Brid- well Collector (USNM); 1 ¢: Glen Echo Md summer 1922, J. C. Bridwell (USNM); 1 3: Glen Echo, VI-28-30 Md/JCBridwell coll. (USNM); 1 d6: Montg. Co. Md, VI- 15-1909/Ernest Shoemaker Collection (USNM); 1 6, 3 @: Washgton DC (USNM); 1 2: Rock Creek, July 13, 1902, Washington DC (USNM); 1 6: Mt. Vernon Va, VI-20-20/Ernest Shoemaker Collection (USNM); 1 ¢: Gt. Falls Va, VI-14-13 (USNM); 2 6: Knoxville, Tenn/G. M. Bentley Coll./June 12, 1916/Quercus/99151 Hopk US (USNM); 1 2: C. Mo./June/Col- lection C. V. Riley (USNM); 1 2: C. Mo/ collection C. V. Riley (USNM); 1 @: Eldon Mo., V-28-1939, E. O. Essig (SEMC); 1 d: Elk City, Kan., 4-2-36, Beamer and San- derson (SEMC); | 2: Ames, Iowa, 7-10- 32, EF Andre (USNM); 1 ¢: Floyd Co., Ind., 6-23-06/6721/Collection E Knab (USNM). Description.—Head: Dark reddish brown; labrum and antenna yellowish brown; densely and coarsely punctate, with short golden appressed pubescence; front with medial line slightly impressed; inter- antennal area moderately concave to almost flat. Pronotum: Dark reddish brown to fus- cous, occasionally with lighter apical and basal margins; densely, coarsely punctate; with fine, dense golden pubescence; width 1.4 times its length; widest just behind mid- dle; sides in dorsal aspect angulate behind middle, almost straight and converging api- cally, strongly rounded behind; anterior transverse depression and lateral impres- sions shallow to obscure. Elytra: Reddish brown; length averaging 1.65 times width, 2.7 times length of pron- otum; basal callosity evident; punctures in regular rows, confused only near scutellum, behind sulcus, and below humeri; punctures nearly as large as width of an interval on disc; intervals costate, especially laterally and at apex; pubescence composed of erect setae in single lines on intervals and very short, obscure setae in punctures. Venter: Dark reddish brown to fuscous; metasternum with sparse moderately coarse punctures on lateral margin; abdomen with- out coarse punctures, with sparse fine punc- tures on last segment; all segments with lat- eral margins entire; last segment of male without large tubercles. Legs: Yellowish brown; all femora with ventral tooth obsolete to absent. Male gen- italia as in Figure 12a. Total length: 3.5 to 4.0 mm. VOLUME 103, NUMBER 1 161 Figs. 1-6. Xanthonia species. 1, Dorsal view of Xanthonia angulata. 2, Dorsal view of X. decmnotata. 3, Abdomen of X. demnotata. 4, Dorsal view of X. furcata. 5, Dorsal view of X. intermedia. 6, Dorsal view Ohexe monticola. Etymology.—From angulatus (Latin), with angles, for the angulate lateral margins of the pronotum. Notes.—This species closely resembles striata but can be distinguished by the more shallow aedeagal emargination, by the di- vergent apices of the aedeagus, and by the more angulate pronotal lateral margins. Plant associations.—Specimens have been collected from Quercus. Distribution DISTRICT OF COLUM- BIA. INDIANA: Floyd Co. IOWA: Story Co.: Ames. KANSAS: Montgomery Co.: Elk City. MARYLAND: Montgomery Co.: Glen Echo. MISSOURI: C. Mo.; Eldon. TENNESSEE: Knox Co.: Knoxville. VIR- 162 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 7-11. Xanthonia species. 7, Dorsal view of X. serrata. 8, Abdomen of X. serrata. 9, Dorsal view of X. stevensi. 10, Dorsal view of X. striata. 11, Dorsal view of X. villosula. GINIA: Fairfax Co.: Great Falls; Mount Md., Oct. 4 14/W L McAtee Collector/ Vernon. Specimen No. 296/NEOTYPE Xanthonia decemnotata (Say) des. C. L. Staines Xanthonia decemnotata (Say) 1999 (USNM)]. (Figs. 2, 3, 12b) Pachnephorus decemnotata: Melsheimer Colaspis decemnotata Say 1824: 445 [neo- LSS35126: type & (here designated): Plummers I., Xanthonia decemnotata: Crotch 1873a: 96, VOLUME 103, NUMBER I 1873b: 34; Gemminger and Harold 1874: 3376; Henshaw 1874: 23; Hubbard and Schwarz 1878a: 640, 1878b: 660; Dury 1879: 172; Harrington 1884: 81; Lefévre 1885: 74; Wickham 1888: 89; Horn 1892: 200; Osborn 1892: 34; Hamilton 1895: 340; Davis 1902: 37; Dury 1902: 165; Ulke 1902: 28; Wickham 1902: 286; Fall and Cockerell 1907: 196; Easton 1909: 53; Blatchley 1910: 1142; Wick- ham 1911: 31; Clavareau 1914: 75; John- son 1915: 316; Chagnon 1917: 240; Leng 1920: 292; Schaeffer 1928: 467; Doug- lass 1929: 11; Johnson 1930: 62; Brimley 1938: 224; Johnson 1941: 99; Hughes 1944: 133; Loéding 1945: 129; Fattig 1948: 19; Wilcox 1954: 401; Dillon and Dillon 1961: 680; Kirk 1969: 92, 1970: 88; Balsbaugh and Hayes 1972: 72; Rouse and Medvedev 1972: 79; Kirk and Balsbaugh 1975: 108; Wilcox 1975: 57; Riley and Enns 1979: 65; Wilcox 1979: 12, 19, 24; LeSage 1991: 309; Dearborn and Donahue 1993: 64; Downie and Ar- nett 1996201835: Description.—Head: Dark reddish brown with vertex darker to black, labrum and basal segments of antenna yellow, api- cal segments reddish brown; densely, coarsely punctate; with silky, golden, close- ly appressed pubescence; front with medial line slightly evident; interantennal area flat. Pronotum: Dark reddish brown with api- cal margin lighter and with very indistinct black macula on each side of middle at base; densely, coarsely punctate; with long, silky, golden, closely appressed pubes- cence; width averaging 1.25 times its length; widest at middle; sides evenly, broadly rounded; anterior transverse de- pression and lateral impressions deep to shallow. Elytra: Reddish brown, each elytron with a black macula at base, at humerus, a linear dorsal one near suture, a row of three across disc, and row of three across apical third, discal and subapical maculae tend to unite transversely; length averaging 1.55 times 163 width, 2.6 times length of pronotum; basal callosity slightly prominent; punctures completely confused on disc and base, tend- ing to form rows only at posterior portion of suture and along lateral margins; punc- tures dense, coarse, and separated by less than the diameter of a puncture; intervals at lateral margins costate; pubescence com- posed of stiff, erect setae scattered between punctures and appressed setae as long as erect setae in punctures. Venter: Black with apex of abdomen lighter; metasternum with very coarse punctures laterally; abdomen with very shallow punctures on last segment, inter- coxal lobe rugose, all segments with lateral margins entire, last segment of male with- out large tubercles. Legs: Femora dark reddish brown, inner surface of meso- and metafemora almost black, tibiae lighter reddish-brown; profe- mur with ventral tooth very small to obso- lete, meso- and metafemora with ventral tooth small to very small. Male genitalia as in Figure 12b. Total length: 3.0 to 3.5 mm. Notes.—This species was described as ‘“‘inhabitating the United States.’ The pre- sent interpretation of decemnotata is based upon the original description which very clearly defines this species. We have been unable to locate the Say type in the Muse- um of Comparative Zoology (Perkins, per. comm.) and presume it to be lost and here- by designate a neotype (Article 75, ICZN 1999). Specimens examined.—CANADA: MANITOBA: Aweme, 16.VII.1923, 30.V1I.1922 (CNC); Brandon (CNC); Ri- ding Mts., May 1912 (CNC); Treesbank, VI-11-14 (CNC); Winnipeg (CNC). NEW BRUNSWICK: Penobsquis, VII-29-26 (CNC). NOVA SCOTIA: Bridgetown (USNM); Digby Co., 2 Sep 18 (USNM). ONTARIO: Almonte, IX-9-1928 (CNC); Arnprior, VIII-28-1938 (CNC); Barry’s Bay; Bell’s Corner, 8-V-1951 (CNC); Consecon, 26-VIII-03 (CNC); Fitzory Harbor, 11-VII- 1938 (CNC); Hastings Co. (CNC); Marmora, 11-8-1952, 21.1V.1952, 14.VIII.1952 (CNC); 164 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Ottawa (CNC); Ridgeway, May 31, 1886, June 25, 1885, 7-5-08 (CASC); Timagami, 20-VITI-1932 (CNC); Toronto (CNC); Tren- ton, 4-IX-10 (CNC). QUEBEC: Aylmer, 25-5-01 (CNC); Burbridge, 27-5-1937 (CNC); Gatineau Park, VIII-11-1957 (CNC); Gracefield, 25-VI-1937 (CNC); Montreal Island (CNC). SASKATCHE- WAN: Oxbow (USNM); Pike Lake, 24-V- 1940 (CNC); Winnipeg (USNM). UNITED STATES: ALABAMA: ?: Barachias, June 9, 1924 (USNM). ARKANSAS: Benton Co.: Bentonville, 21-VI-31 (USNM). CONNECTICUT: (CASC). Fairfield Co.: Stamford (AMNH); Danbury (USNM). Hartford Co.: New Britain (USNM). Litch- field Co.: Colebrook, 21-V-1911, 23-VI- 1914 (AMNH); Litchfield, VI-29-20, VIII- 22-1925, VI-26-1925, VI-27-14, IX/4/11 (AMNH). Middlesex Co.: Killingsworth (USNM). Tolland Co.: Union, 2-IX-952 (USNM). DISTRICT OF COLUMBIA: (USNM); V-19-1956, 18/6, 18-11 (USNM); Rock Creek Park, Apr. 15 ‘07, May 27, 1906 (USNM). ILLINOIS: (USNM). ?: Chicago, 6-19-1920 (USNM); N. Ill. (USNM); Ravinia, VI.25.11, VIII.31 (USNM). Champaign Co.: Urbana, 26 Apr. 1947 (AMNH). Lawrence Co.: V-28-38 (CASC). McLean Co.: Heyworth, VII-31 (USNM). IOWA: Clayton Co.: Guttenberg, 15.XI (USNM). Decatur Co.: Leon, 8-20- 32 (USNM). Dickinson Co.: Lake Okoboji, VII-3-16 (USNM). Howard Co.: Elma, VIII-6-02 (USNM). Johnson Co.: Iowa City, 6.6.1896, V-4, 10-16-15, III-30-189, V-23 (USNM). Mahaska Co.: New Sharon, 8-5-32 (USNM). Monona Co.: Solon, 10-8- 15 (USNM). Pottawattamie Co.: Council Bluffs, XI-19 (USNM). Story Co.: Ames, Sept. 29, 1888 (USNM). Winneshiek Co.: Decorah, XI-19 (USNM). KANSAS: Doug- las Co.: 900 ft. (USNM). Pottawatomie Co.: Onago, June 16, 1901 (USNM). Riley Co.: May 5 (USNM). Shawnee Co.: Topeka, Apr. 27, Apr. 23, Jun. 12 (USNM). KEN- TUCKY: Fulton Co.: Fulton (USNM). LOUISIANA: (USNM). Lincoln Parish: Ruston, HI-23—08 (USNM). MAINE: Ken- nebec Co.: Augusta (UAIC). Penobscot Co.: Orono, 5-1-49, 5-11-49 (UAIC). MARYLAND: Baltimore: VI-24-09, VII- 22, VII-21 (CASC). Montgomery Co.: XI- 21-1912 (USNM); Glen Echo, summer 1922 (USNM); Plummers Island, 3-VI-06, 24-5-08, 30-5-07, Sept. 14, 13, V-25-18, July 20, 26, Oct. 4, 14, 10-6-06, 21 May ‘26, 22.1V.03, 6.9, 3.V.04 (USNM). Prince George’s Co.: Beltsville, IV-18-1920 (USNM). MASSACHUSETTS: (AMNH). ?: Blue Hills. Berkshire Co.: Lenox, June 23, 1891 (AMNH); Mt. Tom, 30 May 95 (USNM); Monterey, July 14, 39 (USNM). Essex Co.: Brookline (AMNH); Cambridge (USNM). Franklin Co.: Sunderland, 8-VII- 18 (CASC). Hampden Co:. Chicopee, May 17, 47 (USNM); Holyoke, Aug. 29, 98 (USNM); Springfield, 28 June 1903 (USNM); West Springfield, 3 May 1903, June 12, 96 (USNM). Norfolk Co.: Natick, XI-24-31 (UAIC). MICHIGAN: (USNM). ?: Detroit (USNM). Cheboygan Co.: 7-30- 39 (SEMC). Easton Co.: East Lansing, 6 May 1940, 7 May 1940, 5-28-90 (UAIC). St. Joseph Co.: Three Rivers (USNM). Washtenaw Co.: Ann Arbor (USNM). MINNESOTA: ?: Alexandria, July 5, 1939, June 30, 1939, 28 June 1940, Aug. 11, 39 (USNM). Saint Louis Co.: Duluth (SEMC). MISSOURI: Adair Co.: Thousand Hills State Park, 13 Jun. 74 (USNM). Boone Co.: Ashland Wildlife Area, 23 Sept. 1977; Co- lumbia, 8 May 78, 25 May 78 (USNM). Randolph Co.: Rudolph Bennett Wildlife Area, 21 May 78 (USNM). St. Louis Co.: Jefferson Barracks, IV-14-1900 (CASC). Stoddard Co.: 3.30.38, Nov. 6, 37 (SEMC). MONTANA: ?: Assiniboine, 21.8, 29.8, 23.8 (USNM). NEBRASKA: Cuming Co.: West Point, 8-87, 4-88 (USNM). NEW HAMPSHIRE: Carroll Co.: North Conway, [X-20-12 (USNM). Grafton Co.: Franconia (AMNH); Hanover (USNM). NEW JER- SEY: (USNM). ?: Big Timber Creek, 11.19.1900 (USNM); Da Costa, VI-30 (CASC); Greenwood Lake, V-18-1919 (USNM); Split Rock, IV-21-01 (USNM). Bergen Co.: Ramsey, [X-28-13 (USNM). VOLUME 103, NUMBER 1 Gloucester Co.: Newfield, IV-30-04 (USNM); Malaga, X-12-14 (USNM). Hun- terdon Co.: Hamden, V-18-14, V-18-10 (AMNH). Morris Co.: Boonton, VI.6.01, VI.02, VI.1.01 (USNM); Chester, VII-5 (AMNH). Passaic Co.: Newfoundland, V- 28-10 (AMNH); Oak Ridge, VI-26-1904 (USNM). Warren Co.: Phillipsburg, VI-19- 14 (CASC). NEW YORK: (USNM). ?: Por- tage, May 30, 88 (CASC); Trenton (USNM). Chenango Co.: Guilford, VII-9- 1925 (USNM). Erie Co.: Colden, 6-7-08 (CASC). Greene Co.: 2500 ft., Oct. 4, 1899 (USNM); New Baltimore, 1880 (AMNHBH). New York City: (USNM); Brooklyn (AMNH); Central Park, V-23—15 (AMNH); Flatbush, 12-V-1894, 30-IV-1894 (AMNH). Orange Co.: West Point, June 9, 1910, May 23. 1910) May 20; 1910; June 13; 1912; June Wi 1911s) May “25, 1908, Apr. 27, PT Maye 10; 1902, Sept. 30> 1909 (USNM). Niagara Co.: Buffalo, June 1896 (CASC); Niagara Falls, 10-15-11 (CASC). Oneida Co.: Remsen, 7-25-1946 (SEMC). St. Lawrence Co.: Rossie, VIII.26.1955 (USNM). Schenectady Co.: Schenectady, VI-19, 26-VIII-O9 (USNM). Suffolk Co.: Yaphank, V-27-17 (AMNH). Tompkins Co.: Ithaca, Mar. 04, May 7, 1939, 17 May 1938, 2 May 1937, 27 May 34, 2 June 1936 (USNM); McLean Bogs Res. (USNM). UI- ster Co.: (USNM); Oliverea, VI-18-18 (USNM). Washington Co.: Lake George, 25: ViIT1894, 30:8.1893, July 27, 14 (AMNH). Wyoming Co.: Pike, 6-4-1900 (USNM). NORTH CAROLINA: ?: Gray- beard Mt., VI-17, V-26 (AMNH). Avery Co.: VI-15-1955 (USNM). Henderson Co.: Black Mountains, VI-VII 1902, [X-24, IX- 20-01 (AMNH, CASC). Jackson Co.: Bal- sam, 20.1V.1955 (USNM). Lincoln Co.: Lincoln, 1 August 1995 (NCSU). Rich- mond Co.: Rockingham, IV-22-1955 (USNM). Surry Co.: Doughton State Park, VI-2-1955 (USNM). Wake Co.: Raleigh, IV-3-1955 (USNM); Schenk Forest, 16- VII-1991 (NCSU). Yancy Co.: Mt. Mitch- ell, 6200 ft., VI-20-1937 (USNM). OHIO: Franklin Co.: Columbus, VI-11, V-30 165 (CASC). Ross Co.: (USNM). PENNSYL- VANIA: (AMNH). ?: Ashbourne, 4-19-17 (USNM); Spring Brook, V-9-45 (USNM); Water Gap (AMNH). Dauphin Co.: VI-20- 96 (USNM). Monroe Co.: Canadensis, VII- X-1926, IX-21-1926 USNM). SOUTH CAROLINA: Florence Co.: Florence, 7 Dec. 1937 (USNM). TENNESSEE: ?: Great Smoky Mts. National Park, V-18- 1957 (CMNC). Knox Co.: Knoxville, VII- 1956 (CMNC). Putnam Co.: Clarksville, 4- 7-08, 4-20-09 (USNM). Smith Co.: Elm- wood (CASC). VERMONT: Lamoille Co.: Stowe, VI-23-13 (USNM). Windham Co.: Brattleboro, Spring 94 (USNM). VIRGIN- IA: ?: Middle Mt., 4000 ft., June 14, 39 (USNM). Arlington Co.: Glencarlyn, Sept. 27, 1914, 1-V-10 (USNM). Augusta Co.: Mt. Elliott, 20-6-34 USNM). Fairfax Co.: Great Falls, June 8, 22, June 30, 1914 (USNM); Vienna, 30-6-43 (USNM). Rock- bridge Co.: Buena Vista, Jul. 26, 1953 (USNM). WEST VIRGINIA: Greenbrier Co.: White Sulphur Springs, July 5, 1910, July 3, 1912 (USNM). Taylor Co.: Grafton (USNM). WISCONSIN: Polk Co. (USNM): July. Total: 585. Xanthonia furcata Staines and Weisman, new species (Figs. 4, 12c) Holotype.—¢d, Union Co., Ill., Jun. 3, *31, 37-21148/wild cherry foliage, W. E Turner, T-5048/Xanthonia sp. HSB ‘37/Ho- lotype Xanthonia furcata Staines and Weis- man [red label] (USNM). Allotype.—, same label data as holo- type/Allotype Xanthonia furcata Staines and Weisman [red label] (USNM). Paratypes.—1 3d, 2 2 with same label data as holotype/Paratype Xanthonia fur- cata Staines and Weisman [red label] (USNM); 3 6, 2 2 Reformed, Missouri, Callaway County/Coll. E. G. Riley, 2 June 1975/Paratype Xanthonia furcata Staines and Weisman [red label] (EGRC). Description.—Head: Reddish brown, labrum and antenna yellow; densely, coarsely punctate; with silky, ashen, re- 166 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 12. Xanthonia spp., dorsal and lateral views of aedeagi. a, X. angulata, b, X. decemnotata. c, X. furcata. cumbent pubescence; front with medial en, recumbent pubescence; width averaging line very weak to completely absent; in- 1.2 times length; widest behind middle; terantennal area flat. sides angulate behind middle, slightly Pronotum: Uniformly reddish brown; rounded and converging apically, strongly densely, coarsely punctate; with silky, ash- rounded behind; anterior transverse depres- VOLUME 103, NUMBER 1 sion very shallow, lateral impressions ob- scure. Elytra: Reddish brown each elytron with dark reddish-brown to blackish macula at middle of base, one on humerus; one on middle of disc, one at apical third, and with a common medial macula in front of mid- dle, markings occasionally indistinct; length averaging 1.35 times width, 2.45 times length of pronotum; basal callosity slightly prominent; punctures in irregular rows which tend to double, confused near scu- tellum, behind sulcus, and below humerus; diameter of a puncture as large as width of an interval on disc; intervals nearly flat on disc, becoming costate on lateral areas and at apex; pubescence composed of erect, stiff setae in single rows on every other interval and appressed setae in punctures nearly as long as setae on intervals. Venter: Dark reddish brown along mid- line, becoming lighter toward apical ab- dominal segments and darker toward lateral margins; metasternum with small punctures at lateral margins; abdomen with a few moderately coarse punctures on intercoxal lobe and on middle of last segment, all seg- ments with lateral margins entire, last seg- ment of male without large tubercles. Legs: Yellow; femora darker to fuscous on inner surface and apex; ventral tooth ab- sent on all femora. Male genitalia as in Fig- ure 12c. Total length: 2.8 to 3.5 mm. Etymology.—From furca (Latin), fork; for the sharply pointed apices of the aedea- gus. Notes.—This species can be distin- guished from all other North American spe- cies by its very distinctive aedeagus which has sharply pointed apices and an acutely angled apical emargination. Plant associations.—Specimens have been collected from foliage of Prunus spp. Distribution.—ILLINOIS: Union Co. MISSOURI: Callaway Co. Xanthonia intermedia Staines and Weisman, new species (Figs. 5, 13a) Holotype.—d, Landae, Chambers Co. Alab/HHSmith coll./Holotype Xanthonia 167 intermedia Staines and Weisman [red label] (USNM). Allotype.—, same label data as holo- type/Allotype Xanthonia intermedia Staines and Weisman [red label] (USNM). Paratypes.—10, each with Paratype Xan- thonia intermedia Staines and Weisman [red label]: 4 d, 3 9: same label data as holo- type; 1 d: Boone Co., Iowa, 4-VII-32/ County #47, IOWA, July 4, 1932, Russell (USNM); 1 3: Wankon, Iowa, VII-2-’ 14, Stoner (USNM); 1 6: Iowa City, Iowa, VII- 12-’10, Stoner (USNM). Description.—Head: Light reddish brown, labrum and antenna yellowish brown; densely, coarsely punctate; with golden, closely appressed pubescence; front with medial line slightly impressed; inter- antennal area varying from moderately to slightly concave. Pronotum: Light reddish brown; densely, coarsely punctate; with short, golden, re- cumbent pubescence; width averaging 1.4 times length; widest behind middle; sides fairly evenly, broadly rounded; anterior transverse depression moderately deep to shallow; lateral impressions obscure. Elytra: Yellowish brown with reddish brown common medial macula, each ely- tron with reddish brown macula at middle of base, one on humerus, a row of three across disc and three across apical third, discal and apical maculae tend to unite lon- gitudinally; length averaging 1.3 times width, 2.85 times length of pronotum; basal callosity slightly prominent; punctures in fairly regular rows, confused near scutel- lum, behind sulcus, and below humerus; punctures as large or larger than width of intervals on disc; intervals costate at lateral margins and apex, often with two costae on each disc; pubescence composed of erect setae in single lines on intervals and an ap- pressed seta nearly as long as erect setae in each puncture. Venter: Reddish brown with apex of ab- domen lighter, abdomen occasionally fus- cous; metasternum with moderately coarse punctures at lateral margins; abdomen with 168 PROCEEDINGS OF TH (| Fig. 13. serrata. Xanthonia spp., dorsal and lateral views moderately coarse punctures on intercoxal lobe, segments four and five with lateral mar- gins very irregularly notched, notches in- creasing in number toward apex; last segment of male with low tubercles each side of mid- dle, area between tubercles flat, depressed. Legs: Yellowish brown; all femora with ventral tooth very small to obsolete. Male genitalia as in Figure 13a. Total length: 2.8 to 3.0 mm. E ENTOMOLOGICAL SOCIETY OF WASHINGTON [| pe () of aedeagi. a, X. intermedia. b, X. monticola. c, X. Etymology.—Combination of inter = be- tween and medius = middle (Latin); for the unifying character of the this species. Notes.—This species is very similar to villosula and can be distinguished by the lateral margins of the last three abdominal segments being notched and the rounded apical lobes of the aedeagus. Distribution ALABAMA: Chambers Co. IOWA: Boone Co. VOLUME 103, NUMBER 1 Xanthonia monticola Staines and Weisman, new species (Figs. 6, 13b) Holotype.—¢d, Avery Co., NC, VI-15- 1955, D. M. Weisman/4000’/Holotype Xan- thonia monticola Staines and Weisman [red label] (USNM). Allotype.—@, same label data as holo- type/Allotype Xanthonia monticola Staines and Weisman [red label] (USNM). Paratypes.—42, each with Paratype Xan- thonia monticola Staines and Wesiman [red label]: 31 36, 9 2—-same label data as ho- lotype; 1 d—Black Mts., VI/10 NC (AMNH); 1 ¢—July/Greene Co. NY, 2500 ft./L. O. Howard Coll. (USNM). Description.—Coloration variable; most males with head black, becoming dark red- dish brown on front, clypeus yellowish brown, pronotum, elytra, and venter black, legs yellowish brown with femora fuscous dorsally and apically; females and one male specimen with head, pronotum, and venter light reddish brown, elytra, labrum, anten- na, and legs yellowish brown. Head: Densely, coarsely punctate; with short, golden, closely appressed pubes- cence; front with medial line moderately impressed, extending to vertex; interanten- nal area deeply to moderately concave. Pronotum: Densely, very coarsely punc- tate; with fine, dense, golden, recumbent pubescence; width averaging 1.4 times length; widest at middle; sides fairly even- ly, moderately rounded; anterior transverse depression and lateral impressions very to moderately deep. Elytra: Length averaging 1.4 times width, 3.0 times length of pronotum; basal callosity moderately prominent; punctures in regular rows, confused only near scutel- lum, behind sulcus, and below humerus; punctures about half the width of an inter- val on disc; intervals flat on disc, becoming costate near lateral margins and apex; pu- bescence composed of very fine, semierect setae in single rows on the intervals and 169 very short, obscure, appressed setae in punctures. Venter: Metasternum with moderately coarse, shallow punctures at lateral mar- gins; abdomen without coarse punctures, with sparse fine punctures on last segment; all segments with lateral margins entire; last segment of male without large tubercles. Legs: Femora with ventral tooth obsolete to absent. Male genitalia as in Figure 13b. Total length: 3.5 to 4.0 mm. Etymology.—From monticulus (Latin) = mountain; for this species apparent restric- tion to mounatinous areas. Notes.—This species is similar to stev- ensi but can be distinguished by the very coarse punctures on the pronotum and head and the male genitalia. This species appears to be limited to the higher mountains of the eastern United States. Distribution.—NEW YORK: Greene Co. NORTH CAROLINA: Avery Co.; Hender- son Co.: Black Mts. Xanthonia serrata Staines and Weisman, new species (Figs. 7, 8, 13c) Holotype.—¢d, Glen Echo, MD, VI-28- 30/JCBridwell coll./Holotype Xanthonia serrata Staines and Weisman [red label] (USNM). Allotype.—¢, same label data as holo- type/Allotype Xanthonia serrata Staines and Weisman [red label] (USNM). Paratypes.—222, each with Paratype Xanthonia serrata Staines and Weisman [red label]: 39—-with same label data as ho- lotype; 2—Glen Echo, MD, summer 1922/ JCBridwell (USNM); 1—Montg. Co. MD, VI-17-1911/Ermest Shoemaker Collection (USNM); 1—Montg. Co. MD, VI-19-1909/ Ernest Shoemaker Collection (USNM); 1— Cabin John Br. MD, 29/VII/14/V. Roberts Coll. (USNM); 1—Plummers Isld MD, 4- 6-08/W. L. McAtee Collector (USNM); 1— Plummers Isld MD, 19-6-06/W. L. McAtee Collector (USNM); 1—Plummers Isld MD, 17-8-14/RCShannon Collector (USNM); 1—Plummers Isld MD, 28-5-08/DHCle- 170 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON mons Collector (USNM); 1—Plummers Isld MD, 13-5-08/DHClemons Collector (USNM); 1—Plummers Isld MD, VI-17- 15/WLMcAtee Collector; 1—Plummers Isld MD, 31-5-09/WLMcAtee Collector (USNM); 1 Plummers Isld MD, 21-VI- 08, E Knab (USNM); 1—Great Falls MD, 27-05/DHClemons Collector/E Knab col- lection (USNM); 2—Langsdale, Chambers Co. Alab/HHSmith coll (USNM); 1—Wad- ley Alab/HHSmith coll (USNM); 1—So. Meriden Ct., 7-6-1914, Harry Johnson/ WSAbbott 1932 thru Bridwell (USNM); 5—Washgtn DC (USNM); 1—D of Col., 6- 19-1919/Ernest Shoemaker Collection (USNM); 1—Washgtn DC, FE Knab, 24-4- 04 (USNM); 1—Washgtn DC, 29-5/Coll. Hubbard and Schwarz (USNM); 1—Wa- shgton DC, 27-6/Coll. Hubbard and Schwarz (USNM); 3—Rosslyn VA/Coll. Chittenden (USNM); 1—Rosslyn VA, 5- 10/Coll. Chittenden (USNM); 1—Rosslyn VA, 6-25/ Coll. Chittenden (USNM); 1— Alex. Co. Va, VI-15-03/Ernest Shoemaker Co. (USNM); 1—Alex. Co. Va, VI-18-10/ Ernest Shoemaker Co. (USNM); 2—Pen- ington Gap VA, 2-7/Collection Hubbard and Schwarz (USNM); 1—Glencarlyn VA, E Knab, 30-V-06 (USNM); 1—Glencarlin, Alex. Co. VA, VI-23-1912/Ermest Shoe- maker Collection (USNM); 1—4-Mile Run VA, June 29-1913/A. Wetmore Collector (USNM); 1—on Solidago/Glencarlyn VA, VI-16-29/Bridwell collector (USNM); 1 Cumdb Mts., Lee Co VA, 6-19/Hubbard and Schwarz Collection (USNM); 1—Mt. Vernon VA, VI-17-1917/Ernest Shoemaker Collection (USNM); 1—Falls Church VA, VI-27-10/L.L. Buchanan Collector (USNM); 1—W. Sulpher W.Va., July 26, 1911, W. Robinson (USNM); 1—Everett, Mass., 1901/WM Mann Coll. 1954 (USNM); 1—Aug. 7, 98, Montgomery, Mass./Fredk Knab Collector (USNM); 1 Montgomery City, Mo., VI-15-1930, Stew- art Clare/Wolf Creek/on hickory (USNM); 2—N.Y./Coll. MLLinell (USNM); 2—N.Y./ Collection JHSmith (USNM); 4—N.Y. (USNM); 2—Xanthonia stevensi Lond id/ W. Robinson bequest (USNM); 1—Ithaca NY/coll. Chittenden (USNM); 2—NJ (USNM); 1—Malaga NJ, VI-16-12/Ernest Shoemaker Collection (USNM); 2—Mala- ga NJ, VI-14-13/Ernest Shoemaker Collec- tion (USNM); 1—Atco NJ, 6-22/Geo- MGreene collection (USNM); 6—Black Mts. NC, VI/11 (AMNH); 1—Black Mts., NC, VI-3 (AMNH); 1—Black Mts., NC, VI-21 (AMNH); 2—Black Mts., NC, VI-29 (AMNH); 2—Black Mts., NC, VI-24 (AMNH); 2—Black Mts., NC, VI-23 (AMNH); 1—Black Mts., NC, 18 July 1916, R. W. Leiby/about 5050 ft. (USNM); 1—Highlands NC/David A. Young, June 5, 1957/on hickory (USNM); 2—Valley of Black Mts., NC, Aug. 9, 1906, W. Beuten- muller (AMNH); 1—Valley of Black Mts., NC, Aug. 5, 1906, W. Beutenmuller (AMNH); 1—Valley of Black Mts., NC, Aug. 6, 1906, W. Beutenmuller (AMNH); 1—Valley of Black Mts., NC, Aug. 16, 1906, W. Beutenmuller (AMNH); 2—L. Toxaway N.C./Collection of Mrs. A. T. Slosson, Acc. 20226 (AMNH); 1—Tryon NC/WFFiske Collector (USNM); 2— Swannanoa Val. NC, V-20-21 (AMNH); I1—Red Knob N.C., 23-6/Coll. Hubbard and Schwarz (USNM); 13—Blowing Rock N.C., VI-27-1955, DM Weisman/white oak (USNM); 1—Robbinsville N.C./David A. Young, July 24, 1958 (USNM); 1—Gray Beard Mt. N.C. ?/17/Collection E Knab (USNM); 1—collection E Knab (USNM); 2- Mon/Coll. Chittenden (USNM); 1—July 1, 94,—town/Fredk Knab Collection (USNM); 50—Gasconade Co., Mo., 1.7 mi S Mt. Sterling, June 10, 78, Coll. E. G. Ril- ey (EGRC); 1—TENN., Dekalb Co., Tech- Aqua Area, at light—15 Aug. 1976, W. M. Beck, Jr. (EGRC); 11—Jct. FF and A, Gas- conade Co., Mo., 26 June ‘74, Coll. E. G. Riley (EGRC); 5- GA, Union Co., Brass- town Bald, Jack’s Knob Trail, 2 July 1983, R. Turnbow (EGRC); 3—LA., E. Feliciana Par., Camp Avondale, 3 mi E. Clinton, V- 22-83, Coll. E. G. Riley (EGRC); 1—VA, Botetourt Co., Craig Creek, ca. 2 mi. W. VOLUME 103, NUMBER I Eagle Rock, VI-14-1991, Coll. E. G. Riley (EGRC). Description.—Head: Reddish brown, la- brum and antenna yellowish brown; dense- ly, coarsely punctate; with short, golden, closely appressed pubescence; front with medial line very weak; interantennal area varying from slightly concave to flat. Pronotum: Reddish brown with darker area in middle of disc; densely, coarsely punctate; with short golden, recumbent pu- bescence; width averaging 1.35 times length; widest at middle; sides evenly broadly rounded; anterior transverse de- pression shallow; lateral impressions ob- scure. Elytra: Light reddish brown with black median macula at suture, each elytron with black macula at middle of base, one on hu- merus, one in middle and one on lateral area of disc, one near suture and one on lateral area of apical third, discal and sub- apical maculae tend to unite; markings of- ten lighter in color and indistinct, occasion- ally reduced in number; length averaging 1.3 times width, 2.45 times length of pron- otum; basal callosity slightly prominent; punctures almost completely confused over disc and base, tending to form rows at su- ture and along lateral margins; often with two longitudinal costae on disc; punctures separated by their diameters; intervals at lateral margins and apex costate; pubes- cence composed of erect setae tending to be in single lines spaced about two punc- tures apart and appressed setae in punctures nearly as long as erect setae. Venter: Dark with apex of abdomen ligh- ter; metasternum with moderately coarse punctures at lateral margins; abdomen with moderately coarse punctures in middle of first and last segments; last three segments with lateral margins finely serrate; last seg- ment of male without large tubercle. Legs: Yellowish brown; profemur with ventral tooth small to very small; meso- and metafemora with ventral tooth very small to obsolete. Male genitalia as in Figure 13c. Total length: 2.4 to 2.8 mm. 171 Etymology.—From serra (Latin) = saw; for the finely serrate lateral margins of the abdomen. Notes.—Some specimens are yellowish- brown with light reddish-brown head and pronotum and without darker markings. The darkest specimens have all discal and subapical maculae completely fused, the dark area on the pronotum extending to and laterally along the apical margin, vertex of head darker than front, venter completely black, and the apices of femora darker. This species is most similar to villosula and can be distinguished by the elytral punctures being confused, the slightly smaller size, and the male genitalia. Plant associations.—Specimens have been collected on Carya, Quercus alba L., and Solidago. Distribution ALABAMA: Chambers Co.: Langsdale; Randolph Co.: Wadley. CONNECTICUT: New Haven Co.: South Meridan. DISTRICT OF COLUMBIA. GEORGIA: Union Co.: Brasstown Bald. LOUISIANA: East Feliciana Parish: Clin- ton. MARYLAND: Montgomery Co.: Cab- in John Bridge; Glen Echo; Great Falls; Plummers Island. MASSACHUSETTS: Es- sex Co.: Everett; Hampden Co.: Montgom- ery. MISSOURI: Gasconade Co.: Mt. Ster- ling. Montgomery Co.: Montgomery City. NEW JERSEY: Gloucester Co.: Malaga; Ocean Co.: Atco. NEW YORK: Tompkins Co.: Ithaca. NORTH CAROLINA: ?: Gray Beard Mt.; Tryon; Red Knob; Buncombe Co.: Swannanoa Valley; Graham Co.: Rob- binsville; Henderson Co.: Black Mts.; Val- ley of Black Mts.; Macon Co.: Highlands; Transylvania Co.: Lake Toxaway; Watauga Co.: Blowing Rock. TENNESSEE: Dekalb Co.: Aqua-Tech Area. VIRGINIA: ?: 4- Mile Run; Alexandria: Mt. Vernon; Arling- ton Co.: Glencarlyn; Botetourt Co: Craig Creek; Fairfax Co.: Falls Church; Rosslyn; Lee Co.: Cumberland Mts.; Penington Gap. WEST VIRGINIA: Greenbrier Co.: White Sulphur Springs. 172 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 14. Xanthonia stevensi, dorsal and lateral view of aedeagus. Xanthonia stevensi Baly (Figs. 9, 14) Xanthonia stevensi Baly 1863: 151 [neo- type d (here designated): Sept./Montreal Que/Moxnote Colln 1940/NEOTYPE Xanthonia stevensi Baly, des. C. L. Staines 1999 (red label) (USNM)]. Crotch 1873a: 96, 1873b: 34; Gemmin- ger and Harold 1874: 3376; Lefévre 1885: 74; Wickham 1888: 89; Osborn 1892: 34. Description.—Variable in coloration but usually with head black, becoming dark reddish brown on front, clypeus yellow; la- brum and antenna yellowish brown; pron- otum, elytra, and venter black; legs yellow- ish brown with femora occasionally fuscous dorsally and apically; specimens ranging from black through dark brown, dark red- dish brown to yellowish brown with light reddish brown head and pronotum. Head: Densely, coarsely punctate; with short, golden, closely appressed pubes- cence; front with medial line faintly im- pressed, extending to vertex; interantennal area deeply to moderately concave. Pronotum: Densely, coarsely punctate; with fine, dense, golden, recumbent pubes- cence; width averaging 1.4 times length; widest at middle; sides evenly, moderately rounded; anterior transverse depression and lateral impressions very to moderately deep. Elytra: Length averaging 1.4 times width, 3.05 times length of pronotum; basal callosity moderately prominent; punctures in regular rows, confused only near scutel- lum, behind sulcus, and below humerus; punctures about half as wide as an interval VOLUME 103, NUMBER 1 on disc; intervals flat on disc, becoming costate near lateral margin and apex; pu- bescence composed of very fine, semierect setae in single rows on intervals and very short, obscure, appressed setae in punc- tures. Venter: Metasternum with moderately coarse, shallow punctures at lateral margin; abdomen without coarse punctures, with sparse, fine punctures on last segment; all segments with lateral margins entire; last segment of male without large tubercles. Legs: Femora with ventral tooth obsolete to absent. Male genitalia as in Figure 14. Total length: 3.4 to 4.2 mm. Notes.—This species was first described from Canada and is the type species of the genus. The types of most of Baly’s species are deposited in The Natural History Mu- seum (London) (Staines and Staines 1999). During a visit to this collection the type of stevensi was not located, nor did Baly’s notes indicate that it was deposited else- where. We presume the type to be lost and designate a neotype from Canada which is deposited in the USNM (Article 75, ICZN 1999). The present interpretation is based on the original description. This species shows considerable varia- tion in external morphology and aedeagal shape. Specimens for additional localities and detailed biological information may ne- cessitat a division of this species. This species is very similar to monticola but can be distinguished by the male geni- talia and the moderately coarse punctation of the head and pronotum. Plant associations.—Adults were collect- ed on Prunus (Roseaceae) and Quercus marilandica Muenchh. (Fagaceae). Specimens examined.—No labels (USNM). CANADA: QUEBEC: Hem- mingford, 5-VII-1925 (CNC); Montreal, no date (USNM); 1-VII-98 (CNC); [X-20-01 (USNM). UNITED STATES: ALABAMA: no further data (AMNH). CONNECTICUT: Litchfield Co.: Colebrook, 23-VI-1914 (CNC); Litchfield, 8/9/12 (AMNH). ILLI- NOIS: ?: Edgebrook, V-30, 7-18-15 173 (USNM). DuPage Co.: S Maywood, VIII- 9-03 (USNM). INDIANA: no further data (USNM). ?: Miller: VI-29-12 (USNM). IOWA: Dickinson Co.: Lake, WVII-22-16 (USNM). KANSAS: Pottawatomie Co.: Onaga, June 16, 1901 (USNM), VI-10-08, VI-15-23 (CASC). Shawnee Co.: Topeka, Jul. (USNM). MARYLAND: Baltimore: VI-24-09, VII-6-09 (CASC). Montgomery Co.: Plummers Island, 19-6-09, 10-VI-08 (USNM), V-30 (CASC). MASSACHU- SETTS: Franklin Co.: Sunderland (Mt. Toby), 8-VII-18 (CASC). Hampden Co.: Montgomery, June 19, 93 (USNM); Spring- field, June 30, 1900 (USNM). Hampshire Co.: Mt. Tom, June 26, 1900 (USNM). MICHIGAN: Eaton Co.: Grand Ledge (USNM). MISSOURI: Girardeau Co.: Dutchtown (USNM). NEBRASKA: Otoe Co.: Nebraska City (USNM). NEW JER- SEY: no further data (USNM). NEW YORK: no further data (AMNH). Tomp- kins Co.: Ithaca, no date, 6/15, 14-VII-1916 (USNM). Yates Co.: Dundee, 13-6 (USNM). NORTH CAROLINA: Cherokee Co.: Murphy (CASC). Henderson Co.: Black Mts., no date, VI-VII, 1902 (CASC). PENNSYLVANIA: ?: Spring Brook, VI- 27-1940 (USNM). Monroe Co.: VI-23- 1924 (USNM). TENNESSEE: Hamilton Co.: Chattanooga, May 31, 56 (CMNC). VIRGINIA: Arlington Co.: Glencarlyn, V- 30-1934 (USNM). Lee Co.: Pennington Gap, 10-7 (USNM). Total: 114. Xanthonia striata Staines and Weisman, new species (Figs. 10, 15a) Holotype.—¢d, Hanover, Pa., 15-6'30, Birdwell and Barber/Holotype Xanthonia striata Staines and Weisman [red label] (USNM). Allotype.—¢, same label data as holo- type/Allotype Xanthonia striata Staines and Weisman [red label] (USNM). Paratypes.—424, each with Paratype Xanthonia striata Staines and Weisman [red label]: 20—same label data as holotype (USNM); 1—Pyziton, Clay Co., Alab/ 174 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 15. HHSmith Coll (USNM); 1—New Haven, Ct., E. BP Van Duzee/Monnett Col. 1940 (USNM); 2—D.C. (USNM); 3—Washgtn DC, VI-30-1956, D. M. Weisman (USNM); 3—Washngtn DC, FKnab, 3-VII-04 (USNM); 1—Washngtn DC, 5 May 95/ EC.Pratt Collector (USNM); 1—&/ Washgnt. DC/Coll. Hubbard and Schwarz (USNM); 1—Brightwood DC, 26-6-05/ DHClemson Collector/Collection FKnab (USNM); 1—Rock Creek, June 11 1906, Washington DC (USNM); 1—Rock Creek June 12 1906, Washington DC (USNM); 1—Rock Creek Pk DC, Jun 14°07, C. E. Burden (USNM); 1-37 mi W Wash DC, ABGurney, 18 Jul 37 (USNM); 1—Yonah Mt., Ga., 6-20-39, PB W. Fattig/2898 (USNM); 1—?/Armuchee, Ga., 6-10-37, P. W. Fattig/2898 (USNM); 1— Kentucky, Ft. Knox, 23 June 1956/ABGurney Colr (USNM); 3—Baltimore Md, VI-26/E E. Xanthonia spp., dorsal and lateral view of aedeagi. a, X. striata. b, X. villosula. Blaisdell, 09 Collector/Blaisdell Collection (CASC); 1—Baltimore Md., VI-18-09/E E. Blaisdell Collector/Blaisdell Collection (CASC); 1—Bladensburg, Md, VII-1-20/ LLBuchanan Collector (USNM); 5—Glen Echo Md, V/28-30/JC Bridwell coll (USNM); 1—Glen Echo Md, June 18 1922/ JRMalloch Collector/Specimen No. 218 (USNM); Grt Falls Md/Washngtn DC, 28- VI-’30, J. C. Bridwell (USNM); 4—So. Mt., Hagerstown Md, Aug 6—13/WSAbbott 1932 thru Bridwell (USNM); 8—South Mts. near Meyersville Md/July 13, 1915/ HLParker Collector (USNM); 5—S. Mts. near Myersville Md/Sept. 2 15/JAHyslop Collector (USNM); 2—South Mountain Md, Jy 6—16/JAHyslop Collector (USNM); 1—Odenton Md, June 11 1922/LLBuchan- an Collector (USNM); 10—Wolfsville Md, Jy 31, 1915/WFPennington Collector/Bush- es (USNM); 1—Aug.17.1902. Mt. Tom, VOLUME 103, NUMBER 1 Mass. Fred K Knab Collector (USNM); 1— Marion Mass, VII 99/Wickham Collection 1933 (USNM); 1—5671, Mo. (USNM); 1—@/N.J./Fred K Knab_- Collector (USNM); 1—N.J./Fred K Knab Collector/ 91 (USNM); 1—N.J./Fred K Knab Collec- tor (USNM); 1—Avalon N.J./July/Kemp/A. Fenyes Collection (CASC); 1—Boonton NJ, VI.4.01/GeoMGreene Collection (USNM); 1—Boonton NJ, VII.24.01/ GeoMGreene Collection (USNM); 1— Boonton NJ, VII.8.01/GeoMGreene Collec- tion (USNM); 1—Boonton NJ, VI.6.01/ GeoMGreene Collection (USNM); 4—Col- ingwd NJ, VI.21/GeoMGreene Collection (USNM); 1—Dundalk NJ, V.31.03/Geo- MGreene Collection (USNM); 1—Dundalk NJ, VII.4.03/GeoMGreene Collection (USNM); 1—Dundee Lake NJ, VIII.31.07/ GeoMGreene Collection (USNM); 2—Fort Lee NJ, JLZabriskie, 25-VII-1894 (AMNH); 1—Malaga NJ, VIII.4.07/ GeoMGreene Collection (USNM); 1— Midvale N.J., VII-24-43/Ernest Shoemaker Collection (USNM); 2—Montclair N.J., VIII-13-21/Ernest Shoemaker Collection (USNM); 1—Oak Ridge N.J., 7-18-194/Er- nest Shoemaker Collection (USNM); 7— Phillipsburg N.J., VIII-2-14/J. W. Green Collector/J. W. Green Collection (CASC); 2—F1047/Ramsey N.J., Aug. 22.09 (AMNH); 1—Riv. NJ, 7.3.99/GeoMGreene Collection (USNM); 1—Sand Hills N.J., VII.21.09/Am. Mus. Nat. Hist. Dept. Invert. Zool. No. 14614 (AMNH); 1—Seal Isle City NJ, V.3.96/Lucarreni Collector/ GeoMGreene Collection (USNM); 7—N.Y. (USNM); 1—New York/Am. Mus. Nat. Hist. (Dept. Invert: Zoo. No. 14613 (AMNH); 3—L.I. (USNM); 2—Long L., N.Y., 6-24-06/Ernest Shoemaker Collection (USNM); 2—Long I., N.Y., 7-1-06/Ernest Shoemaker Collection (USNM); 1—Ag- nedt, LI, VIH-14-12/Ernest Shoemaker Col- lection (USNM); 1—Allegany St. Pk. N.Y., 7-4-40/A. Saunders Collector (USNM); 1 Alps N.Y., VIII.19.09/Charles Schaeffer Collection/HSBarber Bequest 1950 (USNM); 1—Bronxville NY, VI-20°14/ iS Quercus rubra/Collection of L. B. Woodruff Ac. 26824 (AMNH); 1—Bronxville NY, VII-2-’ 14/Quercus rubra/Collection of L. B. Woodruff Ac. 26824 (AMNH); 1—Collec- tion of L. B. Woodruff, Ac. 26824/Xan- thonia villosula, 6/17/11, Bronxville NY (AMNH); 1—Cld. Spg. Harbor L.I., July 31-1900/Ac. 30429/6721 (AMNH); 1— Catskill Mts. NY/Charles Schaeffer Collec- tion/HSBarber Bequest 1950 (USNM); 1— Fishkill NY, July 7 ‘16/ERKalmbach Col- lector (USNM); 1—Lott Wood, Flatbush L.I., J.L.Z., 7-18-90 (AMNH); 1—Lott Wood, Flatbush L.I., J.L.Z., 7-6-91 (AMNH); 3—Water Works, Flatbush L.I., J.L.Z., 10-VII-1893 (AMNH); 1—Water Works, Flatbush L.I., J.L.Z., 7-10-91 (AMNH); VanC Pk NY/Am. Mus. Nat. Hist. Dept. Invert. Zoo. No. 11744 (AMNH); 1—W Chestr Co NY, 8-8/ GeoMGreene Collection/Specimen No. 230 (USNM); 1—West Point, N.Y., June 21, 1910, W. Robinson/Specimen No. 122 (USNM); 1—West Point, N.Y., June 22, 1912, W. Robinson (USNM); 1—Ithaca, N.Y., VI-10-1940, J. N. Belkin (USNM); 4— Alleghany Co., NC, VIII-7-1957, DM Weisman/Cumberland Knob Park (USNM); 1—Alleghany Co., NC, VII-24-1956, DM Weisman/Cumberland Knob Park (USNM); 2—Black Mts., N.C., VII/14 (AMNH); 3 Black Mts., N.C., VI/20 (AMNH); 1— Black Mts., N.C., VI/15 (AMNH); 2— Black Mts., N.C., VI/23 (AMNH); 3— Black Mts., N.C., VI/22 (AMNH); 2— Black Mts., N.C., VI/24 (AMNH); 1— Black Mts., N.C., VI/26 (AMNH); 1— Black Mt., N.C., VII-10-40, J. W. Green/J. W. Green Collection (CASC); 1—Valley of Black Mts., N.C., Aug. 16, 1906, W. Beu- tenmuller (AMNH); 1—Valley of Black Mts., N.C., June 20-30, 1906, W. Beuten- muller (AMNH); 1—Valley of Black Mts., N.C., June 20—23, 1906, W. Beutenmuller (AMNH); 1—77749/Davidson N.C., VI- 1935, Tom Daggy (NCSU); 6—81234/Da- vidson N.C., July 17, 1977, blacklight trap, Tom Daggy (NCSU); 1—Graybeard Mt., N.C., VI/19 (AMNH); 1—Highlands, N.C./ 176 David A. Young, June 5, 1957/on Hickory/ Specimen No. 247 (USNM); 1—Hyde Co., NC, 01-15-1956, DM Weisman (USNM); 2—L. Toxaway, N.C./VI.22/Collection of Mrs. A. T. Slosson, Ac. 26226 (AMNH); 1—N. Carolina, Mt. Pisgah/David A. Young, July 15, 1957 (USNM); 1—Rob- binsville, N.C./David A. Young, July 24, 1958 (USNM); 1—So. Pines N.C., 9-V-19/ A. H. Manee Collector/Specimen No. 71 (USNM); 2—Southern Pines, N.C., VI-2, A. H. Manee (USNM); 3—Ohio (USNM). 2—Blue Ridge Summit Pa./J. A. Hyslop Collector/August 10, 115 (USNM); 3—Del. Water Gap/Collection of Mrs. A. T. Slos- son, Ac. 26226 (AMNH); 2—DuPont Pa, Aug 14-16-44, H. L. Townes (USNM); 1 Glenside Pa, VI.17.06/GeoMGreene Col- lection (USNM); 6—Glenolden Pa, VI-16/ GeoMGreene Collection (USNM); 1— Heckton Mills Pa., 8.VII.10/Coll. By P. R. Myers (USNM); 1—Lehigh Gap Pa, 7.13.1900/GeoMGreene Collection (USNM); 1—Lehigh Gap Pa, VIII.2.02/ GeoMGreene Collection (USNM); 2—Le- high Mt. Pa, V-30/Haimbach Collection (USNM); 1—Lehigh Mt., VI-30/Bethle- hem/G. W. Coffrey Collector/GeoMGreene Collection (USNM); 1—Water Gap Pa/Ac. 5409 Coll Chas..alm (AMNH); 1—Watch Hill, R.L, July 11, 1909, W. Robinson (USNM); 1—Clemson College SC. 8 Jun 1931, J. G. Waters/4 (USNM); °/Bluemont Va, VI-28-1914/Ernest Shoemaker Collec- tion (USNM); 1—@/on walnut, Clifton Va, 5 Jul. 33, Bridwell (USNM); 1—12665t Hopk. U.S./H. B. Kirk Collector/Falls Church Va/Aug 14-14 (USNM); 1—12665i Hopk. U.S./H. B. Kirk Collector/Falls Church Va/July 18-14 (USNM); 1—12665i Hopk. U.S./H. B. Kirk Collector/Falls Church Va/July 16-14 (USNM); 1—Falls Church Va, VI.12.17/GeoMGreene Collec- tion (USNM); 1—E. Falls Church Va., June 24, 17/MANGabrielson Collector (USNM); 1—E. Falls Church Va., 4/VI/14/V. Roberts Coll. (USNM); 2—Glencarlyn Va, FKnab, 30-V-06 (USNM); 1—Glencarlyn Va, FKnab, 11-VI-11 (USNM); 1—Great Falls PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Va, 01¢27:15/C..-f, Gteene* Collector GeoMGreene Collection (USNM); 11— Nelson Co., Va., June 19, 1914, W. Rob- inson (USNM); 1—Nelson Co., Va., June 27, 1915, W. Robinson (USNM); 1—Nel- son Co., Va., July 4, 1915, W. Robinson (USNM); 1—Nelson Co., Va., July 13, 1912, W. Robinson (USNM); 1—Penington Gap Va, 11.7/Coll Hubbard and Schwarz (USNM); 1—Penington Gap Va/Coll Hub- bard and Schwarz (USNM); 1—Rosslyn Va, 7.3/Coll Chittenden (USNM); 1—2500 ft. Alt., 8.4 mi. N. Thorton Gap, Va, 16.6.24, Allard (USNM); 1—Vienna Va, JCBridwell, July 8, 43 (USNM); 1—wW. Sulphur W. Va., July 6, 1911, W. Robinson (USNM); 1—W. Sulphur W. Va., July 7, 1911, W. Robinson (USNM); 1—W. Sul- phur W. Va., July 4, 1912, W. Robinson (USNM); 1—W. Sulphur W. Va., Aug 4, 1912, W. Robinson (USNM); 1—W. Sul- phur W. Va., July 20, 1910, W. Robinson (USNM); 1—@/Montreal Que, 29.VI.02/ Fred] Knab Collector (USNM); 3—6722/ Fred K Knab Collector (USNM); 2—S.I/ Coll M. L. Linell (USNM); 1—Piney Br.? ?, 5-7-05/DHClemons Collector/collection F Knab/Specimens No. 258 (USNM); 16— Randolph Co., Mo., Sugar Creek Lake, Mo- berly, VI-14-80, Coll. E. G. Riley (EGRC); 1-1 M. E Moberly, Mo., Randolph Co., June 8, 72, E. G. Riley (EGRC); 10—Barry Co., Mo., Roaring River St. Pk., VI-8-9- 1979, Coll. E. G. Riley (EGRC); 1—Mis- souri: Barry Co., Scenic Overlook on Sugar Camp rd., 3 mi. SE Roaring River St. Pk./ Mercury vapor and blacklight, VII-3-1986, E. G. Riley (EGRC); 49—Gasconade Co., Mo., 1.7 mi S. Mt. Sterling, June 10, 78, Coll. E. G. Riley (EGRC); 4—USA: MO: Ray Co., Foxglove Nat. H.A., 10-VI-1994, coll. D. G. LeDoux (EGRC); 20—TEXAS: Bastrop Co., Bastrop St. Pk., V-10-1992, Coll. E. G. Riley/beating Quercus marylan- dica (EGRC); 4—TEX: Bastrop Co., Bas- trop St. Pk., [V-19-90, Coll. E. G. Riley (EGRC); 3—USA: TEXAS, Travis Co.- Austin, Univ. Texas Brackenridge Field Lab, 550 ft./11 April 1996, S. L. Staines, VOLUME 103, NUMBER 1 C. R. Nelson, T. C. Robbins (CLSC); 28— Austin, Tex, [V-9-24/J. O. Martin Collector (CASC); 9—Austin, Tex, XII-10-28/J. O. Martin Collector (CASC); 13—Austin, Tex, IV-3-24/J. O. Martin Collector (CASC); 8—TX, Travis Co., Austin, UT-BFL, IV- 25-87, A. Hook/Bio. No. 7-87, A. Hook (USNM). Description.—Head: Dark reddish brown, labrum and antenna yellowish brown; densely, coarsely punctate; with short, closely appressed golden setae; front with medial line obscure to not evident; in- terantennal area moderately concave to flat. Pronotum: Dark reddish brown; densely, coarsely punctate; with fine, dense, recum- bent golden setae; width averaging 1.5 times length; widest just behind middle; sides slightly angulate behind middle, slightly rounded and converging apically, strongly rounded behind; anterior trans- verse depression and lateral impressions moderately deep to shallow. Elytra: Reddish brown; width averaging 1.5 times width, 2.9 times length of pron- otum; basal callosity evident; punctures in regular rows, confused only near scutellum, behind sulcus, and below humeri; punctures nearly as wide as intervals on disc; inter- vals costate especially at sides and apex; pubescence composed of erect setae in sin- gle lines on intervals and very short, ob- scure, appressed setae in punctures. Venter: Dark reddish brown; metaster- num with sparse moderately coarse, shal- low punctures at lateral margins; abdomen without coarse punctures, with sparse, fine punctures on last segment; all segments with lateral margins entire; last segment of male without large tubercles. Legs: Femora with ventral tooth very small to absent. Male genitalia as in Figure 15a. Total length: 3.5 to 4.5 mm. Etymology.—From stria (Latin) = line; for the regular rows of punctures on the el- ytra. Notes.—Some specimens are yellowish brown with light reddish-brown head and pronotum. Others, especially toward the 177 southern end of the range, are darker with head, pronotum, and venter fuscous, elytra dark reddish-brown, labrum, antennae, and legs reddish-brown. Some of the dark spec- imens have suffused darker areas at center or transversely across the pronotal disc and on each elytron at middle of base, on hu- merus, at middle of the elytral suture, and middle of the disc. This species is similar to angulata but can be distinguished by the deeper apical emargination of the male genitalia. Plant associations.—Specimens have been collected on Carya, Juglans, Quercus rubra, and Q. marylandica foliage. Distribution.—CANADA: Quebec: Montreal. UNITED STATES: ALABAMA: Clay Co.: Pyziton. CONNECTICUT: New Haven Co.: New Haven. DISTRICT OF COLUMBIA: Brightwood; Rock Creek Park; 37 mi W. GEORGIA: ?: Yonah Mt.; Floyd Co.: Armuchee. KENTUCKY: Mea- de Co.: Fort Knox. MARYLAND: Anne Arundel Co.: Odenton; Baltimore; Freder- ick Co.: South Mountain; South Mountain near Myersville; Wolfsville; Montgomery Co::) (Glen; Echo; {Great (Falls]| Prince George’s Co.: Bladensburg; Washington Co.: Hagerstown. MASSACHUSETTS: Hampshire Co.: Mt. Tom; Plymouth Co.: Marion. MISSOURI: Barry Co.: Roaring River State Park; Gasconade Co.: Mt. Ster- ling; Randolph Co.: Moberly; Ray Co.: Foxglove Natural Heritage Area. NEW JERSEY: ?: Dundalk; Dundee Lake; Riv.; Sand Hills; Bergen Co.: Fort Lee; Ramsey; Cape May Co.: Avalon; Seal Isle City; Es- sex Co.: Montclair; Gloucester Co.: Col- lingswood; Malaga; Morris Co.: Boonton; Passaic Co.: Midvale; Oak Ridge; Warren Co.: Phillipsburg. NEW YORK: Long Is- land; ?: Agnedt LI; Catskill Mts.; Bronx Co.: Bronxville; Van Cortlandt Park; Cat- taraugus Co.: Allegany State Park; Dutch- ess Co.: Fishkill; Kings Co.: Flatbush, L. 1; Nassau Co.: Cold Spring Harbor; Orange Co.: West Point; Rensselaer Co.: Alps; Tompkins Co.: Ithaca; West Chester Co. NORTH CAROLINA: ?: Graybeard Mt.; 178 Alleghany Co.: Cumberland Knob Park; Graham Co.: Robbinsville; Henderson Co.: Black Mts.; Valley of Black Mts; Mt. Pis- gah; Hyde Co.; Macon Co.: Highlands; Mecklenberg Co.: Davidson; Moore Co.: Southern Pines; Transylvania Co.: Lake Toxaway. OHIO. PENNSYLVANIA: ?: DuPont; Glenside; Heckton Mills; Lehigh Gap; Lehigh Mt.; Delaware Co.: Glenolden; Franklin Co.: Blue Ridge Summit; Monroe Co.: Delaware Water Gap; Northampton Co.: Bethlehem; Lehigh Mt.; York Co.:: Hanover. RHODE ISLAND: Washington Co.: Watch Hill. SOUTH CAROLINA: Pickens Co.: Clemson College. TEXAS: Bastrop Co.: Bastrop State Park; Travis Co.: Austin. VIRGINIA: ?: Clifton; Arlington Co.: Glencarlyn; Fairfax Co.: Falls Church; E. Falls Church; Great Falls; Rosslyn; Vi- enna; Lee Co.: Pennington Gap; Loudoun Co.: Bluemont; Nelson Co.; Prince William Co.: Clifton; Rappahannock Co.: 8.4 mi. N. Thornton Gap. WEST VIRGINIA: Green- brier Co.: White Sulphur Springs. Xanthonia villosula (Melsheimer) (Figs. 11, 15b) Eumolpus villosulus Melsheimer 1847: 169 [neotype 6 (here designated): Lime Rid, Pa/Wickham Collection/NEOTYPE Xan- thonia villosula (Melsheimer) des. C. L. Staines 1999 (USNM)]. Myochrous villosulus: Melsheimer 1853: 25: Xanthonia villosula: Crotch 1873a: 96, 1873b: 34; Gemminger and Harold 1874: 3376; Hubbard and Schwarz 1878a: 640, 1878b: 660; Harrington 1884: 81; Lefév- re 1885: 74; Horn 1892: 200; Hamilton 1895: 340; Davis 1902: 37; Dury 1902: 165; Ulke 1902: 28; Fall and Cockerell 1907: 196; Blatchley 1910: 1142; Wick- ham 1911: 31; Clavareau 1914: 75; An- drews 1916: 100; Johnson 1916: 120; Chagnon 1917: 240; Leng 1920: 292; Wellhouse 1922: 1067; Blatchley 1924: 57; Hatch 1924: 582; Schaeffer 1928: 467; Douglass 1929: 57; Brimley 1938: 224; Johnson 1941: 11; Powell 1941: PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 158; Hughes 1944: 133; Léding 1945: 129; Fattig 1948: 19; Wilcox 1954: 401; Dillon and Dillon 1961: 680; Kirk 1970: 89; Rouse and Medvedev 1972: 79; Kirk and Balsbaugh 1975: 108; Wilcox 1975: 57; Riley and Enns 1979: 65; Wilcox 1979: 19; LeSage 1991: 309; Downie and Arnett 1996: 1333. Myochrous plagiatus (Melsheimer). Melsh- eimer 1853: 125. Nomen nudum. Description.—Head, pronotum, elytra, and venter usually reddish brown with yel- lowish brown labrum, antenna, and legs; some specimens are yellowish brown with reddish brown head and pronotum; others are darker reddish brown, venter fuscous with apex of abdomen lighter, each elytron with dark macula at middle of base, one on humerus, a linear dorsal one near suture, a row of three across disc and three across apical third, the discal and subapical mac- ulae often fused; markings seldom distinct and more often completely absent. Head: Densely, coarsely punctate; with short, golden, closely appressed pubes- cence; front with medial line slightly im- pressed, occasionally extending to vertex; interantennal area moderately concave. Pronotum: Densely, coarsely punctate; with short, golden, recumbent pubescence; width averaging 1.25 times length; widest at middle; sides broadly rounded; anterior transverse depression moderately deep; lat- eral impressions moderately deep to ob- scure. Elytra: Length averaging 1.5 times width, 2.7 times length of pronotum; basal callosity evident; punctures in regular rows, confused only near scutellum, behind sul- cus, and below humerus; punctures as wide as width of elytral interval on disc; inter- vals costate, especially laterally and apical- ly; pubescence composed of erect setae in single lines on intervals and very short, ob- scure, appressed setae at punctures. Venter: Metasternum with moderately coarse punctures at lateral margins; abdo- men with moderately coarse punctures, VOLUME 103, NUMBER 1 sparse to dense on both intercoxal lobe and last segment; last three segments with lat- eral margins finely serrate, occasionally segments three and four irregularly notched and 5 finely serrate; last segment of male without large tubercles. Legs: Femora with ventral tooth very small to obsolete. Male genitalia as in Fig- ure 15b. Total length: 2.8 to 3.5 mm. Notes.—This species was originally de- scribed from Pennsylvania. The present in- terpretation is based on the original descrip- tion. We were unable to locate the Melsh- eimer type in the Museum of Comparative Zoology (Perkins per. comm.) and presume it to be lost, a neotype from Pennsylvania is hereby designated (Article 75, ICZN 1999). Myochrous plagiatus Melsheimer was not accompanied by a description of the species and is a nomen nudum (Article 12, ICZN 1999). Horn (1892) synonymized plagiatus with villosula and mentioned that he had seen Melsheimer’s specimen. No specimens labeled plagiatus were found in the Melsheimer, LeConte or Horn collec- tions (Perkins, pers. comm.) This species is similar to intermedia but can be distinguished by the truncate lobes of the aedeagus and the finely serrate lateral margins of the last abdominal segment. Specimens examined.—??: Cascade, 7.12.08 (CASC); Thatcher’s Pinnacles, 5-V- 57 (USNM). CANADA: ONTARIO: Mar- mora, 20-VIII-1951 (CNC); Toronto (USNM). QUEBEC: Lavoltrle, VI-14-19 (CNC); Montreal (CASC). YUKON: Dawson, June 13, 1917 (USNM). UNIT- ED STATES: ARKANSAS: southwest (AMNH). Hempstead Co.: Hope, VI-11-32, VI-11-1954 (CASC). Polk Co.: Cove, 6-20- 38 (SEMC). COLORADO: (USNM). CONNECTICUT: New Haven Co.: New Haven (CASC). DISTRICT OF COLUM- BIA: 6-19-1909, VI-19-1911, June 30, 19, VI-30-1956, VI-16-1956, 13-9, VII-31-07 (USNM). ILLINOIS: (AMNH, USNM). ?: Chicago, May 30 ‘02 (USNM); Edgebrook, 7-18-15 (USNM); Glendon P’k, VI.28.03 179 (USNM). DuPage Co.: S. Maywood, VIIIL.9.03 (USNM). LaSalle Co.: VI-18- 1938, VI-27-1938, VII-4-1938 (UAIC); Starved Rock State Park, VII 1940 (UAIC). INDIANA: ?: Miller, VI.29.12 (USNM). Floyd Co.: 6-27-03 (USNM). Tippecanoe Col NVIES.1955.--V9E15.1955. CUSNM).- IOWA: Allamakee Co.: Waukon, VII-2-14 (USNM). Appanoose Co.: Centerville, VII- 23-14 (USNM). Boone Co.: VII-32 (USNM), July 4, 32 (CASC). Dickinson Co.: Lake Okoboji, 7-15-16, VI-8-16, VII- 22-16 (USNM). Dubuque Co.: Dubuque, VI-29-14 (USNM). Johnson Co.: Iowa City (CASC), IX-4-14, VI-23-98, [X-3-17, III- 10 (USNM). Story Co.: Ames, 7-10-32 (USNM). KANSAS: Coffey Co.: June 19, 1923, 7-11-39 (SEMC). Doniphan Co.: Walthena, VIII.26, VIII.12 (USNM). Doug- las Co.: 900 ft. (SEMC); Lawrence, 5-31- 1929, June 24, 1919 (SEMC). Leavenworth Co.: 6-30-24 (USNM). Linn Co.: 1915, ele. 862 (SEMC). McPherson Co.: 5-28-23 (USNM). Pottawatomie Co.: Onaga, VI-16- 09, VI-15-23, VI-10-09 (CASC), June 16, 1901 (USNM). Shawnee Co.: 6-13-23 (SEMC); Topeka (SEMC, USNM), Jun 18 (USNM). Wyandotte Co.: 6-23-24 (SEMC). KENTUCKY: Trigg Co.: Cadiz, June 30, 1939 (SEMC). MAINE: Penobscot Co.: Orono, 15 Sept. ‘13 (CASC). MARY- LAND: Baltimore: VI-24-09, VI-26-09, VI- 18-09 (CASC). Baltimore Co.: Sparrows Point, VII-9-31 (CASC). Frederick Co.: 5 mi. W. Thurmont, 16-VII-1971 (USNM). Montgomery Co.: VI-4-01, VI-25-1911 (USNM); Cabin John Bridge, 29-VII-14, July (USNM); Glen Echo, VI.28.30, VI20.30; July 24; 19225) July “12; ‘1922 (USNM); Great Falls, VI.24.11, 12-7-05, VI-25-1911 (USNM); Jackon’s Island, 23- V-13 (USNM); Plummers Island, 14-VI-09, 19-6-05, VIII.5.17, Jly 16, 13, 21-7-07, 20- VI-1912, 15-VII-06, VI.24.17, Jly 9, 1918, 27-6-05, June 11, 27-8-05, 10-6-06, 20-VI- 1912, 13-5-05, 28-6-05, 4-VII-07, V-25- 18, 26-VIII-O1, 25.8.14, 16-VI-1912, 20: ViN9129 13.8.14, 18.6.14, 15.6.02, ly 24 ‘18, 25-8-04, 21-5-02, 26.6.15, 3.8, VII- 180 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 07 (USNM). Prince George’s Co.: Belts- ville, June 4, 14 (USNM). Washington Co.: Hancock, 9-1-16 (USNM). MASSACHU- SETTS: Berkshire Co.: Mt. Tom, Aug. 17, 1902 (USNM). Hampshire Co.: Cumming- ton, Aug 20, 1950 (USNM). Norfolk Co.: Dover (CASC). MISSISSIPPI: Adams Co.: Natchez, V.12.09 (USNM). Oktibbeha Co.: Starkville, Jn 20 (CASC). MISSOURI: (USNM); C. Mo. (USNM). Barry Co.: Roaring River State Park, VI-15-54 (CASC). Boone Co.: Columbia, Apr. 1907 (CASC). Motgomery Co.: Graham Cave State Park, July 6, 1973 (NCSU). St. Louis Co.: St. Louis, 28-V-39 (USNM). NE- BRASKA: ?: Kereoslia (USNM). Otoe Co.: Nebraska City (USNM). NEW HAMP- SHIRE: Belknap Co.: Barnstead, [X-6-28, IX-9-28 (CASC). NEW JERSEY: (USNM). Bergen Co.: Alpine, VII-25-10 (AMNH); Fort Lee, 25-VIII-1894 (AMNH). Morris €o.> Boonton; Vil232085 ( Vile12:01 (USNM); Chester, VII.S (AMNH); Men- dham, July 24, 1916 (USNM). Gloucester Co.: Malaga, VIH-3 (CASC). Ocean Co.: Lakehurst, X-12-14 (USNM). Passaic Co.: Oak Ridge, 6-19-1904 (USNM). Warren Co.: Phillipsburg, VII.19.14 (CASC). NEW YORK: (AMNH, USNM). ?: N. Evans, 5- 14-10 (CASC); Long Island, 7-1-06, VII- 10-1907 (SEMC, USNM). Chautauqua Co.: Dunkirk, 3.7 (USNM). Dutchess Co.: Fish- kill, July 7 ‘16 (USNM). Erie Co.: Colden, 7-5-10 (CASC); East Aurora, 7-16-10 (CASC). Niagara Co.: Buffalo (USNM); Niagara Falls (CASC), 9-9-11 (CASC); Ol- cott, VII-4-18 (CASC). Onondaga Co.: El- bridge, 30.VI.1941 (USNM). Orange Co.: West Point, June 21, 1910 (USNM). Rens- selaer Co.: Alps, VIII.14.09 (USNM). Tompkins Co.: Groton, VIII.12.1955 (USNM); Ithaca, 8/8, 6.15 (USNM). Ulster Co.: Olivera, VI-27->15 (USNM). NORTH CAROLINA: Henderson Co.: Black Mts., VI-VII-1902 (CASC). Wake Co.: Raleigh, June 6, 1974, VI-25-1951 (USNM). OHIO: (USNM). OKLAHOMA: Atoka Co.: Ato- ka, 13.VI.1933 (USNM). Carter Co.: Ard- more, 6-1-09 (USNM). Le Flore Co.: 5-24- 28 (SEMC). Pittsburg Co.: So. McAlester, Ind. T. (USNM). PENNSYLVANIA: (CASC, USNM). ?: Lehigh Gap, July 12.99 (CASC); Water Gap (AMNH). Bucks Co.: Perkasie, VI.18 (USNM). Columbia Co.: Lime Ridge (USNM). Delaware Co.: Glen- olden, VI-16 (USNM). Wayne Co.: Star- light, VI-13-1921 (USNM). SOUTH CAR- OLINA) 2) Mt? Rest, ai duneowoan (USNM). TENNESSEE: Hamilton Co.: Chattanooga, May 31 ‘56 (SEMC). Putnam Co.: Clarksville, 7-4-39 (SEMC). TEXAS: Rockwell Co.: Dallas, [V-26-07, 17 Apr. 07, V-18-07, V-19-07 (USNM). Travis Co.: Austin, XII-10-28, IV-3-24, IV-9-24 (CASC). VIRGINIA: (USNM). ?: Piney Run, VII 1950 (USNM); Scott’s Run, Aug. 2, 14 (USNM). Arlington Co.: Glen- carlyn, V-30-1934 (USNM). Fairfax Co.: VI-17-1913 (USNM); Falls Church, 9.V1I.1918 (USNM); Great Falls, VI-17- 1920 (USNM); Mt. Vernon, VI-20-1929 (USNM), VII-6-18 (SEMC). Loudoun Co.: Bluemont, VI-28-1914 (USNM). Warren Co.: 5 mi N Linden, 3-VH-1971 (USNM). WEST VIRGINIA: Greenbreir Co.: White Sulphur Springs (CASC). WISCONSIN: Dane Co.: Madison, 8/12/27 (USNM). Grant Co.: VI-2-1949 (USNM). Juneau Co.: Sept. 2, 1949 (USNM). Racine Co.: Aug. 5, 1950 (USNM). Wood Co.: Lake Nepco, [X-13-1951 (USNM). Total: 530. ACKNOWLEDGMENTS The following institutions have lent ma- terial for this study, the assistance of the curators responsible is greatly appreciated: American Museum of Natural History, New York (AMNH), M. A. Cazier; California Academy of Sciences, San Francisco (CASC), D. Kavanaugh; Canadian Museum of Nature, Ottawa (CMNC), H. H. Howden; Canadian National Collection, Ottawa (CNC), W. J. Brown; Museum of Compar- ative Zoology, Harvard University, Cam- bridge (MCZC), P. Perkins; E. G. Riley, Texas A & M University, College Stattion (EGRC); Snow Entomological Museum, University of Kansas, Lawrence (SEMO), VOLUME 103, NUMBER 1 G. W. Byers; C. L. Staines (CLSC); Uni- versity of Arizona, Tucson (UAIC), FE G. Werner; National Museum of Natural His- tory, Smithsonian Institution, Washington, DC (USNM), G. B. Vogt. E. G. Riley, Texas A & M University, and A. S. Konstantinov and S. W. Linga- felter, Systematic Entomology Laboratory, USDA commented on an earlier draft of this manuscript. Publication of this manu- script was paid for by the Thomas Lincoln Casey Fund. We thank S. L. Staines for cu- ratorial and editorial assistance. LITERATURE CITED Andrews, A. W. 1916. 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The grape-vine Fidia. Practical En- tomologist 2: 87-88. Weisman, D. M. 1960. A revision of the species of the genus Xanthonia Baly in America, north of Mex- ico. Unpublished Master of Science thesis. North Carolina State University. 58 pp. Wellhouse, W. H. 1919. Xanthonia villosula Mel\sh. in- juring forest trees. Journal of Economic Entomol- ogy 12: 396-397. . 1922. The insect fauna of the genus Cratae- gus. Cornell University Agricultural Experiment Station Memoir 56: 1045-1136. Wickham, H. E 1888. A list of the Coleoptera of Iowa City and vicinity. Bulletin of the Laboratories of Natural History of the State University of Iowa 1: 81-92. 1896. The Chrysomelidae of Ontario and Quebec. Canadian Entomologist 28: 171—174. . 1902. The Coleoptera of Colorado. Bulletin of the Laboratory of Natural History of the State University of Iowa 5: 217-310. . 1911. A list of the Coleoptera of Iowa. Bul- letin of the Laboratory of Natural History of the State University of Iowa 6. 40 pp. Wilcox, J. A. 1954. Leaf beetles of Ohio (Chrysome- lidae: Coleoptera). Ohio Biological Survey Bul- letin 43: 353-506. . 1975. Checklist of the beetles of Canada, United States, Mexico, Central America, and the West Indies. Vol. 1 part 7, the leaf beetles (red version). Biological Research Institute of Ameri- ca. Latham, New York. 166 pp . 1979. Leaf beetle host plants in northeastern North America (Coleoptera: Chrysomelidae). Bi- ological Research Institute, Latham, New York. 30 pp. PROC. ENTOMOL. SOC. WASH. 103(1), 2001, pp. 184-190 HYLAEUS HYALINATUS SMITH, A EUROPEAN BEE NEW TO NORTH AMERICA, WITH NOTES ON OTHER ADVENTIVE BEES (HYMENOPTERA: APOIDEA) JOHN S. ASCHER Department of Entomology, Comstock Hall, Cornell University, Ithaca, NY, 14853- 0901, U.S.A. (e-mail: ja41 @cornell.edu) Abstract.—The common and widespread European bee Hylaeus (Spatulariella) hyali- natus Smith (Hymenoptera: Colletidae) is reported for the first time from the New World. During 1997-2000, specimens were collected in Ithaca, Tompkins County, New York. Diagnostic features are described and photographed to facilitate separation of H. hyali- natus from Nearctic Hylaeus species and from the two other Palearctic Hylaeus species previously reported as adventive in North America. Its geographic distribution, host plants, and seasonal history are summarized from the literature and from collections made in New York. Notes are provided on the occurrence of other adventive bee species including the first records of Hylaeus (Spatulariella) punctatus (Brullé) from northern California and of Megachile (Callomegachile) sculpturalis Smith (Hymenoptera: Megachilidae) from New York state. Key Words: North America Hylaeus hyalinatus is the seventeenth species of bee to be reported as adventive in North America (excluding Apis mellifera L., four other deliberately introduced spe- cies, and Ceratina (Calloceratina) aff. laeta Spinola, which may be adventive in south Texas from the neotropics; Cane, in press). Each of these species is native to the Old World and is thought to have reached North America as a result of accidental anthro- pogenic movement of nests containing dia- pausing bees. Although adventive bees comprise less than 0.5% of North American bee species (Cane, in press), they may com- prise a larger percentage of local faunas, es- pecially in northeastern North America. In Ithaca, Tompkins County, New York, no less than ten species are established and abundant, and these species comprise 3.2% of the total bee fauna of 310 species, and Hymenoptera, Colletidae, Megachilidae, adventive insect, distribution, 4.3% of the 231 pollen-collecting bee spe- cies (John Ascher, unpublished data). More adventive bee species occur in Ithaca, New York than have been reported from any oth- er continental locality in the world. The most diverse fauna of exotic bees is likely that of the Hawaiian Islands. The rate of establishment of exotic bee species appears to have increased in recent decades, espe- cially in the northeastern United States. An- thidium manicatum (L.), Chelostoma fulu- ginosum (Panzer), and Chelostoma cam- panularum (Kirby), were discovered in cen- tral New York in 1963 (Jaycox 1967), 1973 (Eickwort 1980), and 1962 (Eickwort 1980) respectively, while the first North American record of Hoplitis anthocopoides came from Albany County, New York, in 1969 (Eickwort 1970). Two additional Palearctic megachilids have recently become estab- VOLUME 103, NUMBER 1 lished and locally abundant in the eastern United States: Anthidium oblongatum (Illi- ger) (Hoebeke and Wheeler 1999) and Me- gachile sculpturalis Smith (Mangum and Brooks 1997, Batra 1998). RESULTS Hylaeus (Spatulariella) hyalinatus Smith (Figs. 12) During an ongoing survey of the bees of Ithaca, Tompkins County, New York, I col- lected numerous specimens of an unusual hylaeine bee on the Cornell University campus. These proved to be Hylaeus (Spa- tulariella) hyalinatus Smith (Hymenoptera: Colletidae). Although this Palearctic spe- cies is previously unrecorded in the New World, another species of Spatulariella, Hy- laeus (S.) punctatus (Brullé), has been re- ported to be adventive in Los Angeles County, California (Snelling 1983) and Santiago, Chile (Toro et al. 1989). Addi- tional records of H. punctatus are provided below. New York specimens of H. (S.) hyalina- tus hyalinatus agree with the description and figures in Dathe (1980) and match identified specimens in the Cornell Univer- sity Insect Collection (CUIC). Identification was confirmed by Roy R. Snelling of the Los Angeles County Museum (LACM). Voucher specimens collected in Ithaca have been placed in the following collections: CUIC, LACM, the American Museum of Natural History (AMNH), the Snow Ento- mological Museum, University of Kansas (SEM), and the Essig Museum, University of California, Berkeley (UCB). Snelling (1983) modified his key to the Nearctic subgenera of Hylaeus (Snelling 1966) to include the subgenus Spatulariella Popov, which was unknown in the New World prior to his discovery of adventive H. (S.) punctatus in California. Both sexes of H. hyalinatus possess a lamelliform ca- rina between the anterior and lateral faces of the mesepisternum (termed a lamelliform omaulus by Michener 2000), which Snell- 185 ing (1983) suggests is “‘perhaps the best recognition characteristic by which to dif- ferentiate Spatulariella from the native Ne- arctic groups.” In addition, males of H. hy- alinatus, in common with other Spatular- iella, and unlike all native Nearctic species, possess a spatulate eighth sternum that pro- tudes conspicuously from the genital open- ing in combination with long slender gon- ostyli that extend far beyond the apices of the penis valves (see Fig. 1; and Snelling 1983: Fig. 3). Dathe (1980) discusses Spa- tulariella taxonomy and provides illustra- tions and keys for European species. Hylaeus hyalinatus can be separated from H. punctatus using the key provided below. Male A. hyalinatus with black scapes are superficially similar in facial markings to H. (Prosopis) modestus, the most abundant native Hylaeus in the Ithaca area (see figure of H. modestus in Mitchell 1960: Fig. 11). In H. hyalinatus, the ventral surface of the flagellum is paler, the face marks are whitish-yellow rather than bright yellow, the supraclypeal mark does not ex- tend above the level of the ventral margin of the antennal sockets, and the scape is of- ten maculated (Fig. 2). Hylaeus hyalinatus females have patterns of maculation resem- bling several other Hylaeus occuring in Eastern North America, especially the ad- ventive H. bisinuatus Forster, but are read- ily distinguished by the lamelliform carina of the mesepisternum. KEY TO THE SPECIES OF HYLAEUS (SPATULARIELLA) RECORDED AS ADVENTIVE IN THE NEw WORLD Bald Ei caret aa ae rs earths arent ate se Ns oe are eae 2 Female . Eighth metasomal sternum (S8) with distal spatulate process connected to the base by an extremely narrow elongate stalk (Fig. 1; Dathe 1980: Fig. 107D). Supraclypeal area pale; pale lateral face marks extending on the eye margin to well above the antennal bases (Fig. 2; Dathe 1980: Fig. 107B) hyalinatus — S8 with distal spatulate process broadly con- nected to the base (Dathe 1980: Fig. 112D; Snelling 1983: Fig. 5). Supraclypeal area usu- ally black; lateral face marks extending little, wo | 186 Figs. 1-2. process of S8 and elongate, slender gonostyli. 2, Face. if at all, above the level of the epistomal sulcus (Dathe 1980: Fig. 112B; Snelling 1983: Fig. 2) punctatus 3. Mesopleuron and scutum equally finely punc- tate. Propodeum with posterior and lateral fac- es separated by a rounded angle..... hyalinatus — Mesopleuron more coarsely punctate than scu- tum. Propodeum with posterior and lateral fac- es separated by an irregular carina ... punctatus New York specimens and European spec- imens (Dathe 1980) of both male and fe- male H. hyalinatus vary in the extent of yellow markings as is also true of other Hy- laeus species, including H. punctatus (Snelling 1983). As in H. punctatus, some female H. hyalinatus possess a median, preapical clypeal spot whereas others have an entirely black clypeus. Most female H. hyalinatus have well developed lateral face marks as illustrated in Dathe (1980), but one female examined had the lateral face marks reduced to about half their usual size. In 11 of 24 New York males examined, the scape was entirely black, whereas in 13 oth- er males the scape was marked to some de- gree with yellow (Fig. 2). In 6 of the spec- imens, the scape mark consisted of a stripe extending almost its entire length, whereas in the others the mark was less extensive or almost absent. All 11 males collected on or before July 7 lacked pale scape marks, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Hylaeus hyalinatus, male. 1, Apex of metasoma, ventral view; note protruding, spatulate apical whereas all but one male collected after July 22 had scape marks, suggesting that the variable expression of pale markings in Hylaeus hyalinatus may be correlated with environmental conditions and may not re- flect genetic variability. Hylaeus hyalinatus, widely distributed and abundant in north and central Europe, occurs as far north as Finland (Elfving 1951), and ranges from the Pyrenees in the west to the Caucasus and Moscow in the east and to Greece in the south (Dathe 1980). It inhabits the Alps up to an eleva- tion of 1800 m (Beaumont 1958). Two montane subspecies with restricted ranges are recognized by Dathe (1980) in addition to the widespread nominate form. In North America, H. hyalinatus is known only from the vicinity of the Cornell University campus in Ithaca, Tompkins County, New York. The first specimen, a male, was collected 10 June 1997 on the Cornell campus in the Minns Horticulture Garden (MG), a small plot filled with cul- tivated flowers of diverse species. The gar- den is adjacent to a greenhouse conserva- tory containing exotic plants. Females were first noted at MG on 7 July, and by August both sexes were consistently present in numbers at this location. During 1998—2000 VOLUME 103, NUMBER 1 MG remained the center of abundance for the species, but a few specimens were found at other localities within 2.5 km of MG. No specimens were collected further afield, even though I collected dozens of individuals of 7 species of Hylaeus from many localities. No additional specimens could be found among the unsorted Hylaeus in the CUIC. Hylaeus hyalinatus appears to be widely polylectic in Europe, and females are known to forage on flowers of at least seven families (Westrich 1990). In the Minns Gar- den, both males and females have been ob- served visiting plants introduced from the Palearctic region including Lavandula an- gustifolia Mill. [Lamiaceae], Rudbeckia ful- gida Ait. [Asteraceae], Brunnera macro- phylla (Adams) I. M. Johnst. [Boragina- ceae], Alyssum sp. [Brassicaceae], Anethum graveolens L. [Apiaceae], and Foeniculum vulgare Mill. [Apiaceae]. They were found most frequently, and in the largest numbers, on Foeniculum vulgare and Alyssum. Foen- iculum vulgare was the preferred host of H. punctatus in California (Snelling 1983), whereas Alyssum was one of two plants pre- ferred by introduced Hylaeus punctatus in Chile. The only genus of native plant from which H. hyalinatus has been recorded in New York is Solidago (Asteraceae). This is the most widespread and abundant nectar source in Ithaca during late summer and early autumn, and is commonly visited by several Hylaeus species. Westrich (1990) stated that H. hyalinatus is at least partly bivoltine in Germany with a long flight season extending from early May into early October. The Ithaca popu- lation is almost certain to be at least partly bivoltine as well, because specimens have been collected from spring into early au- tumn. Males have been collected from 3 May until 31 August; females from 20 May until 20 September. In Europe, H. hyalinatus nests in pre- formed cavities in a wide range of sub- strates. It inhabits banks of sand, gravel, and loam, dry walls, and house walls up to 187 16 m above gound level. It also uses hollow stems of Rubus, and reuses abandoned nests of mason bees and Trypoxylon (Westrich 1990). It occurs at the edge of forests, as well as in vineyards, orchards, ruderal are- as, and the vicinity of human habitation (Westrich 1990). Given H. hyalinatus’ be- havior and geographic range, its establish- ment in New York is not surprising. Hylaeus hyalinatus is the third Hylaeus species to be reported as adventive in the Americas. Hylaeus (Hylaeus) bisinuatus Forster, native to the Palearctic, was first recorded in Fargo, North Dakota, (as H. stevensi Crawford) around 1912 and is now widespread over much of North America (Hurd 1979). It has also been introduced to the Hawaiian Islands (Hurd 1979), as has the Australian species Hylaeus albonitens (Cockerell) (Hopper and Daly 1995). Hy- laeus bisinuatus is one of eleven Hylaeus species (including H. hyalinatus) known from Tompkins County, New York. Of the nine native species in Tompkins County, two, Hylaeus (Paraprosopis) georgicus (Cockerell), and H. (Cephalylaeus) basalis (Smith), are rare and known from only one, and three specimens respectively, and a third, H. (Hylaeus) saniculae (Robertson), has not been collected locally since 1937. Of the remainder, Hylaeus (Prosopis) mo- destus Say is the most commonly collected, while H. (Prosopis) affinis (Smith), H. (Hy- laeus) mesillae (Cockerell), H. (H.). ellipti- cus (Kirby), H. (H.) verticalis (Cresson), and H. (Prosopis) illinoisensis (Robertson) occur in declining order of abundance. If H. hyalinatus expands in range and increases in abundance, it may compete with these as well as other cavity-nesting Hymenoptera for nest sites (Thorpe 1996, Barthell et al. 1998, Cane, in press). Hylaeus (Spatulariella) punctatus (Brullé) This species was first collected in the New World at Playa del Rey, Los Angeles County, California, in 1981 (Snelling 1983). It was later reported from Chile (Toro et al. 1989). In 1999, I collected 29 188 specimens (1 2, 3 6 on 2 June, 14 2,16 on 21 June, visiting flowers of Aesculus californica (Spach) Nutt. [Hippocastana- ceae] and unidentified Asteraceae and Ro- saceae) on the campus of the University of California, Berkeley, Alameda County, Cal- ifornia. On 20 June 2000, I collected 13 males in Lafayette, Contra Costa, County, California. These are the first northern Cal- ifornia records of Hylaeus punctatus. Ad- ditional records of this species from Pin- nacles National Monument, San Benito County, California in May, 2000 (T. Gris- wold, in litt.), and from localities in south- ern California in addition to Playa del Rey (R. Snelling, personal communication), suggest that H. punctatus is well-estab- lished in California and rather widespread. Voucher specimens have been placed in the CUIC, LACM, and SEM. Anthidium (Proanthidium) oblongatum (Illiger) This species was first detected in North America in Pennsylvania, New Jersey, and Maryland in 1994—1997, and was first not- ed in Tompkins County, New York, in Sep- tember, 1997 (Hoebeke and Wheeler 1999). Since then it appears to have rapidly in- creased in abundance in the Ithaca area. During 1998—2000, I observed and collect- ed abundant specimens of A. oblongatum at several sites in Tompkins County where it was not detected in 1997. Most were vis- iting Sedum or exotic Fabaceae, especially Lotus corniculatus L. On these flowers A. oblongatum consistently outnumbered na- tive megachilids. They were first seen in June and continued activity until at least October 12, when a single male was cap- tured. Megachile (Callomegachile) sculpturalis Smith The giant resin bee Megachile sculptur- alis, native to east Asia, was first reported from North America by Mangum and Brooks (1997) who found it to be well-es- tablished in North Carolina. Batra (1998) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON discussed its biology, and provided addi- tional records from Georgia, Maryland, South Carolina, and Virginia. New records from New York provide the northernmost records from North America and the first records from New York state. In September, 1997, and September, 1999, single heavily worn males were collected in Ithaca (7 September 1997, along the shore of Beebe Lake, adjacent to the Cornell Uni- versity Campus, K. Anderson; and 12 Sep- tember 1999, on the Cornell Plantations, S. Munson). In July, 2000, I discovered a large population of M. sculpturalis in Ithaca, Tompkins County, New York at Penny Lane, Commonland Community (17 @, 2 3, 16 July; 50 2, 83 6, 21 July—12 Au- gust). All females were gathering pollen or nectar from Everlasting Pea, Lathyrus lati- folius L. [Fabaceae], a large-flowered le- gume. Most males were visiting L. latifol- ius, but smaller numbers were found on Melilotus alba Medikus. [Fabaceae] (14 specimens), Asclepias syriaca L. [Asclepia- daceae] (10), Cirsium vulgare (Savi) Te- nore. [Asteraceae] (2), and Coronilla varia L. [Fabaceae] (1). Male M. sculpturalis were observed to enter holes made by car- penter bees (Xylocopa virginica Linnaeus) in a wooden beam beneath the roof of a carport near the Lathyrus patch. These holes presumably contained active nests of M. sculpturalis. Another population of M. sculpturalis was found at a patch of Lath- yrus latifolia 13 miles south of Syracuse, Onondaga County, New York (4 2, 5 Au- gust 2000). Additional records are from the Cornell University Campus (Minns Garden, 4 6, 13-14 August 2000, E. R. Hoebeke and JSA, ex Perovskia artemesioides [Lam- iaceae]) and Staunton, Augusta Co., Virgin- ia (1 2, 1 August 2000, M. E. Carter, ex Verbascum thapsus L. [Scrophulariaceae]). DISCUSSION All of the adventive species mentioned above nest in plant stems or pre-formed cavities, except for Hoplitis anthocopoides, which builds morter-and-pebble nests on VOLUME 103, NUMBER 1 the surfaces of boulders. The habit of build- ing concealed nests above ground in or on portable and sometimes man-made sub- strates, characteristic of many Hylaeinae, Megachilidae, and Xylocopinae (Apidae), facilitates long-distance transport of nests within plants, wood products, and other items of trade. All but one of the species listed by Cane (in press) as accidentally in- troduced to North America belong to these three taxa. The exception is Andrena (Taen- iandrena) wilkella (Kirby), a ground-nest- ing andrenid probably introduced to north- eastern North America from Europe (Mal- loch 1918) and now abundant throughout eastern North America. A. wilkella forages on diverse plants and can therefore be con- sidered as polylectic (Cane, in press), but shows a decided preference for Fabaceae (Westrich 1990). In Ithaca (where it has been present since at least 1904) pollen-col- lecting females are abundant visitors to ex- otic Fabaceae, especially Melilotus alba Desr., from late-spring to mid-summer. Cane (in press) may be correct to assert that nests of fossorial bees are unlikely to enter the United States from other countries due to regulations restricting the importation of soil. However, ground-nesting bee species have been transported long distances within the United States. The ground-nesting hal- ictid, Lasioglossum (Dialictus) imitatum, native to eastern North America, has been reported as adventive in California (Moure and Hurd 1987). In addition, at least two species of Lasioglossum (Dialictus) are now present in Hawaii. More systematic surveys of local bee faunas are needed in order to detect newly adventive species, to monitor the spread of these species, and to assess their impact on the native fauna and flora. Given recent in- creases in transcontinental trade, particular- ly with Asia, North American entomolo- gists should expect many more additions to our insect fauna in the near future, not all of them as benign and attractive as Me- gachile sculpturalis and Hylaeus hyalina- tus. 189 ACKNOWLEDGMENTS I thank E. R. Hoebeke for encouraging me to publish my records of adventive bees, identifying Megachile sculpturalis, and commenting on the manuscript, D. L. Ste- phan for confirming the identity of M. sculpturalis, J. H. Cane for sharing his un- published manuscript about adventive bees, R. R. Snelling for confirming the identity of H. hyalinatus and H. punctatus and for commenting on the manuscript, J. K. Lieb- herr for assisting with the photographs, B. Ehmer for translating German references, and B. N. Danforth and two anonymous re- viewers for commenting on the manuscript. LITERATURE CITED Barthell, J. E, G. W. Frankie, and R. W. Thorp. 1998. Invader effects in a community of cavity nesting megachilid bees (Hymenoptera: Megachilidae). Environmental Entomology 27: 240-247. Batra, S. E. T. 1998. Biology of the Giant Resin Bee, Megachile sculpturalis Smith, a conspicuous new immigrant in Maryland. The Maryland Naturalist 42: 1-3. Beaumont, J. De. 1958. Les Hyménopteres aculéates du Parc National Suisse et des régions limitrophes. Ergebnisse der wissenschaftlichen Unterssuchun- gen der schweizerischen Nationalparks, N. FE 6: 145-235. Cane, J. H. In press. Exotic non-social bees (Hyme- noptera: Apoidea) in North America: ecological implications. /n Strickler, K. and J. H. Cane, eds. For non-native crops, whence pollinators of the future? Thomas Say Publications. Dathe, H. H. 1980. Die Arten der Gattung Hylaeus F in Europa (Hymenoptera: Apoidea, Colletidae). Mitteilungen Zoologisches Museum in Berlin 56: 207-294. Eickwort, G. C. 1970. Hoplitis anthocopoides, a Eu- ropean mason bee established in New York state (Hymenoptera: Megachilidae). Psyche 77: 190— 201. . 1980. Two European species of Chelostoma established in New York state (Hymenoptera: Me- gachilidae). Psyche 87: 315-323. Elfving, R. 1951. Die Gattung Prosopis Fabr. in Finn- land. Notulae Entomologicae 31: 67—92. Hoebeke, E. R. and A. G. Wheeler, Jr. 1999. Anthidium oblongatum (Illiger): an Old World bee (Hyme- noptera: Megachilidae) new to North America, and new North American records for another ad- ventive species, A. manicatum (L.). The Univer- sity of Kansas Natural History Museum Special Publication 24: 21—24. 190 Hopper, D. and H. V. Daly. 1995. Range extensions of Hylaeus (Hymenoptera: Colletidae) on Kauai. Re- cords of the Hawaii Biological Survey 42: 35. Hurd, P. D., Jr. 1979. Superfamily Apoidea, pp. 1741— 2209. In Krombein, K. V., P. D. Hurd, Jr, D. R. Smith, and B. D. Burks, eds. Catalog of Hyme- noptera of America north of Mexico. Volume 2. Smithsonian Institution Press, Washington, D.C. Jaycox, E. R. 1967. An adventive Anthidium in New York state (Hymenoptera: Megachilidae). Journal of the Kansas Entomological Society 40: 124— 126. Malloch, J. R. 1918. Occurrence of a European solitary bee (Andrena wilkella Kirby) in the Eastern Unit- ed States. Proceedings of the Biological Society of Washington 31: 61—64. Mangum, W. A. and R. W. Brooks. 1997. First records of Megachile (Callomegachile) sculpturalis Smith (Hymenoptera: Megachilidae) in the continental United States. Journal of the Kansas Entomolog- ical Society 70: 140-142. Michener, C. D. 2000. Bees of the world. The Johns Hopkins University Press, Baltimore and London. 913 pp. Mitchell, T. B. 1960. Bees of the eastern United States. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Volume 1. North Carolina Agricultural Experi- ment Station Technical Bulletin Number 141, Ra- leigh. 538 pp. Snelling, R. R. 1966. Studies on North American bees of the genus Hylaeus 3. The Nearctic subgenera (Hymenoptera: Colletidae). Bulletin of the South- ern California Academy of Sciences 65: 164-175. . 1983. Studies on North American bees of the genus Hylaeus. 6. an adventive Palearctic species in southern California (Hymenoptera: Colletidae). Bulletin of the Southern California Academy of Sciences 82: 12-16. Thorp, R. W. 1996. Resource overlap among native and introduced bees in California, pp. 143-151. In Matheson, A., S. L. Buchmann, C. O’Toole, P. Westrich, and I. H. Williams, eds. The Conser- vation of Bees. Academic, London. Toro, H., Y. Frederick, and A. Henry. 1989. Hylaeinae (Hymenoptera: Colletidae), a new sub-family of bees for the Chilean fauna. Acta Entomologica Chilena 15: 201—204. Westrich, P. 1990. Die Wildbienen Baden-Wiirttem- bergs. Spezieller Teil: Die Gattungen und Arten. Eugen Ulmer GmbH & Co., Stuttgart. pp. 437— O72: PROC. ENTOMOL. SOC. WASH. 103(1), 2001, pp. 191-206 LIFE HISTORY AND DESCRIPTION OF IMMATURE STAGES OF NEASPILOTA FOOTEI FREIDBERG AND MATHIS (DIPTERA: TEPHRITIDAE) ON ASTER OCCIDENTALIS (NUTTALL) TORREY AND A. GRAY (ASTERACEAE) IN SOUTHERN CALIFORNIA RICHARD D. GOEDEN Department of Entomology, University of California, Riverside, CA 92521, U.S.A. (e- mail: richard.goeden @ucr.edu) Abstract.—Neaspilota footei Freidberg and Mathis is a univoltine, monophagous or nearly monophagous, fruit fly (Diptera: Tephritidae) developing in the flower heads of Aster occidentalis (Nuttall) Torrey and A. Gray (Asteraceae) belonging to the subtribe Asterinae of the tribe Astereae in southern California. It also has been reported from Conyza (as Erigeron) canadensis (L.) (Cronquist) (and as E. pusillus) and Chrysothamnus sp., but both of these host records need confirmation. The egg, first-, second- and third- instar larvae and puparium are described and figured, and these immature stages are compared with those of other Neaspilota. The anterior thoracic spiracles of the second and third instars of N. footei have three papillae. The second and third instars also have an undetermined number of oral ridges with dentate posterior margins in a vertical series lateral to the oral cavity. The appearance and placement of these oral ridges is a distin- guishing generic larval character. The larvae feed mainly on the corollas of florets and ovules as first instars; on corollas, ovules, and soft achenes as second instars; but as third instars, they feed mainly on ovules and soft achenes. A single annual generation is pro- duced on A. occidentalis in southern California. The life cycle is of the aggregative type and overwintering mainly occurs as puparia attached to uneaten achenes and fragments thereof in dead flower heads. Pteromalus sp. (Hymenoptera: Pteromalidae) was reared as a solitary, larval-pupal endoparasitoid of N. footei; Eurytoma sp. (Hymenoptera: Eury- tomidae) and Mesopolobus sp., as probable, solitary, larval-pupal endoparasitoids. Key Words: Insecta, Neaspilota, Aster, Asteraceae, nonfrugivorous Tephritidae, biology, taxonomy of immature stages, allopatry, flower-head feeding, aggregative life cycle, seed predation, parasitoids Revision of the genus Neaspilota (Dip- tera: Tephritidae) by Freidberg and Mathis (1986) facilitated identification of speci- mens reared from California Asteraceae (Goeden 1989) and stimulated several life-history studies, including those on N. viridescens Quisenberry (Goeden and Headrick 1992), N. wilsoni Blanc and Foote (Goeden and Headrick 1999), N. signifera (Coquillett) (Goeden 2000a), N. aenigma Freidberg and Mathis (Goeden 2000b), N. appendiculata Freidberg and Mathis (Goeden 2000c), N. pubescens Freidberg and Mathis (Goeden 2000d), and N. achilleae Johnson (Goeden 2001). This paper describes the immature stages and life history of an eighth species from California, N. footei Freidberg and Math- is, and is the last paper in this series on Neaspilota. 192 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MATERIALS AND METHODS The present study was based in large part on dissections of samples of flower heads of Aster occidentalis (Nuttall) Torrey and A. Gray (Asteraceae) mainly collected on the north shore of Big Bear Lake at 2,020- m elevation, Serrano Meadows Picnic Area, San Bernardino Nat. Forest (N Section), SW San Bernardino Co., during 1996 and 1997. One-liter samples of excised, imma- ture and mature flower heads containing eggs, larvae, and puparia were transported in cold-chests in an air-conditioned vehicle to the laboratory and stored under refrig- eration for subsequent dissection, photog- raphy, description, and measurement. Three eggs, six first-, 14 second-, and 15 third- instar larvae and seven puparia dissected from flower heads were preserved in 70% EtOH for scanning electron microscopy (SEM). Additional prepuparia and puparia were placed in separate, glass shell vials stoppered with absorbant cotton and held in humidity chambers at room temperature for adult and parasitoid emergence. Specimens for SEM were hydrated to distilled water in a decreasing series of acidulated EtOH. They were osmicated for 24 h, dehydrated through an increasing series of acidulated EtOH and two, 1-h immersions in hexa- methyldisilazane (HMDS), mounted on stubs, sputter-coated with a gold-palladium alloy, and studied and photographed with a Philips XL-30 scanning electron micro- scope in the Institute of Geophysics and Planetary Physics, University of California, Riverside. Most adults reared from isolated prepu- paria and puparia were individually caged in 850-ml clear-plastic, screened-top cages with a cotton wick and basal water reser- voir and provisioned with a strip of paper toweling impregnated with yeast hydroly- zate and sucrose. These cages were used for studies of longevity and sexual maturation in the insectary of the Department of En- tomology, University of California, River- side, at 25 + 1°C, and 14/10 (L/D) photo- period. A single pair of a virgin male and female obtained from emergence cages also were held in a clear-plastic, petri dish pro- visioned with a flattened, water-moistened pad of absorbant cotton spotted with honey (Headrick and Goeden 1994) for observa- tions of their courtship and copulation be- havior. Plant names used in this paper follow Hickman (1993) and Bremer (1994); te- phritid names and adult terminology follow Foote et al. (1993). Terminology and tele- graphic format used to describe the imma- ture stages follow Goeden (2000a, b, c, d, 2001), Goeden et al. (1998), Goeden and Headrick (1992, 1999), Goeden and Teerink (1997a, b, 1998, 1999a, b), Teerinkvand Goeden (1999), and our earlier works cited therein. Means + SE are used throughout this paper. Voucher specimens of N. footei immature stages, adults, and parasitoids re- side in my research collections. RESULTS AND DISCUSSION TAXONOMY Adult.—Neaspilota footei was described by Freidberg and Mathis (1986, p. 49-52), who pictured the unpatterned wing, along with drawings (p. 50) of the lateral aspect of the head, male right foretarsus, epan- drium, distiphallus, epandrium and cerci, aculeus and its apex enlarged, and sper- matheca. Immature stages.—The egg, first-, sec- ond- and third-instar larvae, and puparium are described below for the first time. Egg: Fifty-four eggs dissected from field-collected flower heads were white, opaque, smooth, elongate-ellipsoidal, 0.62 + 0.004 (range, 0.56—0.68) mm long, 0.17 + 0.002 (range, 0.16—0.20) mm wide, smoothly rounded at tapered basal end (Fig. 1A); pedicel button-like, 0.02 mm long, cir- cumscribed apically by semicircular or el- liptical aeropyles arranged singly or in two compressed or separate rows parallel to the long axis of the egg (Figs. 1B, C). The egg of N. footei is similar in shape VOLUME 103, NUMBER 1 Acc.Y Spot Magn Det WD -————_——_ 200 uum 10.0kV 3.0 129x SE 25.3 AccV SpotMagn Det WD -F——————+| 104m 10.0kV 3.0 2578 __SE 261 Le) eecae is oe or = =~ Say sg ae ee pe Det WO 10.0 kV 3.0, 2074x SE 2655 AccV Spot Magn ie} we Sa Fig. 1. Egg of Neaspilota footei: (A) habitus, an- terior end to left; (B) pedicel showing pattern of aero- pyles; (C) pedicel of a different egg with its aeropyles. to those of N. viridescens (Goeden and Headrick 1992), N. wilsoni (Goeden and Headrick 1999), and N. appendiculata (Goeden 2000c). The egg of N. footei is about 10% longer on average than the eggs of N. viridescens (Goeden and Headrick 193 1992), but about 30% shorter and narrower than those of N. wilsoni (Goeden and Head- rick 1999) and about 10% shorter than those of N. appendiculata (Goeden 2000c); the latter two species have the largest adults among southern California Neaspilota (Freidberg and Mathis 1986). The aeropyles of N. appendiculata are arranged in one to three rows around the apex of the pedicel (Goeden 2000c), those of N. viridescens are irregularly scattered around the apex (Goe- den and Headrick 1992), and those of N. wilsoni fully cover the pedicel (Goeden and Headrick 1999). First instar: White, elongate-cylindrical, bluntly rounded anteriorly and posteriorly (Fig. 2A); body segments well-defined, nearly free of minute acanthae; gnathoce- phalon smooth, lacking oral ridges, with pair of prominent integumental petals dor- sad of mouthhooks (Figs. 2B-1, C-5); dor- sal sensory organ a well-defined, dome- shaped papilla (Figs. 2B-2, C-1, D-1); an- terior sensory lobe (Figs. 2B-3, C-2, D-2) bears terminal sensory organ (Figs. 2C-3, D-3); lateral sensory organ (Fig. 2D-4), su- pralateral sensory organ (Fig. 2D-5), and pit sensory organ (Fig. 2D-6); stomal sense organ ventrolaterad of anterior sensory lobe (Figs. 2B-4, C-4, D-7), integumental petal (Figs. 2B-1, C-5, D-8) fused laterally with stomal sense organ (Figs. 2B-4, C-4, D-7); mouthhook bidentate (Figs. 2B-5, C-6); median oral lobe laterally flattened (Figs. 2B-6, C-7), posterolateral pair of verruci- form sensilla on gnathocephalon (Figs. 2B- 7, C-8); meso- and metathoracic and ab- dominal lateral spiracular complexes not seen; caudal segment with two stelex sen- silla, dorso- and ventrolaterad of posterior spiracular plate (Fig. 2E-1), neither stelex sensillum basally ringed with minute acan- thae; posterior spiracular plate bears two ovoid rimae, ca. 0.004 mm in length (Fig. 2E-2), and four interspiracular processes, each with two to four branches, longest measuring ca. 0.01 mm (Fig. 2E-3); inter- mediate sensory complex with one stelex 194 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON AccV SpotMagn Det WD /-——————_{ 200m A 10.0kV 3.0 175x SE 26.4 Fig. 2. First instar of Neaspilota footei: (A) habitus, anterior to left; (B) gnathocephalon, frontolateral view, | — integumental petal, 2 — dorsal sensory organ, 3 — anterior sensory lobe, 4 — stomal sense organ, 5 — mouthhook, 6 — median oral lobe, 7 — verruciform sensillum; (C) | — dorsal sensory organ, 2 — anterior sensory lobe, 3 — terminal sensory organ, 4 — stomal sense organ, 5 — integumental petal, 6 — mouthhook, 7 — median oral lobe, 8 — verruciform sensillum; (D) 1 — dorsal sensory organ, 2 — anterior sensory lobe, 3 — terminal sensory organ, 4 — lateral sensory organ, 5 — supralateral sensory organ, 6 — pit sensory organ, 7 — stomal sensory organ, 8 — integumental petal; (E) caudal segment, 1 — posterior spiracular plate, 2 — rima, 3 — interspiracular process; (F) intermediate sensory complex, 1 — stelex sensillum, 2 — medusoid sensillum. VOLUME 103, NUMBER 1 sensillum (Fig. 2F-1) and one medusoid sensillum with short papillae (Fig. 2F-2). The first instar is similar in general hab- itus (Fig. 2A) to that of N. viridescens (Goeden and Headrick 1992), N. wilsoni (Goeden and Headrick 1999), N. signifera (Goeden 2000a), N. aenigma (Goeden 2000b), and N. appendiculata (Goeden 2000c). However, unlike N. viridescens, but like N. wilsoni, N. aenigma, and N. appen- diculata, the dorsal sensory organ of the first instar of N. footei is well defined (Figs. 2B-2, C-1, D-1), as is the anterior sensory lobe (Figs. 2B-3, C-2, D-2) and integumen- tal petal (Figs. 2B-1, C-5, D-8). Also, the pit sensory organ (Fig. 2D-6), not seen in N. viridescens (Goeden and Headrick 1992) and hidden in specimens viewed of N. sig- nifera (Goeden 2000a), is present in N. footei (Fig. 2D-6), as it is in N. wilsoni (Goeden and Headrick 1999), N. aenigma (Goeden 2000b), and N. appendiculata (Goeden 2000c). A fused integumental pet- al and stomal sense organ also was reported in first instars of N. wilsoni (Goeden and Headrick 1999), N. signifera (Goeden 2000a), N. aenigma (Goeden 2000b), and N. appendiculata (Goeden 2000c), as well as Trupanea vicina (Wulp) (Goeden and Teerink 1999b), but these structures were separated in N. viridescens (Goeden and Headrick 1992). Two stelex sensilla dorso- and ventrola- terad of each posterior spiracular plate in the first instar of N. footei, N. aenigma (Goeden 2000b), and N. appendiculata (Goeden 2000c) agreed with the four such sensilla reported to ring the caudal segment of N. wilsoni (Goeden and Headrick 1999), but not the 10 sensilla reported to ring the caudal segment of N. viridescens (Goeden and Headrick 1992). The last number is probably erroneous, as discussed by Goe- den (2000b). Lastly, N. footei lacks the mi- nute acanthae reported to ring basally the lateral stelex sensilla on the caudal segment in N. wilsoni (Goeden and Headrick 1999), where the upright acanthae among them number one to three and are pointed; in N. 195 aenigma (Goeden 2000b), where the up- right acanthus is solitary and rounded api- cally; and in N. appendiculata (Goeden 2000c), where the three basal acanthae are present, but poorly developed. Second instar: White, elongate-cylindri- cal, rounded anteriorly, truncated postero- dorsally (Fig. 3A), body segments well-de- fined, anterior third of meso- and metatho- rax and abdominal segments A-1 through A-6 circumscribed anteriorly by minute acanthae also present on abdominal pleura; dorsal sensory organ not well-defined, flat- tened (Figs. 3B-1, C-1, D-1); anterior sen- sory lobe (Figs. 3C-2, D-2) with terminal sensory organ (Figs. 3C-3, D-3), lateral sensory organ (Fig. 3D-4), supralateral sen- sory organ (Fig. 3D-5), and pit sensory or- gan (Fig. 3D-6); stomal sense organ (Figs. 3C-4, D-7) ventrolaterad of anterior senso- ry lobe; mouthhook bidentate (Figs. 3B-2, C-5, D-8); median oral lobe laterally com- pressed (Fig. 3C-6); four papilliform, inte- gumental petals in each of two separate rows dorsad of each mouthhook (Figs. 3C- 7, D-9); at least eight oral ridges, dentate along posterior margins, in vertical series laterad of oral cavity (Figs. 3B-3, C-8, D- 10); prothorax, circumscribed anteriorly by at least two rows of posteriorly-directed, minute acanthae (Figs. 3B-4, C-9, D-11); anterior thoracic spiracle with three, doli- form papillae (Fig. 3E); lateral spiracular complexes not seen; caudal segment with two stelex sensilla dorsolaterad and ventro- laterad of posterior spiracular plate; poste- rior spiracular plate bears three ovoid rimae (Fig. 3F-1), ca. 0.011 mm long, and four interspiracular processes (Fig. 3F-2), each with two to four, simple branches with one or two, apical teeth, longest branch mea- suring 0.009 mm; intermediate sensory complex with a stelex sensillum (Fig. 3F- 3) and a medusoid sensillum (Fig. 3F-4). The habitus of the second instar of N. footei (Fig. 3A) is more like that of N. wil- soni (Goeden and Headrick 1999), N. sig- nifera (Goeden 2000a), N. aenigma (Goe- den 2000b), N. appendiculata (Goeden 196 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Spot Magn Det WD -F———]_ 204m AccV SpotMagn Det WD 10.0kV 3.0 1859x SE 253 t= ’ Fig. 3. Second instar of Neaspilota footei: (A) habitus, anterior to left; (B) gnathocephalon, lateral view, 1 — dorsal sensory organ, 2 — mouthhook, 3 — oral ridge, 4 — minute acanthae; (C) gnathocephalon, fronto- lateral view, 1 — dorsal sensory organ, 2 — anterior sensory lobe, 3 — terminal sensory organ, 4 — stomal sense organ, 5 — mouthhook, 6 — median oral lobe, 7 — integumental petal, 8 — oral ridge, 9 — minute acanthae, (D) lateral view of gnathocephalon, | — dorsal sensory organ, 2 — anterior sensory lobe, 3 — terminal sensory organ, 4 — lateral sensory organ, 5 — suprlateral sensory organ, 6 — pit sensory organ, 7 — stomal sense organ, 8 — mouthhook, 9 — integumental petal, 10 — oral ridge, 11 — minute acanthae; (E) anterior thoracic spiracle; (F) posterior spiracular plate, 1 — rima, 2 — interspiracular process, 3 — intermediate sensory complex, stelex sensillum, 4 — medusoid sensillum. VOLUME 103, NUMBER 1 2000c), N. pubescens (Goeden 2000d), and N. achilleae (Goeden 2001) than the “‘bar- rel-shaped”’ (doliform) second instar of N. viridescens (Goeden and Headrick 1992). The dorsal sensory organ of N. footei is not well-defined in the second instar (Figs. 3B- 1, C-1, D-1), like N. viridescens (Goeden and Headrick 1992), N. wilsoni (Goeden and Headrick 1999), N. aenigma (Goeden 2000b), and N. achilleae (Goeden 2001), but unlike N. signifera (Goeden 2000a), N. appendiculata (Goeden 2000c), and N. pu- bescens (Goeden 2000d), in which this or- gan is well-defined. The integumental pet- als of the second instars of all eight species studied to date are papilliform, but are only four in number above each mouthhook in N. achilleae (Goeden 2001), like N. signi- fera (Goeden 2000a), but are six in number in N. viridescens (Goeden and Headrick 1992) and N. pubescens (Goeden 2000d), seven in N. appendiculata (Goeden 2000c) and N. wilsoni (Goeden and Headrick 1999), and eight in N. aenigma (Goeden 2000b) and WN. footei (Figs. 3C-7, D-9). Compared to five papillae on the anterior spiracle of the second instar of in N. pu- bescens (Goeden 2000b) and eight in N. ap- pendiculata (Goeden 2000c), N. footei only has three papillae (Fig. 3E), like the three or four papillae reported in second instars of N. viridescens (Goeden and Headrick 1992), N. wilsoni (Goeden and Headrick 1999), N. signifera (Goeden 2000a), N. aenigma (Goeden 2000b), and N. achilleae (Goeden 2001). Finally, the interspiracular processes of N. footei (Fig. 3F-2) each bear two to four branches, like those of N. sig- nifera (Goeden 2000a), but not one to four branches like N. aenigma (Goeden 2000b) and N. pubescens (Goeden 2001), nor five to nine branches like N. viridescens (Goe- den and Headrick 1992), two to six branch- es like N. wilsoni (Goeden and Headrick 1999), nor four branches like N. appendi- culata (Goeden 2000c) and N. achilleae (Goeden 2001). Third instar: Pale yellow, elongate-ellip- soidal, with posterior spiracular plate dark 197 brown to black, tapering and truncated an- teriorly; posterior spiracular plate on caudal segment flattened and upturned dorsally ca. 60° (Fig. 4A), minute acanthae circum- scribe anterior fifth of mesothorax, anterior fourth of metathorax and first abdominal segment (A1), and all of remaining abdom- inal segments, except for pleura of segment A2, posterior half of A7, and most of A8, dorsa of abdominal segments A7 and A8 also covered with minute acanthae; gnath- ocephalon conical (Fig. 4B); dorsal sensory organ defined by a crescentric fold (Figs. 4C-1, D-1) attached dorsally (Figs. 4C-1, D-1) and punctured peripherally by pores (Fig. 4D-2); anterior sensory lobe (Figs. 4B-1, C-2, D-3) bears terminal sensory or- gan (Fig. 4D-4), lateral sensory organ (Fig. 4D-5), supralateral sensory organ (Fig. 4D- 6), and pit sensory organ (Fig. 4D-7); 10 papilliform (including two small, central) or spatulate, integumental petals in two rows above each mouthhook (Figs. 4C-3, E-1); at least seven oral ridges (Fig. 4C-4), toothed ventrally and lateral to oral cavity; stomal sense organ (Figs. 4C-5, D-8) ven- trolaterad of anterior sensory lobe; mouth- hook (Figs. 4B-2, C-6, E-2) tridentate; me- dian oral lobe reduced, laterally flattened, (Fig. 4E-3); gnathocephalon bears at least three verruciform sensilla posteriorad of each anterior sensory lobe and oral ridges (Figs. 4B-3, C-7); prothorax circumscribed by minute acanthae (Fig. 4B-4); verruci- form sensilla circumscribe prothorax pos- teriorad of minute acanthae (Fig. 4B-5); an- terior thoracic spiracle on posterior margin of prothorax bears three doliform papillae (Fig. 4F); mesothoracic and metathoracic lateral spiracular complexes with five and four verruciform sensilla, respectively, each complex alligned vertically, mesothoracic spiracle greatly reduced (not shown) and metathoracic spiracle not found; lateral spi- racular complex of abdominal segment A- 1 with a spiracle (Fig. 5A-1) and four ver- ruciform sensilla, one dorsoposteriorad of, one posteriorad of, and two ventroposter- iorad of the spiracle, the upper and first 198 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON AccV Spot Magn Acc.V Spot Mag 10.0kV 3.0 36x : 10.0kV 30 491, Det WD Fig. 4. Third instar of Neaspilota footei: (A) habitus, anterior to left; (B) gnathocephalon, frontolateral view, | — anterior sensory lobe, 2 — mouthhook, 3 — verruciform sensillum, 4 — minute acanthae, 5 — verruciform sensilla; (C) gnathocephalon, frontolateral view, | — dorsal sensory organ, 2 — anterior sensory lobe, 3 — integumental petal, 4 — oral ridge, 5 — stomal sense organ, 6 — mouthhook, 7 — verruciform sensillum; (D) anterior sensory lobe, | — dorsal sensory organ, 2 — peripheral pore, 3 — anterior sensory lobe, 4 — terminal sensory organ, 5 — lateral sensory organ, 6 — supralateral sensory organ, 7 — pit sensory organ, 8 — stomal sense organ; (E) oral cavity, 1 — integumental petal, 2 — mouthhook, 3 — median oral lobe; (F) anterior thoracic spiracle. VOLUME 103, NUMBER 1 199 a ‘ . Ni pe BccV- “Spot Magn _ Det WD SpohMagn Det WD /}——————_] "5 Eg 10.0 kV 30:31, 0x7 PabO.0KM.0 Tysix “SE. 244 _ TSE: 24.3 | Fig. 5. Third instar of Neaspilota footei, continued: (A) first abdominal, lateral spiracular complex, 1 — spiracle, 2 — verruciform sensilla; (B) caudal segment, 1 — posterior spiracular plate, 2 — dorsolateral stelex sensillum, 3 — lateral stelex sensillum, 4 — ventrolateral stelex sensillum, 5 — intermediate sensory complex; (C) lateral stelex sensillum; (D) ventrolateral stelex sensillum, (E) posterior spiracular plate, | — rima, 2 — interspiracular process, (F) intermediate sensory complex, | — stelex sensillum, 2 — medusoid sensillum. ventroposterior sensilla aligned ventrally 5B-3, C), and ventrolaterad (Figs. 5B-4, D) (Fig. 5SA-2), as are the other two posteriorad of posterior spiracular plate, each sensillum of the first pair (Fig. SA-2); a stelex sensil- surrounded basally by four or five, minute lum dorsolaterad (Fig. 5B-2), laterad (Figs. acanthae (Figs. 5C, D), smaller than other 200 minute acanthae (Figs. 5C, D); each pos- terior spiracular plate (Fig. 5E) bears three ovoid rimae, ca. 0.014 mm in length (Fig. 5E-1), and four, simple or forked interspi- racular processes, each 0.014 mm long (Fig. 5E-2); intermediate sensory complex (Fig. SF) with a stelex sensillum (Fig. 5F-1), sur- rounded basally by reduced minute acan- thae (Fig. 5F-1), and a medusoid sensillum (Fig. 5F-2). 7 The habitus of the third instar of N. footei is like that reported for N. viridescens (Goe- den and Headrick 1992), N. wilsoni (Goe- den and Headrick 1999), N. signifera (Goe- den 2000a), N. aenigma (Goeden 2000b), N. appendiculata (Goeden 2000c), N. pu- bescens (Goeden 2000d), and N. achilleae (Goeden 2001). The pattern of the minute acanthae that circumscribe the body seg- ments is unique for N. footei (see descrip- tion above), but closest to N. achilleae (Goeden 2001) in that the acanthae increase their coverage posteriorly from the anterior fifths of the thoracic segments to include nearly all of the abdominal segments, ex- cept A7 and A& in the former species; whereas, in N. signifera (Goeden 2000a), N. appendiculata (Goeden 2000c) and N. pu- bescens (Goeden 2000d), the anterior part of each body segment is circumscribed, in N. aenigma the anteriors, pleura, and pos- teriors of each segment are circumscribed (Goeden 2000b); in N. wilsoni, all interseg- mental areas and all abdominal segments except the pleura are circumscribed (Goe- den and Headrick 1999); and in N. virides- cens, the intersegmental areas are free of acanthae (Goeden and Headrick 1992). Like N. viridescens (Goeden and Headrick 1992), N. wilsoni (Goeden and Headrick 1999), N. pubescens (Goeden 2000d), and N. achilleae (Goeden 2001), but unlike N. signifera (Goeden 2000a), N. aenigma (Goeden 2000b), and N. appendiculata (Goeden 2000c), the dorsal sensory organ is not well-defined and flattened, and not dome-shaped, in the third instar of N. footei (Figs. 4C-1, D-1). Additional similarities involved the inte- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON gumental petals in the third instars of all six congeners examined to date, all of which are arranged in a double row above each mouthhook and papilliform or spatulate (Goeden and Headrick 1992, 1999; Goeden 2000a, b, c, d, 2001). The integumental pet- als differ in numbers among species and generally increase in number between the last two instars. The stomal sense organ of the third instar of N. footei was obscured by debris in my specimens, and could thus not be compared readily with other congeners (Figs. 4C-5, D-8), but it appears at least as complex as the stomal sense organ of the second instar (Fig. 3D-4). The third instars of all seven species of Neaspilota examined to date have oral ridg- es with dentate ventral margins character- istically arranged in vertical series ventro- laterad of the dorsal sensory organ and lat- erad of the oral cavity (Goeden and Head- rick 1992, 1999; Goeden 2000a, b, c, d, 2001; Fig. 4C-4). Though unfortunately partly hidden and not fully counted in N. footei (i.e., at least seven, Fig. 4C-4) and N. achilleae (Goeden 2001), the oral ridges number eight in N. pubescens (Goeden 2000d), seven or eight in the third instar of N. aenigma (Goeden 2000b), seven in N. appendiculata (Goeden 2000c), but six in the second and third instars of the other three congeners examined to date (Goeden and Headrick 1992, 1999; Goeden 2000a). The appearance and arrangement of these oral ridges appears to be a generic charac- ter; however, Goeden (2000c, d) confirmed that the oral ridges vary in number among third instars of some Neaspilota species. Also, the most ventral, eighth oral ridge of N. pubescens is not ventrally toothed (Goe- den 2000d). The third instars of Trupanea imperfecta (Coquillett), T. jonesi Curran, T. nigricornis (Coquillett), 7. pseudovicina (Hering), T. signata Foote, and T. wheeleri Curran also bear serrated oral ridges (Goe- den and Teerink 1997b, 1998, 1999a; Goe- den et al. 1998; Knio et al. 1996; Teerink and Goeden 1999), but these oral ridges ap- pear to be fewer in number, and are not ar- VOLUME 103, NUMBER 1 A ie ee Acc.V Spot Magn 10.0 kV 3.0 30x Det WD ->————______4 1 SE 244 nn A Fig. 6. Puparium of Neaspilota footei: (A) habitus, anterior to left; (B) anterior end, | — invagination scar, 2 — anterior thoracic spiracle; (C) caudal segment, 1 — rima, 2 — interspiracular process, 3 — intermediate sensory complex. ranged in a more or less regular, vertical row laterad of the mouthhook, as in Neas- pilota. The mouthhooks of the third instars of N. achilleae, N. aenigma, N. appendiculata, 201 N. footei, N. signifera, N. viridescens, and probably N. pubescens, are tridentate (Goe- den and Headrick 1992, Goeden 2000a, b, c, d; Goeden 2001; Figs. 4C-6, E-2); where- as, those of the third instar of N. wilsoni are bidentate (Goeden and Headrick 1999). Such interspecific differences in dentation are supported by our findings that the mouthhooks of third-instar Trupanea vicina (Wulp) are bidentate; whereas, those of 12 other congeners examined from California are tridentate (Goeden and Teerink 1999b and citations therein). The number and appearance of the stelex sensilla surrounding the posterior spiracular plate differ among the Neaspilota species examined to date. These number only four in the first instars of N. wilsoni (Goeden and Headrick 1999), N. aenigma (Goeden 2000b), and N. appendiculata (Goeden 2000c), but, unfortunately, were not ob- served with N. signifera (Goeden 2000a) or N. achilleae (Goeden 2001). This count of stelex sensilla remains at four in the second instars of N. aenigma (Goeden 2000b), N. appendiculata (Goeden 2000c), N. pubes- cens (Goeden 2000d), and N. achilleae (Goeden 2001), and increases to six in third instars of N. wilsoni (Goeden and Headrick 1999), N. aenigma (Goeden 2000b), N. ap- pendiculata (Goeden 2000c), and N. footei (Figs. SC, D). There are inter-instar (intra- specific) and interspecific differences in the incidence and appearance of the minute acanthae that may ring these stelex sensilla basally, but this was not recognized, studied or recorded by my coworkers and me until recently (Goeden 2000b, c, d, 2001). The stelex sensilla of the third instar of N. footei are surrounded by four or five, hemispher- ical, smaller-than-usual, minute acanthae (Fig. 5C, D); however, such basal minute acanthae near these stelex sensilla are lack- ing in the first and second instars (Fig. 2F- 3): Puparium: Mostly gray, with thoracic and posterior two or three segments black- ened, ellipsoidal, and smoothly rounded at both ends (Fig. 6A); anterior end bears the 202 invagination scar (Fig. 6B-1) and anterior thoracic spiracles (Fig. 6B-2); caudal seg- ment circumscribed by minute acanthae; three stelex sensilla, dorsolaterad, laterad, and ventrolaterad of posterior spiracular plates; posterior spiracular plate bears three broadly elliptical rimae (Fig. 6C-1), and four, simple or forked, interspiracular pro- cesses (Fig. 6C-2); intermediate sensory complex with a medusoid sensillum and a stelex sensillum (Fig. 6C-3). Thirty-five pu- paria averaged 3.32 + 0.04 (range, 2.42— 3.70) mm in length; 1.13 + 0.018 (range, 0.94—1.42) mm in width. DISTRIBUTION AND HOSTS Freidberg and Mathis (1986) described the known distribution of N. footei as form- ing, “2.. a wide; inverted .U;, i.e., along both coasts and across southern Canada (Canada: Ontario, Saskatchewan, United States: Alabama, California, Connecticut, Georgia, Maryland, Massachusetts, New York, North Carolina, Ohio, Oregon, Penn- sylvania, Virginia).’’ They and Foote et al. (1993) both mapped this distribution. Freid- berg and Mathis (1986, p. 52) also re- marked that: ‘Although the distribution of this species, as compared to others of the genus, is suspect, we could find no significant differences be- tween the eastern and western populations. The distributional pattern, however, could be an artifact, resulting from insufficient collect- ing in the Midwest and central-south United States.” The last quoted sentence probably also applies to the known distribution of N. achilleae in North America (Freidberg and Mathis 1986; Goeden 1989, 2001). Freidberg and Mathis (1986) listed Chry- sothamnus sp. and Erigeron canadensis L. as unconfirmed host records for single males “‘ex’’ each of these plants in Alpine Co., California, and Nassau Co., New York, respectively. Reed and Hughes (1970) and Holm et al. (1979) followed Cronquist (1943), who tranferred the latter species to Conyza, as C. canadensis (L.) Cronquist, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON commonly called “‘horseweed’’, a native, semi-cosmopolitan weed, common through- out the United States and southern Canada, and now found more or less worldwide (Hickman 1993). My rearing record from flower heads of Aster occidentalis is new (Freidberg and Mathis 1986, Goeden 1989). Aster occidentalis belongs to the subtribe Asterinae of the tribe Astereae, as does C. canadensis (Bremer 1994); however, I have never reared this or any other tephritid from seven samples of mature flower heads of the latter species, nor from three additional samples from C. bonariensis (L.) Cronquist (unpublished data). Chrysothamnus also be- longs to the tribe Astereae, but to a different subtribe, the Solidagininae (Bremer 1994), and only N. viridescens has been reared from this genus to date (Goeden 1989, Goe- den and Headrick 1992). Thus, the records for C. canadensis and Chrysothamnus both need confirmation. BIOLOGY Egg.—In each of 55, closed, preblossom, immature flower heads of Aster occidentalis an average of 1.7 + 0.18 (range, 1—6) eggs of N. footei were inserted separately, pedi- cel-last, through the phyllaries, between them, or among the apically apressed, but separable tips of the phyllaries. Thirty-five eggs (65%) rested with their long axes par- allel to the long axes of the flower heads (Fig. 7A). The remaining eggs rested at an- gles of 15—45°, and 45 of the 55 eggs (82%) were embedded for about half their lengths into the corollas of separate florets (Fig. 7B). An average of 1.5 + 0.24 (range, O- 6) florets/ovules per head were damaged by Oviposition, or about 2% of an average total of 62 (range, 30-85) florets/ovules/achenes counted in 53 preblossom, blossom, or postblossom flower heads. The receptacles of preblossom heads that contained eggs averaged 1.7 + 0.06 (range, 0.85—2.56) mm in diameter. Larva.—Upon eclosion, the 22 first in- stars found feeding in separate, closed, pre- blossom flower heads either tunneled into VOLUME 103, NUMBER 1 Bigw 7. preblossom flower head, (B) two eggs, partly inserted in separate corollas, (C) newly eclosed first instar, (D) second instar feeding on soft achene, (E) early third instar feeding on soft achenes, (F) overwintered puparium in dead flower head. Lines = 1 mm. an ovule, or into a corolla before entering the ovule to which the corolla was basally attached (Fig. 7C). The receptacles aver- aged 2.11 + 0.07 (range, 1.42—2.56) mm in diameter and an average of 3.9 + 0.46 (range, 1—8) florets/ovules was damaged in these 22 flower heads. No receptacle was abraded or pitted by larval feeding. Thus, about 6% (range, 2—-13%) of the average total of 62 florets/ovules/achenes in these flower heads were damaged by first instars. Second instars continued feeding on co- 203 Life stages of Neaspilota footei in Aster occidentalis: (A) egg between small florets in closed, rollas, ovules, or soft achenes in preblos- som and open flower heads. They fed with their bodies parallel, angled, or perpendic- ular to, and their mouthparts directed to- wards or away from the receptacles, but al- ways well above the receptacles (Fig. 7D). Receptacles of 26 flower heads containing second instars averaged 2.25 + 0.07 (range, 1.75—3.13) mm in diameter. These flower heads each contained an average of 1.04 + 0.04 (range, 1—2) larvae that had damaged an average of 8.3 + 1.08 (range, 1—26) flo- 204 rets/ovules/soft achenes, or about 13% (range, 2-42%) of the average total of 62 florets/ovules/soft achenes per flower head. Third instars (Fig. 7E) continued to feed mainly on ovules and soft achenes in pre- blossom, blossom, or postblossom flower heads. Fourteen flower heads that averaged 2.39 + 0.09 (range, 1.75—2.85) mm in di- ameter each contained a single third instar or prepuparium. An average of 31.3 + 2.5 (range, 11—55) of the soft achenes therein were damaged, or about 50% (range, 18— 89%) of the average total of 62 ovules/soft achenes per flower head (Fig. 7E). Third instars in flower heads fed with their long axes oriented parallel to (Fig. 7E) or at various angles to the receptacles. The receptacles were abraded or pitted in only four of a total of 69 (6%) flower heads con- taining third instars or overwintered pupar- ia, thus sap apparently does not constitute a major part of the diet of N. footei. Goeden and Headrick (1992, 1999) and Goeden (2000c, d) described and discussed sap feeding by N. viridescens, N. wilsoni, N. appendiculata, and N. pubescens, respec- tively. Upon completing feeding, the larvae oriented with their anterior ends away from the receptacles, retracted their mouthparts, and formed puparia, which constitute the overwintering stage (Headrick and Goeden 1998). Pupa.—The receptacles of 55 overwin- tered flower heads that contained a single puparium (Fig. 7F) each averaged 3.5 + 0.06 (range, 2.56—4.27) mm in diameter. Neaspilota appendiculata (Goeden 2000c), N. viridescens (Goeden and Headrick 1992), and N. wilsoni (Goeden and Head- rick 1999) overwinter in dead flower heads as prepuparia in protective cells formed of ovule, achene, chaff, pappus, and corolla fragments cemented together by liquid fe- ces and sap that hardened when dry. How- ever, N. footei puparia are partly surrounded and protected only by uneaten achenes and fragments tightly glued to the puparia with- in each flower head (Fig. 7F). The adults PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON emerge from these puparia in early spring (April). Adult.—Adults are probably long-lived in nature, but few were reared from over- wintered puparia, as many contained para- sitoids or were collected after the adults al- ready had emerged. Under insectary con- ditions, a single unmated male lived 104 days, and three virgin females averaged 80 + 15 (range, 52—104) days. Such lengthy longevities compare favorably with average adult longevities reported for adults of N. viridescens (Goeden and Headrick 1992), N. wilsoni (Goeden and Headrick 1999), N. signifera (Goeden 2000a), N. aenigma (Goeden 2000b), N. appendiculata (Goeden 2000c), N. pubescens (Goeden 2000d), and N. achilleae (Goeden 2001). The premating and mating behaviors of N. footei were not studied in the field, but were observed in one petri dish arena of the type found to be so useful with many other nonfrugivorous, tephritid species (Headrick and Goeden 1994). Premating behaviors observed with this pair of N. footei were abdominal pleural distension by males and side-stepping and swaying by both sexes while tracking their opposite sexes (Head- rick and Goeden 1994), and rapid wing ha- mation by both sexes (Headrick and Goe- den 1994). No trophallaxis or nuptial gift presentation was noted as reported with N. viridescens (Goeden and Headrick 1992). A single mating of more than 3-h duration was observed, which began during late af- ternoon at dusk, but was not viewed from beginning to end. This compared with av- erage durations of 153 min reported for N. achilleae (Goeden 2001), 190 min reported for N. aenigma (Goeden 1999b), 235 min reported for N. wilsoni (Goeden and Head- rick 1999), 238 min reported for N. signi- fera (Goeden 2000a), 285 min reported for N. appendiculata (Goeden 2000c), 318 min reported for N. viridescens (Goeden and Headrick 1992), and 1032 min reported for N. pubescens (Goeden 2000d). The copu- latory position was as described in detail for N. achilleae (Goeden 2001). VOLUME 103, NUMBER 1 Seasonal history.—The life cycle of N. footei in southern California follows an ag- gregative pattern (Headrick and Goeden 1994, 1998) in which the puparium is the principal overwintering stage. Come spring (April), adults emerge and aggregate on preblossom shoots of A. occidentalis to mate and subsequently oviposit in the small, newly-formed, closed, preblossom flower heads begining in late May. The lar- vae feed until fully grown, then pupariate in flower heads and overwinter. Neaspilota footei is the only species among eight spe- cies studied by us known to overwinter as puparia in southern California. Neaspilota signifera (Goeden 2000a), N. aenigma (Goeden 2000b), and N. achilleae (Goeden 2001) principally overwinter as adults; whereas, N. viridescens (Goeden and Head- rick 1992), N. wilsoni (Goeden and Head- rick 1999), N. appendiculata (Goeden 2000c), and N. pubescens (Goeden 2000d) also overwinter principally in dead flower heads, but as prepuparia. Natural enemies.—Many individual Pteromalus sp. (Hymenoptera: Pteromali- dae) were reared from puparia of N. footei as solitary, larval-pupal endoparasitoids. A male and female each of Eurytoma sp. (Hy- menoptera: Eurytomidae) and Mesopolobus sp. (Hymenoptera: Pteromalidae) were reared from mature flower heads of Aster occidentalis as probable solitary, larval-pu- pal endoparasitoids. ACKNOWLEDGMENTS I thank Andrew C. Sanders, Curator of the Herbarium, Department of Botany and Plant Sciences, University of California, Riverside, for identifications of plants men- tioned in this paper. Krassimer Bozhilov in the Institute of Geophysics and Planetary Physics, University of California, River- side, greatly facilitated my scanning elec- tron microscopy. The parasitoids were iden- tified by Harry E. Andersen, Huntington Beach, California. I also am grateful to Jeff Teerink and Kristine Gilbert for technical assistance and to David Headrick and Jeff 205 Teerink for their helpful comments on ear- lier drafts of this paper. This was the first manuscript of mine on a tephritid that Louie Blanc, recently deceased, did not preview. I miss him. LITERATURE CITED Bremer, K. 1994. Asteraceae Cladistics & Classifica- tion. Timber Press, Inc., Portland, Oregon. Cronquist, A. 1943. The separation of Erigeron from Conyza. Bull. of the Torrey Botanical Club 70: 629-632. Foote, R. H., EF L. Blanc, and A. L. Norrbom. 1993. Handbook of the Fruit Flies (Diptera: Tephritidae) of America North of Mexico. Cornell University Press, Ithaca, New York. Freidberg, A. and W. N. Mathis. 1986. Studies of Ter- elliinae (Diptera: Tephritidae): A revision of the genus Neaspilota Osten Sacken. Smithsonian Contributions to Zoology 439: 1-75. Goeden, R. D. 1987. Life history of Trupanea con- jJuncta (Adams) on Trixis californica Kellogg in southern California (Diptera: Tephritidae). The Pan-Pacific Entomologist 63: 284—291. . 1989. Host plants of Neaspilota in California (Diptera: Tephritidae). Proceedings of the Ento- mological Society of Washington 91: 164-168. . 2000a. Life history and description of im- mature stages of Neaspilota signifera (Coquillett) (Diptera: Tephritidae) on Hemizonia pungens (Hooker and Arnott) Torrey and A. Gray (Aster- aceae) in southern California. Proceedings of the Entomological Society of Washington 102: 69— 81. . 2000b. Life history and description of im- mature stages of Neaspilota aenigma Freidberg and Mathis (Diptera: Tephritidae) on Erigeron divergens Torrey and Gray (Asteraceae) in south- ern California. Proceedings of the Entomological Society of Washington 102: 384-397. . 2000c. Life history and description of im- mature stages of Neaspilota appendiculata Freid- berg and Mathis (Diptera: Tephritidae) on Ma- chaeranthera canescens (Pursh) A. Gray (Aster- aceae) in southern California. Proceedings of the Entomological Society of Washington 102: 519— 532. . 2000d. Life history and description of im- mature stages of Neaspilota pubescens Freidberg and Mathis (Diptera: Tephritidae) on Lessingia fi- laginifolia (Hooker and Arnott) M. A. Lane (As- teraceae). Proceedings of the Entomological So- ciety of Washington 102: 878-891. . 2001. Life history and description on imma- ture stages of Neaspilota achilleae Johnson (Dip- tera: Tephritidae) on Stephanomeria spp. (Aster- 206 aceae) in southern California. Proceedings of the Entomological Society of Washington 103:60—73. Goeden, R. D. and D. H. Headrick. 1992. Life history and descriptions of immature stages of Neaspilota viridescens Quisenberry (Diptera: Tephritidae) on native Asteraceae in southern California. Proceed- ings of the Entomological Society of Washington 94: 59-77. . 1999. Life history and description of imma- ture stages of Neaspilota wilsoni Blanc and Foote (Diptera: Tephritidae) on Hazardia squarrosa (Hooker and Arnott) E. Greene (Asteraceae). Pro- ceedings of the Entomological Society of Wash- ington 101: 897-909. Goeden, R. D. and J. A. Teerink 1997a. Life history and description of immature stages of Xenochaeta dichromata Snow (Diptera: Tephritidae) on Hier- acium albiflorum Hooker in central and southern California. Proceedings of the Entomological So- ciety of Washington 99: 597-607. . 1997b. Life history and description of im- mature stages of Trupanea signata Foote (Diptera: Tephritidae) on Gnaphalium luteo-album L. in southern California. Proceedings of the Entomo- logical Society of Washington 99: 748-755. . 1998. 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U.S. Department of Agricul- ture, Agricultural Research Service, Agriculture Handbook No. 366. 463 pp. Teerink, J. A. and R. D. Goeden. 1999. Description of the immature stages of Trupanea imperfecta (Co- quillett). Proceedings of the Entomological Soci- ety of Washington 101: 75-85. PROC. ENTOMOL. SOC. WASH. 103(1), 2001, pp. 207-216 THE PERLESTA PLACIDA (HAGEN) COMPLEX (PLECOPTERA: PERLIDAE) IN ILLINOIS, NEW STATE RECORDS, DISTRIBUTIONS, AND AN IDENTIFICATION KEY R. EDWARD DEWALT, DONALD W. WEBB, AND THOMAS N. KOMPARE Illinois Natural History Survey, Center for Biodiversity, 607 E. Peabody Dr., Cham- paign, IL 61820, U.S.A. (e-mail: edewalt @inhs.uiuc.edu) Abstract.—Perlesta placida (Hagen), a complex of perlid stoneflies in eastern and cen- tral North America, currently includes 17 described species. Large collections of this genus exist at the Illinois Natural History Survey, and, as part of an effort to update the status of stoneflies in Illinois, all adult Illinois specimens were reexamined. Perlesta de- cipiens (Walsh) and P. golconda DeWalt and Stark were previously recorded from Illinois, and, with the addition of six new state records, the total number of Perlesta known from Illinois is now eight. The latter include P. cinctipes (Banks), P. nr. teaysia Kirchner and Kondratieff, P. lagoi Stark, P. nelsoni Stark, P. shubuta Stark, and P. xube Stark and Rhodes. For all species but P. cinctipes, ranges are greatly increased from either the southeastern states or the north-central Plains region. Other midwestern states are likely to harbor a speciose assemblage of Perlesta. Distributions and associated ecological con- ditions suggested a wider range of pollution tolerance than is currently known from pub- lished literature. A key to males, females, and eggs is provided to aid with regional identification of Perlesta. Key Words: During much of the 20th century, the ge- nus Perlesta was considered to include a single, highly variable and widespread spe- cies, Perlesta placida (Hagen) (Frison 1935, 1942; Needham and Claassen 1925). Frison (1935) found that only the winter stoneflies, Allocapnia vivipara (Claassen) and Taeniopteryx burksi Ricker and Ross (the latter reported as T. nivalis Fitch by Frison) exceeded its abundance in Illinois. This abundance is true for much of eastern North America. More recently, P. placida has been considered a species complex. Stark (1989), in his revision for the Nearc- tic Region, reported 12 species. He added that his results were preliminary, and that several more species were expected from poorly collected regions. Since then, five Plecoptera, Perlesta placida complex, Illinois, distribution, key additional species have been described (Kondratieff and Baumann 1999, DeWalt et al. 1998, Kirchner and Kondratieff 1997, Stark and Rhodes 1997, Poulton and Stew- art 1991). Frison and his colleagues at the Illinois Natural History Survey (INHS) were pro- lific collectors of aquatic insects from Illh- nois and throughout North America, and nearly all of these specimens are deposited in the INHS insect collection. Illinois Per- lesta have been well collected throughout the century, and adult Perlesta account for 418 of the 18,885 current Plecoptera re- cords at the INHS. These records are ac- cessible in a _ searchable format at www.inhs.uiuc.edu/cbd/EPT/index.html. Frison and others identified the majority of 208 these Perlesta specimens as P. placida be- cause most collections predated Stark’s (1989) revision. Currently, only two species are known from Illinois, Perlesta decipiens (Walsh) (Harris and Webb 1995) and P. golconda DeWalt and Stark (DeWalt et al. 1998). Recent advances in the systematic knowledge of this species complex permit- ted a more thorough evaluation of the Illi- nois fauna using both historical and recent collections. As a result, six additional spe- cies from IIlinois are recorded here. Our objective is to document the species of Perlesta in Illinois by examining existing INHS collections and new material recently collected from throughout the state. To fa- cilitate identification of Illinois Perlesta, we provide a key to the males, females, and eggs. MATERIALS AND METHODS New adult material was collected by sweeping streamside vegetation, by rearing, and by ultra-violet light trapping. Aedeagi were extruded from males in the field or under magnification in the laboratory. The latter worked best and allowed adults time to harden their cuticle and adopt mature coloration before preservation. Using Stark’s (1989) protocol, the last 4—5 ab- dominal segments of both sexes were cleared in order to reveal internal and ex- ternal genitalic characters. Latitude and longitude coordinates for each unique location were determined using a DeLorme Street Atlas USA®, version 4.0. Many records had vague localities, the ex- act locations of which were impossible to obtain. So as not to lose any location data, locations were coded with one of four lev- els of accuracy (Table 1). The four levels of accuracy were denoted on range maps with different sized symbols, usually with closed and open circles of increasing size. Collection locations were stored in elec- tronic format using FileMaker Pro® soft- ware. Cartographic display of these loca- tions was produced using Environmental Systems Research Institute’s ArcView® PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Spatial accuracy codes used to distinguish accuracy of location information for Perlesta localities in Illinois. Spatial Accuracy Code Location Description from Labels 1 Known exact location. 2 Small municipality (<50,000 population) or park, point usually placed at center of town or at waterfront if stream name included on label. 3 Land survey data (principal meridian, town- ship, range, section) or large municipality (>50,000), taken as center of municipality or section. 4 County, plotted as center of county. software. ArcView connected across the INHS’s computer network to the collection location database via FileMaker’s Open Da- tabase Connectivity driver. Traditionally, locality information has been provided in the text of works such as these. We have opted to rely on the Plecop- tera database web site (see introduction for URL) for dissemination of this detailed and highly repetitive information. Those who do not have access to the Internet may re- quest the information from the authors as a text file. RESULTS AND DISCUSSION A total of 3,475 adult Perlesta specimens were identified from 151 unique locations across Illinois (Fig. 1). Eight of the 17 de- scribed Perlesta species were found, six of which are new state records. The Illinois distribution, general ecological require- ments, and additional information on taxo- nomic characters are provided for each spe- cies encountered. Despite the large latitu- dinal gradient, the majority of records in- dicated a mid-June through late July emergence for all species encountered. Perlesta cinctipes (Banks) (Figs. 2, 7, 15) A single female of this species, a new state record, was taken from a small, rocky stream draining limestone bluffs near the 209 VOLUME 103, NUMBER 1 Perlesta spp. @ P. nelsoni foe) P. decipiens ctipes (diamond), P. P. cin Figs. 1-4. Perlesta collection locations in Illinois. 1, All Perlesta collections. 2, s. 4, P. golconda. i (circles). 3, P. decipien (triangle), P. shubuta (squares), P. nelson nr. teaysia P. lagoi Locational Accuracy e | On 2 Owv3 O: Figs. 5-6. Perlesta collection locations in Illinois confluence of the Illinois and Mississippi rivers (Fig. 2). Stark (1989) reported it from regions north and west of the Ozark Moun- tains, where it was collected most frequent- ly from streams of lesser gradient (Poulton and Stewart 1991). Poulton and Stewart (1991) suggested that it could withstand moderate levels of organic enrichment. No detailed life history information was avail- able for this species (Stewart and Stark 1988). Perlesta decipiens (Walsh) (Figs. 3, 8, 16) Perlesta decipiens occurs throughout the northern two-thirds of the state (Fig. 3) and accounted for 48.8% of the records and 47.0% of all adult Perlesta specimens ex- amined. Larger rivers were its principal habitat (DeWalt et al. 1999), where it ac- counted for well over 99% of all Perlesta specimens collected. It was also abundant PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON . 5, P. lagoi. 6, P. xube. in all but the smallest streams in the Ver- milion River basin of eastern Illinois. This is the most widespread of all Per- lesta species. Its distribution spans the Mid- west, east to Pennsylvania, south to Texas, and west to the foothills of Colorado and eastern Wyoming (Stark 1989, Poulton and Stewart 1991). Snellen and Stewart (1979) reported a univoltine-fast life cycle with a lengthy egg diapause. This feature enables the species to inhabit intermittent streams and those with summer low oxygen and high water temperatures. Currently, P. de- cipiens is the only Perlesta species with a published life history. Perlesta nr. teaysia Kirchner and Kondratieff (Figs. 2, 9, 17) One collection consisting of two males and 10 females was taken from a small bluff stream that drained into the Vermilion VOLUME 103, NUMBER 1 River (Fig. 2). This collection represents a new State record. The aedeagi were still in- verted in the two male specimens, but by pulling on the loose ventral extension and unrolling (inside out) the whole structure we were able to discern that the aedeagus was long, thin, and lacked a caecum. These male specimens also had the tenth tergum with two elevated areas with stout sensilla basiconica, and the females had eggs with sessile collars. This description fits well with P. teaysia, except that P. teaysia has eggs with a short, wide collar (Kirchner and Kondratieff 1997). Perlesta frisoni Banks is a similar species, but our specimens lacked the median paraproct spine and the V- shaped subgenital plate notch (U-shaped in our specimens) as described by Stark (1989). It is possible that these specimens actually represent a new species, but with- out additional, properly prepared male specimens, we feel inclined to identify them as near P. teaysia. No detailed life history information is available for this species. Perlesta golconda DeWalt and Stark (Figs. 4, 10, 18) This species was recently described by DeWalt et al. (1998) from the Ohio River in southern Illinois. It was also taken from scattered locations near large rivers, and a single specimen was collected from a rela- tively small reach of the Sangamon River (Fig. 4). This species has also been found in mature, meandering rivers in Nebraska (Nance County, Cedar River, NE hwy 14, N Fullerton, 26 June 1998, 8 males and 8 females; Valley County, Middle Loup Riv- er, NE hwy 70, 25 June 1998, 4 males and 2 females) (B. Kondratieff, personal com- munication). No life history information is available for this species. Perlesta lagoi Stark (Figs. 5, 11, 19) This species represents a new state record and was the second-most abundant Perlesta species taken in Illinois. It contributed 32.8% of all Perlesta records and 44.8% of 21M the specimens. Perlesta lagoi was common throughout most of Illinois (Fig. 5). One major exception was in the Vermilion River basin of eastern Illinois, where only 26 specimens were taken. In contrast, P. de- cipiens contributed 1,106 specimens across all stream sizes in the basin. Perlesta lagoi was the dominant species in small streams throughout the rest of the state, even in slowly flowing streams. The only other published records for this species are for central and northern Mississippi (Stark 1989). No life history studies have been published for this species. Perlesta lagoi is likely widespread throughout the Midwest and has almost cer- tainly been confused with P. decipiens throughout the region. It has a somewhat spatulate paraproct, with a short, medially pointing spur. The spur of P. decipiens is larger and usually pointed directly ventrad. As Stark (1989) noted, the spur and round- ed tip of the paraproct give the appearance of a “‘bird head and beak’”’ in lateral aspect. Eggs of P. lagoi have no collar, while those of P. decipiens have a wide and stout collar. Subgenital plates were truncate in both spe- cies, but the notch was V-shaped in P. la- goi, while distinctly U-shaped in Illinois P. decipiens. Perlesta nelsoni Stark (Figs. 2, 12, 20) Ten percent of the Illinois Perlesta re- cords and 4.6% of the specimens were of P. nelsoni. All collections were from the Shawnee Hills and Ozark Uplands of south- ern Illinois (Fig. 2). Streams in the area contained sandstone or limestone substrates and many dried to pools in the summer. This species was previously known only from North and South Carolina and Ten- nessee (Stark 1989); hence, its presence in Illinois represents a sizable range extension and a new state record. All collections of this species were dated 1977 or later. Im- migration, probably from the neighboring Kentucky highlands, is likely to have oc- curred within the last three decades. Pres- N — N PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON envelope 14 a Fig. 7-14. Male structures of Illinois Perlesta. 7, P. cinctipes (a) lateral aedeagus; (b) lateral paraproct and spine. 8, P. decipiens (a) lateral aedeagus; (b) lateral paraproct; (c) caudal paraproct. 9, P. nr. teaysia (a) lateral aedeagus; (b) lateral tenth tergite and paraproct. 10, P. golconda (a) lateral aedeagus; (b) lateral paraproct; (c) dorsal aedeagus. 11, P. lagoi (a) lateral aedeagus; (b) lateral paraproct:; (c) caudal paraproct. 12, P. nelsoni (a) lateral aedeagus; (b) lateral paraproct. 13, P. shubuta (a) lateral aedeagus; (b) lateral paraproct. 14, P. xube (a) VOLUME 103, NUMBER 1 ently, no life history information is avail- able for this species. Perlesta shubuta Stark (Figs. 2.13, 21) Two records, accounting for 17 speci- mens, came from near the Wisconsin border and along the southeastern Indiana border (Fig. 2). These specimens represent a new state record and a considerable range exten- sion. Previous records include Arkansas, Missouri, Mississippi, and Oklahoma (Poulton and Stewart 1991, Stark 1989). Poulton and Stewart (1991) stated that this species was relegated to larger, permanent streams, and it was the only Perlesta lim- ited to these conditions in their study area. Illinois records were from streams that fit this description. Perlesta xube Stark and Rhodes (Figs. 6, 14, 22) Five percent of the records and 1.5% of the specimens were P. xube. All specimens were taken from small, wooded streams that shrank to pools in the summer, but were seldom completely dry (Fig. 6). Perlesta xube was described from Nebraska (Stark and Rhodes 1997) and also has been re- ported from North Dakota (Kondratieff and Baumann 1999). Its existence in Illinois is a new state record and a sizable range ex- tension. No life history studies have been conducted on P. xube. This species may be confused with P. adena (Stark), but P. xube is distinguished by its long and sinuous ae- deagal tube and the presence of a long dor- sal hair patch running the length of the tube to the large caecum. Perlesta adena has a short, plump aedeagus without a long dorsal patch (Stark 1989). The following charac- teristics of forewing coloration define P. eH 23 xube (Stark and Rhodes 1997): a light mid- dle third of the costal margin, a white patch before the arculus, a white streak along the medial vein, and a light streak in the inter- cubital area, all on a charcoal colored back- ground. Alternatively, the costal margin of P. adena is pale along its entire length, and no other light wing markings are known for this species. DISCUSSION The INHS has a long history of docu- menting the aquatic insect community of the state. The large natural history collec- tion permitted us to re-evaluate a challeng- ing species complex. We have added six new state records for Illinois within this ge- nus alone. These additions increased the number of Illinois stoneflies from 64 (DeWalt et al. 1998, Harris and Webb 1995) to 70. The speciose nature of the P. placida complex will likely hold for other midwest states whose geographic and glacial history is similar to that of Illinois. Additionally, we expect that other perlid complexes, es- pecially Neoperla, will yield additions to the state’s fauna. Frison’s (1935) treatment of Perlesta as a single species led Hilsenhoff (1987) to provide one environmental tolerance value for Perlesta. Lenat (1993), after Stark’s (1989) revision, indicated major differences in environmental tolerance between the rel- atively tolerant P. placida and other more sensitive regional Perlesta species. In find- ing a diverse Perlesta fauna in Illinois, and demonstrating apparently different ecolog- ical occurences, the assignment of one tol- erance value for the entire genus is inap- propriate. To date, the only detailed life his- tory study available is for P. decipiens (Snellen and Stewart 1979). Additional life lateral aedeagus; (b) lateral paraproct. Figs. 7a—9a, 1la, and 12a—13b borrowed or modified, with permission, from Stark (1989). Figs. 14a and 14b borrowed or modified, with permission, from Stark and Rhodes (1997). Abbreviations: c = caecum; dp = dorsal patch; n = nipple; v = ventral extension; Sb = sensilla basiconica. Scale for aedeagi only, 0.15 mm. 214 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 21 a 22 Fig. 15-22. Subgenital plate (a) and egg (b) of Illinois Perlesta females. 15, P. cinctipes. 16, P. decipiens. 17, P. nr. teaysia. 18, P. golconda. 19, P. lagoi. 20, P. nelsoni. 21, P. shubuta. 22, P. xube. Figs. 15a and 15b, 16b, 17b, 19a—21b borrowed or redrawn, with permission, from Stark (1989). Fig. 22a and 22b borrowed or redrawn, with permission, from Stark and Rhodes (1997). Scales for subgenital plates, 0.30 mm, for eggs 0.15 mm. history studies and the association of nymphs with adults would help calibrate tolerance values for species in the genus. A KEY TO MALES, FEMALES, AND EGGS OF THE PERLESTA OF ILLINOIS 1. Head, body, wing, veins, and wing membranes very pale; aedeagal tube with poorly developed caecum (Fig. 10a), dorsal patch broad, with larger lateral setae forming distinct parallel lines (Fig. 10c); subgenital plate lobes rounded (Fig. 18a), notch V-shaped; egg collar sessile, chorion smooth, to slightly granular (Fig. 18b) iss BR agate Aqetren ae eet Ae P. golconda Head and body with areas of pigmentation, wing veins pale to black, wing membrane rang- ing from pale to black; aedeagus with or with- VOLUME 103, NUMBER 1 out caecum; subgenital plate lobes truncate or rounded, notch U- or V-shaped; egg collar ses- sile or stalked, chorion smooth or pitted .... 2 2. Body and wing coloration generally light; males with one or more well-defined patches of sensilla basiconica on tenth tergite (Fig. 9b), aedeagus long, sinuate, lacking caecum (Figs. Same a) ear een ee Are oo ee oe 3 — Body and wing with areas of pigmentation ranging from amber to black; males without well-defined patch of sensilla basiconica, al- though a few scattered single sensilla may be present; aedeagus short or long, but with a cae- Gum (Bigsaeas Sas lla ls anal 4a) eee ee 4 3. Sensilla basiconica in two elevated, circular patches on tenth tergite, paraproct tip pointed and well-sclerotized, without median spine (Fig. 9b); subgenital plate lobes rounded, notch U-shaped (17a); egg chorion with mesal band of inconspicuous shallow pits, collar sessile (Fig. 17b) — Sensilla basiconica in one low, sparse patch; paraproct tip rounded, lightly sclerotized, with median spine well-removed from tip (Fig. 12b); subgenital plate lobes truncate, notch U- shaped (Fig. 20a); egg chorion smooth, collar short and slender eS Gee Okan Eee arer Ons tenet P. nr. teaysia P. nelsoni — Wing coloration amber; body with light brown reticulations on yellowish background 5. Wings with costal area pale throughout, but no other light markings; aedeagus and caecum long and slender, dorsal patch short, not well- developed laterally (Fig. 7a), paraproct spine directed ventrad (Fig. 7b); subgenital plate lobes rounded, notch shallow and V-shaped (Fig. 15a); egg chorion coarsely pitted through- out, collar stalked (Fig. 15b) ...... P. cinctipes — Wings with only middle % to % of costal mar- gin light, light area before arculus, light streaks above medial vein and in intercubital area; ae- deagus and caecum robust, dorsal patch occu- pying most of tube length and extending lat- erally in a wide band on caecum (Fig. 14a), paraproct spine directed medially (Fig. 14b); subgenital plate notch deep, U-shaped (Fig. 22a); egg chorion finely punctate in middle %, collanisessile(Bio22D) ieee ee cen eae P. xube 6. Paraproct rounded or pointed, spine prominent in lateral view and directed ventrally (Fig. 8b, 13b); aedeagus with caecum about two times longer than wide (Fig. 8a, 13a); subgenital plate U- or V-shaped; egg chorion smooth, col- larestalkedi(BiovsllGbs2 lb) ce emeiewen e ee 7 — Paraproct spatulate, spine not prominent in lat- eral view and directed mesally, (Fig. 11b) the latter best seen in caudal view (Fig. 11c); ae- 215 deagus with caecum only as long as wide (Fig. lla); subgenital plate notch V-shaped (Fig. 19a); egg chorion shallowly pitted in middle %, collar sessile (Fig. 19b) P. lagoi 7. Paraproct pointed, face angular (about 45°), spine rotated somewhat medially (Fig. 13b); subgenital plate notch V-shaped (Fig. 21a); egg collar occupying only %4 to % width of pole, anchor pedicel a single thick stalk (Fig. 21b) oy Sadek OF DPA PORCINE MIE P. shubuta — Paraproct rounded, face rounded, spine usually pointed ventrad (Fig. 8b and 8c); subgenital plate notch U-shaped (Fig. 16a); egg collar oc- cupying at least 4% width of pole, anchor ped- icel composed of many filaments (Fig. 16b) P. decipiens ACKNOWLEDGMENTS We thank C. Favret and M. A. Harris for suggesting helpful changes to this manu- script. Several people have been instrumen- tal over the years in collecting and depos- iting material used in this study. These per- sons include A. R. Brigham, W. U. Brigh- am, M. A. Harris, E. A. Lizowski, W. G. Ruesink, and M. J. Wetzel, ail current or former scientists at the INHS. Additionally, B. P. Stark, of Mississippi College, provid- ed encouragement, verification of some species determinations, and inspiration for this project. Our gratitude is also extended to H. P. Boyd, editor of Entomological News, and N. M. Anderson, editor of En- tomologica Scandinavica, for permission to use Perlesta figures from previous issues. Funding for this study was provided by the Illinois Natural History Survey, its Critical Trends and Assessment Program, by a Statewide Biosurvey contract from the II- linois Department of Transportation, and by a National Science Foundation collection computerization grant numbered DBI- 9876756. LITERATURE CITED DeWalt, R. E., D. W. Webb, and M. A. Harris. 1999. Summer Ephemeroptera, Plecoptera, and Trichop- tera (EPT) species richness and community struc- ture in the lower Illinois River basin of Illinois. Great Lakes Entomologist 32: 115-132. DeWalt, R. E., B. P Stark, and M. A. Harris. 1998. 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Pro- ceedings of the Entomological Society Washing- ton 101: 325-331. Lenat, D. R. 1993. A biotic index for the southeastern United States: derivation and list of tolerance val- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ues, with criteria for assigning water-quality rat- ings. Journal of the North American Benthological Society 12: 279-290. Needham, J. H. and P. W. Claassen. 1925. A mono- graph of the Plecoptera or stoneflies of America north of Mexico. Thomas Say Foundation, Ento- mological Society of America 2: 1-397. Poulton, B. C. and K. W. Stewart. 1991. The stoneflies of the Ozark and Ouachita Mountains (Plecop- tera). Memoirs of the American Entomological Society 38: 1-116. Snellen, R. K. and K. W. Stewart. 1979. The life cycle of Perlesta placida (Plecoptera: Perlidae) in an intermittent stream in northern Texas. Annals of the Entomological Society of America 72: 659— 666. Stark, B. P. 1989. Perlesta placida (Hagen), an eastern Nearctic species complex (Plecoptera: Perlidae). Entomologica Scandinavica 20: 263-286. Stark, B. P. and H. A. Rhodes. 1997. Perlesta xube, a new stonefly species from Nebraska (Plecoptera: Perlidae). Entomological News 108: 92-96. Stewart, K. W. and B. P. Stark. 1988. Nymphs of North American stonefly genera (Plecoptera). Thomas Say Foundation, Entomological Society of Amer- ica 12: 1—460. PROC. ENTOMOL. SOC. WASH. 103(1), 2001, pp. 217-221 A NEW SPECIES OF SERICOCEROS KONOW (HYMENOPTERA: ARGIDAE) DAMAGING VILLETANA TREES, TRIPLARIS CARACASANA CHAM. (POLYGONACEAE) IN PARAGUAY DAVID R. SMITH AND EDGAR ARTURO BENITEZ DIiAz (DRS) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. De- partment of Agriculture c/o National Museum of Natural History, Washington, DC 20560- 0168, U.S.A.; (EABD) Divi6n Entomologia, Facultad de Ciencias Agrarias, Universidad Na- cional de Asuncion, Casilla de Correo 1618, Campus Universitario, San Lorenzo, Paraguay Abstract.—A sawfly was discovered severely damaging leaves of villetana trees, Tri- plaris caracasana Cham. (Polygonaceae) in Paraguay. The sawfly, Sericoceros vitellanae Smith, n. sp., is described and separated from other Sericoceros species. Triplaris is a new host record for a species of Sericoceros. Notes on the maternal care, larval feeding habits, and pupation of the sawfly are presented. Key Words: Villetana trees, Triplaris caracasana Cham. (Polygonaceae), are important native shade trees in central Paraguay. They are well adapted to wet soils and are abundant around Asuncion. Damage to the foliage caused by sawfly larvae was reported re- cently at the Facultad de Ciencias Agron- Omicas, Universidad Nacional de Asuncion, in San Lorenzo, 11 km from Asuncion. Adults reared from larvae and adults col- lected ovipositing on the leaves were ob- tained by the senior author, and they rep- resent a new species of Sericoceros Konow. The species is described below and notes are given on its biology. Sericoceros includes 20 species and is distributed from Mexico to northern Argen- tina (Smith 1992). Known hosts are Coc- coloba spp., Lonchocarpus sp., and Chry- sobalanus sp. This occurrence on Triplaris represents a new host record. Sericoceros villetanae Smith, new species (Figs. 1, 3-5) Female.—Length, 8.5 mm. Antenna with scape orange, pedicel brown, flagellum Hymenoptera, Argidae, Sericoceros, new species, Triplaris, Polygonaceae black. Head black with clypeus, supracly- peal area, and interantennal area orange; la- brum and palpi whitish; mandible orange at base, apical % reddish brown. Thorax, ab- domen, and sheath orange. Legs with cox- ae, trochanters, and femora orange; foreti- bia black with inner surface and basal half of outer surface orange; midtibia with inner surface orange, outer surface black and ex- treme apex completely black; hindtibia with inner surface orange, outer surface and api- cal %4 black; tarsi black. Wings lightly, in- fuscate, paler to hyaline at apices; veins and stigma of forewing black except extreme base of stigma, costa, and subcosta orange. Antennal length 1.3 head width. Head and body smooth, shining, impunctate, without surface sculpture. Hindwing with anal cell present. Sheath slender in dorsal view, not expanded or widened at apex. Lancet as in Fig. 1, narrowly constricted at apex and with basal and central annuli sub- parallel and close together. Male.—Length, 6.7—7.0 mm. Antenna black, third segment with inner surface of 218 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-2. basal %4 of each furcation white. Head black with labrum white. Thorax black with pron- otum, mesonotal lateral lobes, and narrow lateral margins of mesoprescutum orange. Abdomen black with sterna whitish. Legs black with outer surface of forefemur and foretibia white. Wings hyaline; veins black. Third antennal segment bifurcate; antennal length subequal to head width. Genitalia as in Figs. 3, 4; inner margin of parapenis nearly truncate, head of penis valve round- ed in lateral view. Larva (from a late instar).—Length, 17.0 mm. Head dark reddish brown, more black Lancets. 1, Sericoceros villetanae. 2, S. calanticatus. around each eye and paler to white on low- er part of frons, clypeus, and labrum. Tho- racic legs yellow brown; body generally pale with black longitudinal band on each side. Spines above spiracles and on dorsum yellow brown, those around and below spi- racles in black band, black. Small plates on venter anterior and posterior to prolegs am- ber. Tenth tergum black, reddish brown an- teriorly. Head with scattered, short setae, no lon- ger than width of antenna. Antenna a single flat segment. Each mandible with 5—7 long setae on outer surface; clypeus with 1—2 se- VOLUME 103, NUMBER 1 Figs. 3-5. italia, lateral view of penis valve. 5, Lateral view of third abdominal segment of larva. tae on each side; labrum with 2-3 setae on each side, anterior margin strongly emar- ginate at center; maxillary palpus 3-seg- mented, with | seta on palpifer and one on first segment; lacinia with 4—5 bifurcate or trifurcate spines; labial palpus 2-segmented, prementum with 3 setae. Prothoracic leg without tarsal claw. Pro- legs on abdominal segments 2—9, small on 2, 8, and 9. Body covered with long spines, the longest being longer than the width of an abdominal annulet. Abdominal segments 1—9 each 3-annulate. Ornamentation of typ- ical abdominal segment as in Fig. 5. First annulus with 3 spines, lower one smallest; second annulet with 3 long spines above spiracle, third annulet with 1 long and 1 short spine; postspiracular area with tuber- cle composed of 4 long spines; surpedal area with 5 long spines, third and fourth from anterior the shortest, and one long blunt, tubular spine with white tip appear- ing to be a glandular opening. Tenth tergum with 20-25 long spines. Venter with 2 its aoe Bay Cr 7 a oe if Anan g: sto iw rE 3 Sericoceros villetanae. 3, Male genitalia, genital capsule, ventral view of left half. 4, Male gen- short, conical spines between prolegs and 2 or 3 smaller spines posterior to these. Host.—Triplaris caracasana Cham. (Po- lygonaceae). Holotype.—Female, labeled ‘‘Paraguay: Central: San Lorenzo, April 1999, E. Ben- {tez coll., on Triplaris caracasana.’’ Depos- ited in the Museo Nacional de Historia Nat- ural del Paraguay, Asuncion. Paratypes.—PARAGUAY: Same data as for holotype (3 females, 4 males); Central, San Lorenzo, 18.11.1999, E. Benitez coll., S/Triplaris sp. (2 females); Asuncion, Dic- 1992, B. Garcete, S/Triplaris sp. (‘‘Ville- tana’’), adult resting over the eggs (1 fe- male); E de Lamora, XII-1997, B. Garcete (1 female). Representatives of each sex de- posited in the Museo Nacional de Historia Natural del Paraguay, Asuncié6n; Facultad de Ciencias Agrarias, Universidad Nacional de Asuncion, San Lorenzo; National Mu- seum of Natural History, Smithsonian In- stitution, Washington, D.C. Other material One pin with eggs at- 220 Pp) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tached on surface of part of a leaf, two pins with cocoons on parts of leaves, and a vial of five larvae have the same data as the holotype. Discussion.—The female coloration of S. villetanae is similar to that of Sericoceros calanticatus (Konow) recorded from the states of Espirito Santo and Rio Grande do Norte, Brazil, and it will run to S. calanti- catus (couplet 17) in Smith’s (1992) key to species. The lancet is very different from that of S. calanticatus, however, as com- pared in Figs. 1 and 2. The lancet of S. villetanae is broad, but narrowly constricted at its apex, the basal and central annuli are close together and subparallel, and there are more distinct teeth. The male of S. calan- ticatus is unknown, but that of S. villetanae can be distinguished from other known males of Sericoceros by comparison of Figs. 3 and 4 with figures 181—190 in Smith (1992). In S. villetanae, the head of the pe- nis valve is more rounded and the inner margin of the parapenis is more truncate than in other species. Larvae for three other species of Seri- coceros are known, S. mexicana (Kirby) (larva described as S. edwardsii (Cresson) by Smith 1972), S. krugii (Cresson), and S. gibba (Klug) (Smith 1972, Kimsey and Smith 1985). The larva of S. villetanae is easily distinguished from those three by the long acute spines (much longer than the width of an abdominal annulet) on the body, presence of 5—7 setae on the outer surface of each mandible, and the general pale coloration with a black longitudinal stripe on each side in the spiracular area. The other three species have short, more conical or blunter protuberances which are shorter than the width of an abdominal an- nulet, have one seta on the outer surface of each mandible, and are generally pale with the tubercles black (Smith 1972, fig. 43; Kimsey and Smith 1985, figs. 24, 25). Biology.—On March 12, 1999, EABD obtained adults and larvae collected from villetana trees on the campus of Universi- dad Nacional de Asuncion, including a fe- male that was sighted protecting an egg batch of about 75 orange eggs on a leaf. The egg batch was symmetrical and round to oval. The eggs were oval and attached on end perpendicular to the leaf surface. The female kept this position over the eggs for several days until the larvae hatched. In the laboratory, larvae were placed in acrylic rearing boxes (33 X 39 X 25 cm) at room temperature (27°C). The larvae were positioned next to each other while feeding, embracing the border of the leaf with the legs and with the abdomen raised and coiled like a question mark. This larval position was described by Costa Lima (1960) for other symphytan larvae. In this position, the larvae remind one of a small monkey, thus locally they are called ““Yso- ka’i’> (monkey worm). In this position, the larvae eat all the leaf except for the central vein. The first cocoons were observed on April 3, 1999. The larvae spin a light brown co- coon made of carton-like matter, barrel-like in shape, with one end rounded and the oth- er end with a flat circular lid. It has a double wall, the inner wall dull and parchment-like and the outer wall irregularly netted, similar to the cocoons of Argidae as described by Wong (1961). The cocoon is firmly attached to the substrate, leaves, or walls of the cage, by one or more sides. Twenty-eight females and 13 males emerged in April 1999. Triplaris has not been recorded as a host for species of Sericoceros. Hosts for other species are Lonchocarpus sp. (Legumino- sae) for S. vumirus Smith; Coccoloba spp. (Polygonaceae) for S. albicollis (Klug), S. gibbus (Klug), S. krugii (Cresson), and S. mexicanus (Kirby); and Chrysobalanus sp. (Rosaceae) for S. krugii. The only references to the biology of Sericoceros species are Martorell (1941) for S. krugii in Puerto Rico and Kimsey and Smith (1985) for S. gibbus in Panama. Mar- torell indicated maternal care for S. krugii by stating that ‘During oviposition the fe- males do not move from the egg-mass even if disturbed”’ and ‘‘... even after the last VOLUME 103, NUMBER 1 egg of a cluster is laid the female stays over the egg-mass as if they were brooding the eggs.’ Kimsey and Smith (1985) did not indicate maternal care for S. gibbus. ACKNOWLEDGMENTS We thank Bolivar Garcete, Museo Na- cional de Historia Natural del Paraguay, Asuncion, for initial identification of the sawfly and help with the manuscript. Linda Lawrence prepared figures | and 2 and Ca- thy Anderson helped with preparation of the plates; both are with the Systematic En- tomology Laboratory, USDA, Washington, DC. Thanks are extended to the following for review of the manuscript: N. M. Schiff, U.S. Forest Service, Stoneville, MS, and M. G. Pogue and E. E. Grissell, Systematic En- tomology Laboratory, USDA, Washington, DEC. Nw N —_ LITERATURE CITED Costa Lima, A. da. 1960. Insetos do Brasil: Hymenop- tera. Primeira Parte. Vol. 11. 368 pp. Kimsey, L. S. and D. R. Smith. 1985. Two new spe- cies, larval descriptions and life history notes of some Panamanian sawflies (Hymenoptera: Argi- dae, Tenthredinidae). Proceedings of the Ento- mological Society of Washington 87: 191-201. Martorell, L. FE 1941. Biological notes on the sea-grape sawfly, Schizocera krugii Cresson, in Puerto Rico. Caribbean Forester 2(3): 141-144. Smith, D. R. 1972. North American sawfly larvae of the family Argidae (Hymenoptera). Transactions of the American Entomological Society 98: 163— 184. . 1992. A synopsis of the sawflies (Hymenop- tera: Symphyta) of America south of the United States: Argidae. Memoirs of the American Ento- mological Society 39, 201 pp. Wong, H. R. 1961. Cocoons of some sawflies that de- foliate forest trees in Manitoba and Saskatchewan. Eighty-Second Annual Report of the Entomolog- ical Society of Ontario, pp. 61—67. PROC. ENTOMOL. SOC. WASH. 103(1), 2001, pp. 222-226 OBSERVATIONS ON THE BEHAVIOR OF MACHIMUS OCCIDENTALIS (HINE) AND MACHIMUS FORMOSUS (HINE) (DIPTERA: ASILIDAE) IN MONTANA KEVIN M. O’ NEILL Department of Entomology, Montana State University, Bozeman, MT 59717, U.S.A. (koneill @ montana.edu) Abstract.—In Montana, Machimus occidentalis (Hine) is a typical free-ranging grass- land asilid, whereas Machimus formosus (Hine) centers its activities around the entrances of small mammal burrows. Their divergent microhabitat preferences are reflected in dif- ferences in their thermoregulatory, mating, and foraging strategies. Both species are op- portunistic predators that take a wide range of prey taxa and sizes. Key Words: On western U.S. grasslands, robber flies of the asilid genus Machimus display two general types of microhabitat preference (James 1941, Lavigne 1968, Dennis and Lavigne 1979, O’Neill and Kemp 1992). Machimus gilvipes (Hine) and M. formosus (Hine) restrict most of their activities to the vicinity of the burrows of small mammals, spending much of the day perched on the walls of the burrows. Machimus callidus (Williston) and M. occidentalis (Hine), in contrast, are more typical grassland asilids that range relatively freely across their hab- itats, perching on bare ground or low plants. In Wyoming, Lavigne and his colleagues have studied all four species, with particular attention to M. gilvipes (Rogers and Lavig- ne 1972, Lavigne 1968, Schreiber and La- vigne 1986) and M. callidus (Dennis and Lavigne 1979). Working in Montana, O’Neill (1994, 1995) reported several prey records for M. occidentalis, and O’Neill and Kemp (1992) reported on the thermo- regulatory behavior of M. formosus and M. occidentalis. Here, I report on further be- havioral observations and prey records of these two species. robber fly, Asilidae, prey, mating METHODS Machimus occidentalis, a gray robber fly 13-17 mm long, was studied on 36 days during June and July of 1988-1994. Most observations were made in several gullies 14 km south of Three Forks, MT. One gully was dry, with vegetation dominated by grasses such as Stipa comata Trin. and Rupr., Bouteloua gracilis (H.B.K) Lag., and Bromus tectorum L., with scattered shrubs (especially Rhus trilobata Nutt.). The other gully, about 1 km to the south was more mesic, and contained a large patch of lupine (Lupinus sp.) mixed with grasses. Machi- mus formosus, a yellowish-brown species 11-15 mm long, was observed on 13 days from late June-early August in 1988-1991 at a grassland site 10 km south of Three Forks dominated by crested wheatgrass (Agropyron cristatum L. (Gaertn)). Here, the sizes of the robber flies are reported as maximum head width, whereas those of prey are given as body length or as the product of body length and thorax width. The latter was used in analyses as a simple two-dimensional index of prey size that ac- counts for variation in prey shape. Prey VOLUME 103, NUMBER 1 mass was not used because prey were col- lected after the robber flies had fed for un- known periods of time when prey were col- lected. RESULTS AND DISCUSSION Machimus occidentalis (Hine) Machimus occidentalis ambushed prey in mid-air after flights about 10-30 cm long initiated from perches on the ground or on low plants. The 125 prey collected included insects from 29 families in eight orders (Fig. 1) (sample sizes are given where N > 1): ERPHEMEROPTERA, Leptophlebitidae: Paraleptophlebia sp.; Baetidae: unidenti- fied sp.; ORTHOPTERA, Acrididae: Me- lanoplus sanguinipes (F) (nymph), Phlibos- troma quadrimaculatum (Thomas) (nymph); HOMOPTERA, Cicadellidae: Philaronia bilineata (Say), Prairiana subta Ball (3), Psammotettix lividellus (Zettinger) (2), unidentified sp. (2 nymphs); Aphididae: unidentified sp.; HEMIPTERA, Miridae: Leptopterna ferrugata (Fallen), Litomiris debilus (Uhler); LEPIDOPTERA, Geome- tridae: unidentified sp.; Lycaenidae: Ly- caenides sp.; Pyralidae: Crambus sp.; CO- LEOPTERA, Chrysomelidae: Cryptoce- phalus notatus EF; Scarabaeidae: Serica an- thracina LeConte, Aphodius vittatus Say, Aphodius distinctus (Mueller), Dichelonyx backii (Kirby) (15); Elateridae: Ctenicera glauca (Germar); DIPTERA, Anthomyi- idae: Paregle cinerella (Fallen); Asilidae: Machimus occidentalis Hine (5); Atherici- dae: Atherix pachypus Bigot; Calliphoridae: Phormia regina (Meigen); Chironomidae: unidentified sp. (3); Milichiidae: Pholeo- myia indecora (Loew), Sarcophagidae: Ac- ridophaga spp. (4), Arachnomyia sp. (2), Blaesoxipha opifera (Coquillett), Protodex- ia hunteri (Hough), Ravinia iherminieri (Robineau-Desvoidy) (2), unidentified sp.; Sepsidae: Sepsis sp.; Simuliidae: Simulium sp. (3); Stratiomyiidae: Sargus cuprarius (L.); Tabanidae: Chrysops mitis Osten Sacken; Tachinidae: Besseria brevipennis (Loew), Chetogena tachinomoides (Town- 223 send), Frontiniella parancilla Townsend, Ptilodexia rufipennis (Macquart), unidenti- fied sp. (2); Tipulidae: Tipula sp.; uniden- tified family; HYMENOPTERA, Formici- dae (all alates): Formica ciliata Mayr, For- mica subpolita Mayr (40), Formica sp. (3), Lasius sp. (2); Halictidae: Lasioglossum sp.; Ichneumonidae: Anomalon reticulatum (Cresson), unidentified sp.; Mutillidae: un- identified sp.. Nineteen of the ant prey and six of the cannibalism records were previ- ously reported in O’Neill (1992, 1994). Machimus occidentalis took a wide range of prey sizes relative to their own body size, from an aphid just over 1 mm long, to flies and ants 10—16 mm long (Fig. 1), but there was no correlation between predator and prey size (rz = 0.05, P = 0.59, N = 106). Prey records from a particular site and time were often quite uniform. For example, on 8, 9, and 12 June 1992 in the south gully, 25 of 31 prey were either large tachinids and sarcophagids (N = 10) or the beetle Dichelonyx backii (N = 15). However, on 10, 19, and 21 June 1992 in the north gully, 21 of 26 prey were alates of the ant For- mica subpolita, males of which commonly formed mating swarms in the vicinity and provided a flush of resources for predators in the area (O’Neill 1994). As with foraging, the mating strategy of male M. occidentalis consisted of ambush- ing females passing in flight, without con- spicuous courtship. Each coupled pair im- mediately descended to low plants, the male always remaining mounted dorsally on the female during copulation. Males (mean + SE head width = 3.14 + 0.02 mm) tended to be larger than their mates (mean = 2.96 + 0.02) (N = 43, Wilcoxon signed rank test, P<. 0:001), Machimus formosus (Hine) As in Colorado (James 1941) and Wyo- ming (Lavigne 1968), M. formosus centers its activity around burrow entrances of small mammals, which at the Montana site were made by badgers (Taxidea taxus) and ground squirrels (Citellus sp.). When the 224 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Ga VV. occidentalis | M. formosus & 2 Re O ©: O — Oo _ T ol i Tae S Ss Le) 9 atin) (a Sesa SaaS e ¢ s Prey size (mm?) Cc Oo 5 O oO O = O Fig. 1. Top: range of prey sizes (area in 5 mm? increments) used by the two species; bottom: frequency distributions of prey orders used by M. occidentalis and M. formosus. VOLUME 103, NUMBER 1 flies first became active each day, they perched on fully insolated level soil outside of burrow entrances, but as soil surface temperatures rose above about 32°C, they began perching on the walls of burrows, down to depths of 14 cm (O’Neill and Kemp 1992). During several surveys of the surrounding grasslands, I observed an M. formosus away from the burrows only once. However, they were often common at bur- rows. On 16 July 1989, 38 of 100 burrows were occupied by individual M. formosus. Two days later, 42 of 100 burrows were oc- cupied, 21 by a single male, six by a single female, 13 by flies of undetermined sex, and two by pairs in copula. Occupied bur- rows were sometimes within 1 m of one another, but often separated by tens of me- ters. I made no quantitative assessment of distribution of potential burrows, but sev- eral general factors seemed to be involved in their occupation. The flies were not pre- sent in older burrows partially filled with soil, or at burrows with entrances blocked by spider webs. In addition, burrows ori- ented so that they provided no shade in the afternoon were often abandoned during hot- ter portions of the day. In Wyoming, burrow-dwelling Machimus gilvipes appear to take some prey directly from the walls of the burrow (Schreiber and Lavigne 1986), but in all 16 predations that I observed, prey were taken in mid-air above the burrow. The 43 prey of M. for- mosus included insects from ten families in six orders (Fig. 1): HOMOPTERA, Cica- dellidae: Aceratagallia sp. (2), Commellus sexvittatus (Van Duzee) (3), Psammotettix lividellus (Zettinger), unidentified sp. (2 nymphs, 1 adult); HEMIPTERA, Miridae: Adelphocoris lineolatus (Goeze); COLE- OPTERA, Carabidae: Harpalus sp.; LEPI- DOPTERA, Pyralidae: unidentified sp.; Tortricidae: unidentified sp. (3); DIPTERA, Anthomyiidae: unidentified sp., Dolicho- podidae: Medetera vittata Van Duzee (12) (this represents a new state record for this dolichopodid for Montana; Bickel 1985), Scatopsidae: unidentified sp. (6); Sepsidae: 225 Saltella sphondylii (Schrank); HY MENOP- TERA, Formicidae (all alates): Dorymyr- mex insana (Buckley), Formica neogagates Emery, Formica obtusipilosa Emery, For- mica subpolita Mayr. There seems to be no particular reason for some of the prey (e.g., leafhoppers, moths, and ants) to be associated with mammal burrows, so they probably just happened to come within foraging range of the resident fly. However, the scatopsids and dolichopodids increased their vulnera- bility by swarming above the burrows which they perhaps used as landmarks. As with M. occidentalis, there was evidence of temporal variability in prey records for M. formosus. On 18 July 1989, six of seven prey were an unidentified scatopsid fly that swarmed above burrows, whereas on 24 and 25 July, when scatopsids were absent from prey, eight of 11 were dolichopodid flies. Although the size distributions of adults of the two Machimus species over- lapped, M. formosus generally took smaller prey (mean size = 5.7 + 0.9 mm’, N = 41) than M. occidentalis (15.8 + 1.0 mm’, N = 106; Mann-Whitney test, P < 0.001) (Fig. 1). Except during mating, the flies apparent- ly occupied burrows alone, tolerating the presence of conspecifics only during mating interactions. Each of the seven times that I observed two non-mating flies at a burrow, one departed when approached by the other, once when butted by the resident. Nine mating pairs were observed on the walls of burrows, all coupled end-to-end. The single complete mating observed was _ initiated when a female entered a male’s burrow and was caught in mid-air by the male; the pair then descended in copula to the lip of the burrow entrance where they mated for 7 min and 20 s. Like many asilids, M. occidentalis and M. formosus are opportunistic predators whose diets seem primarily determined by their body sizes (relative to potential prey) and by the vagaries of local prey availabil- ity. The biology of M. formosus is partic- 226 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ularly constrained by the peculiar micro- habitat preference that it shares with M. gil- vipes and M. polyphemi Bullington and Beck (Bullington and Beck 1991). The use of burrows by M. formosus affects their thermoregulatory strategy (O’Neill and Kemp 1992), as well as their foraging strat- egy (because their prey base is restricted) and their mating strategy (because mates are available only in one spatially circum- scribed microhabitat). However, by occu- pying burrows, M. formosus may minimize it’s contacts with larger robber flies, such as Efferia staminea (Williston), which often prey on smaller asilids, including M. occi- dentalis (O’ Neill 1992). ACKNOWLEDGMENTS I thank Ruth O’Neill for assisting in the field, C. Riley Nelson for identifying the asilids, and James Bess, Richard Hurley, T. Keith Philips, and Catherine Seibert, for identifying prey. Ruth O’Neill, Peter Jen- sen, and Richard Hurley reviewed the man- uscript. This work was supported by grants from USDA-ARS and USDA-APHIS-PPQ and by the Montana Agricultural Experi- ment Station (publication #2000-2). LITERATURE CITED Bickel, D. J. 1985. A revision of the Nearctic Mede- tera (Diptera: Dolichopodidae). United States De- partment of Agriculture, Technical Bulletin No. 1692, pp. 1-109. Bullington, S. W. and A. F Beck. 1991. A new species of Machimus Loew (Diptera: Asilidae) from bur- rows of Gopherus polyphemus (Testudines: Tes- tudinidae). Annals of the Entomological Society of America 84: 590-595. Dennis, D. S. and R. J. Lavigne. 1979. Ethology of Machimus callidus with incidental observations on M. occidentalis in Wyoming (Diptera: Asili- dae). Pan-Pacific Entomologist 55: 208-221. James, M. T. 1941. The robber flies of Colorado (Dip- tera: Asilidae). Journal of the Kansas Entomolog- ical Society 14: 27-53. Lavigne, R. J. 1968. Notes on two species of Asilus (Diptera: Asilidae) associated with animal bur- rows, with a redescription of Asilus gilvipes Hine. Journal of the Kansas Entomological Society 41: 334-339. O’ Neill, K. M. 1992. Body size asymmetries in pred- atory interactions among robber flies. Annals of the Entomological Society of America 85: 34—38. . 1994. The male mating strategy of the ant Formica subpolita Mayr (Hymenoptera: Formici- dae): Swarming, mating, and predation risk. Psy- che 102: 93-108. . 1995. Digger wasps (Hymenoptera: Spheci- dae) and robber flies (Diptera: Asilidae) as pred- ators of grasshoppers (Orthoptera: Acrididae) on Montana rangeland. Pan-Pacific Entomologist 71: 248-250. O’ Neill, K. M. and W. P. Kemp. 1992. Behavioral ther- moregulation in two species of robber flies occu- pying different grassland microhabitats. Journal of Thermal Biology 17: 323-331. Rogers, L. E. and R. J. Lavigne. 1972. Asilidae of the Pawnee National Grasslands in northeastern Col- orado. University of Wyoming Agricultural Ex- periment Station. Science Monograph 25:1—35. Schreiber, E. T. and R. J. Lavigne. 1986. Ethology of Asilus gilvipes (Hine) (Diptera: Asilidae) associ- ated with small mammal burrows in southeastern Wyoming. Proceedings of the Entomological So- ciety of Washington 88: 711-719. PROC. ENTOMOL. SOC. WASH. 103(1), 2001, pp. 227-248 ANATOMICAL AND FUNCTIONAL DATA ON FEMALE AND MALE REPRODUCTIVE SYSTEMS OF SOME DUNG BEETLE SPECIES OF APHODIINAE AND EUPARIINAE OF MEXICO (COLEOPTERA: SCARABAEOIDEA: APHODIIDAE) IMELDA MARTINEZ M., CUAUHTEMOC DELOYA, AND MARCO DELLACASA (IMM) Departamento de Ecologia y Comportamiento Animal, Instituto de Ecologia, A.C. Apartado Postal 63, 91000 Xalapa, Veracruz, México; (CD) Departamento de En- tomologia, Instituto de Ecologia, A.C. Apartado Postal 63, 91000 Xalapa, Veracruz, Méx- ico; (MD) Museo di Storia Naturale e del Territorio Universita de Pisa, Via Roma 79, 56011, Calci (PI), Italy Abstract.—Female and male reproductive systems of two species of Aphodiinae, Aphodius opisthius and Cephalocyclus hogei, as well as of one species of Eupriinae, Ataenius apicalis are described. Comparisons are then made between 11 species of both subfamilies. The reproductive systems of both sexes of Aphodiinae and Eupariinae are similar, but there are differences between species of the two Aphodiinae genera and be- tween the two subfamilies. In females, ovarioles number in each ovary varies among species. The genital chamber of Aphodius and Ataenius is similar, but seems different in Cephalocyclus. The spermathecae are variable according to species. The spermathecal duct and spermathecal glandular duct originates in the same place in Aphodius, but not in Ataenius and Cephalocyclus. Males of Aphodius have 7, Cephalocyclus have 6, and Ataen- ius have 2 testicular follicles. The seminal vesicle only is present in Cephalocyclus. The glandular reservoirs are similar in Aphodius and Cephalocyclus, but differentet in Ataen- ius. The anterior part of ejaculatory duct and the internal sac differ among species of the two subfamilies. The preputial ventral gland only is present in Aphodius and Cephalo- cyclus. Resumen.—Se describe el aparato reproductor de hembras y machos de dos especies de Aphodiinae, Aphodius opisthius y Cephalocyclus hogei, y de una especie de Eupriinae, Ataenius apicalis, y se hacen las comparaciones entre las 11 especies examinadas de las dos subfamilias. El aparato reproductor de las hembras y los machos de Aphodiinae y Eupariinae estan conformados anat6émicamente de manera semejante. Sin embargo, existen diferencias entre las especies de los dos géneros de Aphodiinae y de las dos subfamilias. En las hembras el numero de ovariolas por ovario es variable segtin la especie. La camara genital es semejante en Aphodius y Ataenius, en Cephalocyclus es diferente. Las esper- matecas son diferentes de especie a especie. El] conducto espermatecal y el conducto de la glandula de la espermateca se originan en el mismo sitio en Aphodius; no asi en Ataenius y en Cephalocyclus. En los machos de Aphodius hay 7 foliculos testiculares, Cephalocyclus tiene 6 y Ataenius tiene 2. La vesicula seminal sdlo existe en Cephalo- cyclus. Los reservorios glandulares son semejantes en Aphodius y Cephalocyclus, en Ataenius son diferentes. La parte anterior del conducto eyaculador y el saco interno di- fieren entre las especies de las dos subfamilias. La glandula prepucial ventral sdélo se presenta en Aphodius y Cephalocyclus. 228 Key Words: phore The family Aphodiidae (Scarabaeoidea) (sensu Balthasar 1963) is composed of 8 tribes and 29 genera, containing 3,400 spe- cies worldwide. The subfamily Aphodiinae consists of about 1,750 species, of which 94% belong to the genus Aphodius (Della- casa 1987). The species of this genus are commonly refered to as aphodiine dung beetles. Because the focus of this paper is morphology, we have chosen to use the tra- ditional taxonomic classification, although the subgenera of Aphodius have been raised to the rank of genus by Dellacasa et al. (2000). The subfamily Eupariinae contains 593 species, of which 54% belong to the genus Ataenius which contains about 228 American species (Deloya 1994). Aphodi- idae species are characteristic of temperate regions in the Northern Hemisphere, al- though some important species are found in subtropical and tropical regions (Hanski and Cambefort 1991). The reproductive system of Aphodiidae has not been well studied. Knowledge of the female anatomy consists solely of some very generalized schemes (Willimzik 1930, Lumaret 1980, Yoshida 1994). The number of ovarioles per ovary is known for only 29 species of Aphodiinae, 2 of Eupariinae, and 3 of Psammodiinae (Stein 1847, Willimzik 1930, Ritcher and Baker 1974, Yasuda 1987, Yosida 1994, Gittings and Giller LOOM): The male reproductive system has been described for only 15 species of Aphodiinae and 2 of Eupariinae (Bordas 1900, Pluot- Sigwalt and Martinez 1998). The histolog- ical structure of the testes is known for 9 species of Aphodius and 2 of Ataenius, and of spermatozoa for 2 species of Aphodius and 4 of Ataenius (Virkki 1957, Martinez and Cruz 1999). Spermatogenesis has been studied in 10 Aphodius species (Virkki 95m): PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Aphodius, Cephalocyclus, Ataenius, females, males, anatomy, spermato- The paucity of literature on reproductive system anatomy in Aphodiidae species, and the need to characterize it in order to un- derstand their reproductive cycles, motivat- ed for this morphological study. MATERIALS AND METHODS Most of the species examined were col- lected in dung pats in various pastures, largely in the state of Veracruz, Mexico: Aphodiinae: Aphodius (Trichaphodius) op- isthius Bates, from Cuiyachapa, Veracruz; A. (Planolinus) vittatus Say, from Las Vi- gas, Veracruz; A. (Bodillus) sallei Harold, from La Mancha, Veracruz; A. (Labarrus) pseudolividus Balthasar, from Alto Lucero, Veracruz; A. (Nialaphodius) nigrita Fabri- cius, from Quiahuistlan, Veracruz; Cephal- ocyclus hogei Bates, from Cuiyachapa, Ve- racruz. Eupariinae: Ataenius apicalis (Hin- ton) and At. sculptor Harold, from Los Lir- ios, Actopan, Veracruz; At. cribrithorax Bates, from Los Tuxtlas, Veracruz; At. par- aperforatus Deloya and Ibafnez-Bernal (2000), from Ohuapan, Veracruz; and At. setiger Bates, from Tuxpan, Iguala, Guer- rero (caught in a light trap at night). For each species, 5 to 30 individuals of each sex were examined. The reproductive system was dissected in Ringer solution. Subsequently, the material was fixed in AFATD (96% ethanol-formaldehyde-trich- loroacetic acid-dimethylsulfoxide), in which it was extended, and then stored in 96% ethanol. In some cases, the reproduc- tive system was completely stained, using the Feulgen-green light technique and mounted whole in Canada balsam (Gabe 1968, Martinez 1999). To observe cuticular structures, the reproductive system was stained with chlorazol black after macera- tion with 5% potassium hydroxide (Cara- yon 1969). During dissection and after fix- ation and staining, the reproductive system VOLUME 103, NUMBER 1 was drawn to scale with the help of a cam- era lucida. In the present study, we follow the anatomical terminology of Snodgrass (1935) and Matzuda (1976). RESULTS Females The reproductive system of Aphodius, Cephalocyclus, and Ataenius females con- sists of two ovaries, two lateral oviducts, one common oviduct, a genital chamber, and the spermatheca along with its gland. Aphodius opisthius (Fig. 1A).—Each ovary is generally formed of 9 telotrophic ovarioles, although sometimes there are only 8. Each ovariole has a terminal fila- ment, germarium, vitellarium, and pedicel. The ovarioles for each ovary are covered in a fine membrane, the peritoneal sheath, which at its apex forms the ovary’s long terminal filament. The vitellarium has oo- cytes of two notably different sizes. The pedicels of each ovary lead into the respec- tive lateral oviduct. Basal oocytes of any ovariole mature simultaneously. The lateral oviducts lead into the com- mon oviduct, which is slightly longer than the lateral oviducts. The exterior walls of all oviducts are formed of circular muscles. The common oviduct narrows before join- ing the anterior part of the genital chamber. Dissection reveals the genital chamber to be a relatively bulky organ surrounded by numerous, strong muscles that run circular- ly and longitudinally. After treatment with the potassium hydroxide solution and stain- ing, this organ is found to consist of two quite different regions: an anterior part that corresponds to the bursa copulatrix, and a posterior part that might be considered as some type of vaginal sac (Fig. 2A). The bursa copulatrix is wider where the com- mon oviduct joins it; it narrows until it meets the vulva, at about the same height as the vaginal sac. The spermathecal duct ends medio-ventrally, very close to the end of the oviduct. The vaginal sac, ventral to the bursa copulatrix, is wide and rounded, 229 and shorter that the bursa copulatrix; it nar- rows until it terminates in the vulva or gon- opore. The vulva and the anus, ventral to the vulva, end in a common space that might be thought of as a cloaca. The bursa copulatrix has a thick cuticular intima with many longitudinal folds. The folds are especially numerous in the area where the common oviduct and spermathe- cal duct terminate. The vaginal sac has a very thick cuticular intima, with little fold- ing, and lateral, circular, highly sclerotized structures that have spines throughout their expanse (Fig. 2A). The spermatheca or spermathecal capsule is also sclerotized, C-shaped, with a broad, relatively flat base and a narrow apex, which is more rounded and less sclerotized than the base. Muscles are found between the apex and the base. The spermathecal duct issues from the base of the spermathe- ca, is thin and up to 4 to 5 mm long, and curls around itself before joining the bursa copulatrix. Dissection reveals the sperma- thecal gland to be voluminous, almost spherical, and with a rough wall formed by the epidermal glands (Fig. 1A). After treat- ing the spermatheca with potassium hy- droxide, the glandular reservoir appears long, with a very thin, cuticular wall and a short duct that joins the base of the sper- matheca beside the beginning of the sper- mathecal duct (Fig. 3). Comparison of Aphodius females.—The number of ovarioles per ovary varies, 8 or 9 in A. opisthius, but only 5 in A. vittatus, A. sallei, A. pseudolividus, and A. nigrita. All Aphodius species have 2 oocytes in the vitellarium, except for A. pseudolividus, which has 3. The size of the mature basal oocyte, with chorion, varies with the spe- cies (Table 1). Basal oocyte maturation is simultaneous in all ovarioles in individuals of all species examined. The genital chamber is formed by the bursa copulatrix and vaginal sac in all Aphodius species. The anterior part of the genital chamber, where the common ovi- duct joins, is highly folded and relatively 230 gale vittatus. D, A. nigrita. E, A. pseudolividus. F, Cephalocyclus hogei. (Abbreviations: bo = basal oocyte; co = common oviduct; sd = spermathecal duct; g = germarium; ge = genital chamber; lo = lateral oviduct; ol = ovarial ligament; ov = ovary; pe = pedicel; s = spermatheca; sg = spermathecal gland; vu = vulva or gonopore). enlarged in A. opisthius and A. sallei, and less folded and enlarged in A. vittatus, A. nigrita, and A. pseudolividus (Figs. 2A—E). In most Aphodius species, the bursa copu- latrix and vaginal sac are separated, and emerge nearly independently very close to the vulva (Figs. 2A—D), except in A. pseu- dolividus, in which the bursa copulatrix and vaginal sac are not entirely separate and join much before reaching the vulva (Fig. 2E). In the bursa copulatrix, the spermathe- cal duct emerges very close to the opening of the common oviduct in all species except A. pseudolividus, in which the spermathecal duct emerges farther from the common ovi- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ov Reproductive system schemes in Aphodiinae females. A, Aphodius opisthius. B, A. sallei. C, A. duct (Fig. 2E). The cuticular intima in this region varies depending on species; the ter- mination of the spermathecal duct is found only among the folds of the anterior region, as in A. opisthius and A. sallei (Figs. 2A, B) or at the back of a folded structure hav- ing a funnel-like form, as in A. vittatus, A. nigrita, and A. pseudolividus (Figs. 2C—E). The vaginal sac is shorter than the bursa copulatrix in all Aphodius species, but its structure varies among the species. The cu- ticular intima is smooth and thick in A. op- isthius, A. sallei, and A. nigrita (Figs. 2A, B, D), slightly folded in A. pseudolividus (Fig. 2E), and very folded in A. vittatus VOLUME 103, NUMBER 1 231 0.5mm Fig. 2. Female Aphodiinae genital chambers after treatment with potassium hydroxide and chlorazol black. A, Aphodius opisthius. B, A. sallei. C, A. vittatus. D, A. nigrita. E, A. pseudolividus. FE, Cephalocyclus hogei. (Abbreviations: be = bursa copulatrix; co = common oviduct; rgs = reservoir of spermatheca gland; s = spermatheca; sd = spermathecal duct; vs = vaginal sac; vu = vulva or gonopore). Table 1. Anatomical data on the female reproductive system of Aphodiinae and Eupariinae (n, number of the females per species examined in the present study). Mature in Vitellarium Oocytes Cephalo-caudal Ovarioles per Ovary Species Individuals (n) Length (mm) (number) (number) (mm) APHODIINAE Aphodius opisthius 30 4.5 9_-9* 2 0.90 A. vittatus 20 4.0 5-5 2 0.85 A. sallei 20 5.0 5-5 2 0.80 A. nigrita 6 35) 5-5 2 0.50 A. pseudolividus 25 5.0 5-5 3 0.65 Cephalocyclus hogei 30 6.0 5-5 2) 0.90 EUPARIINAE Ataenius apicalis 30 6.0 8—8* 2 0.75 At. sculptor 30 6.0 9-9 3 0.80 At. cribrithorax 20 4.0 3-3 4 0.60 At. setiger 10 4.5 3-—3* 3) 0.55 At. paraperforatus 10 6.0 11-11 3 0.50 * Number of ovarioles per ovary varies within and among females of the same species. A B ign 3: PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON C Spermatheca schemes in Aphodiinae females. A, Aphodius opisthius. B, A. sallei. C, A. vittatus. D, A. nigrita. E, A. pseudolividus. R Cephalocyclus hogei. (Abbreviations: sd = spermathecal duct; rgs = reservoir of spermatheca gland; sc = spermathecal capsule). (Fig. 2C). Further, some species show cu- ticular areas that are very sclerotized, [as in A. opisthius (Fig. 2A)], less sclerotized, [as in A. vittatus (Fig. 2C)], or not sclerotized at all [as in A. sallei, A. nigrita, and A. pseudolividus (Figs. 2B, D, E)]. The form of the spermatheca and the cu- ticular reservoir of the spermathecal gland differs from species to species, but the sper- mathecal duct is long in all species (Figs. 2, 3). In all Aphodius species, the sperma- theca is curved, with a flat and sclerotized base. The apex may be rounded, as in A. opisthius, more spherical, as in A. sallei, A. vittatus, and A. nigrita, or relatively point- ed, as in A. pseudolividus. The spermathe- cal gland reservoir joins the base of the spermatheca beside the spermathecal duct in all species. However, its form varies, be- ing long in A. opisthius and A. sallei, almost spherical in A. vittatus and A. nigrita, and rounded in A. pseudolividus (Fig. 3). Cephalocyclus hogei (Fig. 1F).—The re- productive system anatomy is similar to those of Aphodius species, but their genital chamber is different. Each ovary has 5 ovarioles. Each ovariole contains 2 oocytes, which are of two notably different sizes. Basal oocytes mature simultaneously in all oviaroles. Each ovary has a lateral oviduct attached, and these 2 oviducts join the short common oviduct. A distinctive character of this species is that the genital chamber is a sac-like vagina, very elongated, with thick walls, enlarged and with numerous folds in its anterior region; the genital chamber then narrows markedly when it approaches and terminates in the vulva. There is no sepa- ration between the bursa copulatrix and the vaginal sac. The common oviduct joins the VOLUME 103, NUMBER 1 233 Fig. 4. Reproductive system schemes in Eupariinae females. A, Ataenius apicalis. B, At. sculptor. C, At. cribrithorax. D, At. setiger. E, At. paraperforatus. (Abbreviations: bo = basal oocyte; co = common oviduct; sd = spermathecal duct; g = germarium; gc = genital chamber; lo = lateral oviduct; ol = ovarial ligament; ov = ovary; pe = pedicel; s = spermatheca; sg = spermathecal gland; vu = vulva or gonopore). chamber laterally, beneath the anterior swelling of the vagina. The spermathecal duct joins the medio-ventral region where the vagina narrows, somewhat distant from the entry of the common oviduct. The cu- ticular intima of the genital chamber is thick and shows numerous longitudinal folds, more numerous folds than those of the anterior swelling (Fig. 2F). The spermatheca is curved, with a round- ed base which is slightly wider than the apex. From the basal region, the long sper- mathecal duct, measuring up to 4 mm, emerges and then curls for its entire trajec- tory until it joins the medio-ventral region of the vagina. The spermathecal gland is oval with a rugged surface. After treatment with potassium hydroxide, the cuticular res- ervoir is seen to be fusiform and pointed (Fig. 2F), and its duct joins the medio-ven- tral region of the base of the spermatheca (Figs 3B); Ataenius apicalis (Fig. 4A).—The ova- ries have a variable number of ovarioles, most (n = 20) have 8 ovarioles per ovary, while a minority (n = 7) have 7 or 8 in the same female and a few (n = 3) have 8 or 9. The ovarioles in each ovary are wrapped in a peritoneal sheath that ends in the ovar- ial ligament. Each ovariarole is telotrophic, having a terminal filament, germarium, vi- tellarium, and pedicel. In mature females, each vitellarium has two oocytes. Pedicels join their respective lateral oviducts, which 234 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 5. A, Ataenius apicalis. B, At. sculptor. C, At. cribrithorax. D, At. setiger. E, At. paraperforatus. (Abbreviations: be = bursa copulatrix; co = common oviduct; rgs = Genital chambers in Eupariinae females after treatment with potassium hydroxide and chlorazol black. reservoir of spermatheca gland; s = spermatheca; sd = spermathecal duct; vs = vaginal sac; vu = vulva or gonopore). continue into the common oviduct, which in turn joins the genital chamber. Basal oo- cytes mature simultaneously in all ovari- oles. The genital chamber is bulky and _ sur- rounded by strong, numerous muscles (Fig. 4A). After treatment with potassium hy- droxide, the bursa copulatrix is clearly seen dorsal to the vaginal sac (Fig. 5A). The bur- sa copulatrix is short and enlarged in its an- terior region where the common oviduct joins it; subsequently it narrows until it joins the dorsal face of the vaginal sac, much before the vulva. The cuticular intima has thin walls and longitudinal folds. The spermathecal duct joins its medio-ventral anterior region, very close to the termina- tion of the common oviduct. The vaginal sac is fan shaped, with its anterior region much broader than its pos- terior region, from which it narrows until it terminates in the vulva. The cuticular inti- ma is thicker here for the vaginal sac than for the bursa copulatrix, with a rough sur- face and longitudinal folds. In the mid-re- gion of the broadest part, toward the ventral surface of the bursa copulatrix, there is a spine-like cuticular projection (Fig. 5A). The spermatheca is wider and more sclerotized in the basal region, while its api- cal region is much thinner, pointed, and less sclerotized. From the basal region emerges a short spermathecal duct, which empties into the bursa copulatrix. The spermathecal gland reservoir is long and slender, and joins the medio-ventral anterior region of the spermatheca (Fig. 6A). Comparison of Ataenius females.—The number of ovarioles per ovary differs from one Ataenius species to another, with 3 in At. cribrithorax and At. setiger, usually 8 in At. apicalis, 9 in At. sculptor, and 11 in At. paraperforatus. The number of ovarioles can also vary within any individual female VOLUME 103, NUMBER 1 of the same species. In At. apicalis, while there are generally 8 ovarioles in each ova- ry, there may be 7 or 8, or 8 and 9; and in At. setiger, which generally has 3 ovarioles per ovary, on rare occasions there may be 4 (Table 1). The number of oocytes in the vitellarium varies according to species, with 2 in At. apicalis, 3 in At. sculptor, or 4 in At. cri- brithorax. A mature basal oocyte measures 0.50 to 0.80 mm, depending on the species (Table 1). It seems that in all species oocyte maturation is simultaneous in all ovarioles of any female. The bursa copulatrix leads into the dorsal surface of the vaginal sac in all species (Fig. 5). The anterior region of the cuticular intima is thin with few folds in At. sculptor, At. cribrithorax, and At. parapeforatus (Figs. 5B, C, D), while it is broader and shows more folds in At. apicalis and At. setiger (Figs. 5A, E). The vaginal sac is long, except in At. apicalis, in which is fan shaped. The cuticular intima of the vaginal sac 1s much thicker than that of the bursa copulatrix; the cuticular intima has a rough surface and numerous longitudinal folds. The folds are not so notable in At. sculptor (Fig. 5B), but very notable in At. apicalis, At. cribrithorax, At. paraperforatus, and At. setiger (Figs. 5A, C, D, E). A spine-like structure is found on the dorsal surface of the vaginal sac only for At. apicalis, while a spherical structure similarly is found in At. paraperforatus (Figs. 5A, D). The spermatheca is relatively wide at its base and its apex is relatively pointed in all species studied. The base is more rounded and sclerotized in At. sculptor (Fig. 6B). The spermathecal duct is short in all spe- cies, measuring about 1 mm, and begins at the base of the spermatheca. On dissection, the spermathecal gland is large, with a rough surface (Fig. 4). The cuticular reser- voir of the spermathecal gland is long in all species, although its size varies from one species to another (Fig. 6). The spermathe- cal gland duct terminates in the medio-ven- DBS tral anterior region of the spermatheca in all species (Fig. 6). Comparison of Aphodius, Cephalocyclus, and Ataenius females.—Aphodius, Cephal- ocyclus, and Ataenius females all show the same general scheme of reproductive sys- tem, though some differences are found. The number of ovarioles varies among spe- cies in the same genus, from one subfamily to another, and in some cases intraspecifi- cally as well (Table 1). These characteristics are also found in other species of Aphodi- inae, Eupariinae, and Psamodiinae, insofar as the number of ovarioles is known (Table 2D): The common oviduct leads into the an- terior section of the genital chamber in Aphodius and Ataenius species, while in C. hogei the common oviduct joins the vagina (Pigs.25*5)3 The genital chamber along with bursa co- pulatrix and vaginal sac are present in Aphodius and Ataenius species, but not in Cephalocyclus, in which only a sacular gen- ital chamber or vagina is observed (Figs. 2, 5). The termination of the bursa copulatrix and of the vaginal sac are at about the same height in Aphodius; in Ataenius, the termi- nation of the bursa copulatrix is in the dor- sal wall of the vaginal sac. Most likely the vaginal sac serves to restrain the internal sac during copulation so that the ejacula- tory duct penetrates as far as the anterior section of the bursa copulatrix at the same height as the spermathecal duct opening. The spermathecal duct terminates close to the termination of the common oviduct in a highly folded structure in Aphodius and an unfolded structure in Ataenius; while in C. hogei the spermathecal duct simply ter- minates in the vaginal wall, relatively far from the end of the common oviduct (Figs. DES)! The form of the spermathecal capsule differs among the various species of Aphod- ius and Ataenius and C. hogei. The sper- mathecal duct is very long in Aphodius spe- cies and Cephalocyclus, but relatively short in Ataenius species. The spermathecal A B Fig. 6. gland varies among species, as does the form of its reservoir and the placement of its opening in the spermatheca. The sper- mathecal gland terminates in the base of the spermatheca beside the opening of the sper- mathecal duct in Aphodius, while in Ataen- ius species and Cephalocyclus it terminates in the medio-ventral anterior region of the spermatheca (Figs. 2, 3, 5). Males The reproductive system in individual Aphodius, Cephalocyclus, and Ataenius males consists of 2 testes and 2 vasa defer- entia, 2 accessory glands with their respec- tive glandular reservoirs and glandular ducts, and an ejaculatory bulb that termi- nates in the aedeagus (Fig. 7). Aphodius ophistius (Fig. 7A).—Each tes- tis has 7 spherical follicles of the same size; the corresponding vas efferens is slender and short. The vas efferens terminate in Spermatheca schemes in Eupariinae females PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Cc . A, Ataenius apicalis. B, At. sculptor. C, At. cribrithorax. D, At. setiger. E, At. paraperforatus. (Abbreviations: sd = spermathecal duct; rgs = reservoir of the spermatheca gland; sc = spermathecal capsule). their respective vas deferens. The vasa de- ferentia are curled up beneath the testis and continue until they terminate in the anterior region of the ejaculatory bulb. Accessory glands appear as very long fil- aments. Each gland terminates in a fusiform glandular reservoir, from which the glan- dular duct emerges medio-ventrally. The glandular duct is narrow and terminates parallel to the vas deferens in the anterior region of the ejaculatory bulb. The ejaculatory bulb itself is relatively large and complex, with a very thick, ex- ternal muscular wall. This wall houses not only the ejaculatory duct itself but also the anterior section of the internal sac. The ejaculatory duct is found in the anterior re- gion of the bulb where the vas deferens and glandular ducts terminate, has its own mus- cular wall, and folds back onto itself until it terminates in the anterior part of the in- ternal sac. The anterior section of the inter- VOLUME 103, NUMBER 1 Table 2. as reported in the literature. 237 Number of ovarioles per ovary in females of Aphodiinae, Eupariinae, and Psammodiinae species, _—_———— Species Ovarioles (number)* Authors —— eee APHODIINAE Aphodius (Aphodius) fimetarius 7-1 Stein 1847, Willimzik 1930, Gittings & Giller 1997 A. (Aphodius) elegans 7-7 Yoshida 1994 A. (Teuchestes) fossor WV! Willimzik 1930, Ritcher & Baker 1974, Gittings & Giller 1997 A. (Othophorus) haemorrhoidalis =) Ritcher & Baker 1974, Yoshida 1994 A. (Othophorus) brachysomus 7-71 Yoshida 1994 A. (Colobopterus) quadratus 5-5 Yoshida 1994 A. (Colobopterus) erraticus 5-5 Gittings & Giller 1997 A. (Agrilinus) pratensis 5-5 Yoshida 1994 A. (Agrilinus) breviusculus 5-5 Yoshida 1994 A. (Agrilinus) sordidus Ss) Yoshida 1994 A. (Agrilinus) ater 5-5 Gittings & Giller 1997 A. (Agrilinus) rufus 5-5 Gittings & Giller 1997 A. (Acrossus) depressus 11-11 Gittings & Giller 1997 A. (Acrossus) rufipes 7-71 Gittings & Giller 1997 A. (Melinopterus) prodromus 5) Gittings & Giller 1997 A. (Melinopterus) sphacelatus 4/5—4/5* Gittings & Giller 1997 A. (Chilothorax) inquinatus 5-5 Willimzik 1930 (=distinctus Muller) A. (Chilotorax) distinctus 5-5 Ritcher & Baker 1974 A. (Pseudogolius) coloradensis 5=5"5 Ritcher & Baker 1974 A. (Tetraclipeodes) denticulatus 5-5 Ritcher & Baker 1974 A. (Calamosternus) granarius 5-5 Ritcher & Baker 1974; Gittings & Giller 1997 A. (Labarrus) lividus 5-5 Ritcher & Baker 1974 A. (Koshantschikovius) fucosus 6-6 Ritcher & Baker 1974 A. (Cinacanthus) hirsutus 6—6* Ritcher & Baker 1974 A. (Esymus) pusillus 3=5 Yoshida 1994 A. (Phaeaphodius) rectus 7-71 Yoshida 1994 Xeropsammobeus desertus 5-5 Ritcher & Baker 1974 Aegialia blanchardi 3-3 Ritcher & Baker 1974 EUPARIINAE Ataenius cognatus 3-3 Ritcher & Baker 1974 At. deserta 3-3 Ritcher & Baker 1974 PSAMMODIINAE Psammodius oregonensis 3-—3* Ritcher & Baker 1974 Trichiorhyssemus riparius 2-2 Ritcher & Baker 1974 Pleurophorus caesus 2-2 Ritcher & Baker 1974 * Number of ovarioles per ovary can vary in females of the same species. nal sac occupies the posterior half of the ejaculatory bulb; on dissection, it can clear- ly be seen, given its highly ornamented cu- ticular form. The posterior part of the ejac- ulatory bulb is found within the aedeagus. The ventral preputial gland is found at the same level as the aedeagus parameres ba- ses. After treating the ejaculatory bulb with potassium hydroxide and staining it with chlorazol black, the ejaculatory duct’s cu- ticular intima and the internal sac can be clearly seen. The ejaculatory duct is long— when extended it measures longer than the total length of the ejaculatory bulb—and is divided into two regions. The anterior part, Fig, 7. in which the vasa deferenentia and glan- dular ducts terminate, is wide, taking the form of a funnel with low-cut borders, hav- ing a smooth, sclerotized cuticular intima. The posterior part, which terminates in the internal sac is long and narrow; its cuticular intima is smooth, slightly folded, and less sclerotized. The anterior part of the internal sac, which is found within the ejaculatory bulb, is wide, very sclerotized, and very or- namented; its posterior part, which is found in the aedeagus, is narrow, sclerotized, and ornamented (Figs. 8A, 9A). Comparison of Aphodius males.— Aphodius pseudolividus males have been described in detail by Pluot-Sigwalt and Martinez (1998); therefore we omit related histological details, considering that few differences are found at this level. In fact, A. pseudolividus has been included in this PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Schematic of reproductive system in Aphodiinae males. A, Aphodius opisthius. B, A. sallei. C, A. vittatus. D, A. nigrita. E, A. pseudolividus. EF Cephalocyclus hogei. (Abbreviations: ae = aedeagus; ag = ac- cessory glands; eb = ejaculatory bulb, including the ejaculatory duct and the anterior part of the internal sac; gd = glandular duct; gr = glandular reservoir; pg = ventral preputial gland; pa = paramere; ph = phallobase; sg = spiculum gastrale; t = testis; tf = testicular follicles; sy = seminal vesicle; vd = vas deferens). study largely as a point of comparison. We found that this species shares some char- acters the reproductive system with the oth- er species examined, but it presents differ- ences as well. Aphodius species have 7 testicular folli- cles. In most, the testicular follicles are of different sizes within an individual of any species: A. vittatus, A. sallei, A. nigrita, and A. pseudolividus all have 2 large follicles and 5 small ones; an exception to this rule is A. opisthius, which has testicular follicles of the same size. Accessory glands are long filaments in all species examined. A saculiform glan- dular reservoir is found in all species. The glandular duct begins in the posterior part of the glandular reservoir in most species, except in A. opisthius, in which it begins in the medio-ventral region (Fig. 7). VOLUME 103, NUMBER 1 A B Fig. 8. The ejaculatory bulb is found in all spe- cies, but the ejaculatory duct and internal sac differ from one species to another. The ejaculatory duct is longer than the ejacula- tory bulb in all species studied, though it is very long in A. opisthius and A. pseudoli- vidus, and relatively short in A. sallei, A. vittatus, and A. nigrita (Figs. 8, 9). The form of borders in the anterior part of the ejaculatory duct differs among species, while the cuticular intima may be smooth and without spines, as in A. opisthius, A. sallei, and A. nigrita (Figs. 8A, B, D), or have spines, as in A. vittatus and A. pseu- dolividus (Figs. 8C, E). The anterior region of the internal sac, which is observed inter- nally in the posterior area of the ejaculatory duct, is highly cuticular and presents orna- mentation that differs from one species to 239 Schematics of ejaculatory ducts in Aphodiinae males. A, Aphodius opisthius. B, A. sallei. C, A. vittatus. D, A. nigrita. E, A. pseudolividus. F, Cephalocyclus hogei. (Abbreviations: aed = anterior section of the ejaculatory duct, where the vas deferens and glandular ducts terminate; ped = posterior section of the ejaculatory duct, which terminates in the anterior part of the internal sac). another (Fig. 9). All Aphodius species have a ventral preputial gland (Fig. 7). Cephalocyclus hogei (Fig. 7F).—Each testis has 6 follicles of different sizes, with 2 of the follicles being larger than the other 4. The vas deferens show three different re- gions: (1) The anterior part, which is short, thick, and rolled up beneath the testicle; (2) the intermediate part, which is a saculiform vesicle that narrows at its base, and which continues, forming; and (3) the posterior re- gion of the vas deferens, which runs into the ejaculatory duct. Most likely, the inter- mediate region functions as a seminal ves- icle. The accessory glands form long fila- ments, running into the saculiform glandu- lar reservoir, which in turn terminates in the last third of the glandular duct ventrally. > — Fig. 9. D, A. nigrita. E, A. pseudolividus. F, Cephalocyclus hogei. (Abbreviations: ed = ejaculatory duct; gd = glandular duct; is = internal sac [anterior section]; ph = phallobase; vd = vas deferens; ms = muscular sheath). The glandular ducts terminate parallel to the vas deferens, in the anterior section of the ejaculatory bulb. The ejaculatory bulb is similar in many ways to that seen in Aphodius species. After treatment with potassium hydroxide and staining, the ejaculatory duct, when extend- ed, is seen to be longer than the ejaculatory bulb. The anterior section of the ejaculatory duct, where the vasa deferentia and glan- dular ducts terminate, is wide, with low-cut borders bearing ornamentation and spines, and a highly sclerotized cuticular intima. The posterior region is much narrower, less sclerotized, and folded until it reaches the anterior part of the internal sac (Fig. 8F). The internal sac is strongly sclerotized, pre- senting long ornamentations (Fig. 9F). A ventral preputial gland is present (Fig. 7F). Ataenius apicalis (Fig. 10A).—Each tes- tis is formed of two oval follicles with their TT get tel, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Da, ig, arity OSG wy SS TH ew ( el hy Diagrams of ejaculatory bulbs in Aphodiinae males. A, Aphodius opisthius. B, A. sallei. C, A. vittatus. respective vas efferens. The two vasa effer- entia terminate in the corresponding vas de- ferens. The vasa deferentia have a thin wall, with the slendor anterior section folded back on itself, after which these organs thin further as they continue to an unfolded pos- terior region of much greater diameter, but which thins considerably as it terminates in the ejaculatory bulb. The accessory glands are long filaments, continuing with the glandular reservoir, which itself has two regions: (1) The long anterior region, of greater diameter than the glands; and (2) the posterior region which doubles back to form a blocked sac. At the base of the sac is a short, narrow duct that terminates in the anterior region of the glan- dular duct, which in turn narrows and then terminates in the anterior part of the ejac- ulatory bulb. Glandular secretions can be found in the anterior part of the reservoir, VOLUME 103, NUMBER 1 Fig. 10. Schematics of Eupariinae male reproductive system. A, Ataenius apicalis. B, At. sculptor. C, At. cribrithorax. D, At. setiger. E, At. paraperforatus. (Abbreviations: ae = aedeagus; ag = accessory glands; eb = ejaculatory bulb, including the ejaculatory duct and internal sac; gd = glandular duct; gr = glandular reservoir; pa = paramere; ph = phallobase; sg = spiculum gastrale; t = testis; tf = testicular follicles; vd = vas deferens). which apparently functions as a duct secre- tion collector, and in the anterior part of the glandular duct, but they are mainly accu- mulated in the blind sac that functions as a true reservoir. The ejaculatory bulb has a thick muscu- lar wall. The two vasa deferentia and two glandular ducts terminate in the bulb’s an- terior part, and the base of the aedeagus ter- minates in its posterior part. In its interior, the ejaculatory duct and anterior section of the internal sac are found. The ejaculatory duct has its own muscular wall, and it ter- minates in the anterior section of the inter- nal sac, which is highly sclerotized and or- namented. After treatment with potassium hydrox- ide, the ejaculatory bulb can be clearly ob- served. It is a relatively short and folded duct, which ends by joining the internal sac. It consists of two parts: the anterior section, in which the vasa deferentia and glandular ducts terminate, is funnel-shaped, with a thick and spiny cuticular intima; and the posterior section, which terminates in the internal sac, consists of a narrower duct with a thin and very slighly folded cuticular intima (Fig. 11A). The internal sac is highly sclerotized and ornamented (Figs. 11A, 12A). Comparison of Ataenius males.—The same general scheme of reproductive sys- tem is present in all species examined. All species have two follicles per testicle, though their size varies with species (Table 3). Accessory glands, with their reservoirs and ducts, show the same general pattern anatomically. Ejaculatory ducts vary from species to species, above all the morphol- ogy of the anterior section, though in all species examined the cuticular intima in this region presented cuticular spines (Figs. 11A, B,; C, D). The internal sac differs among species also, including its ornamen- 242 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 0.5 mm Fig. 11. Diagrams of Eupariinae male ejaculatory ducts: A, Ataenius apicalis. B, At. sculptor. C, At. cri- brithorax. D, At. setiger. E, At. paraperforatus. (Abbreviations: aed = anterior region of the ejaculatory duct, where the vas deferens and glandular ducts terminate; ped = posterior part of the ejaculatory duct which terminates in the anterior region of the internal sac). Table 3. Anatomical data on the male reproductive system in Aphodiinae and Eupariinae (n, number of the individual males examined per species in the present study). Testis Follicles Preputial Glands Species Individuals (n) ~ Total Number + (Number) of a Given Size (mm) Presence (X) APHODIINAE Aphodius opisthius 30 7 (7) 0.26 x A. vittatus 20 7 (2) 0.38 + (5) 0.15 xX A. sallei 20 q (2) 0.32 + (5) 0.16 xX A. nigrita 6 V (2) 0.26 + (5) 0.13 xX A. pseudolividus 25 7 (2) 0.44 + (5) 0.22 Xx Cephalocyclus hogei 30 6 (2) 0.38 + (4) 0.26 xX EUPARIINAE Ataenius apicalis 30 2 (2) 0.60 — At. sculptor 30 2 (2) 0.96 — At. cribrithorax 20 2 (2) 0.40 — At. setiger 10 2) (2) 0.45 — At. paraperforatus 5 2) (2) 0.40 — VOLUME 103, NUMBER 1 243 Table 4. Anatomical data on the male reproductive system in Aphodiinae and Eupariinae species as reported in the literature. Follicles per Testis Preputial Of Different Gland Species Of Same Size Size Presence (X) Authors APHODIINAE Aphodius (Aphodius) fimetarius 6 — — Bordas 1900 —_ 5) ae 2 xX Pluot-Sigwalt & Martinez 1998 A. (Aphodius) coniugatus 7 = = Bordas 1900 A. (Phalacronotus) quadrimaculatus 6 — — Bordas 1900 A. (Nialus) varians 6 = _ Bordas 1900 A. (Agrilinus) ater — 5) ap 2 x Pluot-Sigwalt & Martinez 1998 A. (Biralus) satellitius — 5) Pp 2 x Pluot-Sigwalt & Martinez 1998 A. (Bodilus) sallei — Sap 2 Xx Pluot-Sigwalt & Martinez 1998 A. (Calamosternus) granarius aap 2 Xx Pluot-Sigwalt & Martinez 1998 A. (Colobopterus) erraticus —- D> aw xX Pluot-Sigwalt & Martinez 1998 A. (Coprimorphus) scrutator — 5) ae 2 Xx Pluot-Sigwalt & Martinez 1998 A. (Labarrus) pseudolividus oo 3) ar 2 x Pluot-Sigwalt & Martinez 1998 A. (Platyderides) fuliginosus* — S) ar 2 x Pluot-Sigwalt & Martinez 1998 A. (Teuchestes) fossor 6 = — Bordas 1900 7 — Xx Pluot-Sigwalt & Martinez 1998 A. (Trichaphodius) opisthius 7 -- x Pluot-Sigwalt & Martinez 1998 EUPARIINAE Ataenius cribrithorax 2 a — Pluot-Sigwalt & Martinez 1998 At. sculptor 2 -- _— Pluot-Sigwalt & Martinez 1998 * Recently this species was transferred to the genus Cephalocyclus (Dellacasa, M. et al., 1998). tation patterns (Fig. 12). None of these spe- from the glandular reservoir begins in the cies has a preputial gland. posterior section in Aphodius, Cephalocy- Comparison of Aphodius, Cephalocyclus, clus, and Ataenius, though in A. opisthius it and Ataenius males.—The number of tes- emerges medio-ventrally (Figs. 7, 10). ticular follicles per testis is 7 for Aphodius, The ejaculatory bulb has a thick muscu- 6 for Cephalocyclus, and 2 for Ataenius, ar wall that contains the ejaculatory duct consistent with other observations of and also the anterior section of the internal Aphodius and Ataenius species (Tables 3 ac in Aphodius, Cephalocyclus, and Ataen- and 4). In Aphodius fossor and A. opisthius, 5. species (Figs. 9, 12). The ejaculatory the 7 follicles are of the same size, while in duct is longer in Aphodius and Cephalocy- other Aphodius species 2 follicles are larger : : f clus species than in Afaenius, but in all than the other 5, and in Cephalocyclus 2 ; Airs 3) ; : three genera the anterior section is relative- follicles are larger than the other 4 (Tables ; ‘ ; Bead) ly sclerotized, with ornamentation or The seminal vesicle is found only in C. spines. In Aphodius, Cephalocyclus, and hogei; none is found in Aphodius or Ataen- Ataenius, the cuticular intima of the ejacu- ius species (Figs. 7 and 10) latory duct’s anterior region has a mor- Accessory glands are longer in species of phology that is characteristic for any partic- Aphodius and Cephalocyclus than in those ular species, while the posterior region has of Ataenius. The glandular reservoir is 4 thin, smooth cuticular intima, on occasion straight in Aphodius and Cephalocyclus, but with folds (Figs. 8, 11). The internal sac has in Ataenius it has a blind sac in the poste- a highly sclerotized and ornamented cutic- rior region. The glandular duct thatemerges ular wall in all species studied, though the Fig 12. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON gd Schematics of ejaculatory bulbs in Eupariinae males. A, Ataenius apicalis. B, At. sculptor. C. At. cribrithorax. D, At. setiger. E, At. paraperforatus. (Abbreviations: ed = ejaculatory duct; gd = glandular duct; is = internal sac [anterior section]; ph = phallobase; vd = vas deferens; ms = muscular sheath). specific type of ornamentions varies accord- ing to the species (Figs. 9, 12). A ventral preputial gland is found in Aphodius and Cephalocyclus, but not in Ataenius (Figs. 7, 10). Male insects form the spermatophore during copulation. The spermatophore con- sists principally of secretions of the acces- sory glands and vasa deferentia together with spermatozoa (Mann 1984). Until now, these features have not been examined for any Aphodiidae species. In the present study, observations of the genital chamber of females that recently copulated were im- mature, and without oocytes in Ataenius apicalis, At. sculptor, At. cribrithorax, and At. paraperforatus, and of some Cephalo- cyclus hogei females with small, maturing oocytes. No spermatophore was observed in any Aphodius species. In Ataenius species and in C. hogei, the spermatophore is a large, whitish structure, which when recently formed completely distends the genital chamber, occupying a volume almost half as large as the female’s abdominal cavity (Fig. 13). DISCUSSION The reproductive system of females and males in the Aphodiinae and Eupariinae species examined are similar for each sex. However, there are differences among the species of the two subfamilies and between the two genera of Aphodiinae. Females.—The number of ovarioles per ovary varies among Aphodius, Cephalocy- clus, and Ataenius species, and they can vary among the females of the same spe- cies, just as in the other 33 species of these genera that have been studied (Table 2). However, is it not known whether the num- ber of ovarioles is related to egg-laying be- VOLUME 103, NUMBER 1 Figs 13: paraperforatus. E, C. hogei. Only in the first species is the spermatophore shown in the genital chamber, though the spermatophore structure is found in all these species (Abbreviations: co = common oviduct; ov = ovary; sp = spermatophore). havior or to the fecundity of the species for any Aphodiinae species. Basal oocytes mature simultaneously in all Aphodius, Cephalocyclus, and Ataenius species studied. However until now, little has been known about whether one egg is laid, or several simultaneously, or about the fecundity (total number of eggs laid by the female of any particular species over her life span) of any of these species, data that are important in understanding reproductive cycles. Egg-laying behavior is known for only 20 Aphodius species. Some lay groups of eggs, the remainder single eggs. De- pending on the species, the number of eggs laid at any one time varies from | to 18 (Willimzik 1930, White 1960, Rojewski 1983, Yoshida and Katakura 1992, Yoshida 1994, Palestrini and Barbero 1994, Gittings and Giller 1997). Fecundity is known for only 5 Aphodius species varying from 8 to 138 eggs laid per individual over a lifetime in females that have 5 or 7 ovarioles per 245 1mm Diagrams of spermatophores in A, Ataenius apicalis. B, At. sculptor. C, At. cribrithorax. D, At. ovary (Yasuda 1987, Yoshida 1994). Only Gittings and Giller (1997), based on the study of 10 Aphodius species of 8 subgen- era, suggested a possible relationship be- tween taxonomy and egg-laying behavior. The bursa copulatrix and vaginal sac were here seen in Aphodius and Ataenius species, but not in Cephalocyclus, which simply has a sacular vagina. Willimzik (1930) described the vagina for Aphodius fossor, A. fimetarius, and A. inquinatus ( = A. distinctus Muller), adding that in the two last species the dorsal region of the vagina is highly sclerotized and expanded; how- ever, he did not indicate the separation be- tween the bursa copulatrix and vaginal sac that surely exists in these species. Males.—The number of testicular folli- cles found in Aphodius and Ataenius spe- cies is consistent with the observations of Pluot-Sigwalt and Martinez (1998) for other species in the same genera. Aphodius spe- cies always have 7 follicles in each testis. 246 Most of the species studied have testicular follicles of two sizes in any individual, while a minority have follicles of all one size. In contrast, Cephalocyclus has only 6 follicles of two different sizes per testis. Observations in other species and other genera show that differences may be found. Bordas (1900) noted that A. fossor, A. fi- metarius, A. quadrimaculatus, and A. vari- ans have only 6 testicular follicles, though these data need to be verified. Later obser- vations in A. fossor and A. fimetarius, showed that all individuals of the species appear to have 7 follicles per testis (Pluot- Sigwalt and Martinez 1998). In the present study, the seminal vesicle was observed only in C. hogei; this struc- ture was not found in any species of Aphod- ius or Ataenius. In another comparative study of Scarabaeidae, Geotrupidae, and Aphodiidae males (Pluot-Sigwalt and Mar- tinez 1998), the seminal vesicle was simi- larly not observed in any species. Cephal- ocyclus hogei males need to be studied to determine what is the significance of the seminal vesicle presence in the reproductive strategy of the species. Accessory glands are commonly seen in males of the two subfamilies studied here, although they are longer in Aphodiinae than in Eupariinae species. Together the acces- sory gland secretions and the spermatozoa from the testis form the spermatophore dur- ing copulation. However, the spermato- phore was observed only in several Ataen- ius species and Cephalocyclus hogei; ap- parently, Aphodius species do not form this structure. Questions still remain about the nature of the spermatophore. Its formation might be related to the size of the spermatozoa. In Aphodius, Cephalocyclus, and Ataenius the spermatozoa are stored in the spermatheca (Imelda Marinez M., personal observations) to be used during fertilization of the oo- cytes, but the timing of copulation and the means by which the spermatozoa arrive at the spermatheca are not known. In the Aphodiinae species examined, the sperma- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tozoa were large and of two sizes; these species also have two different sizes of tes- ticular follicles. In Aphodius, the sperma- tozoa measure from 600 to 2,000 pm, de- pending on the size of the testicular follicle, while in Ataenius they measure from 120 to 160 wm (Martinez and Cruz 1999). In C. hogei, they are 800 to 1,500 ym (M. Cruz, unpublished observations). In Ataenius, the size of the spermatozoa would not present an obstacle to the for- mation of the spermatophore, nor to the storage of the spermatheca. In contrast, in Aphodius, spermatozoa size could raise dif- ficulties for both these processes, in partic- ular, these species’ spermatheca measures only between 200 and 300 pm. If in fact no spermatophore is formed in Aphodius, what function is played by the abundant se- cretions of the accessory glands? Research is needed on the role of secretions and the behavior of spermatozoa during and after copulation. The ejaculatory bulb [in both the Apho- diinae and Eupariinae species] described in detail by Pluot-Sigwalt and Martinez (1998), is an organ with a strong muscular wall that contains the ejaculatory bulb proper and the anterior part of the internal sac. The bulb functions as a muscular pump to emit secretions and spermatozoa during copulation. The internal sac has a very sclerotized cuticular wall, which shows or- namentions in its anterior section that are characteristic of each species. Investiga- tions are needed to ascertain whether the ornamentations have some relation to the morphology of the vaginal sac in females of the same species. The preputial gland, seen only in Apho- diinae males, has been well described (Pluot-Sigwalt 1995, Pluot-Sigwalt and Martinez 1998). It apparently serves a lu- bricating function during copulation (Mar- tinez and Linares 1994). ACKNOWLEDGMENTS We are grateful for the help of Magda- lena Cruz R., Jesis Sosa M., and Mayvi VOLUME 103, NUMBER 1 Alvarado O. for insect collections; for the valuable technical contributions of A. Cle- mentina Gonzalez and Olvia M. Rodriguez; for Ann Covalt’s English translation of the manuscript; and R. E. Woodruff and anoth- er anonymous reviewer for their valuable suggestions. This work was carried out with the support of the project “Biologia y re- produccion en escarabajos coproéfagos”’ (Coleoptera Scarabaeidae: Aphodiidae) (CONACYT 27940 N) and the Departa- mento de Ecologia y Comportamiento An- imal of the Instituto de Ecologia A.C. (902— 38). LITERATURE CITED Balthasar, V. 1963. Monographie der Scarabaeidae und Aphodiidae der palearktischen und orientalischen Region. Coleoptera: Lamellicornia. III Aphodi- idae. Prague: Tschechoslowakischen Akademie der Wissenschaften. 652 pp. Bordas, L. 1900. Recherches sur les organes reprod- ucteurs males des Coleoptéres. Annales des Sci- ences Naturelles, Zoologie et Biologie Animale 11: 283-448. Carayon, J. 1969. Emploi du noir chlorazol en anato- mie microscopique des Insectes. Annales de la So- ciété Entomologique de France. (n.s.) 5: 179-193. Dellacasa, G., P. Bordat, and M. Dellacasa (2000). A revisional essay of world genus-group taxa of Aphodiinae. Memorie Societa entomologica Itali- ana. Genova. (in press). 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WASH. 103(1), 2001, pp. 249-256 CLARIFICATION OF SOME TAXONOMIC PROBLEMS IN ANISOSCELINI AND LEPTOSCELINI (HEMIPTERA: COREIDAE: COREINAE) RICHARD J. PACKAUSKAS AND CARL W. SCHAEFER Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs CT 06269-3043, U.S.A. (e-mail: schaefer@uconnvm.uconn.edu); RJP current address: Associate Curator of Entomology (Sternberg Museum), Department of Biological Scienc- es, Fort Hays State University, Hays, KS 67601, U.S.A. Abstract.—Genera previously created for species of the genera Leptoglossus Guérin- Ménéville and Anisoscelis Latreille (Anisoscelini) are given species-group status. These species groups are keyed, their included species listed with synonymies, and their dis- tributions given. Dallacoris Osuna (Leptoscelini) is a name never published and therefore invalid; its single species is restored to the genus Phthia Stal as Phthia picta (Drury). The species groups in Leptoglosus (and the number of included species) are the dilaticollis Species group (3 species), gonagra species group (1), harpagon species group (3), cinc- tipes species group (3), lineosus species group (3), and zonatus species group (24). Key Words: species groups In two theses, Osuna revised the coreine tribes Anisoscelini (1977) and Leptoscelini (1981). He made numerous taxonomic and nomenclatorial changes in these theses, in- cluding the proposing of several new gen- era for existing species. However, only one of these theses (1977, Anisoscelini) was published (Osuna 1984), and many of the changes made in the thesis (1977) were not included in the publication (1984). Cons- equently, many of Osuna’s nomenclatorial and taxonomic changes are not published and, therefore, invalid. Some have never- theless crept into the literature, and others (such as the status of the anisosceline genus Veneza) are causing confusion, because workers on the economically important Leptoglossus zonatus (Dallas) are unsure of the correct generic name (R. Zucchi, per- sonal communication). Here we discuss Osuna’s changes, reduce many of his proposed new genera to species Anisoscelini, Leptoscelini, Leptoglossus, Anisoscelis, Phthia synonyms, groups, indicate to which of these groups the various species belong (something Osu- na neglected to do in his publication [1984]), key the species groups, and give their currently known distributions and syn- onymies. Anisoscelini In 1984, Osuna published as a mono- graph part of his dissertation on the coreine tribe Anisoscelini (as Anisoscelidini; because the combining form of the Greek “‘-scelis”’ is “*-scelin-,” ‘‘Anisoscelini’” is correct [see Packauskas 1994a]). In this monograph, Osu- na (1984) created several new genera for species hitherto included in Leptoglossus Guérin-Ménéville and Anisoscelis Latreille. Unfortunately, he did not indicate what spe- cies formerly in those two genera were to be placed in the new ones except for a type species designated for each genus. This leaves many Leptoglossus species ipso fac- 250 to remaining in Leptoglossus, whereas their (presumably) close relatives are type spe- cies of other genera; the same is true of Anisoscelis species. Moreover, this mono- graph appears to be unobtainable in the United States (our copies are copies of The Natural History Museum, London, copy) and the author himself has not responded to inquiries. In an unpublished revision of the Ani- soscelini and related tribes Packauskas (1994b) has examined Leptoglossus and Anisoscelis (both sensubus latis). Packaus- kas has also studied the new genera of Osu- na (1984) as well as the species groupings in Allen’s (1969) revision of Leptoglossus, many of which Osuna proposed be raised to generic rank. We do not agree that the new genera for Leptoglossus species, or those for Anisos- celis species, are worthy of generic rank, and we discuss the reasons here. We also consider additional nomenclatorial matters. Leptoglossus Guérin-Ménéville Leptoglossus Guérin-Ménéville 1831 (1838): pl. 12. Fig. 9. Anisoscelis: Spinola 1837: 200. Theognis Stal 1862: 294. New synonymy. Theognis: Kiritschenko 1935: 191. Theognis: Hussey 1953: 33. Microphyllia Stal 1870: 167. New synon- ymy. Microphyilia (sic): Gibson & Holdridge 1918: 4. Haeckelia Kirkaldy 1904: 280. n. n. for Mi- crophyllia New synonymy. Nannophyllia Bergroth 1913: 143. n. n. for Haeckelia New synonymy. Fabrictilis Osuna 1984: 112. New synon- ymy. Leptoglossus Osuna 1984: 115. New syn- onymy. Nannophyllia: Osuna 1984: 113. Stalifera Osuna 1984: 108. New synony- my. Theognis: Osuna 1984: 111. Veneza Osuna 1984: 117. New synonymy. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Type species: Leptoglossus dilaticollis Guérin-Ménéville 1831. Allen (1969) revised the genus Lepto- glossus, described five new species, and provided a key to the 38 species known at that time. Since then, nine new species (Brailovsky 1976, 1990; Alayo and Grillo 1977; Yonke 1981; Brailovsky and Barrera 1994) have been described. In addition, Osuna (1984) transferred two species from Leptoglossus to the genus Nannophyllia Bergroth (= Microphyllia Stal). In his re- vision, Allen (1969) reviewed the genus Theognis Stal and discussed reasons for re- taining Stal’s own proposition to synony- mize this genus under Leptoglossus Guérin. Allen also proposed two large species groups, each containing four smaller spe- cies groups. In 1984, Osuna published a monograph on the tribe Anisoscelini (as ‘“‘Anisosceli- dini’’), in which he claimed to have sepa- rated the species within the genus Lepto- glossus and to have placed them in six gen- era. Three of these were new, one was an existing genus, and one was a resurrection: Leptoglossus Guérin-Ménéville sensu stricto (5 spp.); Nannophyllia Bergroth, an existing genus (7 spp.); Fabrictilis Osuna (2 spp.); Stalifera Osuna (4 spp.); Theognis Stal, a resurrected genus (3 spp.); and Ve- neza Osuna (26 spp.). The numbers in par- enthesis are taken directly from the abstract of Osuna’s paper. Nowhere in this published paper does Osuna explicitly move species into each of the newly created genera, with the exception of the type species given for each of the genera. These were the follow- ing: Cimex gonagra Fabricius (for Fabric- tilis), Theognis fasciolatus Stal (for Nan- nophyllia), Anisoscelis cincta Herrich- Schaeffer (for Stalifera), Theognis lineosus Stal (for Theognis), and Anisoscelis zonata Dallas (for Veneza). Because a type species has been designated for each of these four new genera, they are valid, but each con- tains only a single species. In order to determine what Osuna has VOLUME 103, NUMBER 1 done here, one needs to go to Osuna’s (1977) original dissertation work on the Anisoscelini, which we emphasize is un- published and whose new names are invalid under the Rules ICZN 1985, Art 9(11)). Osuna (1977), in this work, stated into which of these new genera the other Lep- toglossus species were to be placed. Ex- amining this work, one can see that Veneza Osuna consists of Allen’s (1969) Division B. Osuna’s other four genera, including Leptoglossus Guérin (sensu stricto), are all elevations of Allen’s Division A species groups to generic status. Stalifera Osuna consists of Allen’s cinctus group. Fabrictil- is Osuna may consist of Allen’s australis group. Two of the members of Allen’s harpagon group (L. harpagon and L. fla- vosignatus) were transferred to the genus Nannophyllia Bergroth, and the other three species (L. alatus, L. lineosus, and L. sub- auratus) were placed in the resurrected ge- nus Theognis Stal. This left Leptoglossus Guérin (sensu stricto) containing but three species, L. dilaticollis, L. fulvicornis, and L. rubrescens. In his monograph, Osuna referred to “generic equivalence” as the reason for the creation of four new genera, but we believe his reasoning is flawed. We do not find good external structural characters that val- idate this creation of four new genera. The main reason for doing so appears to be the contention that Stal’s Microphyllia (the name of which had been changed to Nan- nophyllia by Bergroth [1913] via Kirkaldy [1904]: see generic synonymy) is a genus and, therefore, perhaps other species groups should be also (= generic equivalence’). Nannophyllia is a problematic genus with close affinities to Leptoglossus (sensu lato) and is not separated from Leptoglossus by any distinctive characters, other than color differences. In Stal’s original description the genus was differentiated by the shape of the first antennal segment and the tiny tibial expan- sion (hence his ‘‘Microphyllia’’). Osuna’s placement (in his thesis [1977]) of two 251 more species into the genus destroyed the only good external differentiating character (small tibial expansion, which the added species do not possess), although the spe- cies are still grouped by color differences. Because there are no consistent external structural characters separating Osuna’s new genera, or Nannophyllia from Lepto- glossus sensu lato, and, moreover, because all Osuna’s new genera are based only on color characters, and otherwise have more characters in common than they have dif- ferences, we propose that his new genera and the genera Nannophyllia and resurrect- ed Theognis be considered synonyms of Leptoglossus and be reduced to species- group status. This tightens the generic lim- its of Leptoglossus and does not radically change the current taxonomy, as no other authors have followed Osuna since publi- cation of his monograph. We also believe that the addition of new genera within a tribe where very few taxa are identified or differentiated is unwarranted and can lead to confusion; instead, we recognize these units as species groups. Allen (1969) separates Leptoglossus spe- cies into two Divisions, one of which he divides further into Groups. Five of our six species groups are Groups of Allen’s Divi- sion A; the sixth of our groups (zonatus species group) is all of Allen’s Division B. Allen (1969) named four of his five Divi- sion A Groups; one of these, australis Group, we rename the gonagra species group, because Leptoglossus australis has been synonymized with L. gonagra by Slat- er and Baranowski (1986). The following key will illustrate that most of the differences are those of color- ation, and will serve to distinguish the six species groups: Key to the species groups of Leptoglossus 1. Thoracic pleura orange to dark reddish brown, without strongly contrasting yellowish mark- ings; thoracic and abdominal venter usually with numerous small piceous spots ........ 7 — Thoracic pleura with at least three, usually more strongly contrasting, yellowish maculae, fasciae; or venter widely yellow.......... 3 . Abdominal tergites with a median pale yellow longitudinal fascia... . dilaticollis species group — Abdominal tergites without a median pale yel- low longitudinal fascia ... zonatus species group 3. Clavus and corium dark brown with strongly contrasting yellow or ochraceous veins .. . lineosus species group — Clavus and corium dark, veins usually concol- orous, Or, at most, bright red, yellow ...... 4 4. Pronotal disk with narrow, arcuate, pale trans- verse fascia; thoracic pleura with 10—12 pale yellow maculae on each side; abdominal venter with 6-7 complete or maculate longitudinal LASCIAC CH ee A ee eae gonagra species group — Pronotal disc without narrow, transverse fascia, usually unicolorous or with round pale spots; pleura never with more than 6 yellowish mac- ulae on each side; abdominal venter without omenialiMell TASCO, g65 6060 ce00 cosa 508 5 5. Pronotum with 4 small yellowish spots, two on anterior disk and two on posterior marginal area (some fusion may occur); lateral margins of pronotum entire, without serrations or teeth harpagon species group — Pronotum without spots, disk entirely yellow, contrasting with remainder of pronotum and hemelytra; lateral margins of pronotum serrate cinctus species group i) List of current species of Leptoglossus Guérin-Ménéville with species-group placements and distributions dilaticollis species group Leptoglossus dilaticollis Guérin-Ménéville 1831(1838): pl. 12; Brazil, Mexico, Pan- ama. Leptoglossus fulvicornis Westwood 1842: 17; eastern U.S. Leptoglossus magnoliae Heidemann 1910: 191, syn. by Allen 1969. Theognis fulvicornis Hussey 1953: 30. Leptoglossus rubrescens (Walker), 1871: 135; Brazil. gonagra species group Leptoglossus gonagra (Fabricius) 1775: 708; world tropics and subtropics Anisoscelis precipua Walker 1871: 128, syn. by Allen 1969. Leptoglossus australis (Fabricius) PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1775: 708, syn. by Baranowski & Slater 1986. harpagon species group Leptoglossus fasciolatus (Stal) 1862: 295; Colombia. new combination. Leptoglossus flavosignatus Bléte 1936: 28; Peru. Leptoglossus harpagon (Fabricius) 1775: 101; Brazil. cinctus species group Leptoglossus cinctus (Herrich-Schaeffer) 1836: 91: Argentina, Brazil, Bolivia, Co- lombia, Costa Rica, Cuba, Fr. Guiana, Guyana, Honduras, Mexico, Panama, Paraguay, Venezuela. Leptoglossus crassicornis (Dallas) 1852: 454; Argentina, Bolivia, Colombia, Par- aguay, Uruguay. Leptoglossus fasciatus (Westwood) 1842: 17; Argentina, Brazil. lineosus species group Leptoglossus alatus (Walker) 1871: 129; Brazil. Leptoglossus lineosus (Stal) 1862: 295; Mexico. Leptoglossus subauratus Distant 1881: 126; El Salvador, Guatemala, Nicaragua. zonatus species group Leptoglossus ashmeadi Heidemann 1909: 237 ASHE SUES: Leptoglossus balteatus (Linnaeus) 1771: 534; Bahamas, Jamaica, Cuba, Dom. Re- public, Puerto Rico, St. Thomas, Greater Antilles. Anisoscelis selecta Walker 1871: 127, syn. by Allen 1969. Leptoglossus brevirostris Barber 1918: 35: southwestern U.S., Mexico. Leptoglossus chilensis (Spinola) 1852: 172; Argentina, Brazil, Chile, Paraguay, Uru- guay. Leptoglossus chilensis concaviusculus (Berg) 1892: 70, subspecific status by Allen 1969. Leptoglossus clypealis Heidemann 1910: VOLUME 103, NUMBER 1 195; Mexico, central & southwestern UES: Leptoglossus corculus (Say) 1832: 12 (326); eastern U.S. Leptoglossus concolor (Walker) 1871: 128; Belize, Costa Rica, Cuba, Dominican Re- public, Haiti, Guatemala, Panama, Puerto Rico, Mexico, Virgin Islands. Leptoglossus conspersus Stal 1870: 163; Brazil, Colombia, Mexico. Leptoglossus dentatus Berg 1892: 68; Ar- gentina, Uruguay. Leptoglossus grenadensis Allen 1969: 108; Grenadines. Leptoglossus humeralis Allen 1969: 126; Guyana, French Guiana. Leptoglossus ingens (Mayr) 1865: 434; Ar- gentina, Bolivia, Brazil, Paraguay. Anisoscelis santaremus Walker 1871: 129, syn. by Allen 1969. Leptoglossus impictipennis Stal 1870: 163; Bolivia, Guyana, Brazil, Colombia. Leptoglossus impictus (Stal) 1859: 233; Ar- gentina, Uruguay. Leptoglossus lonchoides Allen 1969: 124; Brazil, Peru. Leptoglossus macrophyllus Stal 1870: 162; Colombia, Venezuela. Leptoglossus neovexillatus Allen 1969: 113; Argentina, Bolivia, Brazil, Para- guay, Peru, Uruguay. Leptoglossus occidentalis Heidemann 1910: 196; formerly western U.S., but has moved east quickly; see McPherson et al. (1990) and Gall (1992), Mexico. Leptoglossus oppositus (Say) 1832: 12; Mexico, eastern U.S. Leptoglossus_ pallidivenosus Allen 1969: 128; Panama. Leptoglossus quadricollis (Westwood) 1842: 17; Argentina, Brazil. Leptoglossus impressicollis 1892: 69, syn. by Allen 1969. Leptoglossus phyllopus (Linnaeus) 1767: 731; U.S., Mexico, Costa Rica, Guate- mala. Leptoglossus stigma (Herbst) 1784: 258; Brazil, Ecuador, Paraguay, Surinam. Leptoglossus zonatus (Dallas) 1852: 452; Berg DS Argentina, Bolivia, Brazil, Colombia, Costa Rica, Ecuador, El Salvador, Gua- temala, Honduras, Mexico, Nicaragua, Panama, Peru, southwestern U.S., Vene- zuela. Anisoscelis vexillatus Stal 1855: 485. syn. by Allen 1969. Species of uncertain species-group placement (not examined) Leptoglossus confusus Alayo & Grillo 1977797 -"Cuba. Leptoglossus dearmasi Alayo and Grillo 1977= 101 Cuba: Leptoglossus dialeptos Brailovsky & Bar- rera 1994: 57; Venezuala. Leptoglossus digitiformis Brailovsky 1990: 121; Brazil. Leptoglossus jacquelinae Brailovsky 1976: 36; Mexico. Leptoglossus nigropearlei Yonke 1981: 213; Panama. Leptoglossus tetranotata Brailovsky & Bar- rera 1994: 60; French Guiana. Leptoglossus usingeri Yonke 1981: 217; Mexico. Leptoglossus venustus Alayo & Grillo S77 99> Cuba: Anisoscelis Latreille Anisoscelis Latreille 1829: 197. Anisoscelis: Osuna (in part) 1984: New synonymy. Bitta Osuna (in part) 1984: 104. New syn- onymy. 102. Type species: Lygaeus foliaceus Fabricius 1803: 210, designated by Laporte 1833: 31. Osuna (1984) divided Anisoscelis into two genera: Anisoscelis (sensu stricto) con- taining only the type species (A. foliacea); and a new genus, Bitta, naming only one species in the genus (the type, A. affinis). In the monograph (1984) he does not men- tion the other species included in Aniso- scelis (sensu lato), although he does so in his dissertation (1977); again, these genera (Anisoscelis [sensu stricto] and Bitta) each validly contains only a single species. 254 Osuna (1984) does not discuss the dif- ferences between his two genera. However differences are given in his key. The first character in his key—‘‘first antennal seg- ment less than two times length of head (Anisoscelis)” versus “‘first antennal seg- ment two times length of head (Bitta)’’— varies among species within both groups on either side of his division, often being near- ly to more than twice the length of the head (Packauskas 1994b). His second charac- ter—‘“‘metallic green to blue on scutellum and pronotum (except in A. discolor and A. foliacea marginella, where pale maroon) (Anisoscelis), versus pronotum and scutel- lum having yellow to white spots (Bit- ta)’’—alone does not distinguish groupings at the generic level. We see no good char- acters or valid apomorphies on which to base the raising of these groupings of spe- cies in Anisoscelis (sensu lato) to generic levels. Therefore, we consider Osuna’s gen- era to be synonyms and recognize them only as species groups, as there are more and stronger characters which unite these two groups and distinguish them as a unit from other anisosceline genera (Packauskas 1994b). Key to the species groups of Anisoscelis 1. Hind tibial dilation long, extending to apex of tibia; genital capsule of male with a U-shaped notch medially ........ foliaceus species group — Hind tibial dilation short, not extending to apex of tibia; genital capsule of male with a medial PLOCESSIINS Soyo oe es Ses affinis species group List of current species of Anisoscelis with species-group placement and distributions foliaceus species group Anisoscelis (Anisoscelis) foliaceus (Fabri- cius) 1803: 210; Brazil, Peru, Surinam. Anisoscelis (Anisoscelis) foliaceus marginellus (Dallas) 1852: 457; Brazil. Subspecific status by Osuna (1984). Anisoscelis (Anisoscelis) discolor Stal 1854: 235; Taiti? [sic] PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Anisoscelis (Anisoscelis) scutellaris Stal 1870: 159; Colombia. affinis species group Anisoscelis (Bitta) affinis Westwood 1842: 275; Mexico. Anisoscelis (Bitta) flavolineatus Blanchard 1849: pl. 6; Colombia. Anisoscelis (Bitta) gradadius Distant 1880: 122; Guatemala. Anisoscelis (Bitta) hymenipherus Westwood 1840: 275; Mexico. Anisoscelis (Bitta) podalicus Brailovsky & Mayorga 1995: 198, New combination; Costa Rica. Species of uncertain species-group placement (not examined) Anisoscelis alipes Guérin-Ménéville 1831: pl. 75; Mexico. Leptoscelini Phthia Stal Phthia Stal 1862: 294. Dallacoris Osuna 1981: dum. 75. Nomen nu- Osuna proposed the genus Dallacoris for this species, but did so in another disserta- tion (1981); he never published the name. It is therefore invalid, although Henry and Froeschner (1992) included it and the com- bination Dallacoris pictus as additions to their Catalog (1988). Phthia picta is a wide- spread (see below), mainly neotropical in- sect, and is a pest on many crops (Mitchell 2000). Phthia picta (Drury) 1770: 107; Argentina, Brazil, Colombia, Cuba, Honduras, Mex- ico, Puerto Rico, St. Martin, Surinam, Uruguay, Venezuela. ACKNOWLEDGMENTS We thank T. J. Henry for valuable advice; and we thank an anonymous reviewer for the amusing suggestion that this paper was written as an act of self-aggrandizement. VOLUME 103, NUMBER 1 LITERATURE CITED Alayo, D. P. and H. R. Grillo. 1977. Los hemipteros de Cuba—XVI. El género Leptoglossus Guérin (Hemiptera: Coreidae) en Cuba. Centro Agricul- tura 1977: 91-111. Allen, R. C. 1969. A revision of the genus Leptoglos- sus Guérin (Hemiptera: Coreidae). Entomologica Americana 45: 35-140. Baranowski, R. M. and J. A. Slater. 1986. Coreidae of Florida. Arthropods of Florida and Neighboring Land Areas. Vol. 12. Florida Department of Ag- riculture and Consumer Services. Gainesville, Florida. 82 pp. Barber, H. G. 1918. A new species of Leptoglossus: A new Blissus and varieties. Bulletin of the Brook- lyn Entomological Society 13: 35-39. Berg, C. 1892. Nova Hemiptera faunarum Argentinae et Uruguayensis. Bonariae: Pauli E. Coni et Fil- iorum. 112 pp. (reprint) Bergroth, E. 1913. Supplementum catalogi heteropter- orum Bruxellensis I. Coreidae, Pyrrhocoridae, Colobathristidae, Neididae. Mémoires de la Socie- té Entomologique de Belgique 22: 125-183. Blanchard, E. 1849. Hémiptéres. In D’Orbigny, C. Dictionnaire Universal d’ Histoire Naturelle 6. Blote, H. C. 1936. Catalogue of the Coreidae in the Rijksmuseum van Natuurlijke Historie. Part II. Coreinae, second part. Zoologische Mededeelin- gen (Leiden) 19: 23-66. Brailovsky, H. 1976. Contribucién al estudio de los Hemiptera-Heteroptera de México: VIII. Una nue- va especie de Leptoglossus Guérin (Coreidae-Cor- einae) y datos sobre distribucidn de las especies Mexicanas del género. Anales de Instituto de Biologia Universidad Nacional Autonomia Méxi- co (Serie Zoologica) 47: 35—42. . 1990. Generos nuevos y especies nuevas de coreidos neotropicales (Hemiptera-Heteroptera- Coreidae: Acanthocerini, Leptoscelidini y Anisos- celidini. Anales de Instituto de Biologia Univer- sidad Nacional Aut6nomia México (Serie Zoolo- gica) 61: 107-123. Brailovsky, H. and E. Barrera. 1994. Descripcién de cuatro especies y una subespecie nuevas de la tri- bu Anisoscelidini (Hemiptera-Heteroptera: Corei- dae). Anales de Instituto de Biologia Universidad Nacional Aut6nomia México (Serie Zoologica) 65: 45-62. Brailovsky, H. and C. Mayorga. 1995. Especie y sub- especie nuevas del género Bitta (Hemiptera-Het- eroptera: Coreidae: Anisoscelidini). Anales de In- stituto de Biologia Universidad Nacional Autén- omia México (Serie Zoologica) 66: 197—204. Dallas, W. S. 1852. List of the specimens of hemip- terous insects in the collection of the British Mus- eum. Catalog of Hemiptera. Part II. London. Pp. 369-592. Distant, W. L. 1880-1893. Insecta. Rhynchota. Hem- 255 iptera-Heteroptera. Vol. 1. Jn Godman and Salvin, eds. Biologia Centrali-Americana. London. 462 Pp. Drury, D. 1770-1782. Illustrations of Natural History, Wherein are Exhibited Upwards of Two Hundred and Forty Figures of Exotic Insects, According to their Different Genera. B. White, London. 3 vols. Fabricius, J. C. 1775. Systema entomologia, sistens in- sectorum classes, ordines, genera, species, adjectis synonymis, locis, descriptionibus, observationi- bus. Flensburgi et Lipsiae, Korte. 832 pp. . 1803. Systema Rhyngotorum secundum ord- ines, genera, species, adjectis synonymis, locis, observationibus, descriptionibus Brunsvigae, Car- olum Reichard. 314 pp. Gall, W. K. 1992. Further eastern range extension and host records for Leptoglossus occidentalis (Het- eroptera: Coreidae): well-documented dispersal of a household nuisance. Great Lakes Entomologist 25: 159-171. Gibson, E. H. and A. Holdridge. 1918. The genus Nar- nia Stal and a key to the genera of Anisoscelini A. & S. (Coreidae: Heteroptera). Psyche 25: 1—4. Guérin-Ménéville, E E. 1831 (1838). Hémiptéres In Duperrey’s Voyage Autour du Monde Executé par Ordre du Roi, sur la Corvette de la Majeste La Coquille, Pendant les Annees 1822, 1824, et 1825. Zoologie (plates X—XII, 1831). Heidemann, O. 1909. New species of Tingitidae and descriptions of a new Leptoglossus. Bulletin of the Buffalo Society of Natural Science 9: 231-238. 1910. New species of Leptoglossus from North America (Hemiptera-Coreidae). Proceed- ings of the Entomological Society of Washington 12: 191-197. Henry, T. J. and R. C. Froeschner. 1988. Catalog of the Heteroptera, or True Bugs of Canada and the Continental United States. E. J. Brill, Leiden, The Netherlands. . 1992. Corrections and additions to the “‘Cat- alog of the Heteroptera, or True Bugs of Canada and the Continental United States.”’ Proceedings of the Entomological Society of Washington 94: 263-272. Herbst, J. E W. 1780-1789. Gemeinnutzige Natur- geschichte des Thierreichs. Berlin and Stralsund. Vol. 6. Herrich-Schaeffer, G. H. 1836-1853. Die Wanzenarti- gen Insekten. C. H. Zeh’schen Buchhandlung, Niirnberg. 3: 33-114 (1836); 7: 1-16 (1842); 9: 257-348 (1851); 9 [“‘historischer Ubersicht’’ and “Index”’]: 1-210 (1853) {Dates and pagination follow Bergroth 1919, Entomologische Mittei- lungen 8: 188-189}. Hussey, R. E 1953. Concerning some North American Coreidae (Hemiptera). Bulletin of the Brooklyn Entomological Society 47: 29-34. ICZN (International Code of Zoological Nomencla- ture). 1985. Third edition. International Trust for Zoological Nomenclature, London. 338 pp. Kiritschenko, A. N. 1935. Uber die Gattungen Lepto- glossus Guérin und Theognis Stal (Hemiptera, Coreidae). Konowia 14: 191. Kirkaldy, G. W. 1904. Bibliographical and nomencla- torial notes on the Hemiptera. Entomologist 37: 279-283. Laporte, F L. de. 1833. Essai d’une classification sys- tématique de l’ordre des Hémiptéres. (Hémiptéres Hétéropteres, Latr.) Family Anisoscélites, Lygéi- tes, and Coréites. Magazin de Zoologie 52: 24— 47. Latreiile, M. 1829. La Regne Animal Distribué d’ aprés de Base a |’Histoire Naturelle des Animaux et d’Introduction a |!’Anatomie Comparée. Tom. V. Paris. Linnaeus, C. 1767. Museum Ludovicae Ulricae Regi- nae Svecorum gothorum, vandolorumque Insecta & Conchilia. Laurentio Salvii, Holmiae (Sweden). . 1771. Mantissa plantarum altera generum. Ed- itionis II. Holmiae (Sweden). impensis direct. L. Salvi. pp 143-587. Mayr, G. L. 1865. Diagnosen neuer Hemipteren II. Verhandlungen der Kaiserlich-Koniglichen Zool- ogisch-Botanischen Gesellschaft in Wien 15: 429-446. McPherson, J. E., R. J. Packauskas, S. J. Taylor and M. FE O’Brien. 1990. Eastern range extension of Leptoglossus occidentalis with a key to Lepto- glossus species of America north of Mexico (Het- eroptera: Coreidae). Great Lakes Entomologist 23: 99-104. Mitchell, P L. 2000. Economic importance of leaf- footed bugs (Coreidae). Jn Schaefer, C. W. and A. R. Panizzi, eds. Heteroptera of Economic Impor- tance. CRC Press, Boca Raton, Florida. Osuna, E. 1977. Revision generica de la tribu Anisos- celidini (Hemiptera-Heteroptera, Coreidae). Univ- ersidad Central de Venezuela, Facultad de Agron- omia, Instituto de Zoologia Agricola. Maracay, Venezuela. 166 pp. (unpublished) . 1981. Revision generica de la tribu Leptos- celidini (Hemiptera-Heteroptera-Coreidae). Univ- ersidad Central de Venezuela, Facultad de Agron- omia, Instituto de Zoologia Agricola. Maracay, Venezuela. 113 pp. (unpublished) . 1984. Monografia de la tribu Anisoscelidini (Hemiptera, Heteroptera Coreidae) I. Revision PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Genérica. Boletin Entomologia Venezolana 3: 77— 148. Packauskas, R. J. 1994a. Key to the subfamilies and tribes of the New World Coreidae (Hemiptera), with a checklist of published keys to genera and species. Proceedings of the Entomological Society of Washington 96: 44—53. . 1994b. A revision of the tribes Acanthoce- phalini, Anisoscelini, and Leptoscelini (Heterop- tera: Coreidae: Coreinae). Ph.D. Dissertation, Uni- versity of Connecticut, Storrs, Connecticut. 340 pp. (unpublished) Say, T. 1832. Descriptions of new species of hetero- pterous Hemiptera of North America. New Har- mony. 39 pp. (Reprinted 1859 by K. L. Leconte.) Spinola, M. M. 1837. Essai sur les Genres d’Insectes Apparenanta a l’ordre des Hémipteres, Lin. ou Rhyngote, Fabr. et a la section de Héteropteres, Dufour. Chez Yves Gravier, Génes. 383 pp. . 1852. Orden VII. Hemipteros. Jn Gay, His- toire Fisca y Politica de Chile. Zoologica. 7: 113— 320. Paris, Imprenta de Maulde y Renou. Stal, C. 1854. Nya Hemiptera. Ofversigt af Kongliga Svenska Vetenskaps-Akademiens Forhandlingar 11(8): 231-255. . 1855. Nya Hemiptera. Ofversigt al Kongliga Svenska Vetenskaps-Akademiens Forhandlingar 12(4): 181-192. 1859. Hemiptera species novas descripsit. Konglika Svenska Fregattens Eugenies Resa Omkring Jorden. III. (Zoologi, Insekter). Pp. 219— 298. . 1862. Hemiptera mexicana enumeravit spe- ciesque novas descripsit. Stettin Entomologische Zeitung 23: 289-325. . 1870. Enumeratio Hemipterorum. Bidrag till en f6reteckning Ofver alla hittils kinda Hemiptera, jemte systematiska meddelanden. Part 1. Konglika Svenska Vetenskaps-Akademiens Handlingar O(l) I 23 2: Walker, E 1871. Catalogue of Hemiptera—Heteroptera in the collection of the British Museum. British Museum, London. Part IV. Westwood. J. O. 1842. Catalogue of Hemiptera in the collection of the Rev. E W. Hope. Part II. J. C. Bridgewater. London. Yonke, T. R. 1981. Descriptions of two new species of neotropical Leptoglossus Guérin (Hemiptera: Cor- eidae). Proceedings of the Entomological Society of Washington 83: 213-221. PROC. ENTOMOL. SOC. WASH. 103(1), 2001, pp. 257-259 NOTE Seed Feeding by a Multispecies Swarm of Flea Beetles (Coleoptera: Chrysomelidae: Galerucinae: Alticini) The large and colorful flea beetle Omo- phoita aequinoctialis (L.) is a familiar sight in many Central American habitats, from forest edges and clearings to agricultural ar- eas. However, despite its abundance, very little is known of its life cycle and behavior. In July 1999, a large swarm of these beetles was encountered in a forest fragment in southern Costa Rica at the Wilson Botanical Garden, Las Cruces, Puntarenas Province. The beetles were concentrated along a short length of trail, and many were seen briefly alighting on the leaves of understory plants. No feeding was apparent on any of these plants; their only use by the beetles was for brief rest periods. Many other individuals in the swarm were seen crawling on the sur- face of the leaf litter on the trail, or flying just above the ground. At the slightest dis- turbance, most of those on the ground also took wing and flew back up to the under- story plants. The beetles in the leaf litter were observed feeding in the interior of broken seeds (Figs. 1, 2). These seeds, which were abundant in the litter along a 20 m stretch of the trail, came from a single canopy tree which was identified as Aegi- phila anomala Pittier (Verbenaceae). Seen from the ground, there appeared to be nu- merous seedpods on the branches of the tree, but few intact seedpods were on the ground and all those were very old and de- cayed. Seeds of this tree, however, have very hard coats and persist after the rest of the fruit has disappeared. Most of the seeds had large holes or were broken in half. In contrast to the apparent nervousness of the beetles perched on plants or crawling on top of the leaf litter, beetles actively feeding in- side the seeds were not readily disturbed and fed for several minutes at a time. The beetle swarm was not composed en- tirely of O. aequinoctialis; about ten per- cent were Asphaera nobilitata (Fabr.) and a single individual of Omophoita championi Jacoby was also observed. The Asphaera were also seen seeking out and feeding on Aegiphila seeds. Both Asphaera and Omo- phoita belong to the subtribe Oedionychina within the Alticini, and are probably con- generic, although no phylogenetic analysis has yet been attempted. Fifteen specimens of O. aequinoctialis were collected and are deposited as vouchers in the Instituto Na- cional de Biodiversidad (INBio) in Costa Rica. Despite its abundance in Central America and northern South America, there are ap- parently no explicit host plant citations for O. aequinoctialis. In his description of O. punctulata, Bechyné and Bechyné (1963. Beitrage zur Kenntnis der Salvadorischen Chrysomeloidea. Iheringia—Zoologia 31: 62) reported that species as ‘“‘found on Ver- benaceae’”’. Jolivet and Hawkeswood (1995. Host-plants of Chrysomelidae of the world. Backhuys Publishers, Leiden. 281 pp.) re- ported the genus Omophoita as associated with Verbenaceae, but without giving any specific references. On the other hand, a third species, O. simulans Jacoby, has been observed feeding on Convolvulaceae, as has A. nobilitata (Flowers and Janzen, 1997. Florida Entomologist 80: 334-366). The three Omophoita species aequinoctial- is, punctulata, and simulans are all very similar in appearence, differing only in punctation and the shape of one of the pairs of elytral spots. Except for the subfamily Bruchinae (whose larvae feed on seeds), feeding on seeds by any of the Chrysomelidae has not been previously reported. However, the as- sociation of the Las Cruces swarm with 258 Verbenaceae is consistent with what little has been reported in the literature concern- ing the host plants of Omophoita. The Ae- giphila tree that produced the seeds had rel- atively few leaves, and these were all full of small holes consistent with Alticini feed- ing damage. Due to the height of the tree, it was not possible to see if there were any insects feeding on the leaves at the moment. Gregarious behavior in leaf beetles is not uncommon and usually involves groups of adults and/or larvae exploiting a single or a group of host plants. Resulting aggregations are generally modest in size and involve a complex interplay of beetle responses to conspecifics, to their host plants, and ten- dencies to random movements (Morris et al. 1996. In P. H. A. Jolivet, and M. L. Cox, eds., Chrysomelidae Biology Volume 2: Ecological Studies. SPB Academic Publish- ing, Amsterdam, The Netherlands, pp. 303— 322.). However, two species of the flea bee- tle genus Macrohaltica (Galerucinae, Alti- cini) form stable aggregations of as many as 5,000 individuals which move together from one food plant to the next, as well as staying in non-feeding swarms (Eberhard et al. 1993. Psyche 100: 93-119). In some Cassidini and Chrysomelinae, larvae form defensive aggregations, a phenomenon called cycloalexy (Vasconcellos-Neto and Jolivet. 1988. Bulletin de la Société ento- mologique de France 92(9—10): 291—299), while in a number of species the females stay with and protect their larval broods (Jolivet. 1999. In M. L. Cox, ed., Advances in Chrysomelidae Biology 1. Backhuys Publishers, Leiden, pp. 391—409; Windsor and Choe. 1994. In P. H. Jolivet, M. L. Cox, and E. Petitpierre, eds., Novel Aspects of the Biology of the Chrysomelidae. Kluwer Academic Publishers, The Netherlands, pp. 111—117). Other Cassidini also form aggre- gations during dormancy in tropical dry forests (Flowers. 1991. Biotropica 23: 308— 310). Swarming behavior with a different ob- jective has been documented in the Clytrini (Monros. 1953. Acta Zoolégica Lilloana PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 14: 5-274; Flowers et al. 1994. Coleopter- ists’ Bulletin 48: 79-89; Flowers and Jan- zen 1997) and Alticini (Flowers and Tiffer. 1994. Brenesia. 37:135—136). Here the ac- tivity is apparently a part of mating behav- ior, although feeding by individuals not ac- tively copulating can result in substantial plant damage. Taxa where this form of mat- ing behavior has been observed include the Clytrini genera Anomoea (Flowers et al. 1994), Babia (Flowers and Janzen 1997), Megalostomis, and Temnodachrys (Monrés 1953), and an unidentified species in the monoplatine flea beetle genus Hypolampsis (Alticini; Flowers and Tiffer 1994). The swarming in Omophoita and As- phaera observed at Las Cruces does not fit into any of the types of gregariousness known to date in the Chrysomelidae. No mating activity was observed, and the seed feeding was well outside the normal leaf feeding behavior of Alticini. A further com- plicating aspect is that while beetles were seen feeding on opened seeds, none were seen chewing through the seed hulls. Thus, the opened seeds could have been a bonan- za for the beetles inadvertently provided by some vertebrate, possibly a rodent (Luis Di- ego Gomez, personal communication), and the beetles may have been opportunistically feeding on a part of their normal host plant that is not normally accessible. Such op- portunistic feeding in Chrysomelidae has also been observed in another flea beetle which feeds on Euphorbia elata (Euphor- biaceae). If a stalk of this plant is broken, the sap flowing from the cut stem rapidly attracts large numbers of Centralaphthona nr. lessmanni Bechyné & Bechyné (Chry- somelidae: Galerucinae: Alticini; Flowers and Janzen 1997). It appears that in at least these few chrysomelid species, secondary plant compounds in parts other than leaves can stimulate strong feeding responses, en- abling the leaf beetles to occasionally ex- ploit non-leaf parts of their host plants. I sincerely thank Dr. Luis Diego Gomez and the staff of the Las Cruces field station for their assistance and hospitality, and Dr. VOLUME 103, NUMBER 1 Figs. 1-2. omala. Barry Hammel (Missouri Botanical Gar- den) for his help in identifying the plant. I also thank the staff of INBio for transpor- tation and other assistance during this pe- riod. This research was supported in part by grants from INBio and the Biodiversity Re- sources Development Project, GEF/World Bank, and by a grant (FLAX 91005) from Omophoita aequinoctialis. 1, On seeds of Aegiphila anomala. 2, Feeding inside seed of A. an- CSREES, USDA, to Florida A&M Univer- sity. R. Wills Flowers, Agricultural Research Programs, Florida A&M University, Tal- lahassee, FL 32307, U.S.A. (e-mail: r. flowers @ famu.edu) PROC. ENTOMOL. SOC. WASH. 103(1), 2001, pp. 260-261 BooK REVIEWS Hawkmoths of the World. An Annotated and Illustrated Revisionary Checklist (Lepidoptera: Sphingidae). By Ian J. Kitching and Jean-Marie Cadiou. Cornell University Press, Ithaca, New York. 2000. 226 pp. Hard bound, 8.5 X 11 in. ISBN 0-8014-3734-2. The Sphingidae, commonly referred to as hawkmoths, are among the best known and most easily recognized insect families. Their large size, unique role in pollination, various unusual morphological adaptations, worldwide distribution, and the relative ease with which most can be sampled, all ensure that they will attract the attention of collectors and scientists alike. In addition, with about 1,050 species, they constitute a relatively manageable world fauna, com- pared to many insect families. Within the last century alone, the world sphingid fauna has been treated four times: Rothschild and Jordan’s (1903) classic systematic revision; the combined contributions to Seitz’s Gross-Schmetterlinge der Erde (Jordan 1911-12, Denso 1912, Hering 1927, Seitz 1928-29, Draudt 1931, Gehlern 1932); the beautifully illustrated Sphingidae Mundi of d’ Abrerra (1987); and Bridges’ (1993) thor- ough taxonomic catalogue of the family. The present work makes five. The authors of this checklist, Ian Kitch- ing and Jean-Marie Cadiou, are possibly the two most knowledgeable workers on the family Sphingidae in the world today - Kitching, curator of the extensive sphingid collection at The Natural History Museum, London, and Cadiou, curator of the largest personal collection of sphingids in the world, his own. Together they have pro- duced a book that represents the absolute state-of-the-art knowledge of the taxonomy and nomenclature of the family. The introductory chapter provides an ex- traordinarily thorough background descrip- tion of the family, with detailed sections on morphology, biology, pollination, economic impact, rarity, conservation, classification, regional distribution patterns, faunas, and a review of previous catalogue/checklists. It is chock-full of interesting facts (even if you're not a sphingid enthusiast) and ref- erences to additional resources. It is one of the most thorough and eclectic reviews of any family of Lepidoptera I have ever read. As a closet conservation biologist, I was surprised to read that, according to the au- thors, not one species of Sphingidae truly can be regarded as “endangered.” A\l- though two species formerly were consid- ered extinct (Hawaiian endemics), and one species was listed as “‘endangered”’ by the U.S. Fish and Wildlife Service, all of these subsequently have been discovered in na- ture in numbers that appear to represent sta- ble and viable populations. These three are among the sphingids that are not easily in- ventoried using standard collecting tech- niques. The checklist itself, which occupies only 17% of the entire text (i.e., 37 pages), is organized alphabetically by genera, species within each genus, subspecies within each species, and synonyms within each appro- priate taxonomic category. Although close- ly related taxa end up in very different plac- es in the checklist, the practice of arranging names alphabetically allows one to find taxa easily with little or no use of the ex- tensive index to names. All newly proposed nomenclatural changes are marked clearly with references to explanations (i.e., Notes). In the middle of the checklist, there are 8 color plates of various adult sphingids—6 plates of pinned animals and 2 plates of live animals in typical resting posture. The plates are excellent, for the most part, but reproduction of a couple of the photographs of live animals is not of the highest quality. I am a little puzzled at how the authors de- termined which species to illustrate. Maybe it is explained in the text, and I missed it. VOLUME 103, NUMBER 1 The majority of the text (112 pages) is dedicated to the extensive “‘Notes”’ section (627 of them), where all nomenclatural changes, corrections of previous mistakes, and other tidbits of information are pre- sented in a clear and well organized man- ner. This is the real “‘meat’’ of the book, and it is where the authors’ in-depth knowl- edge of the group really shines. In an attempt to make the checklist as complete as possible, the authors include an appendix in which two new species and two new subspecies are described, an unusual contribution to a checklist. Following the appendix is an addenda, apparently for in- formation discovered by the authors, or published subsequent to, the page-proof stage of the main text. For example, listed in the addenda are several species described in 1999 and 2000. This is a good indication of just how comprehensive and up-to-date the included information is. Because the family Sphingidae already is relatively well known, this treatment does not represent a giant leap forward in our understanding of the group. It does, how- ever, present the most modern, up-to-date, and thorough review of the nomenclature of the family, reflecting a high level of schol- arship and scientific knowledge typical of the authors. The overall physical quality of the book is high as well, with a solid cover, good binding, and good quality paper, typ- ical of Cornell University Press. For those interested in collecting and studying Sphingidae, this volume will be a must-have. For curators responsible for or- Revision of the Nearctic Species of the Genus Polypedilum Kieffer (Diptera: Chironomidae) in the Subgenera P. (Po- lypedilum) and P. (Uresipedilum) Oyewo and Sether. David E. Maschwitz and Ed- win E Cook, 2000. Bulletin of the Ohio 261 ganizing Lepidoptera collections, this book will be an invaluable source of current no- menclature on the family. For those who are collectors of quality books on Lepidoptera, I highly recommend this checklist, which is so much more than a checklist. LITERATURE CITED Bridges, C. A. 1993. Catalogue of the family-group, genus-group and species-group names of the Sphingidae of the world. C. A. Bridges, Urbana, Ill. 282 pp. d’Abrerra, B. 1987 (1986). Sphingidae Mundi. Hawk moths of the world. E. W. Classey, Faringdon, U.K. 226 pp. Denso, P. 1912. Appendix. Hybrids of Sphingidae, pp. 260-270. In Seitz, A., ed. Gross-Schmetterlinge der Erde 2. Draudt, M. 1931. Family: Sphingidae, pp. 845—900. In Seitz, A., ed. Gross-Schmetterlinge der Erde 6. Gehlern, B. 1932. Family: Sphingidae, hawk moths, pp. 137-166. In Seitz, A., ed. Gross-Schmetter- linge der Erde 2 (suppl.). Hering, M. 1927. Sphingidae, pp. 357-386. In Seitz, A., ed. Gross-Schmetterlinge der Erde 14. Jordan, K. 1911-12. 16. Family: Sphingidae, pp. 229— 260. In Seitz, A., ed. Gross-Schmetterlinge der Erde 2. Rothschild, L. W. and K. Jordan. 1903. A revision of the lepidopterous family Sphingidae. Novitates Zoologicae 9 (suppl.): 1-972. Seitz, A. 1928-29. 16. Family: Sphingidae, pp. 523- 576. In Seitz, A., ed. Gross-Schmetterlinge der Erde 10. John W. Brown, Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, % National Museum of Natural History, Washington, DC 20560-0168 U.S.A. (e- mail: jbrown@ sel.barc.usda. gov) PROC. ENTOMOL. SOC. WASH. 103(1), 2001, pp. 261—263 Biological Survey New Series volume 12, No 3, vi + 135 pp. ISBN 0-86727- 135-3. A not infrequent observation concerns the quantity of unfinished or unpublished taxonomic/systematic work from United States graduate students. As a newcomer to the academic system here it will be inter- esting to test this impression, and if con- firmed, identify what causes the prob- lems—after all, National Science Founda- tion dissertation improvement grants are unique. David Maschwitz’s taxonomic study of the chironomid genus Polypedil- um, completed a quarter of a century ago, belonged in this completed but unpublished category until this year. Although its con- tents have been available through Disser- tation Abstracts, taxonomic decisions, es- pecially regarding the nine new species but including many new life histories, have re- mained essentially unavailable to the as- semblage of chironomidologists. In the intervening period this active group of tax- onomists mostly have refrained from pub- lishing on North American Polypedilum. However the genus is speciose, almost global in distribution, and with immature stages encountered frequently in all kinds of waters, from glacial lakes through major rivers to small container habitats. Inevitably contributions to the literature have been made in the past 25 years, necessitating modification and updating of Maschwitz’s thesis to produce the bulletin reviewed here. In this task Maschwitz has been aided ably by Ole Szther, whose co-publications have necessitated several alterations to the no- menclature, John Epler, free-lance chiron- omidologist and aquatic coleopterist from Florida, and Brian Armitage, the director of the publishers of the volume, the Ohio Bi- ological Survey. The introduction to this work shows how valuable and innovative Maschwitz’s pro- ject was. For example, immature stage mor- phology is crucial in the identification of chironomids—in fact the principal demand for means of identifications come from aquatic biologists, and for life history stag- es other than the adult. Maschwitz under- took a labor-intensive but essential tech- nique that is not always followed in con- temporary chironomid studies, namely a PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON program of rearing individual field-collect- ed larvae to adult (preferably the male) stage. The success rate of 17% larvae reared to adult is enviable, providing the project with pupal associations for 17, and larvae for 15 of the species included. Per- haps rearing may be less difficult in this genus, since Polypedilum includes several tolerant and eurythermic species, with some gaining nuisance as swarming adults asso- ciated with nutrient-enriched habitats of the immature stages. Notably the genus in- cludes the famous cryptobiotic, west Afri- can P. vanderplanki that Hinton showed survived extraordinary extremes of temper- ature under desiccation. Although North America lacks such entomological record breakers, at least one central Californian species appears somewhat desiccation tol- erant. Following the tradition of Cook and his students, the authors critically look at the terminology used in chironomid morphol- ogy, finding cause to doubt some male hy- popygial terminology, and by extension, the homology with other nematocerans. The authors make a particularly strong review of the homologies of the structures of the larval labium and labrum and provide a sound synonymy of the terminology as used by the major students of chironomid larval morphology. Although some scepticism is cast on the homologies of “‘premandible”’ and “‘mentum’’, these and some other un- certain terms are retained, presumably due to their virtual ubiquitous use by students of Chironomidae. Keys are presented to the subgenera (mostly doubtfully monophyletic) for adult males, but it is acknowledged that the lar- vae and pupae cannot be allocated unequiv- ocally, despite some constructs by other authors. Species groups are recognised in Polypedilum sensu stricto, and it is here that the intervening years have had effects, for example, as the ‘“‘cultellatum or convictum” group has been elevated to subgeneric rank (Uresipedilum Oyewo and Sezther). The in- corporation of this into the text, and foot- VOLUME 103, NUMBER 1 noted recognition of the recent elevation of the “ontario” group as subgenus Cerobreg- ma Sether and Sundal, is less than seam- less. The descriptive text that follows is slightly less extensive than has become the case for Chironomidae, and is none the worse for this, with diagnostic features clearly portrayed. The figures are the min- imum required to illustrate the diagnostic features of each stage. Unfortunately, at the reduction employed for the figure of the complete male hypopygium, the precise shape and setation of the very important su- perior volsella cannot always be detected and only rarely is a better magnified image provided. One Nearctic species (P. flavum (Johan- nsen)) is rescued tentatively from synony- my with the widespread Palearctic P. con- victum Walker, and four species are recog- nized as Holarctic in distribution (P. cultel- latum Goetghebuer, P. nubeculosum Meigen, P. laetum Meigen and P. pedestre Meigen). The European type material for these names may be difficult to obtain, or even be lost, but reliance on selected con- temporary European interpretation of these taxa without examination of types may yet cause nomenclatural instability. Although the publication is authored jointly by Mas- chwitz and his graduate supervisor Edwin Cook (now retired from the University of Minnesota), the author of the new species is the senior author alone, providing the somewhat cumbersome attribution of au- thorship of Maschwitz in Maschwitz and Cook. The study includes discussion concerning the problems of immature stages failing to fit categories based on adult males, and cas- es in which similar larvae give rise to pupae and/or adults of differing species groups, or even subgenera. Perhaps here lies evidence of the vintage: the work lacks the prereq- uisite to understand any apparent incongru- ence between life history stages—an esti- mate of phylogeny. Seventy-five years ago 263 the great English dipterist EW. Edwards stated: “It is inconceivable that, rightly under- stood, the evidence as to ancestry provided by the larvae should be in conflict with that given by the adult forms. Where such facts appear to be the case, it can only be due to a misreading of the facts’? (Edwards, EW. 1926. Proc. II Int. Congr. Entomol. Zurich, p. 112). Edwards, far ahead of his time as ever, distinguished between ‘archaic or palingenetic and newer or coenogenetic characters” in assessing degrees of rela- tionship. He illustrated a case of false sim- ilarity as deriving from “parallel retention . of numerous archaic features” and an- other of non-resemblance of related taxa as due to “affinities ... obscured in a confus- ing complexity of secondary adaptations’. Although the former case related to larvae and the latter to adults, we now understand that any life history stages may converge, or unpredictably fail to reflect phylogeny, and as Edwards concluded, ‘‘lJarval and adult characters should be treated as com- plementary to one another, and the evi- dence derived from each stage considered together.”’ It is an opportunity missed—the message of Edwards, presented with such clarity and reinforced and elaborated later by Hennig, did not enlighten the publica- tion reviewed here. The phylogenetic rela- tionships of Polypedilum, despite the atten- tions of the Norwegian/Ghanaian school under Ole Szther, thus remain confused, with a morass of para- and even polyphyla (inter alia subgenus Pentapedilum). At least now we can identify the Nearctic species as adults, and often as immature stages, even if we remain uncertain about allocation of all stages to higher ranks—for this we should thank the author and the Ohio Bio- logical Survey for delivering this useful work after such a lengthy gestation. Peter S. Cranston, Evert and Marion Schlinger Chair in Insect Systematics, De- partment of Entomology, University of Cal- ifornia, Davis, CA 95616-8584, U.S.A. e- mail: pscranston @ ucdavis.edu PROC. ENTOMOL. SOC. WASH. 103(1), 2001, p. 264 SOCIETY MEETINGS, 2000—2001 Meetings of the Society are held at 7:00 PM in the Cathy Kerby Seminar Room (East Court—340) of the National Museum of Natural History, Smithsonian Institution, 10th and Constitution NW, Washington, DC. October 5, 2000—Arthur V. Evans, Research Associate, National Museum of Natural History, Smithsonian Institution, ‘““Spineless in Richmond: An entomologist explores Virginia” November 2, 2000—Jonathan R. Mawdsley, Smithsonian Institution Postdoctoral Re- search Fellow, “‘Biogeography of checkered beetles (Coleoptera: Cleridae) in the ‘sky islands’ of southeastern Arizona” December 14, 2000—David W. Roubik, Smithsonian Tropical Research Institute, ‘““Truth or consequences: What we know about bee population dynamics and diversity in eco- logical time”’ January 4, 2001—David G. Furth, Department of Entomology, Smithsonian Institution, ‘Searching for sumacs and flea beetles; From African poison arrows to Mexican poison ivy” February 1, 2001—Brian Wiegmann, Department of Entomology, North Carolina State University. Title to be announced. March 1, 2001—Dolores Savignano, U.S. Fish and Wildlife Service, “‘Potential impacts of pesticides on pollinators” April 5, 2001—Rex Cocroft, Division of Biological Sciences, University of Missouri, ‘“Communication and cooperation in group-living insects” May 3, 2001—Steve Gaimari, Department of Entomology, Smithsonian Institution. Title to be announced. Notice OF NEw PUBLICATION “Revision of the Net-Winged Midges of the Genus Blepharicera MacQuart (Diptera: Blephariceridae) of Eastern North America” by Gregory W. Courtney Memoirs of The Entomological Society of Washington, Number 23 (See inside back cover for ordering information) PUBLICATIONS FOR SALE BY THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MISCELLANEOUS PUBLICATIONS A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera), by E. Eric Grissell and Michael E. OPTS Sa at te La i Mod SiO oe NS em Es SR i ANT PRES En lar ae A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera): Second Edition, Revised, by E. Eric Peeelatic (iCHaeL.D. Seuatit, 6 pp. 1997 to) Wes a MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Memoirs 2, 3, 7, 9, 10, 11, and 13 are no longer available. No. 1. No. 23. The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939 A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 1952 A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp. 1957 __.. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi (CLV STP REASTUN 1) ogul [es ek RY stl ERR Ue SARL San ee ee. Sec Se. ART SD The North American Predaceous Midges of the Genus Palpomyia Meigen (Diptera: Cerato- pogonidae), by W. L. Grogan, Jr. and W. W. Wirth. 125 pp. 1979 _____-- . The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff. Fo CS SNES NR AN Ie RN a a RAD SO ae as 2 is LAE NAD 7 ae . Biology and Phylogeny of Curculionoidea, edited by R. S. Anderson and C. H. C. Lyal. 174 Pepa ad pce eer ANE tp PR ete ck ul ma Ah TCS TR Pas atch el) A nL he a Ree . A Revision of the Genus Ceratopogon Meigen (Diptera: Ceratopogonidae), by A. Borkent FEvayabN Au ba Gs ayetz) ciel peta 2TH oy 0) al AS 0° Es Wee A Mae Ao A BIA, CURSE AN es Ea Ut ROCESS Wa EULA . The Genera of Beridinae (Diptera: Stratiomyidae), by Norman E. Woodley. 231 pp. 1995 ___ . Contributions on Hymenoptera and Associated Insects, Dedicated to Karl V. Krombein, edited bypass NOTden anGeAy So Menkes 2 Tl Gyprpy i 199 Ges eae nee ae ee ae Jd eee . Contributions on Diptera, Dedicated to Willis W. Wirth, edited by Wayne N. Mathis and Wallanides Groran.Jt29 7 sop lOO 7 ves AoE ee estes ent eos aE he IA oe . Monograph of the Stilt Bugs, or Berytidae (Heteroptera), of the Western Hemisphere, by moomasi. tenrywWh49 pp yl O07 «ssh eee ALT NRA ear tas ON Sel SiR adiiee tne nye Ua ee . The Genera of Elaphidiini Thomson 1864 (Coleoptera: Cerambycidae), by Steven W. Lin- AS Rete a US pope al 99 Bees. eee et Se MND gS a AEN aU aM EP RAL Wail AU uD oe eytnee ne eae an . New World Blepharida Chevrolat 1836 (Coleoptera: Chrysomelidae: Alticinae), by David G. iver SPIRO 2 RG AS cf: Sean ee eae OSS AY Licliat, N-bee Se eNO oe > ber me UN Nb UBe! Notice apne a atc . Systematics of the North American Species of Trichogramma Westwood (Hymenoptera: ricnoprammatidae). by Joon DePintor28i/pp- 1999s see Se eee Revision of the Net-Winged Midges of the Genus Blepharicera Macquart (Diptera: Blepha- riceridae) of Eastern North America, by Gregory W. Courtney. 99 pp. 2000 ____..__-____- $10.00 15.00 $15.00 15.00 15.00 15.00 12.00 5.00 25.00 25.00 25.00 25.00 25.00 18.00 12.00 12.00 28.00 14.00 Back issues of the Proceedings of the Entomological Society of Washington are available at $60.00 per volume to non-members and $25.00 per volume to members of the Society. Prices quoted are U.S. currency. Postage extra except on prepaid orders. Dealers are allowed a discount of 10 percent on all items, including annual subscriptions, that are paid in advance. All orders should be placed with the Treasurer, Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Washington, D.C. 20560-0168. CONTENTS (Continued from front cover) OSWALD, JOHN D. and NATHAN M. SCHIFF—A new species of the genus Dilar Rambur (Neuraptera? Dilanidae) from: Borneo oo iii Bae Sat satetss Sieve clels thie lates tre at ete lake PACKAUSKAS, RICHARD J. and CARL W. SCHAEFER—Clarification of some taxonomic problems in Anisoscelini and Leptoscelini (Hemiptera: Coreidae: Coreinae) .................. QIAO, G. X. and G. X. ZHANG—A taxonomic study of the genus Anomalosiphum Takahashi from China (Homoptera: Aphididae: Greenideinae) ................ 00. cece cece cece eee ete eeeee ROBBINS, RICHARD G., STEVEN G. PLATT, THOMAS R. RAINWATER, and WENDY WEISMAN-—Siatistical measures of association between Amblyomma sabanerae Stoll (Acari: Ixodida: Ixodidae) and the furrowed wood turtle, Rhinoclemmys areolata (Dumeéril and Bibron)!(Testudimes: Emiydidae), 1nnorthern Belaze 74-2: oe one nee ee eee SMITH, DAVID R. and EDGAR ARTURO BENITEZ DiIAZ—A new species of Sericoceros Konow (Hymenoptera: Arigidae) damaging villetana trees, Triplaris caracasana Cham. (Polysonaceae) in’ Paraguay’: .(03 soos Sas cece aicion sala raul: SMe sea ta Ae aaah SPINELLI, GUSTAVO R. and WILLIAM L. GROGAN, JR.—A revision of the Patagonian preda- ceous midge genus Borkenthelea Spinelli and Grogan (Diptera: Ceratopogonidae) .......... STAINES, C. L.—A new genus and two new species of Eumolpinae (Coleoptera: Chrysomelidae) from tropical America ine... tess Selah Ne alee eet) Leal CML IO RE ich As dae STAINES, C. L. and D. M. WEISMAN—The species of Xanthonia Baly 1863 (Coleoptera: Chrysomelidae: Eumolpinae) in North America east of the Mississippi River ................ WHEELER, aS G., JR. and CRAIG A. STOOPS—Cacopsylla peregrina (Foerster) (Sternor- rhyncha: Psylloidea: Psyllidae): First U.S. records of an Old World specialist on hawthorns CE Tata SUS SPP!) 28h rare atadlirs tees ADs etre lle era lalla sale Gala Mer so cartt’ alee Aha au Rea aA SP Ree Ne NOTE FLOWERS, R. WILLS—Seed feeding by a multispecies swarm of flea beetles (Coleoptera: Chrysomelidaes Galerucinae: Alticimt) Sic aes paiden oh calc ook Set oe nese Bi ae ene BOOK REVIEWS BROWN, JOHN W.—Hawkmoths of the World. An Annotated and Illustrated Revisionary Checklist (Lepidoptera: Sphingidae), by Ian J. Kitching and Jean-Marie Cadiou.............. CRANSTON, PETER S.—Revision of the Nearctic Species of the Genus Polypedilum Kieffer (Diptera: Chironomidae) in the Subgenera P. (Polypedilum) and P. (Uresipedilum) Oyewo and Seether, by David E. Maschwitz and Edwin F. Cook ................... 0 cece e eee eee eens MISCELLANEOUS Society Meetings, 2000-2001 i merase se see eee oe eeice Santis tak 5 acide aoe eRe eerste ai alate Sane ‘Notice of New Publication v3.77 ose ce he ei leet ee ee Ge ae ne aS eT ee Die ee 74 249 143 54 alg 147 98 157 103 257. 260 vot.103 te! APRIL 2001 NO. 2 E64 x (ISSN 0013-8797) a ia Ba PROCEEDINGS of the ENTOMOLOGICAL SOCIETY of WASHINGTON PUBLISHED YVUARTERLY ARCE-PEREZ, ROBERTO and RODOLFO NOVELO-GUTIERREZ—A new genus and species of Psepheninae (Coleoptera: Dryopoidea: Psephenidae) from Mexico ....................-... 389 BALCAZAR-LARA, MANUEL A. and CARLOS R. BEUTELSPACHER—A new species of Adhemarius Oiticica (Lepidoptera: Sphingidae) from Mexico .................0cccccceececeee 312 BRENNER, GREGORY J. and JOHN D. LATTIN—Notes on three species of Anthocoridae (Hemiptera: Heteroptera) from Hawaii, including the first record of Buchananiella continua G12 ges ra SPA SoZ HONy irs ane SA er 1 Pe OAC UE AR ROS a BN ee es 386 COOK, JERRY L.—Review and first New World endemic of the strepsipteran genus Corioxenos Blur (sirepsiptera: Corioxenidac: Corioxeninde) -),. 0.0). 009 ccs de Ss beeen 396 DAVIS, DONALD R.—A new species of Prototheora from Malawi, with additional notes on the distribution and morphology of the genus (Lepidoptera: Prototheoridae) .................... 452 FERGUSON, DOUGLAS C. and SEI-WOONG CHOI—A new species of Eulithis Hiibner Geemiogptera. Geometricae) trom Calitorma: yi). She ete ee lok ee 367 GRISWOLD, TERRY—Two new species of trap-nesting Anthidiini (Hymenoptera: Megachilidae) Merit an ATEN ais CRN ite Oe or U gk gen Baan eiTeaktcas pie atta ah oa mien ea wie Re Rea GER 269 HASTRITER, MICHAEL W., MAURICIO E. ALARCON, and MICHAEL F. WHITING—A col- lection of fleas (Siphonaptera) from the San Martin Reserve, Valdivia Province, Chile ...... 437 HUERTA, HERON—A new species of the genus Atrichopogon Kieffer (Diptera: Ceratopogoni- Peedi RATT MERTON eta get en lh Asia 5 ea ck eR ame SME EA ewes Bed Coe GE Malet CL Ie 373 KOTEJA, JAN and GEORGE O. POINAR, JR.—A new family, genus, and species of scale insect (Hemiptera: Coccinea: Kukaspididae, new family) from Cretaceous Alaskan amber .......... 356 KROMBEIN, KARL V. and BETH B. NORDEN—Notes on trap-nesting Sri Lankan wasps and bees (Hymenoptera: Vespidae, Pompilidae, Sphecidae, Colletidae, Megachilidae) ............ 274 LATTIN, JOHN D. and TAMERA LEWIS—Amphiareus constrictus (Stal) (Hemiptera: Heteroptera: Anthocoridae) from California: Clarification of previous record and citation .... 334 LIU, ZHIWEI and HORACIO BONFIL—Differential performance of a Contarinia gall midge (Diptera: Cecidomyiidae) on ant-defended Acacia cornigera (Fabaceae) .................... 376 (Continued on back cover) THE ENTOMOLOGICAL SOCIETY OF WASHINGTON OFFICERS FOR 2001 JOHN W. Brown, President MICHAEL G. POGUE, Treasurer GABRIELA CHAVARRIA, President-Elect RONALD A. OcHoa, Program Chair Stuart H. McKamey, Recording Secretary STEVEN W. LINGAFELTER, Membership Chair Ho iis B. WILLIAMS, Corresponding Secretary Davip G. Furtu, Past President Jon A. LEwis, Custodian Davin R. Situ, Editor Publications Committee RAYMOND J. GAGNE THOMAS J. HENRY Wayne N. Martuis Honorary President Louise M. RUSSELL Honorary Members KARL V. KROMBEIN RONALD W. HopGEs DONALD M. ANDERSON WILLIAM E. BICKLEY All correspondence concerning Society business should be mailed to the appropriate officer at the following address: Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Wash- ington, D.C. 20560-0168. MEETINGS.—Regular meetings of the Society are held in the Natural History Building, Smithsonian Institu- tion, on the first Thursday of each month from October to June, inclusive, at 7:00 P.M. Minutes of meetings are published regularly in the Proceedings. MEMBERSHIP.—Members shall be persons who have demonstrated interest in the science of entomology. Annual dues for members are $25.00 (U.S. currency). PROCEEDINGS.—The Proceedings of the Entomological Society of Washington (ISSN 0013-8797) are pub- lished quarterly beginning in January by The Entomological Society of Washington. POSTMASTER: Send address changes to the Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Washington, D.C. 20560-0168. Members in good standing receive the Proceedings of the Entomo- logical Society of Washington. Nonmember U.S. subscriptions are $60.00 per year and foreign subscriptions are $70.00 per year, payable (U.S. currency) in advance. Foreign delivery cannot be guaranteed. All remittances should be made payable to The Entomological Society of Washington. The Society does not exchange its publications for those of other societies. PLEASE SEE PP. 771-772 OF THE JULY 2000 ISSUE FOR INFORMATION REGARDING PREPARATION OF MANUSCRIPTS. STATEMENT OF OWNERSHIP Title of Publication: Proceedings of the Entomological Society of Washington. Frequency of Issue: Quarterly (January, April, July, October). Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash- ington, D.C. 20560-0168. Editor: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Books for Review: David R. Smith, Systematic Entomology Laboratory, ARS, USDA, % Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560-0168. Managing Editor and Known Bondholders or other Security Holders: none. This issue was mailed 27 March 2001 Periodicals Postage Paid at Washington, D.C. and additional mailing office. PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 265-268 FIRST REPORT OF IXODES AURITULUS NEUMANN (ACARI: IXODIDA: IXODIDAE) FROM THE BLACKISH CINCLODES, CINCLODES ANTARCTICUS (GARNOT) (AVES: PASSERIFORMES: FURNARIIDAE), WITH ADDITIONAL RECORDS OF PARASITISM OF CINCLODES SPP. BY THIS TICK SPECIES RICHARD G. ROBBINS, LouIs N. SORKIN, AND FRANCOIS VUILLEUMIER (RGR) Armed Forces Pest Management Board, Walter Reed Army Medical Center, Washington, DC 20307-5001, U.S.A. (e-mail: robbinrg@acq.osd.mil); (LNS) Division of Invertebrate Zoology, Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024-5192, U.S.A. (e-mail: sorkin@amnh.org); (FV) Division of Vertebrate Zoology, Ornithology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024-5192, U.S.A. (e-mail: vuill@amnh.org) Abstract.—An engorged adult female of the tick /xodes auritulus (Acari: Ixodida: Ix- odidae) is reported from a nestling of Cinclodes antarcticus (Aves: Passeriformes: Fur- nariidae) on Isla Gonzalo, Diego Ramirez Archipelago, off Chile’s southernmost coast. Additional collections of /. auritulus are described from C. fuscus in Pert and southern Chile (Navarino Island), and from C. patagonicus in southern Argentina (Tierra del Fuego). These are the first published records of any tick species from members of the genus Cinclodes, which comprises about 13 species of ground- or hole-nesting ovenbirds in the Andes and Patagonia. Key Words: The American Museum of Natural His- tory (AMNH) has acquired a single fully engorged (overall length 8.5 mm, width 4.5 mm) female tick specimen that had been removed by one of us (FV) on 1 December 1985 from the neck of a dead nestling blackish cinclodes, Cinclodes antarcticus (Garnot) (Aves: Passeriformes: Furnari- idae), found away from its nest at the bot- tom of a nesting burrow on Isla Gonzalo, the second largest of southernmost Chile’s Islas Diego Ramirez (56.30S, 68.44W). The geography, vascular flora, and vertebrate fauna of these remote islands have been de- scribed by Pisano (1972) and Pisano and Schlatter (198la, b). This was the only C. antarcticus nest found by us on Isla Gon- zalo, and it contained three nestlings, now Ixodes, Acari, tick, Cinclodes, Aves, Diego Ramirez, Chile in the ornithological collections of the AMNH (numbers 817070—817072). Be- cause their gonads were not detectable an- atomically, the sex of these nestlings could not be determined by dissection. Two nest- lings (AMNH 817070, body mass 63.4 g; AMNH 817071, body mass 60.5 g) were alive when collected and had been actively fed by both parent birds. However, the par- asitized nestling (AMNH 817072; body mass 47.5 g) apparently succumbed to star- vation after wandering from the nest to the burrow, where the parents may have failed to feed it. The tick specimen’s unique suite of char- acters (hypostomal dentition 5/5 apically; palpal segment I with large, anteriorly di- rected process; scutum with sparse, shallow 266 punctations) enabled us to immediately identify it as Ixodes (Multidentatus) auri- tulus Neumann, a regular parasite of pas- seriform, galliform and, less often, ciconi- iform (sensu Monroe and Sibley 1993) birds throughout the Western Hemisphere (Cooley and Kohls 1945, Arzua et al. 1994, Foster et al. 1996, Arzua and Barros-Bat- testi 1999). The specimens in the type se- ries of J. auritulus came from Punta Arenas, southern Chile, but the host bird was not identified (Neumann 1904). Among_ the passeriform birds parasitized by J. auritulus in South America are species in the subos- cine families Conopophagidae, Formicari- idae, Furnariidae and Tyrannidae, and the oscine families Parulidae, Thraupidae and Turdidae (Arzua and Barros-Battesti 1999, who followed the familial nomenclature of Meyer de Schauensee ‘1983’? [1982]). Outside the Americas, J. auritulus has been recorded from Antarctica (Gressitt and We- ber 1959), Australia (Roberts 1970), and New Zealand (Dumbleton 1961). To date, I. auritulus has not been reported from con- tinental Africa, though Arthur (1965) ar- gued that, in light of its very wide occur- rence in the Southern Hemisphere, “‘it would seem reasonable to believe”’ that this tick will one day be found there. In this regard, Theiler (1959) cited two records of I. auritulus from Marion Island (46.52S, 37.51E), which lies over 1,900 km south- east of Cape Town, South Africa, but is ad- ministered by that country. This is the first published report of J. au- ritulus—or any tick species—from Cinclo- des. As such, it begs the question of wheth- er I. auritulus parasitizes other members of this genus, which comprises about 13 spe- cies of ground-foraging and hole- or bur- row-nesting South American ovenbirds (sometimes called shaketails) found in open habitats. Species of Cinclodes chiefly in- habit high-altitude streamsides in the Andes Mountains and Patagonia, and similar streamsides at lower elevations as well as seacoasts in the Tierra del Fuego and Cape Horn Archipelagos and the Falkland Islands PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (Sibley and Monroe 1990, Ridgely and Tu- dor 1994, Vuilleumier, personal observa- tion). Two subspecies of C. antarcticus have been described: nominate C. a. an- tarcticus from the Falkland Islands, where it is now most abundant on outer islands (Woods 1988, Strange 1992, Vuilleumier 1996); and C. a. maculirostris Dabbene, which is localized, and scarce to rare, from southernmost South America, where it is largely restricted to the archipelagos of Ti- erra del Fuego, Cape Horn, and Diego Ra- mirez (Hellmayr 1925, Ridgely and Tudor 1994, Vuilleumier, personal observation). Both subspecies occur mostly along sea- shores, where they forage for invertebrates, such as marine amphipods and mollusks, in the dense mats of seaweed and kelp (Ma- crocystis spp.) of the intertidal zone and in the peaty soil among tussocks of Poa fla- bellata (Lamarck) Raspail (Poaceae), a grass characteristic of the subantarctic zone (Moore 1983). Cinclodes antarcticus is of- ten found in or near breeding colonies or resting aggregations of seabirds, especially penguins (Spheniscidae), albatrosses (Di- omedeidae), diving-petrels (Pelecanoididae) and petrels (Procellariidae), as well as such marine mammals as sea lions and fur seals (Otariidae) and elephant seals (Phocidae). Indeed, C. antarcticus can be regarded as a commensal of seals and seabirds in that it both lives among them and consumes their excreta. It is probably no coincidence that I. auritulus has been recorded from the same subantarctic areas where these sea- birds and mammals breed or congregate in nonbreeding groups. Thus, in New Zealand, Dumbleton (1953) reported J. auritulus from the nest material of the diving-petrel Pelecanoides urinatrix (Gmelin), a species that is also abundant in the Diego Ramirez Islands (Schlatter 1984, Vuilleumier, per- sonal observation). We have located only three additional collections of I. auritulus from Cinclodes, all from species other than C. antarcticus: 1 2 ex C. fuscus (Vieillot), the bar-winged cinclodes, Pert: Cercapuquio (12.23S, VOLUME 103, NUMBER 2 75.19W), 7.VI.1969, W.E. Dale, Rocky Mountain Laboratories (RML) No. 59286; bor .-2 nympns cx. C6. juscus, Chile: Isla Navarino (55.05S, 67.40W), 19.1.1964, G.E. Watson, RML 117504; and 1 nymph ex C. patagonicus (Gmelin), the dark-bel- lied cinclodes, Argentina: Bahia Buen Su- ceso (54.49S, 65.13W), 24.IV.1971, collec- tor unknown (but likely either G.E. Watson or J.P. Angle of the U.S. National Museum of Natural History, or David Bridge, as all three men collected birds at Bahia Buen Su- ceso between 22 and 26 April 1971 (Payn- ter 1995)), RML 118083. Cinclodes fuscus, either a single species or a group of sister species, has the largest range of any mem- ber of its genus, occurring from northern Colombia and Venezuela to the Fuegian and Cape Horn archipelagos. It is probably also the most common Cinclodes, breeding chief- ly at high elevations in the Andes and Pa- tagonia, where it favors streamside habitats and moorlands (Vuilleumier, personal ob- servation). Two individuals of C. fuscus, apparently paired, were studied by FV on Isla Gonzalo on 1 December 1985, but no evidence of nesting was detected. Cinclodes patagonicus is a locally common species that occurs in southern Chile and Argentina, including Tierra del Fuego and Cape Horn, where it may live sympatrically and syn- topically with C. antarcticus, to which it may also be phylogenetically close. Al- though both C. patagonicus and C. antarc- ticus live along seacoasts, the latter is de- cidedly more maritime and is more inti- mately associated with marine birds and mammals (Vuilleumier, personal observa- tion). The records summarized here suggest that 7. auritulus is capable of parasitizing any species of Cinclodes throughout the vast collective range of this genus. Unfor- tunately, they also suggest that, at the dawn of the twenty-first century, we have suc- ceeded in sampling only a minority of suit- able hosts and have barely begun to under- stand the ecological relationships between birds and their ectoparasites. 267 ACKNOWLEDGMENTS For their constructive comments on an earlier version of this work, we thank our colleagues at the Armed Forces Pest Man- agement Board: Commander George W. Schultz, U.S. Navy, and Lieutenant Colonel Richard N. Johnson, U.S. Army. We are es- pecially grateful to Lance A. Durden, Insti- tute of Arthropodology and Parasitology, Georgia Southern University, Statesboro, for generously supplying all records of 1. auritulus in the U.S. National (formerly Rocky Mountain Laboratories) Tick Collec- tion, where he is Associate Curator, and for providing copies of several otherwise in- accessible publications. Francois Vuilleu- mier’s visit to Islas Diego Ramirez was made possible through the courtesy of the Chilean Navy, and especially Comandante Marco Groetaers T., Governor of the Chi- lean Antarctic Province in Puerto Williams, Navarino Island, and Comandante Eduardo Garcia D., Master of the Yelcho. In addi- tion, Vuilleumier thanks Dr. Tobias Salathé for field assistance and gratefully acknowl- edges the financial support received from the National Science Foundation, the Na- tional Geographic Society, and the Sanford Fund for his field work in South America generally and southern South America and the Falkland Islands in particular. The opin- ions and assertions advanced herein are those of the authors and are not to be con- strued as official or reflecting the views of the U.S. Departments of the Army or De- fense. LITERATURE CITED Arthur, D. R. 1965. Ticks of the Genus /xodes in Af- rica. The Athlone Press, University of London. Arzua, M., D. M. Barros, P. M. Linardi, and J. R. Bo- telho. 1994. Noteworthy records of Ixodes auri- tulus Neumann, 1904 (Acari, Ixodida) on birds from Parana, southern Brazil. Memorias do Insti- tuto Oswaldo Cruz 89: 129. Arzua, M. and D. M. Barros-Battesti. 1999. Parasitism of lxodes (Multidentatus) auritulus Neumann (Ac- ari: Ixodidae) on birds from the City of Curitiba, State of Parana, southern Brazil. Memorias do In- stituto Oswaldo Cruz 94: 597—603. 268 Cooley, R. A. and G. M. Kohls. 1945. The genus /x- odes in North America. National Institute of Health Bulletin 184: 1—246. Dumbleton, L. J. 1953. The ticks (Ixodoidea) of the New Zealand sub-region. Cape Expedition Series Bulletin 14: 1—28. . 1961. The ticks (Acarina: Ixodoidea) of sea birds in New Zealand waters. New Zealand Jour- nal of Science 4: 760—769. Foster, G. W., J. M. Kinsella, R. D. Price, J. W. Mer- tins, and D. J. Forrester. 1996. Parasitic helminths and arthropods of greater shearwaters (Puffinus gravis) from Florida. Journal of the Helminthol- ogical Society of Washington 63: 83-88. Gressitt, J. L. and N. A. Weber. 1959. Bibliographic introduction to Antarctic-Subantarctic entomolo- gy. Pacific Insects 1: 441—480. Hellmayr, C. E. 1925. Catalogue of birds of the Amer- icas. Part IV. Furnariidae-Dendrocolaptidae. Field Museum of Natural History Publication 234, Zoo- logical Series, 13: i-iv, 1-390. Meyer de Schauensee, R. 1982. A Guide to the Birds of South America. Reprint with additions by In- tercollegiate Press, Philadelphia. Monroe, B. L. and C. G. Sibley. 1993. A World Check- list of Birds. Yale University Press, New Haven, Connecticut. Moore, D. M. 1983. Flora of Tierra del Fuego. An- thony Nelson, Oswestry, United Kingdom, and Missouri Botanical Garden, St. Louis. Neumann, L. G. 1904. Notes sur les ixodidés. H. Ar- chives de Parasitologie 8: 444—464. Paynter, R. A. 1995. Ornithological gazetteer of Ar- gentina. Second edition. Museum of Comparative Zoology, Harvard University, Cambridge, Mas- sachusetts. Pisano, V. E. 1972. Observaciones fito-ecolégicas en PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON las Islas Diego Ramirez. Anales del Instituto de la Patagonia, Punta Arenas 3: 161—169. Pisano, V. E. and R. P. Schlatter. 198la. Vegetacion y flora de las Islas Diego Ramirez (Chile). I. Car- acteristicas y relaciones de la flora vascular. An- ales del Instituto de la Patagonia, Punta Arenas 12: 183-194. . 1981b. Vegetacion y flora de las Islas Diego Ramirez (Chile). Il. Comunidades vegetales vas- culares. Anales del Instituto de la Patagonia, Punta Arenas 12: 195—204. Ridgely, R. S. and G. Tudor. 1994. The Birds of South America, Vol. II: The Suboscine Passerines. Uni- versity of Texas Press, Austin. Roberts, EF H. S. 1970. Australian Ticks. CSIRO, Mel- bourne, Australia. Schlatter, R. P 1984. The status and conservation of seabirds in Chile, pp. 261—269. In Crozxall, J. P., P. G. H. Evans and R. W. Schreiber, eds., Status and Conservation of the World’s Seabirds. Inter- national Council for Bird Preservation Technical Publication No. 2, Cambridge, United Kingdom. Sibley, C. G. and B. L. Monroe. 1990. Distribution and Taxonomy of Birds of the World. Yale University Press, New Haven, Connecticut. Strange, I. J. 1992. A Field Guide to the Wildlife of the Falkland Islands and South Georgia. Harper Collins Publishers, London. Theiler, G. 1959. African ticks and birds. Proceedings of the lst Pan-African Ornithological Congress, Ostrich Supplement 3: 353-378. Vuilleumier, E 1996. Is the avifauna of the Falkland (Malvinas) Islands impoverished or at equilibri- um? Southern Connection Newsletter (10): 22-33. Woods, R. W. 1988. Guide to the Birds of the Falkland Islands. Anthony Nelson, Oswestry, United King- dom. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 269-273 TWO NEW SPECIES OF TRAP-NESTING ANTHIDIINI (HYMENOPTERA: MEGACHILIDAE) FROM SRI LANKA TERRY GRISWOLD USDA-ARS Bee Biology and Systematics Laboratory, Utah State University, Logan, UT 84322-5310, U.S.A. (e-mail: tgris@biology.usu.edu) Abstract.—Two new trap-nesting species of Anthidiellum (Pycnanthidium), A. butarsis and A. krombeini, are described from Sri Lanka. Comparisons are made with other known Oriental species of the subgenus. Key Words: The known bee fauna of Sri Lanka sug- gests that the tribe Anthidiini (Megachili- dae) is poorly represented. Only Anthidiel- lum (Pycnanthidium) ramakrishnae (Cock- erell) (Pasteels 1972), Pseudoanthidium (Exanthidium) rotundiventre (Pasteels 1987), and the parasitic Euaspis edentata Baker (Pasteels 1980 as E. carbonaria Smith, Baker 1995) are recorded from Sri Lanka. Evidence that our knowledge of the anthidiines of Sri Lanka is far from com- plete is demonstrated by recent collections of megachilid nests by Karl Krombein and Beth Norden which yielded two new spe- cies of Anthidiellum. Here these new spe- cies are described to make the names avail- able for descriptions of their nesting biol- ogies (Krombein and Norden 2001). Terminology in the descriptions follows that of Michener (2000). The following ab- breviations are used: T1, T2, ... to repre- sent the tergal segments of the apparent me- tasoma; S1, S2, ... similarly for the sterna, and Fl, F2, ... for the flagellar segments of the antenna. The order of female and male descriptions is reversed between the two species because the holotypes are not of the same sex. Ordinarily species in the same subgenus would be described from the same sex to facilitate comparisons be- bees, Megachilidae, Anthidiini, Anthidiellum, Sri Lanka, trap-nest tween the species. In this case the two spe- cies belong to different species groups, in one of which the females apparently bear the most diagnostic features, in the other, the males. Further, types for all described species of the former group are females. Thus the decision for holotypes of the op- posite sex. Anthidiellum (Pycnanthidium) butarsis Griswold, new species (Figs: 13; 5) Female.—Length, 6 mm; forewing length, 4.5 mm. Black except as follows: white marks on basal two-thirds of mandi- ble, T-shaped mark on clypeus, paraocular stripe to level of midocellus, basal spot on fore- and midtibiae, ventral stripe on fore- femur, outer face of hind tibia except for discal black spot, outer faces of basitarsi; lemon yellow marks obliquely mesal to an- tennal socket, stripe on gena, spot on pron- otal lobe, longitudinal stripe laterally on scutum, spot on axilla, medially interrupted posterior mark on scutellum, metapleural spot, tergal marks: lateral spot on T1 be- coming broader and more nearly joined on succeeding terga to simple quadrate mark on T6; light red predominantly on femora, inner surfaces of tibiae. Wings hyaline. Pu- 270 bescence white. Scopa yellow. Punctation of head, thorax dense except on anterior surface of mesopleuron; coarser on clypeus dorsally, supraclypeal area, lateral face of mesopleuron dorsally; very fine, contiguous on scutum, scutellum. Anterior face of T1 matt without punctation; T1-5 with narrow impunctate apical margins approximately 1- 2 punctures wide; T1 densely, nearly con- tiguously punctate; T2-5 less densely punc- tate especially basomedially; T6 contigu- ously punctate. Mouthparts not exceeding fossa in re- pose. Mandible four-toothed, third tooth ob- scure. Clypeal margin with four denticles. Subantennal suture slightly arcuate. Prono- tal lobe narrowly lamellate, anterior margin straight. Scutal-scutellar suture narrowly fo- veolate, not divided medially. Scutellum rounded posteriorly with shallow notch me- dially. Omaular carina complete, reaching ventral midline. Hind basitarsus one and one half times as long as broad (Fig. 3). Second hind tarsus broader than long. Preapical ridge of T6 slightly notched me- dially in dorsal view, in posterior view up- turned medially. Scopal hair blunt apically. Male.—Length, 5 mm; forewing length, 4 mm. Markings as in female except: entire lower half of face yellow except for small interantennal spot, T3-5 with complete yel- low bands, margins translucent. Fl as broad as long. F2-10 one and one half times as long as broad. Hind basitarsus twice as long as broad. Margin of T7 with median angle in ventral view. Margin of S4 with blunt submedian black spine. S5 with wide V-shaped posterior margin, margin with black comb well separated medially (Fig. 5). Genitalia as in Fig. 1. Types.—Holotype 2. SRI LANKA, Kandy District, Anniewatte, Kandy, 7°18’N 80°38’E, nest collected 5 Apr 1997, nest 18, K. V. Krombein, B. B. Norden, A. W. Nor- den, J. W. Norden. Reared paratypes: 2 2, same data as holotype; 1 2, same data ex- cept foundress. Non-reared paratypes, all same locality data and collectors: 1 2, 1- 31 Mar 1997; 1 3d, 1 2, 9-30 Apr 1997; 3 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ?, 1-8 May 1997. Holotype in the National Museum of Natural History, Smithsonian Institution, Washington DC. Paratypes at Washington and Utah State University, Lo- gan. Discussion.—Anthidiellum butarsis be- longs in a group of species that differ from other A. (Pycnanthidium) by the omaular carina reaching the ventral midline, the ab- sence of a longitudinal carina on the hind tibia and basitarsus and by the enlarged hind basitarsus of the female. This group has in the past been accorded subgeneric status (Pygnanthidiellum Mavromoustakis) but was synonymized by Michener and Griswold (1994). They are well represented in Africa, but the only known Oriental Re- gion representatives are A. latipes (Bing- ham), A. ramakrishnae (Cockerell), and A. rasorium (Smith). Anthidiellum butarsis can be distinguished from A. /Jatipes and A. rasorium by the greatly enlarged hind bas- itarsus. It further differs from A. rasorium from southern India in the fine, contiguous scutal punctation and the hindtarsal seg- ment two of the female broader than long. None of the African species I have studied has the fine scutal punctation either. Anthidiellum butarsis appears close to A. ramakrishnae from southern India. An evaluation of its relationship is complicated by the condition of the type of A. ramak- rishnae, the only known specimen of this species. The female holotype lacks the di- agnostic hind legs, the metasoma is glued to the mesosoma so that the propodeum is not visible, and the central portion of the scutum is missing. Differences between A. butarsis and the type of A. ramakrishnae include anterior face of mesopleuron with- out impunctate area ventrally, lateral face of mesopleuron more coarsely and less dense- ly punctate dorsally, posterior margin of scutellum more narrowly rounded with shallow but distinct emargination medially, T3 punctation more coarse, punctures sparse basomedially, and apical carina of T6 shallowly incurved in dorsal view, in posterior view bent upward medially. The VOLUME 103, NUMBER 2 271 Figs. 1-5. 1, 3, 5, Anthidiellum butarsis. 2, 4, A. krombeini. 1, 2, Male Genitalia, dorsal view. 3, Female hind tibia and tarsus. 4, 5, Male SS. 272 markings on the face also differ. In A. bu- tarsis there is an oblique mark along the inside of the antennal socket. Markings are variable in some anthidiines, but the other characters listed here are usually diagnostic. For this reason, and in the absence of qual- ity comparative material of A. ramakrish- nae, this species is here recognized as a new species. ; Etymology.—From the Latin bu, large or huge, for the greatly enlarged hind basitar- sus. Anthidiellum (Pycnanthidium) krombeini Griswold, new species (Figs. 2, 4) Male.—Length, 4 mm; forewing length, 3.5 mm. Black except as follows: white marks on mandible except apically, ventral stripe on forefemur, hind tibia distally, bas- itarsi, narrow apical bands on T4-5, entirely on T7; lemon yellow on clypeus, paraocular area to level of antennal socket, most of su- bantennal area, elongate spot on gena near summit of eye, spot on pronotal lobe, trans- verse spot on anterior margin of scutum midway between lateral margin and mid- line, medially interrupted posterior mark on scutellum, longitudinal stripe on fore- and midtibiae, posterolateral spot on T1, trans- verse bands on T3-6, slightly interrupted on T3; amber on distal tarsi, terga laterally, sterna. Wings dusky; veins including stigma black. Pubescence white. Punctation of head, dorsum of thorax coarse, contiguous except slightly separated medially on clyp- eus; finer on metanotum, metapleuron, lat- eral face of propodeum; punctation of pron- otum, tegula, anterior face of mesopleuron very fine, sparse on anterior face of meso- pleuron; posterior face of propodeum below narrow pitted basal shelf impunctate. An- terior face of T1 polished without puncta- tion; T1-4 with very narrow impunctate margins less than puncture width; T1-2 densely, nearly contiguously punctate; T3- 7 contiguously punctate; punctures of T1- 2, T6-7 finer than on scutum. Mandible tridentate. Scape three times as PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON long as broad. F1 broader than long. F2-10 slightly longer than broad. Mouthparts scarcely exceeding fossa in repose. Suban- tennal suture strongly arcuate. Pronotal lobe narrowly lamellate, anterior margin straight. Scutal-scutellar suture distinctly foveolate, divided medially. Scutellum rounded pos- teriorly with very shallow median notch. Omaular carina incomplete, not quite reach- ing ventral midline. Hindbasitarsus more than three times as long as broad. Margin of T7 without median angle in ventral view. S2 thickened preapically. Margin of S3 with fringe of long, medially curved hair. Margin of S4 without submedian black spine. S5 with shallow, evenly curved pos- terior emargination, margin with black comb scarcely divided medially (Fig. 4). Genitalia as in Fig. 2. Female.—Length, 4.5 mm; forewing length, 4 mm. Markings as in male except: mandible, subantennal area, basitarsi, hind- tibia dark; clypeal, paraocular marks white; frons with round lemon yellow spot; fore, mid tibial marks reduced or absent; white apical bands of T4-5 absent; mark on T6 quadrate. Mandible tridentate, with long cutting edge and obscure dorsal tooth above two lower teeth. Clypeal margin with four ob- scure denticles. Hind basitarsus more than two times as long as broad. Second hind tarsus as long as broad. Preapical ridge of T6 not notched medially in dorsal view, in posterior view not upturned medially. Sco- pa off-white, hair blunt apically. Types.—Holotype ¢. SRI LANKA, Kandy District, Anniewatte, Kandy, 7°18’N 80°38’E, 31 May — 3 Jun 1997; nest 5; K. V. Krombein, B. B. Norden, A. W. Norden, J. W. Norden. Reared paratypes: 1 6, 1 2 same data; 2 2, same except nest 21. Non- reared paratype, same locality data and col- lectors: 1 2, 1-31 Mar 1997. Holotype in the National Museum of Natural History, Smithsonian Institution, Washington DC. Paratypes at Washington and Utah State University, Logan. Discussion.—Anthidiellum krombeini is VOLUME 103, NUMBER 2 one of the smallest A. (Pycnanthidium). It is similar in size to A. smithii (Ritsema) of Indonesia and Malaysia (Pasteels 1972) dif- fering in the coarse punctation of the frons, scutum and terga and the shiny posterior face of the propodeum. In addition, the male of A. krombeini lacks a small central comb on S5 anterior to the apical comb, F1 is broader than long, and the hind tibia is more slender. In the female F1 is not longer than broad. Etymology.—It is a great pleasure to name this species after Dr. Karl Krombein, who has invested so much of his life in the study of Sri Lankan Aculeata. It 1s addi- tionally appropriate to name this trap-nest- ing species for Dr. Krombein, who has made a major contribution to our knowl- edge of aculeate nesting biology including the first comprehensive study of trap-nest- ing bees and wasps (Krombein 1967). ACKNOWLEDGMENTS I thank Karl Krombein and Beth Norden for the opportunity to describe these new species and Christine Taylor, The Natural History Museum, London, for the loan of 273 types. Susanna Messinger kindly agreed to produce the illustrations. My thanks to Wil- ford Hanson, Frank Parker, and Robbin Thorp for reviews of the manuscript. LITERATURE CITED Baker, D. B. 1995. A review of Asian species of the genus Euaspis Gerstacker. Zoologische Medede- lingen 69: 281-302. Krombein, K. V. 1967. Trap-nesting wasps and bees: Life histories, nests and associates. Smithsonian Press, Washington, D.C. 570 pp. Krombein, K. V. and B. B. Norden. 2001. Notes on trap-nesting Sri Lankan wasps and bees (Hyme- noptera: Vespidae, Pompilidae, Sphecidae, Colle- tidae, Megachilidae). Proceedings of the Ento- mological Society of Washington 103:274—281. Michener, C. D. 2000. The Bees of the World. John Hopkins University Press, Baltimore. 913 pp. Michener, C. D. and T. L. Griswold. 1994. The clas- sification of Old World Anthidiini. The University of Kansas Science Bulletin 55: 299-327. Pasteels, J. J. 1972. Révision des Anthidiinae de la région Indo-Malaise. Bulletin et Annales de la So- ciété Royale Belge d’Entomologie 108: 72—128. . 1980. Révision du genre Euaspis Gerstaecker. Bulletin et Annales de la Société Royale Belge d’Entomologie 116: 73-89. . 1987. Megachilidae récoltes par le Dr. K. V. Krombein et son equipe au Sri Lanka. Bulletin et Annales de la Société Royale Belge d’Entomologie 123: 225-235. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 274-281 NOTES ON TRAP-NESTING SRI LANKAN WASPS AND BEES (HYMENOPTERA: VESPIDAE, POMPILIDAE, SPHECIDAE, COLLETIDAE, MEGACHILIDAE) KARL V. KROMBEIN AND BETH B. NORDEN Department of Entomology, National Museum of Natural History, Smithsonian Insti- tution, Washington, DC 20560-0188, U.S.A. Abstract.—A trap-nesting study was conducted in Kandy, Sri Lanka, during February to June 1997. Wooden pieces ca. 17 cm long with central grooves of either 3 or 5 mm in diameter were placed outside our residence, providing nesting sites for stem-nesting Hymenoptera. A diversity of solitary wasps and bees, including two new species of An- thidiellum, were found nesting in these “artificial stems.’’ Evidence of supersedure was noted, and we give the first report of a probable new subgenus of Hylaeus that uses a mixture of resin and sand for cell construction. Notes on nest architecture are provided for the observed trap-nesters belonging to the families Vespidae, Pompilidae, Sphecidae, Colletidae, and Megachilidae. Key Words: Sri Lanka, trap-nest, stem-nesting, Hymenoptera, Vespidae, Pompilidae, Sphecidae, Colletidae, Megachilidae We spent about four months in Sri Lan- ka, February to June 1997, and were based in the city of Kandy. During the period 9 February through 8 April we rented a large home of relatively recent construction in the Anniewatte suburb near the Botanical Garden at Peradeniya. The house was on a single level, and as is customary in tropical housing, there were louvers above the ex- terior doors and windows to promote air circulation and to exclude vertebrates. A di- verse population of solitary wasps and bees nested in borings in the wooden framing of louvers and the wooden pieces that formed the vanes of the louvers. Cooper (1953) and Krombein (1967) demonstrated the effectiveness of wooden trap-nests in attracting twig-nesting solitary wasps and bees. These traps were made in blocks of straight grained wood of variable sizes, being 75-152 mm long and 20-25 mm wide. A hole 64-152 mm long was drilled in the center of each block to a dis- tance 13 mm from the end of the block. When nests were completed, they were split longitudinally along the midline so that the nest architecture, prey and development of brood could be noted. Medler and Fye (1956) successfully used as traps, holes drilled in sumac (Rhus) stems. We had some wooden traps of different construction that had been made by the late Paul D. Hurd, Jr. His traps were made from wooden pieces glued together (Figs. 1, 2). We set out a total of 25 traps between a pair of vanes in the louvers, and in a dozen bundles of four, two of each diameter rout- ing, in the crotches of tree branches on the property. The routings in the traps were 3 to 5 mm wide and about as deep. The rout- ings were made in pieces of wood 15.2 cm long, 12.7 mm deep and 19 mm wide; the VOLUME 103, NUMBER 2 routing was made down the middle of the 19 mm side. A strip of wood 16.5 cm long, 19 mm wide and 6.4 mm thick was glued above the routing so that it protruded 12.7 mm beyond the end of the piece containing the routing; a square of wood 19.1 xX 19.1 mm and 6.4 mm thick was glued to the oth- er end to seal the basal end of the trap. This kind of trap had not previously been tested in the field. We examined the traps daily when we were in Kandy. Five traps were sealed at the entrance between 16 March and | April. We sealed each trap in a plastic Whirl-pak® in which we made pin pricks for ventila- tion. In preparation for our move to new hous- ing in Kandy we gathered all the traps on 4—5 April. We probed each with a grass stem to determine occupancy, and sealed in individual Whirl-paks eight traps that were partially filled. Nests were completed in most of the eight traps but dates of nest completion are unknown. Findings from this study are detailed below. Family Vespidae Subancistrocerus sichelii (Saussure) Nest #15 was only partially filled on 5 April when we removed it from one of the louvers. We kept it in a sealed Whirl-pak until we opened the trap on 5 May. The routing was 3 mm wide and 2.5 mm deep. The basal 10.2 cm of the routing was empty and capped by a mud partition 2 mm thick. The single cell was 17 mm long including a mud cap | mm thick. It contained a silken cocoon spun against the cell walls and a newly enclosed, fully colored, 9.4 mm long male of S. sichelii. Apparently the vespid foundress had been superseded by an anthidiine bee, be- cause there was a pollen mass 3 mm long and traces of resin between the capped cell of S. sichelii and the nest entrance. Hylodynerus wickwari (Meade-Waldo) We found a female of H. wickwari, 7.5 mm long, in the apical part of a routing in POTS) nest #13 that also contained a single cell of Aylaeus sedens Snelling (see below). There was no evidence of nest construction by the vespid, but she had spent the previous night in the trap. Rhynchium brunneum (Fabricius) D. K. Wijayasinghe placed some of our traps on trees at her family home in Elle- pola, Palapathwela. She reared a large fe- male of Rhynchium brunneum from a trap placed horizontally on a cashew tree 1.3 m above the ground. It had an enlarged rout- ing 13 mm wide and deep with a rounded bottom. Family Pompilidae Auplopus mutabilis (Smith) We noted that nest #1 had been capped at the entrance by 16 March. We placed the nest in a sealed Whirl-pak. Two pairs of A. mutabilis were crawling rapidly around within the Whirl-pak on 1 April. We noted one male trying to mate with a female. Upon opening the trap we found the rout- ing to be 3 mm wide and deep. The basal end was empty, and the foundress had placed a mud plug 6 mm thick 7.2 cm from the basal end. There was then a series of four cells, each containing silken cocoon remnants; the cells from the innermost were 10, 7, 12, and 9 mm long including the mud seals capping the cells which were 2 mm thick in cell 1, and 1 mm thick in the others. Beyond the provisioned cells was a vestib- ular cell of 24 mm including a mud plug 2 mm thick, and finally a terminal cell of 6 mm including the 0.5 mm thick plug at the routing entrance. Considering the smaller size of males, it seems probable that fe- males developed in cells 1 and 3, males in 2 and 4. We picked up nests #8 and #23 on 5 April; neither was capped at the entrance but both had been plugged some distance from the basal end of the routing. The nests were placed unopened in individual sealed Whirl-paks. Three males of A. mutabilis emerged 276 from nest #23 into the Whirl-pak on 16 April. The routing was 3 mm wide and deep, and there was a mud plug 2 mm thick at the base. There followed a series of three cells containing cocoon fragments. Each cell was 10 mm long, including closing mud plugs 2 mm thick. A fourth cell con- tiguous to cell 3 was 18 mm long with a mud plug 8 mm thick; it contained a cocoon that held a shriveled, post-defecated larva. Contiguous to cell 4 were two empty cells, the innermost was 11 mm long including a mud plug 3 mm thick, the outermost was 10 mm long of which 2 mm was the final mud plug. The apical 9 cm of the routing was empty. The largely undamaged silken cocoon in cell | was 6 mm long, closely woven, creamy, ovoid anteriorly and taper- ing to the narrow posterior 0.5 mm apex containing the meconium. A female of A. mutabilis emerged 26 April from the third nest, #8. The routing was 4 mm wide by 3 mm deep, the basal end was empty and there was a plug of mud 2 mm thick 10 cm from the base. Beyond this plug was the only provisioned cell, 9 mm long, of which 1.5 mm was the mud partition that sealed the cell. Next was a vestibular cell 9 mm long including the 1.5 mm thick mud plug. The rest of the routing was empty. The foundress had done one thing omitted by the makers of nests #1 and #23-she had put a thin layer of mud on the ceiling just above the routing. She was pos- sibly reacting to some glue used to seal the trap. Family Sphecidae Pison punctifrons Shuckard Four nests were occupied by this species. The foundress sealed the entrance to nest #2 with mud on 27 March, and we placed it in a plastic Whirl-pak on that date. A pair of P. punctifrons emerged 27—28 April while we were in the field. We opened the nest on 30 April. The routing was 3 mm wide and deep; the basal 5 mm of the rout- ing was empty and capped by a mud par- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tition 0.5 mm thick. The first cell was 13 mm long, including a thin mud cap 0.5 mm long; it contained an ovoid, delicate, light brown, silken cocoon 9 mm long, appar- ently that of a species of Trypoxylon, and some dry spider fragments beyond the apex of the cocoon. There followed an empty in- tercalary cell 7 mm long, capped by a thin mud partition 0.5 mm thick; we do not know whether this cell was capped by the foundress Trypoxylon, or by a female of P. punctifrons to mark its supersedure of the nest. Cell 3 was 12 mm long including the 0.5 mm mud cap, and contained brittle Pi- son cocoon fragments. Cell 4 was an empty intercalary cell 6 mm long with the usual thin mud plug. Cell 5 was 13 mm long with a mud cap 0.5 mm thick; it contained brittle fragments of a Pison cocoon. Cell 6 was an empty intercalary cell 20 mm long with a thin mud plug. Cell 7 was an empty vestib- ular cell 11.3 cm long with 1.0 mm mud plug at the nest entrance. We picked up nest #9 on 4 April, probed it with a grass stem and found it blocked below the entrance. We placed it in a Whirl- pak® and split it open on 5 May. The rout- ing was 4 mm wide and 5 mm deep and contained a mud plug 1.5 mm thick at the basal end. Cell 1 was 11 mm long including the 0.5 mm thick mud plug; it contained only dead spiders. Cells 2 and 3 were 7 mm long including a 0.5 mm closing plug; each contained a brittle Pison cocoon. Cell 4 was partially stored with dead spiders; some spi- ders dropped out of the open end of the nest and were lost. A female and male P. punc- tifrons emerged from the cocoons on 8 May. Nest #14 was blocked 7.2 cm inside the trap entrance when we picked it up on 4—5 April. Four P. punctifrons, three females and one male, emerged from the nest the morning of 3 May. There was a mud plug 1 mm thick at the basal end of the routing. The positions of the cell partitions had been obliterated by movements of the occupants before their emergence. Nest #19 was only partially filled when VOLUME 103, NUMBER 2 we picked it up and placed it in a Whirl- pak on 5 April. We opened the nest 5 May and found a mud plug | mm thick 5.5 cm from the basal end. There were two com- pleted cells, each with a Pison cocoon, and a third incomplete cell with a few dead spi- ders. Cell 1 was 15 mm long including the 1 mm plug, and the cocoon was 9 mm long. Cell 2 was 8 mm long with a 1 mm cap, and the cocoon was 6 mm long. A female wasp emerged 10 May, and a second wasp escaped from the Whirl-pak. The dead, sometimes damaged spiders from cells 1 and 4 of nest #9 were identified by W. P. Wijesinghe as follows: Theridiidae ?—1 juvenile, gen. & sp. indet. Araneidae—1 juvenile of a species of Cyr- tophora Simon ? 1 juvenile of a species of Neo- scona Simon 2 adult and 1 subadult males, gen. & sp. indet. 7 juveniles and juvenile females, possibly conspecific with spe- cies above 1 fragmented juvenile, gen. & sp. indet. 2 juveniles, gen. & sp. indet. Oxyopidae—1 juvenile, gen. & sp. indet. Salticidae—1 juvenile of a species of Brettus Thorell 2 juveniles of species 1 of Rhene Thorell 2 juveniles of species 2 of Rhene Thorell 2 juveniles, possibly different taxa Family indet.—1 juvenile, gen. & sp. indet. Wijesinghe commented (in litt.) that the families represented suggest that the nest foundress ‘‘... had hunted for prey among vegetation.” Trypoxylon buddha Cameron This slender wasp, 8-10 mm long, was reared from two nests. Nest #3 was com- pleted on 27 March and placed in a sealed Whirl-pak. We opened the trap on 5 May to ascertain nest structure and development. ZTE ; , sy Bh eit EE ger are mt * Se ee sie Figs. 1-2. Hurd trap nest. 1, Component parts: end piece 19 mm square and 6 mm thick to seal inner end of trap; routing 3 mm wide in bottom piece 15.2 cm long; top piece 16.5 cm long to seal top of routing. 2, Two assembled traps oblique from beneath with rout- ings 5 mm wide (left) and 3 mm wide (right). The routing was 4 mm wide and 3 mm deep. The basal end was empty for 5.5 cm and sealed by a mud partition 0.5 mm thick. Cell 1 was 15 mm long including the 0.5 mm thick cap; it contained a light brown, silken, ovoid cocoon 9 mm long attached to the base of the cell. Next was an empty intercalary cell 15 mm long including a mud cap | mm thick. The second brood cell was 20 mm long including the 1 mm thick closing cap; it contained a cocoon of similar size, shape and color as that in cell 1. The remainder of the routing contained a vestib- ular cell 4.8 cm long including the closing mud cap 0.5 mm thick at the routing en- trance. Nest #4, in a routing 3 mm wide, was completely sealed on 30 March and opened on 5 May. The basal end was empty for 15.5 cm with a 0.5 mm thick seal capping it. Two partial mud seals were 18 and 22 mm beyond this inner mud seal, then a complete 1 mm seal and an empty cell 6 278 mm long with a 1 mm thick seal. The next cell was 12 mm long including the 1 mm cap; it contained a light brown, ovoid silken cocoon 9 mm long. Beyond that cell was a vestibular cell 3.4 cm long including the 1 mm thick mud cap at the routing entrance. Another 7rypoxylon nest, nest #2, has al- ready been discussed under Pison punctif- rons. The nest had been founded by a spe- cies of Trypoxylon which had capped the first cell and the nest was then superseded by P. punctifrons. The first cell contained a light brown, silken Trypoxylon cocoon, 9 mm long. It is probable that this nest was made by T. buddha. The cocoon is the same size and color. Furthermore, 7. buddha was the more common species that we collected around our home; our total catch of Trypoxylon by net around the house was nine females of the slender, elongate 7. buddha and one fe- male of the somewhat stockier, shorter T. errans Saussure. Colletidae Hylaeus sedens Snelling (Figs. 3, 5) This nest, #13 was placed in the louver above one of the doors. We shone a flash- light into the trap on 4 April and saw the head of a wasp near the entrance. We gath- ered the trap on 5 April; probing with a grass stem indicated that the bore hole was occupied to within 2 cm of the entrance. We placed the nest in a Whirl-pak, and later that day a brightly marked female vespid wasp, 7.5 mm long, Hylodynerus wickwari (Meade-Waldo), emerged into the Whirl- pak. The nest was kept in the Whirl-pak and a female Hylaeus sedens, 6.5 mm long, emerged on 4 May. The unicellular nest was in a 5 mm rout- ing. There was a 2 mm thick plug of resin and sand grains 7 cm from the basal end. The cell was 15 mm long including a 1.5 mm thick closing plug of resin and sand grains with some tiny pebbles. The base, apex and walls of the single cell were coat- ed with a delicate transparent membrane. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON This was not a silken cocoon spun by the mature larva, but a secretion, probably silk, deposited by the foundress before she re- gurgitated the liquid nectar-pollen mixture that constituted the larval food (Krombein 1967): A vestibular cell 4.5 cm long was par- tially closed by a 2.5 mm thick mass of resin and sand on the bottom of the routing. The partial completion of the vestibular cell suggests the possibility that the female Hy- lodynerus wickwari may have superseded the Hylaeus sedens. Discussion.—The type series of H. se- dens came from two localities in southern India, but Snelling (1980) included as this taxon one female from Peak View Motel, Kandy, which differed from the type series in having the mandibles blackish brown rather than ferruginous. The single female that we reared from Kandy also has dark rather than reddish mandibles. The closest relative appears to be H. eur- ygnathus Snelling (1980) also from two lo- calities in southern India and also known only from females. Males are unknown, but Snelling (1980) described three species from India with en- larged mandibles, two of which may be the opposite sex of the two female species men- tioned above. The species has a broader, flatter man- dible than is typical of Hylaeus, a distinc- tion that it shares with H. eurygnathus. In comparison with the North American H. modestus Say, a frontal view of the head shows the relatively large mandibles of H. sedens clearly exposed (Fig. 3) while the lower edge of left mandible of H. modestus is barely visible (Fig. 4). When the head of H. sedens (Fig. 5) is tipped backward, the thin, broadened mandible with small, weak apical tooth, oblique margin above the api- cal tooth, and close, fine striae on lower half of H. sedens (Fig. 5) are a contrast with the narrow, bidentate mandible lacking a cutting edge and striae of H. modestus (Fig. 6). The labrum is also markedly different in the two species. Snelling (personal com- VOLUME 103, NUMBER 2 Figs. 3-6. munication) suggested that the broadened blade-like mandibles in H. sedens may be used in gathering resin for nest construc- tion. Such nesting behavior certainly is an additional strong argument for assigning it to a new subgenus. This is the first report of a species of Hy- laeus using a mixture of resin and sand in nest construction. Hylaeus (H.) modestus usually makes a series of cells whose walls and partitions are made of a delicate, trans- parent membrane secreted by the foundress, although occasionally the cells may be sep- 279 Hylaeus species. 3, H. sedens, frontal view of head. 4, H. modestus, frontal view of head. 5, H. sedens, external view of mandibles. 6, H. modestus, external view of mandibles. arated by a thin partition of compacted wood fibers (Krombein 1967). Megachilidae Anthidiellum butarsis Griswold (Bic. 7) We obtained only one nest #18 of this species from a bundle of traps placed in a crotch of a small clove tree. Probing with a grass stem on 5 April showed that this 3 mm wide routing was partially filled, so we placed the trap in a sealed Whirl-pak. The female emerged from the trap the next day. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON eax. sat sal, Fig. 7. We split open the trap on 5 May. The basal 3.4 cm of the routing was empty and capped partially by a narrow, small bit of mud (Fig. 7, m) and then completely by a thin partition of resin (Fig. 7, p); apparently, the bee had superseded a mason wasp or bee. The next 5 cm contained a series of seven cocoons each separated by a thin par- tition of resin, and the cell walls also were coated with resin. Numerous small drops of resin were on the walls of the next 3.7 cm and the outer 3 cm contained no resin. We opened the cocoons in cells 1, 6 and 7 on 30 May and found a viable, curled up, post-defecated larva in each. The rodlike fecal pellets were voided at the outer end of the cell and formed a crown around the apex of the cocoon (Fig. 7, f). The anterior end of the cocoon had a small median nip- ple. Three females were collected from this nest in addition to the foundress. Anthidiellum krombeini Griswold This small anthidiine occupied two traps. The entrance of nest #5 was sealed with resin on 1 April and placed in a sealed Whirl-pak. We split it open on 5 May and found that there was a 6 mm thick plug of resin mixed with debris at the basal end of m Anthidiellum butarsis, cell 1 (m, mud; p, partition of resin; f, fecal pellets) and basal half of cell 2. the routing. The next 11 cm of the routing was empty and sealed by a plug of resin 7 mm thick. Then there were four cells, each containing a cocoon 5 mm long. The cell walls were covered with a thin coat of resin; the partitions between the cells were thin septa of resin except that the outermost cell had a plug of resin 5 mm thick. The re- mainder of the routing consisted of an emp- ty vestibular cell 7 mm long including the 1 mm thick entrance seal. The routing was 3 mm wide and 5 mm deep. There was no emergence by 31 May, but on 3 June two males and a female had emerged and were dead in the Whirl-pak. The second nest #21 was gathered on 5 April and by probing with a grass stem was found to be partially occupied. We split open the trap on 5 May and found a parti- tion of resin 0.5 mm thick 8.8 cm from the empty basal end of the 4 mm wide routing. The first cell was 7 mm long and contained a moribund bee larva that had consumed only half of the mass of pollen and nectar provided. The next two cells, 8 and 7 mm long, contained light tan, shiny cocoons 5 mm long. As in the first nest, the cell walls were thinly coated with resin, and the cells were capped by thin partitions of resin. VOLUME 103, NUMBER 2 There were bits of resin on the walls be- tween cell 3 and the entrance suggesting that we gathered the nest while the foun- dress was still actively nesting. As in the first nest, adult emergence occurred be- tween 31 May and 3 June. There was a dead male, undoubtedly from cell 2, in the Whirl-pak on the latter date. Within the nest was a fragmented, fully eclosed male, the undoubted occupant of cell 3, that had been disarticulated by the occupant of cell 2 in its efforts to leave the nest. Adult emergence through the partitions capping the cells made it impossible to de- termine whether the cocoon had a median nipple and whether the feces formed a crown around the anterior end of the co- coon. ACKNOWLEDGMENTS We thank T. L. Griswold, Bee Biology and Systematics Laboratory, Utah State University, Logan, UT, for describing An- thidiellum butarsis and A. krombeini that we had reared or collected by net in Kandy. We are indebted to John Norden for his help in setting out and tending traps that were placed too high for us to reach easily. Ar- nold Norden assisted with this effort and with some collecting by hand net and is thanked as well. Also, we appreciate the ef- forts of D. Wijayasinghe who placed sev- eral traps at her home in Palapathwela. We are grateful to R. R. Snelling, Natural History Museum of Los Angeles County, CA, for identifying Hylaeus sedens Snell- ing, and to W. P. Wijesinghe, City College of New York for identification of spiders. 281 Identifications of wasps are by KVK as fol- lows: Vespidae by comparison with speci- mens identified by the late J. van der Vecht; Pompilidae by comparison with specimens identified by R. Wahis; and Sphecidae by comparison with specimens identified by A. V. Antropov (Pison) and the late K. Tsuneki (Trypoxylon). We thank Suzanne W. T. Ba- tra and Roy R. Snelling for their very help- ful reviews of a draft of this contribution. The scanning electron micrographs were prepared by BBN. Within the Smithsonian we thank Susann G. Braden, Scanning Electron Microscope Laboratory, for tech- nical assistance, and George L. Venable for skillful preparation of the plates. This work was supported in part by the Smithsonian Krombein Endowment Fund and a Research Opportunity Fund award to KVK, and a Senior Fulbright Fellowship award to BBN. LITERATURE CITED Cooper, K. W. 1953. Biology of eumenine wasps. I. The ecology, predation and competition of Ancis- trocerus antilope (Panzer). Transactions of the American Entomological Society 79: 13-35. Griswold, T. 2001. Two new species of trap-nesting Anthidiini (Hymenoptera: Megachilidae) from Sri Lanka. Proceedings of the Entomological Society of Washington 103:269—273. Krombein, K. V. 1967. Trap-nesting wasps and bees: life histories, nests and associates. Smithsonian Press, Washington. 570 pp. Medler, J. T. and R. E. Fye. 1956. Biology of Ancis- trocerus antilope (Panzer) in trap-nests in Wis- consin. Annals of the Entomological Society of America 49: 97-102. Snelling, R. R. 1980. New bees of the genus Hylaeus from Sri Lanka and India (Hymenoptera: Colleti- dae). Contributions in Science, Natural History Museum of Los Angeles County, No. 328. 18 pp. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 282-290 A REVIEW OF THE SOUTHERN SOUTH AMERICAN GENUS GYRETRIA ENDERLEIN (DIPTERA: SPHAEROCERIDAE: LIMOSININAE) S. A. MARSHALL Department of Environmental Biology, University of Guelph, Guelph, Ontario, Canada N1G 2W1 Abstract.—Gyretria melanogaster (Thomson) and Gyretria biseta (Duda), new com- binations, are redescribed. The genus Gyretria Enderlein, previously treated as a synonym of Chaetopodella, is resurrected for these two species. Gyretria is redescribed and its relationships to other Limosininae discussed. The presence of eversible abdominal sacs, found in males of both Gyretria species, is noted for the first time in the Sphaeroceridae. A lectotype is designated for Leptocera (Chaetopodella) biseta Duda, and Sclerocoelus rectangularis (Malloch) is given as a new combination. Key Words: designation, distribution Several species of New World Limosini- nae are characterized by velvety black prui- nosity, usually with patterns of stripes and silver spots at least around some bristle ba- ses. Some of these species are in well-de- fined genera such as Poecilosomella Duda, Pterogramma Spuler, Opacifrons Duda, and Leptocera Olivier, but others have been artificially grouped within the similarly pat- terned Old World genus Chaetopodella Duda. Of the four New World species pre- viously placed in Chaetopodella, one (rec- tangularis Malloch) belongs in Sclerocoe- lus Marshall (n. comb.), and one (tonsa Duda) has little in common with Chaeto- podella except color. Chaetopodella tonsa (Duda) is one of two velvety black species in an undescribed genus of mostly brown species. The two remaining New World species usually placed in Chaetopodella (melanogaster Thomson and biseta Duda) are similar to Old World Chaetopodella in most non-abdominal characters including pigmentation and chaetotaxy of the female legs and thorax. Despite these similarities, Diptera, Sphaeroceridae, Gyretria restituted, new combinations, lectotype the two New World species differ widely from Old World Chaetopodella in abdomi- nal characters and in several unique sexu- ally dimorphic characters of the head and legs, and warrant treatment as a separate ge- nus. Previous authors have recognized the distinctness of melanogaster and _ biseta, and genera have been proposed with each of these species as type. Since one of the generic names available for this group is preoccupied, the remaining generic name, Gyretria Enderlein, originally coined for the single species binodatipes Enderlein (= melanogaster Thomson), is resurrected and redefined to include Gyretria melanogaster and Gyretria biseta. Gyretria Enderlein Gyretria Enderlein 1938: 652. Type species Gyretria binodatipes Enderlein 1938 (original designation) = Limosina melan- ogaster Thomson 1869. Alima Duda 1938: 5, 95 (preoccupied by Alima Leach 1818 [Crustacea]). Type VOLUME 103, NUMBER 2 species, Leptocera (Chaetopodella) bise- ta Duda 1925 (monotypy). Alma Duda; Richards 1967: 11 (misspell- ing; also preoccupied by Alma Grube 1855 [Oligochaeta]). Diagnosis.—Members of the genus Gyr- etria are among the easiest sphaerocerids to recognise to the generic level. Their velvety black integument combined with milky or Opaque-brownish wing membranes gives them a distinctive habitus, and the presence of ventral bristles on both the mid tibia and tarsomere one of the mid leg distinguish Gyretria from the few other South Ameri- can species with similar colors. Unlike Old World Chaetopodella, Gyretria have legs that are sexually dimorphic in both species, the fore and mid femora bearing stout ven- tral bristles in the male and tarsomeres 1 and 2 of the hind leg flattened and greatly enlarged in the male. The most unusual character shared by both Gyretria species is a pair of eversible sacs that extend anteri- orly into the male abdomen from openings behind the fourth sternite. Generic description.—Body length 2-3 mm, body dark brown to black with a heavy silvery pruinosity, pruinosity of scu- tum broken by three relatively bare longi- tudinal strips. Frons longitudinally ridged, orbits and interfrontal strips raised. Frons with silvery pollinose interfrontal strips, back of head with 2 bright silvery spots. Postocellar and postvertical bristles small, inner and outer occipital bristles large, sub- equal to upper orbital bristle; inner and out- er vertical bristles very large. Prosternum linear, bare. Mid coxa and trochanter each with a very long anterior preapical bristle. Mid tibia of female with a large mid ventral and apical ventral bristle; mid tibia of male without a mid ventral bristle but with a dense row of about 20 long, fine ventral bristles on distal half. Dorsal surface of mid tibia in both sexes with preapical dorsal bristles and 4—5 anterodorsal and 4—6 pos- terodorsal bristles. Tarsomere one of mid leg with a mid ventral bristle. Hind leg of 283 male with first two tarsomeres greatly en- larged, tarsomeres 2-5 strongly flattened, hind leg of female with tarsomere 2 un- modified and tarsomere 1 short but not strongly flattened or enlarged. Scutum with 3 pairs of postsutural dorsocentral bristles (sometimes with a small presutural pair), three pairs of intra-alar bristles, 2 supra-alar bristles and a very long postalar bristle reaching beyond apex of scutellum. Scutel- lum with 4 long bristles, each basally sur- rounded by a small patch of silver pruinos- ity. Acrostichal setulae long, in 6-8 rows between anterior dorsocentral bristles. Wing long, at least slightly opaque, alula broad; halter white. Male abdomen conspicuously constricted behind segment three, with narrower pos- terior part of abdomen usually deflexed; ter- gite 5 (normally telescoped under tergite 4) short, almost transverse dorsally. Mem- brane between sternite 4 and sternite 5 with two eversible sacs behind sternite 4, sacs often everted in critical-point dried speci- mens (Fig. 1). Sternite 5 small, divided into longidudinal anterior and transverse poste- rior parts. Synsternite 6+7 free from ster- nite 8, broad ventrally, closely associated with posterior part of sternite 5, expanded on right side and continuing across dorsum to form a complete loop; ring sclerite large. Epandrium with stout posterolateral bristles and a group of long ventral bristles; ventral angles of epandrium widely separated and extending ventral to cerci; cercus pale, flat; subepandrial sclerite with heavily scleroti- sed, dorsally projecting lateral processes. Surstylus broad, pale, setulose. Hypandrium heavily sclerotised, with a short, dark an- terior apodeme; arms fused to anterolateral corners of epandrium. Distiphallus very broad. Female abdomen short, not telescop- ing; tergite 8 divided into two lateral scler- ites, tergite 10 setulose, with 2—3 small dor- sal bristles; cerci very short, setulose, with some longer distal bristles. Spermathecae spherical, single unpaired spermatheca larg- er than the 2 paired ones, sclerotised parts of ducts short. SQ Figs 1: Gyretria biseta, Male. Relationships and similar genera.—Gyr- etria species have a number of superficial similarities to Chaetopodella, including the ventral bristle on tarsomere one of the mid leg, three dorsocentral bristles, opaque wing membrane, non-telescoping female termin- alia, and similar chaetotaxy on the female mid tibia. None of the many unusual male genital characters and sexually dimorphic leg characters are shared between Gyretria and Chaetopodella. Similar color characters are seen in two species of the northern Neo- tropical tonsa group, but these species lack a ventral bristle on tarsomere one, lack a mid ventral mid tibial bristle in both sexes, and appear to belong to a larger clade with- in which the color characters are derived independently of either Gyretria or the Old World Chaetopodella. Most of the abdom- inal characters of Gyretria are so derived they are difficult to compare with homolo- gous characters in other clades. On the basis of non-abdominal characters Gyretria 1s PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON most similar to the Old World genus Chae- topodella, and it is possible that some of the shared color and chaetotaxy characters do reflect close relationships between Old World Chaetopodella and the southern South American clade discussed here. Nonetheless, the southern South American clade is geographically isolated, there are no abdominal synapomorphies linking the New World and Old World species, and there are many distinct defining characters for the southern South American clade. These facts argue strongly for the treatment of the southern South American clade as a separate genus Gyretria rather than treating it as part of the Old World genus Chaeto- podella. KEY TO SPECIES OF GYRETRIA — Head with a single orbital bristle. Wing mem- brane brownish. Hind tarsomeres brown a ne Sop rsat like Bi lye Grind, avd SH G. biseta (Duda) — Head with two orbital bristles. Wing membrane white. Tarsomeres 3—5 of hind leg white, con- VOLUME 103, NUMBER 2 trasting with brown tarsomere 1. ........ G. melanogaster (Thomson) Gyretria biseta (Duda 1925), new combination (Figs. 1-11) Leptocera (Chaetopodella) biseta Duda 1925: 149. Type locality: Bolivia, Mapiri, Lorenzopata. Chaetopodella biseta: Duda 1929: 34 [ge- neric combination]. Alima biseta: Duda 1938: 95 [generic com- bination]. Description.—Body length ca. 2.5 mm. Pruinosity of scutum broken by three rela- tively bare longitudinal strips, middle strip wider than lateral ones, scutum bare be- tween posterior two pairs of dorsocentral bristles. Interfrontal area 1.5 as broad as high, flanked by three equal interfrontal bristles. Orbital bristles stout, in a single long pair. Male face silvery yellow, almost white; female face light brown; antenna and gena light brown, arista short-haired,as long as head width; eye bare, twice as high as genal height; gena with 3—4 rows of small bristles. Palp with two long preapical ven- tral bristles, stipes heavily sclerotized, with a conspicuous row of about 8 long, stout bristles in male (Fig. 1); stipes with shorter, thin bristles in female. Fore leg brown in female; male tibia bi- colored with basal half pale brown to yel- low, distal half black. Fore femur of male with 2 long, stout ventral bristles; fore tibia of male swollen distally. Mid leg brown, femur paler than tibia. Mid femur of male with a short and a long ventral bristle in basal third; mid femur of female without ventral bristles. Dorsal surface of mid tibia in both sexes with preapical dorsal bristles and 5 anterodorsal and 4 posterodorsal bris- tles, distal posterior surface of male mid tibia with a dense patch of about 10 stout bristles (Fig.1); dorsal preapical bristle of male mid tibia stout and distictly bent at middle. Tarsomere one of male curved, ex- panded distally and with row of stout an- terior bristles on distal two-thirds. Hind legs 285 mostly pale brown, indistinct rings of dark- er brown present distally on femur and both proximally and distally on tibia; tarsomeres dark brown. Katepisternum with 2 long bristles, posterior dorsal bristle reaching wing base, anterodorsal bristle one third as long. Wing slightly opaque, brownish; viens pale brown; second costal sector 1.6 as long as third costal sector, costa extend- ing slightly beyond end of R4+5; alula broad. Halter white. Male abdomen with first two tergites (syntergite 1+2 and tergite 3) twice as long and almost twice as broad as tergite 4, syn- tergite 1+2 velvety black, tergites 3 and 4 velvety black except for large, quadrate, pale, posteromedial sections, tergite 5 (nor- mally telescoped under tergite 4) reduced to a U-shaped anterior margin, narrow antero- lateral lobes, and a long, narrow, postero- medial apodeme (Fig. 10). Sternite 4 nar- row, subquadrate, not modified, membrane between sternite 4 and sternite 5 with two very large membranous everted lobes or in- vaginations covered with long, pale setae (everted lobes in critical-point dried speci- mens, invaginations in air-dried specimens) (Figs. 1, 10). Sternite 5 with a dark, sub- quadrate, anteriorly emarginate anterior piece and a transverse posterior piece with posterolateral corners extended into pale, multifurcate lobes (Fig. 9). Epandrium with conspicuously long ventral and postero- ventral bristles, posterodorsal bristle shorter than largest ventral bristles (Figs. 6-7). Sur- stylus broad, pale, setulose, with a narrow, distally expanded lateral lobe near mid ven- tral margin (Fig. 6). Postgonite broad, dis- tally bifurcate, basiphallus short, with a ta- pered epiphallus (Fig. 8). Female abdomen (Figs. 2—3, 5) short, sternite 8 small, simple; no visible internal genital sclerotisation. Sternite 10 large, se- tulose, medially pale. Spermathecae spher- ical; surface smooth (Fig. 4) Type material examined.—Lectotype (male, here designated to fix the current in- terpretation of this name and to ensure sta- bility and uniformity in its future interpre- 286 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 2-10. Gyretria biseta. 2-5, Female. 2, Terminalia, left lateral. 3, Terminalia, ventral. 4, Spermathecae. 5, Terminalia, dorsal. 6-11, Male. 6, Terminalia, left lateral. 7, Terminalia, posterior. 8, Aedeagus and associated structures. 9, Terminalia and sternite 5, ventral. 10, Posterior part of abdomen, left lateral. Abbreviations: T5, tergite 5; T7, tergite 7; T10, tergite 10; PG, postgonite; S6+7, synsternite 6+7; S5, sternite 5; SUR, surstylus; EPAN, epandrium. VOLUME 103, NUMBER 2 tation). BOLIVIA. Mapiri, Lorenzopata “7.v.03°> (SMTD). (Other specimens in type series in very bad condition). Other material examined (approx. 250 specimens).—ARGENTINA. Jujuy, Calile- gua National Park, 18—28.xii.1987, S. Peck. Salta, Campo Quijano, 30 km E Salta, 20.11.1992, dung traps and swept over dung, S.A. Marshall; Rosario de Lerma, 29.11.1992, swept from ditch, S.A. Marshall. Misiones, Iguazu, 4—10.1x.1927, RC & EM Shannon (USNM). Catamarca, El Pintado, S. La Vifia, 650 m, 27—29.1x.1968, L. Pefia (CNC). BOLIVIA. Santa Cruz, Potrerillo de Cruerda, 4—7.1.1998, H. Howden. BRA- ZIL. Rio de Janeiro, Nova Friburgo, sweep dirty area near water falls, 26.1.1990, S.A. Marshall; Minas Gerais, 1km E Lavras, dung traps in ditch, 18—20.11.1990, S. A. Marshall; 20 km SW Barbacena, dung traps, 25.11.1990, S.A. Marshall; Prado, sweep along river, 21.11.1990, S.A. Mar- shall; Tiradentes, malaise trap, 15— 17.11.1990, S.A. Marshall. Parana, Curitiba, University campus, sweep in forest, 10.11.1990, S.A. Marshall; Morretes, 9.11.1990, S.A. Marshall; Curitiba, dung traps 30 km S BR 277, 6—9.11.1990, S.A. Marshall; Curitiba, intercept trap in woods behind Natural History Museum; Londrina, Parque Arthur Thomas, sweep, 1.11.1990, S.A. Marshall. PARAGUAY. Parana, SW Saltos delsGuaira, xi1.8.1971. L. Pefia (AMNH). Distribution.—Argentina, Bolivia, Bra- zil, Paraguay, Comments.—This highly autapomorphic species is one of the most easily recognised species of Sphaeroceridae. The loss of one orbital bristle, reflected in the specific name, is unusual, as are the sexual dimor- phisms of face color and leg chaetotaxy. The very large, apparently inflatable ab- dominal pouches opening behind sternite 4 are of special interest. Many Diptera (in- cluding some Ramphomyia females [Em- pidae] and many Tephritidae males) puff out their abdominal pleura to either increase their body profile or to expose pheromone 287 glands. Eversible pouches are found in a few acalyptrates other than Gyretria, in- cluding the chloropid Thaumatomyia notata Meigen, some Otitidae (including Xantho- crona Wulp) and the tephritid Anastrepha ludens (Loew). The latter species is known to use the everted pouches to aid in dissem- ination of sexual attractant pheromones, and Gyretria species probably use their pouches in the same fashion. Gyretria melanogaster (Thomson 1869), new combination (Figs. 11-18) Limosina melanogaster Thomson 1869: 603. Type locality: Argentina, Buenos Aires. Leptocera (Chaetopodella) melanogaster: Richards 1963: 238 [generic combina- tion]. Chaetopodella melanogaster: Rohaéek 1983: 114 [generic combination]; Mar- shall 1997: 168. Leptocera (Chaetopodella) pulchripes Duda 1925: 151. Type locality: Paraguay; Richards 1963: 238 [synonymy]. Caetopodella pulchripes: Duda 1929: 34 [lapsus, generic combination]. Leptocera (Chaetopodella) pulchripes var. griseithorax Richards 1931: 79; Richards 1967: 11 [synonymy]. Gyretria binodatipes Enderlein 1938: 655; Type locality: Chile, Juan Fernandez Is.; Richards 1967: 11 [synonymy]. Description.—Body length ca. 2.0 mm, body dark brown to black with a heavy sil- very pruinosity, pruinosity of scutum bro- ken by three relatively bare longitudinal strips, middle strip wider than lateral ones. Interfrontal area as broad as high, flanked by three equal interfrontal bristles. Orbital bristles stout, in two pairs, lower pair half as long as upper pair. Face, outer surface of antenna and gena light brown, inner face of first flagellomere rufous at least basally; arista short-haired, as long as head width; eye bare, twice as high as genal height; gena with 3—4 rows of small bristles. Palp 288 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 11-18. Gyretria melanogaster. 11-13, Female. 11, Terminalia, left lateral. 12, Terminalia, ventral.. 13, Spermathecae. 14—18, Male. 14, Terminalia, left lateral. 15, Terminalia, posterior. 16, Aedeagus and associated structures. 17, Posterior part of abdomen, lateral. 18, Posterior part of abdomen, ventral. Abbreviations: T5, tergite 5; PG, postgonite; S6+7, synsternite 6+7; S5, sternite 5; S8, sternite 8; S10, sternite 10; SUR, surstylus; CER, cercus. VOLUME 103, NUMBER 2 with two long preapical ventral bristles, stipes with 3 long, thin bristles. Foreleg brown, male fore femur with a long, stout ventral bristle, male fore tibia with a pro- nounced swelling in distal third. Mid coxa, femur, and tibia except apex brown, apex of tibia and tarsomeres yellow to pale brown. Mid femur of male with two long ventral bristles in basal third; mid femur of female without ventral bristles. Dorsal surface of mid tibia in both sexes with four pairs of anterodorsal-posterodorsal bristles, the third anterodorsal bristle displaced almost to an- terior surface. Hind leg with tip of tarso- mere 2 and all of tarsomeres 3—5 white, sharply contrasting with brown tarsomere 1; hind leg of female similarly pigmented, but tarsomere 2 unmodified and tarsomere 1 small. Katepisternum with 2 long bristles, posterior dorsal bristle reaching wing base, anterodorsal bristle half as long. Wing milky-white, viens except costa very pale, costa pale brown; second costal sector 1.2— 1.3X as long as third costal sector, costa not extending beyond end of R4+5. Male abdomen with first two tergites (syntergite 1+2 and tergite 3) twice as long and almost twice as broad as tergite 4, syn- tergite 1+2 velvety black, tergite 3 velvety black except for large, quadrate, pale pos- teromedial section, tergite 4 brown, tergite 5 (normally telescoped under tergite 4) pale and desclerotised except for transverse and weakly U-shaped anterior margin and dark anterolateral lobes (Figs. 17—18). Sternite 4 subquadrate, not modified, membrane be- tween sternite 4 and sternite 5 with two small, membranous swellings or invagina- tions covered with long, pale setae (swell- ings in critical-point dried specimens, in- Vaginations in air-dried specimens). Sternite 5 very narrow, slightly expanded at anterior and posterior end, posterolateral corners ex- tended into pale, multifurcate lobes (Fig. 18). Epandrium with long ventral and pos- teroventral bristles and a very long poster- odorsal bristle (Fig. 15). Surstylus broad, pale, setulose, with a narrow, distally ex- panded lateral lobe near mid ventral margin 289 (Fig. 14). Postgonite broad, apex simple; basiphallus short (Fig. 16). Female abdomen short, sternite 8 small, simple; a small convex sclerite near genital opening and internal genital sclerotisation present (Figs. 11-12). Sternite 10 large, se- tulose, medially desclerotised. Spermathe- cae spherical, with surface striate (Fig. 13). Type material examined.—Holotype (male, headless) and 5 paratypes (3 females, 2 males) of Chaetopodella pulchripes var. griseithorax Richards. ARGENTINA. Buenos Aires, “21.x.26’> (BMNH). Holo- type (female) of Limosina melanogaster Thompson. ARGENTINA. Buenos Aires (labelled with three printed labels: “‘Buen/ Ayr.,” ““Kinb.”’ and ‘“‘Typus”’ and an old red loan tag “*245/60.’’) (NHRS); examined by Dr. Thomas Pape (NHRS). The type of G. binodatipes Enderlein was not exam- ined, but Enderlein’s figures leave no doubt about the identity of this species, which does not resemble anything else found on Juan Fernandez Islands. Other material examined (approx. 300 specimens).—ARGENTINA. Buenos Ai- res, San Isidro, 25.viii.1929, Shannon & Shannon (USNM); Salta, Palo Pintado, N. San Carlos, 1,900 m, 6.x.1968, L. Pefia (CNC); Catamarca, 18 km N La Merced, 1,000 m, 26.1x.1968, L. Pea (CNC). BRA- ZIL. Sao Paulo, Sao José de Barreiro, 1,650 m, 1.1969, malaise, M. Alvarenga; Rio de Janeiro, Teresopolis, Pedra do Sino, 2,000 m, human dung, 14.11.1990, S. Marshall; Santa Catarina, Nova Teuténia, 300—500 m, x1.1966, E Plaumann (CNC); Minas Gerais, Tiradentes, Serra de Tiradentes, human dung, 16.11.1990, S. Marshall. CHILE. Juan Fernandez Islands, Robinson Crusoe Island, San Juan Bautista, English Bay, Plazoleta, 1.1992, 1.1993, dung, sweep, and pans, S.A. Marshall; Santiago, Penalolen, x.1953, L. Pena (CNC); Hacienda Illapel, Coquimbo, 1—6.x1.1954, 500 m, L. Pena (CNC); Rio Clarillo, 10 km S Santiago, 3.x1.1989, S.A. Marshall. Distribution.—Argentina, Brazil, Para- guay, Uruguay, Chile. 290 Comments.—This easily recognized spe- cies is one of the most common insects on the Juan Fernandez Islands, where it is abundant on equine feces. Gyretria melan- ogaster is also locally abundant in mainland Chile, but where it is sympatric with G. bis- eta in Brazil and Argentina the latter spe- cies seems to be much more abundant. ACKNOWLEDGMENTS Thanks to the following museums who provided specimens used in this study: American Museum of Natural History, New York (AMNH); Canadian National Collec- tion of Insects, Ottawa (CNC); National Museum of Natural History, Smithsonian Institution, Washington, DC (USNM); Cal- ifornia Academy of Sciences, San Francis- co (CAS); Staatliches Museum fiir Tierkun- de, Dresden (SMTD). Dr. Thomas Pape, Naturhistoriska Riksmuseet, Stockhom, Sweden (NHRS) kindly examined the type of G. melanogaster. Thanks also to Rebec- ca Langstaff for her help with the illustra- tions, and to Matthias Buck, Alan Norrbom and Jindrich Rohacéek for their helpful com- ments on the manuscript. Unless otherwise stated, specimens are in the University of Guelph Collection. Funding for this work was provided by NSERC. LITERATURE CITED Duda, O. 1925. Die aufereuropéischen Arten der Gat- tung Leptocera Olivier-Limosina Macquart (Dip- teren) mit Berticksichtigung der europadischen Ar- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ten. Archiv fiir Naturgeschichte, Berlin, Abteilung A, 90(11)(1924): 5-215. . 1929. Die Ausbeute der deutschen Chako-Ex- pedition 1925/1926 (Diptera). VI. Sepsidae, VII. Piophilidae, VIII. Cypselidae, IX. Drosophilidae und X. Chloropidae. Konowia 8(1): 33-50. . 1938. Sphaeroceridae (Cypselidae). Jn Lind- ner, E., ed., Die Fliegen der palaearktischen Re- gion, Vol. 6, 182 pp., E. Schweizerbart’sche Ver- lagsbuchhandlung, Stuttgart. Enderlein, G. 1938. Die Dipterenfauna der Juan Fer- nandez Inseln und der Oster Insel, pp. 643-680. In Skottsberg, C., ed., The Natural History of Juan Fernandez and Easter Island, 3 (Zool.), Uppsala. Marshall, S. A. 1997. Sphaeroceridae of the Juan Fer- nandez Islands, Chile. Studia Dipterologica 4: 165-171. Richards, O. W. 1931. Sphaeroceridae (Borboridae). Jn Diptera of Patagonia and south Chile, based main- ly on material in the British Museum (Natural His- tory) 6(2): 62-84. . 1963. Sphaerocerid flies from South and Cen- tral America in the collection of the California Academy of Sciences (Diptera). Pan-Pacific En- tomologist 39: 231—246. . 1967. 72. Family Sphaeroceridae (Borbori- dae). In Vanzolini. E. P. and N. Papavero, eds., A Catalogue of the Diptera of the Americas South of the United States, Vol. 72, 28 pp., Departa- mento de Zoologia, Secretaria da Agricultura, Sao Paulo. Rohaéek, J. 1983. A monograph and re-classification of the previous genus Limosina Macquart (Dip- tera, Sphaeroceridae) of Europe. Part II. Beitrage zur Entomologie, Berlin 33: 3-195. Thomson, C. G. 1869. 6. Diptera. Species nova des- cripsit. In Kongliga svenska fregatten Eugenies Resa omkring jorden under befal af C. A. Virgin, aren 1851—1853. pt. 2 (Zoologie), Sec.1 (Insecter), 617 pp. (p. 443-614), Kongliga Svenska Veten- skaps-Akademien, P. A. Norstedt & S6ner, Stock- holm. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 291-307 REVISION OF NORTH AMERICAN ALEIODES WESMAEL (PART 6): THE GASTERATOR (JURINE) AND UNIPUNCTATOR (THUNBERG) SPECIES-GROUPS (HYMENOPTERA: BRACONIDAE: ROGADINAE) PAUL M. MARSH AND Scott R. SHAW (PMM) Cooperating Scientist, Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, % National Museum of Natural His- tory, Washington, D.C. 20560-0168, U.S.A.; current address: PO. Box 384, North New- ton, KS 67117, U.S.A. (e-mail: swampy @southwind.net); (SRS) University of Wyoming Insect Museum, Department of Renewable Resources, University of Wyoming, Laramie, WY 82071-3354, U.S.A. (e-mail: braconid@uwo.edu) Abstract.—The Aleiodes gasterator (Jurine) species-group is defined to include several Palaearctic species and the following species from North America: atricornis (Cresson), n. comb. (= ferrugineus Enderlein, n. syn.); brachyphlebus, n. sp.; bucculentus, n. sp.; burrus Cresson (= fulvus Cresson, n. syn., nigricoxis Viereck, n. syn., cockerelli Viereck, n. syn., fusicaudus, n. nud., waldeni, n. syn.); carinatus (Ashmead) n. comb. (= pubes- cens Ashmead, n. syn., nasonii Ashmead, n. nud.); medicinebowensis, n. sp.; smithi, n. sp.; townesorum, n. sp.; vierecki, n. sp. The unipunctator (Thunberg) species-group also includes several Palaearctic species and the following North American species: harrimani (Ashmead), n. comb.; pseudoterminalis, n. sp.; terminalis Cresson. Both groups have small ocelli and have the marginal cell of the hind wing gradually widening to the wing apex. Species in the gasterator group are usually unicolored yellow or orange and have the malar space longer than the basal width of the mandible; species in the unipunctator group are bicolored black and red and have the malar space about equal to the basal width of the mandible. Key Words: Braconidae, Aleiodes, parasitoids, revision The rogadine braconid genus Aleiodes Wesmael is worldwide in distribution, but is particularly species-rich in the Holarctic Region. Aleiodes is well diversified in North America, with at least 90 species in the United States and Canada (S. Shaw et al. 1997). This study is the sixth in a series of planned papers on Aleiodes species- groups, intended to provide a complete re- vision of the genus for North America (see S. Shaw et al. 1997, 1998a, b; Marsh and S. Shaw 1998, 1999). The species treated in this paper, the gas- terator (Jurine) and unipunctator (Thun- berg) species-groups, include some of the largest, commonest and most distinctive species in eastern North America. For ex- ample, A. terminalis Cresson is the most abundant species in collections, and one of the few that can be reliably identified, based on color patterns, even without magnifica- tion. All species are distinguished by small ocelli and the marginal cell of the hind wing gradually widening to the apex of the wing, narrowest at base. Species in the gas- terator group are distinguished by the malar space being longer than the basal width of the mandible and by their generally brown 292 or orange unicolored body; the unipuncta- tor group has a shorter malar space and the body is bicolored black and red. In their phylogenetic analysis of Aleiodes, Fortier and Shaw (1999) included three species- subgroups, gasterator, atricornis and uni- punctator, within the gasterator species- group. We consider the unipunctator group as a distinct species-group as presented in the species-group key of S. Shaw et al. (1997). Because our main intent is to pro- vide a revision of the North American spe- cies, species treatments are limited to the Nearctic fauna. Aleiodes species are koinobiont endopar- asitoids of lepidopterous larvae, especially macrolepidoptera of the superfamilies Noc- tuoidea and Geometroidea, and to a lesser extent, Arctioidea, Sphingoidea and Papi- lionoidea (S. Shaw et al. 1997). Very little is known about the biology of the species of the groups included in this paper, but the few records indicate parasitism of noctuid larvae. The method of parasitism, unique to the tribe Rogadini, is noteworthy: the Al- eiodes larva completes its feeding and pu- pates within the shrunken and mummified remains of the host caterpillar. In all known cases, the form of the mummy caused by a particular Aleiodes species is characteristic for that host and parasitoid, so mummified remains are of considerable diagnostic val- ue and should be retained with the parasit- oid when reared. For a more complete dis- cussion of Aleiodes biology, readers may refer to M. Shaw (1983, 1994), M. Shaw and Huddleston (1991), S. Shaw (1995) and S. Shaw et al. (1997). METHODS Species covered in this paper can be identified as members of the subfamily Ro- gadinae using the keys of S. Shaw (1995), M. Shaw and Huddleston (1991) or Whar- ton et al. (1997). Our definition of Aleiodes follows that of S. Shaw (1995), S. Shaw et al. (1997) and van Achterberg (1991). Specimens can be determined as Aleiodes using the keys of van Achterberg (1991) or PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON S. Shaw (1997). The species-groups of North American Aleiodes can be keyed us- ing the key provided in S. Shaw et al. (1997). Fortier and Shaw (1999) provided a cladistic analysis of the Aleiodes species- groups. Terminology follows that used for Aleio- des by S. Shaw et al. (1997), S. Shaw (1995) and Marsh (1989). Microsculpture terminology follows that of Harris (1979). Wing vein terminology agrees with the sys- tem adopted by Sharkey and Wharton (1997) and agrees closely with that of Hub- er and Sharkey (1993). A labeled diagram of wing veins was provided by S. Shaw et ale (1997): Acronyms for collections where type ma- terial is deposited are as follows: ABS (Archbold Biological Station, Lake Placid, FL), AEI (American Entomological Insti- tute, Gainesville, FL), ANSP (Academy of Natural Sciences, Philadelphia, PA), BMNH (The Natural History Museum, London, UK), CNC (Canadian National Collection, Ottawa, Ontario, Canada), CSU (Colorado State University, Fort Collins, CO), INHS (illinois Natural History Sur- vey, Champaign, IL), MCZ (Museum of Comparative Zoology, Harvard University, Cambridge, MA), MISS (Mississippi State University, Mississippi State, MS), MSU (Michigan State University, East Lansing, MI), NCDA (North Carolina Department of Agriculture, Raleigh, NC); NCSU (North Carolina State University, Raleigh, NOC), NNML (Nationaal Natuurhistorisch Muse- um, Leiden, The Netherlands), RMSEL (Rocky Mountain Systematic Entomology Laboratory, University of Wyoming, Lara- mie, WY), TAMU (Texas A&M University, College Station, TX), UCD (University of California, Davis, CA), UK (University of Kansas, Lawrence, KS), USNM (National Museum of Natural History, Smithsonian Institution, Washington, DC). ALEIODES GASTERATOR SPECIES-GROUP Included species.—gasterator (Jurine) 1807, n. comb., Europe; grassator (Thun- VOLUME 103, NUMBER 2 berg) 1822, n. comb., Europe; burrus Cres- son 1869, North America; atricornis (Cres- son) 1872, n. comb., North America; car- inatus (Ashmead) 1889, n. comb., North America; rufipes (Thomson) 1891, n. comb., Europe; ecuadoriensis (Brues) 1926, n. comb., Ecuador; pallidistigmus (Telenga) 1941, n. comb., Europe; confor- mis (Muesebeck) 1960, n. comb., Uruguay; fortis (Muesebeck) 1960, n. comb., Uru- guay; brachyphlebus, n. sp.; bucculentus, n. sp.; medicinebowensis, n. sp.; smithi, n. sp.; muesebecki, n. sp.; townesorum, Nn. sp. Diagnostic characters.—Oral opening oval, diameter equal to or slightly greater than malar space; malar space at least equal to basal width of mandible, usually longer (Fig. 8); hind wing marginal cell narrowest at base, widening to wing apex, vein RS not sinuate. Comments.—This is a large group distin- guished by the small ocelli, hind wing mar- ginal cell narrowest at base and the non- pectinate tarsal claws. It is distinguished from the unipunctator species-group by the wider malar space and the generally uni- colored body (see key to species-groups in S. Shaw et al. 1997). KEY TO THE NORTH AMERICAN SPECIES OF THE GASTERATOR SPECIES-GROUP ile Fore wing vein |cu-a beyond 1M by dis- tance less than or equal to length of Icu-a, rarely slightly longer (Fig. 4) - Fore wing vein Icu-a beyond IM by dis- tance distinctly greater than length of Icu- Ey) (Lor 2.011) A ee ee ear a a 4 Body bicolored, head and mesosoma black, metasoma red carinatus (Ashmead) _ Body unicolored orange or yellow, at most apical metasomal terga black Fore wing vein 3RSb curved and reaching wing margin well before apex of wing, marginal cell short (Fig. 4); antenna with more than 40 antennomeres 15 cin top ew ooo ae brachyphlebus, new species - Fore wing vein 3RSb straight and reaching wing margin near wing apex, marginal cell longer; antenna with 40 antennomeres or less medicinebowensis, new species Antenna black with middle third of flagel- vin WaM saeco nese dee smithi, new species 4(1). 293 - Antenna entirely black, brown or yellow, or yellow at base gradually darkening to apex, without white annulus 5(4). First metasomal tergum coarsely rugose, without any indication of costae (Fig. 9) wise wile Fpsegiaes vogeiytie eee et eae burrus Cresson - First metasomal tergum costate or costate rug ose) (Pig: 10) ees eee eens 6 6(5). Antenna with less than 40 antennomeres: metasomal terga 1—2 orange, terga 3-7 DIAC AR Orc Ae x nt See nee aches 7 - Antenna with more than 40 antennomeres; body entirely honey yellow or orange ... 8 7(6). Hind femur black on apical %; malar space usually orange townesorum, new species — Hind femur orange; malar space black .. . ead P EPEC EES SAE bucculentus, new species 8(6) Diameter of lateral ocellus equal to or slightly greater than ocell-ocular distance; hind wing vein RS parallel to anterior wing margin on basal % and distinctly curving downward toward wing apex (Fig. 3); tem- ples in dorsal view narrow, less than eye Wid thine ae eee vierecki, new species ~ Diameter of lateral ocellus less than ocell- ocular distance; hind wing vein RS more or less straight to wing apex; temples in dorsal view as wide as eye . atricornis (Cresson) Aleiodes atricornis (Cresson), new combination (Figs: 2:, 10) Rogas atricornis Cresson 1872:188. Rhogas ferrugineus Enderlein 1920 (1918): 156. N. syn. Diagnosis.—Body unicolored honey yel- low to yellow orange, mesosoma occasion- ally marked with black in male, apical tar- somere brown, antenna with scape and ped- icle honey yellow and flagellum either en- tirely brown or orange on basal half darkening to brown on apical half, wings hyaline, veins yellow to brown; body length, 6.0—9.5 mm; 58—68 flagellomeres; length of malar space greater than basal width of mandible; diameter of oral open- ing greater than basal width of mandible and equal to malar space; face costate-co- riaceous, frons, vertex and temple coria- ceous; pronotum coriaceous, rugose later- ally, mesonotum and scutellum coriaceous, mesopleuron coriaceous, subalar sulcus and sternaulus weakly rugulose, propodeum ru- 294 Figs. 1-6. A. smithi. 6, A. terminalis. gose coriaceous, median carina usually complete, occasionally absent apically; first and second metasomal terga costate coria- ceous, median carina complete, third ter- gum costate on basal half, median carina absent (Fig. 10); fore wing with vein Icu-a beyond IM by distance nearly twice length of lcu-a; hind wing with marginal cell gradually widening, vein RS straight, vein m-cu short and weak (Fig. 2); tarsal claws PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Wings of Aleiodes species. 1, A. burrus. 2, A. atricornis. 3, A. vierecki. 4, A. brachyphlebus. 5, not pectinate but with 2—3 thin spines at extreme base. Type material examined.—Rogas atri- cornis Cresson, lectotype female (designat- ed by Cresson 1916), Texas [Philadelphia]. Rhogas ferrugineus Enderlein, lectotype fe- male (here designated to fix the current in- terpretation of this name and to ensure sta- bility and uniformity in its future interpre- tation), Mexico, Chiapas, L. Conradt S. VOLUME 103, NUMBER 2 [Warsaw]; paralectotype male (here desig- nated), same data. The status of the type series of atricornis is very confused because of the existence of labeled type specimens at both Philadel- phia (7 specimens) and Washington D.C. (4 specimens) that exceed the total number of specimens in the original syntype series (4 specimens). Cresson (1872) described atri- cornis based on four female specimens from Texas (Belfrage collection). In the in- troduction of that paper he stated that ‘“‘un- less otherwise indicated” the types “are to be found in the collection of the American Entomological Society’’ (Philadelphia). Since he did indicate “‘Belfrage’’ as the source of these specimens, presumably part of the syntype series may have been re- turned to Mr. Belfrage after the completion of the 1872 work. However, it is not clear, either from statements in the 1872 paper or from examination of the specimens in Phil- adelphia and Washington, DC, which four specimens actually comprise the original syntype series. Mr. Belfrage died in 1882, and subsequently his collection was donat- ed to the Smithsonian Institution. Among the specimens accessioned at the Smithson- ian were exactly four female specimens now with ‘Texas, Belfrage’’ labels. By 1896 these four specimens had been as- signed the USNM type number 1622, in- dicating that they were presumed to be the type series at that point in time. In 1916 Cresson published his paper on the Cresson types of Hymenoptera, wherein a lectotype of atricornis was designated from the ma- terial at Philadelphia and assigned type number 1672 (although E. T. Cresson, Jr. assisted with the preparation of this manu- Script, it was published by E. T. Cresson, Sr.). This data indicates a single specimen from Texas in good condition, but Belfrage is not listed on the label data, nor are other specimens listed. Cresson (1916) stated that “‘unique’’ specimens were returned to Mr. Belfrage. Since atricornis was described based on four specimens (not unique), we must assume that at least one syntype was 295 kept in Philadelphia. Clearly, more speci- mens were later collected by Mr. Belfrage and donated to both the American Ento- mological Society and the Smithsonian In- stitution. These specimens have now be- come mixed with the syntype series such that it is now impossible for us to determine with certainty the identity of the four syn- types. Given that Cresson (1916) recog- nized the Belfrage collection as being in Washington, DC, and designated other lec- totypes based on the USNM material, we must assume that he correctly picked an original syntype in Philadelphia as the lec- totype in this case. However, since the iden- tity of the remaining three syntype speci- mens is uncertain, it would be fruitless to attempt to identify the paralectotypes at this time. Distribution.—Specimens examined were from South Dakota, Nebraska, Kansas, Oklahoma, Texas, New Mexico, Arizona and Mexico. Biology.—Several noctuid hosts are list- ed in the literature but we have seen no specimens reared from hosts; these records need to be confirmed. Comments.—Aleiodes atricornis was previously distinguished from ferrugineus only by the color of the antenna, that of atricornis being entirely brown and that of ferrugineus being orange basally changing to brown apically. Otherwise they are iden- tical in morphology and distribution and we consider them to be the same species. Del- fin and Wharton (2000) transferred ferru- gineus to Aleiodes. Aleiodes brachyphlebus Marsh and Shaw, new species (Fig. 4) Female.—Body color: dark honey yel- low or brown with black marking along no- tauli, sternaulus, sides of scutellum, metan- otum, propodeum and first metasomal ter- gum; antenna brown basally, becoming black toward apex; wings hyaline, veins brown. Body length: 5—7 mm. Head: eyes and ocelli normal size; 46 antennomeres, 296 first flagellomere longer than second, basal half of flagellomeres about as wide as long; malar space % eye height and equal in length to basal width of mandible; temple about % eye width; occipital carina not meeting hypostomal carina; oral space broad and oval, width greater than malar space and face length; clypeus protruding; ocelli small, ocellocular distance slightly greater than diameter of lateral ocellus; face, frons and vertex rugose, temple smooth; maxillary palpus not swollen; man- dibles large, tips crossing when closed. Me- sosoma: pronotum rugose laterally; meson- otum weakly coriaceous and dull anteriorly, shining and smooth posteriorly, scutellum smooth and shining; notauli weakly scro- biculate, meeting posteriorly in small tri- angular rugose area; mesopleuron weakly punctate and shining, subalar sulcus and sternaulus rugose; propodeum rugose, me- dian carina present only weakly at base. Legs: tarsal claws long and slender, not pectinate at apex, with 4—5 short slender spines at extreme base; inner spur of hind tibia about as long as % hind basitarsus; hind coxa weakly punctate and shining dor- sally. Wings (Fig. 4): fore wing vein r % length of 3RSa and of m-cu, marginal cell short, vein 3RSb meeting wing margin well before wing apex, vein Icu-a beyond 1M by distance slightly less than length of Icu- a, vein 1CUa % length of 1CUb; hind wing with vein RS straight on basal half, bent downward on apical half so marginal cell suddenly wider at apex, vein r-m slightly shorter than 1M, vein M+CU longer than 1M, vein m-cu indicated only by short clear raised line. Metasoma: first tergum costate- coriaceous, length about equal to apical width, median carina complete; second ter- gum costate-coriaceous, median carina complete; third tergum costate on basal half, smooth on apical half, median carina absent; remainder of terga smooth; ovipos- itor about % length of hind basitarsus. Male.—Essentially as in female. Holotype.—?: WASHINGTON, Ritz- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ville, May 10, 1922, M. C. Lane col. De- posited in USNM. Paratypes.—BRITISH COLUMBIA: 1 ?, Stone Mt. Pk., 3,800 ft., July 13, 1973, H. and M. Townes. COLORADO: 2 @&, Mesa Co., Big Wash, May 4, 1992, Kon- dratieff, Cranshaw & H. Knuttel. NE- BRASKA: 2 @, Valentine Refuge, June 4, 1972, H. and M. Townes. OREGON: 3 °&, 4 S, Riley, April 29 and May 18, 1976, H. and M. Townes. WASHINGTON: 1 2, same data as holotype. WYOMING: 1 4, Albany Co., TISN R73W, 7,500’, May 15, 1978, C. D. Ferris. Deposited in USNM, RMSEL, CSU, AEI. Biology.—Unknown. Comments.—This species is superficially similar to burrus and atricornis but is dis- tinguished by the short fore wing marginal cell, the short distance separating vein lcu- a from 1M, and the smooth mesonotum. Etymology.—The specific name is from the Greek brachys meaning short and the Greek phlebos meaning vein, referring to the short radial vein and radial cell in the fore wing. Aleiodes bucculentus Marsh and Shaw, new species Female.—Body color: head orange ex- cept temple and malar space which are black, antenna orange, apical % of flagellum brown; mesosoma orange, mesopleuron, metanotum and sides of propodeum black; first and second metasomal terga orange, third tergum varying from entirely black to black on apical % and orange on basal %, remainder of terga black, metasomal venter orange; legs orange, apex of hind femur and tibia marked with brown, apical tarsomere of hind leg sometimes black; wings dusky, veins brown. Body length: 5.5—6.0 mm. Head: eye small, malar space 7, eye height and longer than basal width of mandible; temple as wide as eye, swollen in dorsal view; 36-38 antennomeres, nearly all fla- gellomeres as wide as long; occipital carina not distinctly meeting hypostomal carina; oral space circular, slightly wider than basal VOLUME 103, NUMBER 2 width of mandible; ocelli small, ocellocular distance about twice diameter of lateral ocellus; face rugulose-coriaceous, frons ru- gose, vertex and temple coriaceous; maxil- lary palpus short and slightly swollen. Me- sosoma: pronotum rugose laterally; meson- otum and scutellum coriaceous, notauli weakly scrobiculate, area where they meet obscured by pin; mesopleuron smooth me- dially except for hair pits, subalar sulcus and sternaulus rugose; propodeum rugose, median carina complete. Legs: tarsal claws not pectinate; hind coxa somewhat rugulose dorsally. Wings: fore wing with vein r 4 length of 3RSa and % length of m-cu, vein lcu-a beyond 1M by distance greater than length of Icu-a, vein 1CUa % length of 1CUb; hind wing marginal cell gradually broadening to apex, vein RS straight, vein M-+CU twice length of 1M, vein r-m about ¥, length of 1M. Metasoma: first tergum wider at apex than long, costate-rugose, median carina complete; second tergum costate, median carina complete, third ter- gum costate on basal %, smooth on apical ’%2; remainder of terga smooth; ovipositor short, less than % length of hind basitarsus. Male.—Unknown. Holotype.—?: ALBERTA, Kananaskis, Bor Exp. Sta. Seebe, June 15, 1968, H. J. Teskey, Malaise trap. Deposited in CNC. Paratypes.—ALBERTA: 2 9°, Bilby, June 7, 27, 1924, George Salt. MANITO- BA: 1 2, Churchill, June 29, 1956. Depos- ited in USNM, RMSEL, BMNH. Biology.—Unknown. Comments.—The swollen temples are characteristic for this species which, along with the color of the head and hind legs, will distinguish it from townesorum. Etymology.—The specific name is from the Latin bucculentus meaning full cheeks in reference to the swollen temples. Aleiodes burrus Cresson (Figs. 1, 8, 9) Aleiodes burrus Cresson 1869:381. Aleiodes fulvus Cresson 1869:381. N. syn. Rhogas nigricoxis Viereck 1903:97. N. syn. 297 Rhogas cockerelli Viereck 1905:266. N. syn. Rhogas fuscicaudus Viereck, in Withington 1909-329) N:; nud: Aleiodes waldeni Viereck 1917(1916):236. N. syn. Rogas burrus: Marsh 1979:179. Aleiodes burrus: Delfin and Wharton 2000: 58. Diagnosis.—Body unicolored in female and some males, honey yellow, occasion- ally with black markings on mesosoma, an- tenna orange basally to black at apex; in many males, body often bicolored orange and black, head and mesosoma black, me- tasoma orange or orange and black, wings hyaline, veins brown, vein C+Sc+R in fore wing of female often yellow; body length, 6.0—-8.0 mm; 45-52 antennomeres in fe- male, 55—60 in male; malar space longer than basal width of mandible (Fig. 8); face, frons and vertex rugose to rugose-areolate, temple sometimes nearly smooth; ocello- cular distance greater than diameter of lat- eral ocellus; pronotum rugose; mesonotum coriaceous-punctate, punctures deep and conspicuous in male, mesopleuron rugose except punctate shining medially; propo- deum rugose, median carina complete; first and second metasomal terga coarsely ru- gose, first tergum always and second usu- ally without any or only weak indications of costae, median carina complete, third ter- gum costate on basal 4%—% median carina absent (Fig. 9); vein Icu-a of fore wing be- yond IM by distance greater than length of Icu-a; hind wing marginal cell gradually widening, vein RS straight or bent slightly downward at middle, vein m-cu present but weak (Fig. 1); tarsal claws not pectinate. Type material examined.—Aleiodes bur- rus Cresson, lectotype female, I[llinois [ANSP]. Aleiodes fulvus Cresson, lectotype male, Canada [ANSP]. Rhogas nigricoxis Viereck, holotype male, Beulah, New Mex- ico [ANSP]. Rhogas cockerelli Viereck, ho- lotype male, Douglas County, Kansas [UK]. 298 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 7-12. species. 9, A. burrus. 10, A. atricornis. 11, A. harrimani. 12, A. pseudoterminalis. Aleiodes waldeni Viereck, holotype male, Stonington, Connecticut [USNM]. Distribution.—Eastern Canada south to Virginia, west to Alberta and Arizona. Biology.—Reared from the noctuids Ac- ronicta hasta Guenée, A. lobeliae Guenée, and Feltia subgathica (Haworth). Comments.—This large honey-yellow and moderately common species is distinc- tive by its coarsely rugose first metasomal tergum and the usually bicolored black and orange male. Previously, the exact identities of the species fulvus, nigricoxis, cockerelli 7-8, Head of Aleiodes species. 7, A. terminalis. 8, A. burrus. 9-12, Metasomal terga of Aleiodes and waldeni, all described from males, were problematic. After closer examination, it is clear that they all represent the males of this dimorphically colored species. For many years the identities of cockerelli Viereck and fuscicaudus Viereck were not certain because no holotypes were found. Recently a specimen was found in the Snow Ento- mological Museum, University of Kansas, labeled as the type of fuscicaudus but which agrees precisely with the original descrip- tion and locality data of cockerelli. The name fuscicaudus appears only in a list of VOLUME 103, NUMBER 2 types in the Snow Museum published by Withington (1909). We suspect that Viereck described the species as fuscicaudus in his original manuscript, changed the name to cockerelli in the published version, but failed to change the type label on the spec- imen from which Withington made his en- try. Aleiodes carinatus (Ashmead), new combination Rhysipolis carinatus Ashmead 1889 (1888): O25. Rhogas pubescens Ashmead 1889 (1888): 632. N. syn. Aleiodes nasonii Ashmead: Nason 1905: 298. N. nud. Diagnosis.—Body bicolored, head, me- sosoma and coxae black, tegula yellow, pal- pi, mandibles, remainder of legs and me- tasoma mostly orange, wings dusky hya- line, stigma black; body length 9 mm; 66— 68 antennomeres; malar space short, equal to or less than basal width of mandible; face and frons rugose, temple and vertex rugose- coriaceous; mesonotum and scutellum co- riaceous; pronotum rugose; mesopleuron rugose, smooth above episternal scrobe; propodeum coarsely rugose dorsally, ru- gose-coriaceous laterally, median carina complete; first and second metasomal terga rugose, median carinae complete; third ter- gum striate on basal half, coriaceous on api- cal half, median carina on basal half only; remainder of terga coriaceous; vein lcu-a of fore wing beyond 1M by distance less than length of Icu-a, hind wing marginal cell narrowest basally, gradually widening apically, vein RS straight; tarsal claws with 2-3 short spines at base, hind coxa rugulose dorsally. Type material examined.—Rhysipolis carinatus Ashmead, holotype male, Texas [USNM]; Rhogas pubescens Ashmead, ho- lotype male, Wisconsin [USNM]. Distribution.— Wisconsin, [llinois, and Texas. Biology.—Host unknown. 299 Comments.—At present known only from the male. This species is likely the male of another species but we have not found any females with the combination of characters that distinguish carinatus, name- ly the bicolored body with entirely orange metasoma, the short malar space, and vein Icu-a of the fore wing positioned close to the vein 1M. Aleiodes medicinebowensis Marsh and Shaw, new species Female.—Body color: dark honey yel- low; antenna honey yellow on basal half, brown on apical half; maxillary palpomeres 1 and 2 and labial palpi entirely black; me- sosternum black; wings dusky, veins dark brown, tegula yellow. Body length: 7.0 mm. Head: eyes and ocelli normal size, not covering most of head; 38—39 antennom- eres, all flagellomeres as wide as long; ma- lar space longer than basal width of man- dible and about % eye height; temple broad, nearly equal to eye width; occipital carina not reaching hypostomal carina; oral space circular, width slightly greater than basal width of mandible and about %4 face height; clypeus swollen; ocelli small, ocellocular distance greater than diameter of lateral ocellus; face and frons rugose, vertex and temples rugulose; maxillary palpus not swollen; mandible small, tips not crossing when closed. Mesosoma: pronotum rugose; mesonotum and scutellum punctate, notauli weakly scrobiculate; mesopleuron punctate medially, smooth above episternal scrobe, subalar sulcus and sternaulus rugose; pro- podeum areolate, median carina sometimes absent on apical half. Legs: tarsal claws not pectinate, with few spines at extreme base; hind coxa rugulose dorsally. Wings: fore wing with vein r about 12 length of 3RSa and % length of m-cu, vein Icu-a beyond 1M by distance equal to or very slight greater than length of Icu-a, vein 1CUa % length of 1CUb; hind wing vein RS straight, marginal cell gradually broadening to wing apex, vein r-m shorter than 1M, vein 1M about %4 length of M+CU, vein m- 300 cu weak and indistinct. Metasoma: first ter- gum costate rugose, apical width about equal to length, median carina complete; second tergum costate rugose, median ca- rina complete; third tergum costate on basal 4, median carina absent; remainder of terga smooth; ovipositor % length of hind basi- tarsus. Male.—Unknown. Holotype.—?: WYOMING, Albany Co., 1.5 mi. W of Centennial, Medicine Bow Natl. Forest, Snowy Range, mixed forest, Malaise, Mian, July 14-27, 1991. Deposited in RMSEL. Paratypes—WYOMING: 1 2, Albany Co., Medicine Bow Nat. Forest, 1 mi. N. Lincoln Monument, mixed forest nr. sage- brush, Mian June 20—26, 1990, Malaise trap; | 2, Albany Co., Medicine Bow Nat. Forest, 2 mi. N on Rd. 705, July 15-19, 1991, willow bog Malaise trap. Deposited in RMSEL, USNM. Biology.—Unknown. Comments.—This species is similar to burrus but can be distinguished by its short- er antennae and the dark colored labial pal- pi. Etymology.—Named for the type locality of Medicine Bow National Forest. Aleiodes smithi Marsh and Shaw, new species (Fig. 5) Female.—Body color: entirely honey yellow, mesonotum occasionally marked with brown; antenna with scape and pedicel yellow, flagellum black except for middle 5—12 flagellomeres which are white; wings lightly infumated, veins brown, stigma with yellow spot at base, tegula yellow. Body length: 6-8 mm. Head: eyes and ocelli normal size; 44-50 antennomeres, flagel- lomeres beyond middle as long as wide; first and second flagellomeres equal in length, apical flagellomere bluntly pointed; malar space long, % eye height and 2 times basal width of mandible; temple moderate, ¥, eye width; occipital carina not always meeting hypostomal carina; oral opening PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON moderate, width equal to 7 face height and ¥%, malar space; clypeus narrow and not pro- truding; ocelli small, ocellocular distance 2 times diameter of lateral ocellus; face, frons, vertex and temple rugose-coriaceous, face with short median ridge between an- tennae; maxillary palpus somewhat swol- len; mandibles small, tips not crossing when closed. Mesosoma: pronotum rugose; mesonotum and scutellum coriaceous; no- tauli scrobiculate, meeting posteriorly in wide rugose area; mesopleuron coriaceous and dull medially, shining above episternal scrobe, subalar sulcus and sternaulus strongly rugose; propodeum strongly ru- gose, median carina not complete, obscured on apical half of propodeum. Legs: tarsal claws not pectinate, with only 2—3 slender spines at extreme base; inner spur of hind tibia less than half length of hind basitarsus; hind coxa granular dorsally at base, rugose dorsally at apex. Wings: lightly infumated; fore wing with vein r short, about / length of 3RSa, vein Icu-a beyond IM by distance slightly more than length of Icu-a, vein 1CUa nearly % length of 1CUb; hind wing vein RS nearly straight, marginal cell grad- ually widening to apex, vein r-m shorter than 1M, vein M+CU slightly longer than 1M, vein m-cu indicated by short weakly infuscated line. Metasoma: first tergum strigate-rugose, slightly longer than apical width, median carina complete; second ter- gum strigate-rugose, median carina usually complete, sometimes indistinct apically; third tergum strigate on basal %, coriaceous on apical %, rarely a short stub of median carina present when carina on second ter- gum complete; remainder of terga finely co- riaceous and shining; ovipositor short, % length of hind basitarsus. Male.—Essentially as in female except antennae may be slightly longer, with up to 56 antennomeres. Holotype.—?: VIRGINIA, Clarke Co., U. Va. Blandy Exp. Farm, 2 mi. S Boyce, July 25—August 7, 1990, Malaise trap, Da- vid R. Smith. Deposited in USNM. Paratypes.—FLORIDA: 1 @, Enterprise; VOLUME 103, NUMBER 2 1 2, Alachua Co., E. Gainesville, February 18, 1975, H. Greenbaum, Malaise trap; 1 3, Archbold Biol. Sta., Lake Placid, High- lands Co., May 5, 1989, M. Deyrup. GEORGIA: 1 <6, Pine Mtn., Raburn Co. 1,400’, May 15, 1957, W. R. M. Mason; 2 3, Forsyth, May 20-31, 1968, G. Heinrich. ILLINOIS: | 3, N. Illinois, Andreas Bolter; 2 6, Hart collection #171 (no other data). KANSAS: 2 2, Onaga, Crevecoeur. KEN- TUCKY: 1 ¢, Boone Co., Big Bone Lick St. Park, June 22, 1982, R. Wharton; 3 ¢, Locust Grove, Louisville, Jefferson Co., August 13, 1978 and September, 1982, S. Riegler MARYLAND: 6 @, 20 6, Cabin John, dates ranging from May 1, 1916 to June 24, 1920 and mostly collected by R. MsFouts; lady College: Park,»PGx,Go., June 21, 1981, S. R. Shaw; 4 2, 6 ¢. Glen Echo, July, 1917 and July 20, 1919, R. M. Fouts; 1 2, Patuxent, July, 1980; 1 2, Co- lesville, Montg. Co., June 8, 1975, A. S. Menke. MASSACHUSETTS: 1 2, Marthas Vineyard, August 10, 1931, C. W. Johnson; 1 3d, Nashawena Is., Elizabeth Is., July 7, 1971, C. T. Parsons. MICHIGAN: 1 @, Lansing, July 17, 1956, H. Niemezyk. MIS- SISSIPPI: 1 2, Oktibbeha County, Stark- ville, July 11, 1987, M. Ludlow, black light trap; 1 2, Jefferson Davis County, July 18, 1973, J. R. McCoy, light trap. MISSOURI: 3 2, Williamsville, July 5-September 10, 1969, J. T. Becker; 1 2, Columbia, Boone Co., October 1969, E D. Parker, collector. ONTARIO: 1 @, Pt. Pelee, July 17, 1962, S. M. Clark. NEW JERSEY: 1 ¢, Colling- wood, July 17, 1904, G. M. Greene; 1 @, Moorestown, August 1, 1939, H. & M. Townes. NEW YORK: 1 6, Cold Spring Harbor, C. T. Brues. NORTH CAROLINA: 7 2, 7 6, Johnston Co., Clayton, August 29 and September 7, 1972, ex. Lascoria ambigualis, W. M. Brooks (Soybean Re- search Voucher Specimens); 1 6, Washing- ton Co., Plymouth, September 7, 1972, W. M. Brooks (Soybean Research Voucher Specimens); 1 2, L. Junaluska, July 28, 1937. Ha VaWeems;Jn;d1--°:, 7d, Orange Co., Chapel Hill, September 9, 1975 to 301 September 4, 1976, Malaise trap; 1 2, Swain Co., Smokemont, July 16, 1977, Malaise trap; 2 2, Bertie Co. near Cahaba, July 27, 1976 and October 31, 1978, Mal- aise trap; 1 2, Mecklenburg Co., Charolet- te, August 22, 1979, Malaise trap; 8 &, 1 3, Martin Co. near Williamston, June 16— July 29, 1975, Malaise trap. PENNSYL- VANIA: 1 2, North East, June 1917. TEN- NESSEE: 1 ¢, Lexington, Natchez Trace S. P, June 15-19, 1972, G. Heinrich. VIR- GINIA: 100 2, 104 d, same data as holo- type with dates ranging from May 1, 1990 to October 22, 1990; 7 2, 2 3, Louisa Co., 4 mi S Cuckoo, dates ranging from May 28, 1987 to June 18, 1988, J. Koke and D. R. Smith, Malaise trap; 1 ¢, Arlington, Coun- try Club Hills, October 10, 1942, R. A. Cushman. WISCONSIN: 1 6, West Bend, Washington Co., August 13—14, 1966, H. E. Evans. MEXICO: 1 ¢, Colima, 9 mi NE Comala, July 17-18, 1983, Kovarik, Har- rison, Schaffner; 1 2, Morelos, 4.4 mi E Cuernavaca, July 27-29, 1976, Malaise trap, Peigler, Gruetzmacher, R. & M. Mur- ray, Schffner; 1 2, Oaxaca, 10.8 mi S El Punto, July 19, 1987, R. Wharton. COSTA RICA: 1 @, San José Prov., Ciudad Colon, 800 m, December 1989—January 1990, Luis Fournier; 1 ¢@ Guanacaste Prov., Cerro 1 Hacha, nw Volcan Orosi, 300 m, 1988; 1 2, Cartago Prov., La Cangreja, 1,950 m, November—December, 1992, Paul Hanson; 4 2, Cartago Prov., Dulce Nombres, Vivero Linda Vista, 1,400 m, June-August, 1993, Paul Hanson; 7 2, 7 d, San José Prov., Zurqui de Moravia, 1,600 m, January—Feb- ruary, 1989, July, 1990, March—May, 1992, February, 1996, Paul Hanson. Deposited in USNM, RMSEL, TAMU, MCZ, BMNH, CNG. MISS; UCD; NGSU, «NCDA; NNML, INHS, ABS, MSU. Biology.—Three specimens were reared from Lascoria ambigualis Walker (Noctui- dae). Comments.—This species is distinct from all other North American species by the white annulus on the antennae. Also distinctive is the very short vein r in the 302 fore wing. It resembles the Neotropical spe- cies ecuadoriensis Brues which also has a white annulus on the antennae, but that spe- cies has darker hind tibiae, a black head and black prothoracic area, and a much deeper and coarsely scrobiculate groove between metasomal terga 2 and 3. Etymology.—This species is named for the collector of most of the type series from Virginia, sawfly specialist David R. Smith. Aleiodes townesorum Marsh and Shaw, new species Female.—Body color: head varying from orange to light brown, antenna with scape, pedicel and basal % of flagellum yel- low, apical % of flagellum brown; mesoso- ma orange, mesosternum black; first and second metasomal terga orange, third and following terga entirely black; fore and middle legs honey yellow, middle tibia and tarsus sometimes darker, hind coxa and tro- chanters honey yellow, hind femur honey yellow on basal % or less, black on apical *—%4, hind tibia honey yellow on basal %, black on apical %, hind tarsus black; wings dusky, veins brown. Body length: 5.0—6.0 mm. Head: 35-38 antennomeres, nearly all flagellomeres as wide as long; eye small, malar space 4% eye height and longer than basal width of mandible; temple slightly less than width of eye, not swollen in dorsal view; occipital carina not meeting hypos- tomal carina; oral space circular, slightly wider than basal width of mandible; ocelli small, ocellocular distance about twice di- ameter of lateral ocellus; face costate punc- tate, malar space punctate, temple punctu- late and shining, vertex costate punctate, frons rugulose and shining. Mesosoma: pronotum rugose; mesonotum and scutel- lum punctate, notauli weakly scrobiculate, meeting in triangular rugose area before scutellum; mesopleuron minutely punctate and shining, subalar sulcus and sternaulus rugose; propodeum rugose, median carina distinct only on basal half. Legs: tarsal claws not pectinate; hind coxa somewhat rugulose dorsally. Wings: fore wing with PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON vein r % length 3RSa and % length of m- cu, vein Icu-a beyond 1M by distance greater than length of Icu-a, vein 1CUa % length of 1CUb; hind wing marginal cell gradually broadening to apex, vein RS straight, M+CU 1.5 times length of 1M, vein r-m about ¥; length of 1M. Metasoma: first tergum wider at apex than long, cos- tate-rugose, median carina complete; sec- ond tergum costate-rugose, median carina complete, third tergum costate on basal %, smooth on apical 4%; remainder of terga smooth; ovipositor short, less than 4% length of hind basitarsus. Male.—Essentially as in female; head, flagellum and mesosoma black; 45—47 an- tennomeres. Holotype.—?: ARIZONA, Portal, Au- gust 23, 1987, H. and M. Townes. Depos- ited in AEI. Paratypes.—ARIZONA: 8 @&, 13 6, same data as holotype with date of August 23—September 12, 1987; 1 2, 5 mi. W. Por- tal, Chiricahua Mtns., August 18, 1958, R. E. Rice. Deposited in USNM, RMSEL, AEI. Biology.—Unknown. Comments.—This species is very similar to bucculentus but can be distinguished by the darker hind leg, lighter head color, and the temple which is not swollen in dorsal view. Etymology.—The specific name is in honor of our friends and colleagues, Mar- jorie and her late husband Henry Townes. Aleiodes vierecki Marsh and Shaw, new species (Fig. 3) Female.—Body color: entirely honey yellow, apical flagellomeres occasionally light brown; wings hyaline, fore wing veins brown, yellow at wing base, tegula yellow, hind wing veins yellow. Body length: 7.0— 8.5 mm. Head: 56—65 antennomeres, basal flagellomeres about as wide as long; malar space equal to basal width of mandible and about % eye height; temple narrow, about % eye width; occipital carina not meeting VOLUME 103, NUMBER 2 hypostomal carina; oral opening small, cir- cular, diameter about equal to basal width of mandible and slightly less than eye height; ocelli moderate sized, diameter of lateral ocellus equal to or slightly greater than ocell-ocular distance; head entirely ru- gulose; palpi not swollen; mandibles small, tips not crossing when closed. Mesosoma: pronotum rugose laterally; mesonotum and scutellum coriaceous, notauli deep, scrobic- ulate, meeting in triangular rugose area be- fore scutellum; mesopleuron smooth dor- sally, rugulose below and at subalar sulcus and sternaulus; propodeum rugose dorsally, coriaceous laterally, median carina ob- scured on apical %. Legs: tarsal claws not pectinate, hind coxa smooth dorsally. Wings (Fig. 3): fore wing with vein r % length of 3RSa and about % length of m-cu, vein 1cu- a beyond IM by distance slightly greater than length of Icu-a, vein 1CUa % length of 1CUb; basal % of hind wing vein RS nearly parallel to apical wing margin, then curving downward, marginal cell suddenly widening, vein r-m about % length of 1M, vein 1M about 7; length of M+CU, vein m- cu short but distinct, often arising from 2M slight distad of r-m. Metasoma: first tergum rugulose costate, apical width twice basal width, median carina complete; second ter- gum rugulose costate, median carina com- plete, third tergum rugulose costate on basal %, remainder weakly coriaceous and shin- ing; remainder of terga weakly coriaceous and shining; ovipositor about *%4 length of basitarsus. Male.—Essentially as in female. Holotype.—?: KANSAS, Lawrence, April 1956, A. R. Barr, collected at light. Deposited in USNM. Paratypes.—KANSAS: 18 2, 1 36, same data as holotype. MICHIGAN: 2 92, 1 6, Midland Co., June 6, 1939, June 9, 1940, June 23, 1941, R. R. Dreisbach. OKLAHOMA: | 2, Latimer Co., Red Oak, May 7-14, 1985, K. Stephan. Deposited in USNM, RMSEL, AEI. Biology.—Unknown. Comments.—This species is very close 303 to atricornis but is separated by the larger ocelli, slightly curved hind wing vein RS and narrower temple. The short malar space places it near species in the unipunctator group, but those species have the malar space even shorter and bodies bicolored black and red. Etymology.—Named for H. L. Viereck who was a student of the Braconidae and other parasitic Hymenoptera in the early 1900’s. ALEIODES UNIPUNCTATOR SPECIES-GROUP Included species.—unipunctator (Thun- berg) 1822, n. comb., Europe; ruficornis (Herrich-Schaeffer) 1838, n. comb., Eu- rope; terminalis Cresson 1869, North America; hirtus (Thomson) 1891, n. comb., Europe; harrimani (Ashmead) 1902, n. comb., North America; eurinus (Telenga) 1941, n. comb., Europe; turkestanicus (Te- lenga) 1941, n. comb., Europe; alboannu- latus (Belokogyl’skij) 1988, n. comb., Eu- rope; pseudoterminalis n. sp. Diagnostic characters.—Body bicolored red or orange and black; malar space short- er than or equal to basal width of mandible (Fig. 7); hind wing vein RS straight, mar- ginal cell narrowest at base and widening toward wing apex. Comments.—This species-group is dis- tinguished from the gasterator group by its bicolored body and shorter malar space. KEY TO THE NORTH AMERICAN SPECIES OF THE UNIPUNCTATOR SPECIES-GROUP ie Third metasomal tergum costate on basal %4, remainder of tergum smooth and shining (Fig. 11) harrimani (Ashmead) = Third metasomal tergum costate on at least basal 4%, remainder of tergum coriaceous or Strate andydulli(Eiosl2)i Sian si eer-. see ee 2, 2(1). Second metasomal tergum always orange or red; fore and middle femora entirely orange; length of ovipositor at most % length of hind baSitansus#®.-4) sets aes sil terminalis Cresson = Second metasomal tergum black, rarely par- tially or entirely orange; fore and middle femora orange with black at apex; length of ovipositor at least *4 length of hind basitar- sus, often equal in length Pere oe a Woes pseudoterminalis, new species 304 Aleiodes harrimani (Ashmead), new combination (Fig. 11) Rhogas harrimani Ashmead 1902:253. Diagnosis.—Body bicolored, head, an- tenna, mesosoma black, first and second metasomal terga orange, remainder black, mandible, tegula, fore and middle legs or- ange except apical tarsomere black, hind coxa, trochanter and femur orange, hind tib- ia yellow on basal half, black on apical half, hind tarsus black; malar space short, less than basal width of mandible; face rugu- lose-coriaceous, frons rugose, vertex and temple coriaceous; mesopleuron smooth medially, subalar sulcus and sternaulus ru- gose; propodeum rugose, median carina complete; first and second metasomal terga costate, median carina complete, third me- tasomal tergum costate on basal % or less, remainder of tergum smooth and shining; vein Icu-a of fore wing beyond vein 1M by distance equal to 1.5 length of Icu-a, hind wing marginal cell narrowest basally, grad- ually widening apically, vein RS straight in apical % (Fig. 11); tarsal claws not pecti- nate. Type material examined.—Rhogas har- rimani Ashmead, holotype male, Fox Point, Alaska, Harriman Expedition [USNM]. Distribution.—Alaska, Utah, Washing- ton, South Dakota, Colorado, Montana and Wyoming. Biology.—Unknown. Comments.—Although previously re- corded only from Alaska, harrimani also occurs at high elevations in northwestern United States. It is attracted to lights and has been taken in Malaise traps. It is su- perficially similar to terminalis in size and color, but can be recognized by its shorter malar space, hind tibia without a black base, and the smooth apex of the third me- tasomal tergum. Aleiodes pseudoterminalis Marsh and Shaw, new species (Fig. 12) Female.—Body color: head and meso- soma black; antenna brown; first metasomal PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON tergum red, remainder of terga black, venter of metasoma orange on basal half; legs or- ange, apex of fore and middle femur black, apical segment of fore and middle tarsus dark brown; apex of hind femur black, base and apex of hind tibia black, orange medi- ally, hind tarsus dark brown; wings hyaline, veins dark brown, tegula yellow. Body length: 7.0-—8.0 mm. Head: 56—59 anten- nomeres, all flagellomeres slightly longer than wide, first and second flagellomeres about equal in length; malar space equal to or slightly less than basal width of mandible and % to %4 eye height; temple %2 eye width; occipital carina obscured at apical ends, not quite meeting hypostomal carina; oral open- ing small, circular, about equal in length to malar space; clypeus somewhat swollen; ocelli moderate in size, ocellocular distance equal to diameter of lateral ocellus; face coarsely rugose with smooth raised median ridge below antennae which usually termi- nates in several carinae near clypeus; frons costate; vertex and temple coarsely coria- ceous; occiput smooth and shining; maxil- lary palpus not swollen; mandible small. Mesosoma: pronotum rugose costate; me- sonotum and scutellum coriaceous; notauli shallow, scrobiculate, meeting posteriorly in Narrow rugose area; mesopleuron smooth and shining medically with scattered punc- tures, subalar sulcus and sternaulus rugose; propodeum coarsely rugose, median carina usually complete, occasionally obsolete api- cally. Legs: tarsal claws not pectinate api- cally, with a few stout spines at base; inner spur of hind tibia about % length of hind basitarsus; hind coxa smooth dorsally. Wings: hyaline; fore wing vein r % length of 3RSa and % length of m-cu, vein Icu-a beyond IM by distance twice length of Icu- a, vein 1CUa nearly % length of 1CUb; hind wing with vein RS straight but not par- allel with wing margin, marginal cell wid- ening toward apex, vein r-m slightly shorter than 1M, vein M+CU slightly longer than 1M, vein m-cu short and weakly indicated. Metasoma (Fig. 12): first tergum costate, length equal to basal width, median carina VOLUME 103, NUMBER 2 complete; second tergum costate, median carina complete; third tergum costate on basal half, median carina absent; remainder of terga coriaceous; suture between second and third terga deep and strongly scrobic- ulate; ovipositor at least % length of hind basitarsus, sometimes equal in length. Male.—Essentially as in female. Holotype.—?: VIRGINIA, Arlington, May 25, 1938, J. EK G. Clarke, Ex. Eupsilia devia. Deposited in USNM. Paratypes—MARYLAND: 1 2, Glen Echo. June, >, 1938. J. E.G. ‘Clarke. ex. Eupsilia devia; 12 2, 4 3, Silver Spring, May 19-28, 1938, J. E G. Clarke, ex. Eup- silia devia; 1 2, Burtonsville, June 29, 1980, A. S. Menke. MICHIGAN: 1 2, 1 6, Midland Co., July 11, 1936, July 2, 1938, R. R. Dreisbach; 1 2, Washtenaw Co., Ann Arbor, July 12-21, 1982, R. Wharton; 2 &, Ann Arbor, June 1976, I. and P. Gauld. NEW HAMPSHIRE: 1 ¢, Hanover, C. M. Weed. NORTH CAROLINA: 2 2, 25 ¢, Swain Co., Smokemont, June 6—August 26, 1977, Malaise trap. PENNSYLVANIA: 1 ?, Enola, July 20, 1908. TENNESSEE: 1 3, Elkmont GSMN Park, June 15, 1946, R. R. Dreisbach. VIRGINIA: 5 2, same data as holotype with additional dates of May 25225, 20, 1938, June 1, 1938; Vienna, | 9, August 9, 1936, J. C. Bridwell. WIS- CONSINe 1) 2, Fond du: lac Co T13N, RI9E, S23, September 4, 1975, Gypsy mot MI: 1 2, 16, Jackson Co., T21N, R4W, S27, May 27—September 22, 1975, Gypsy moth M.T:;; 1 ¢, T21N, R4W, S33, August 9-16, 1976, Gypsy Moth M.T:; 1 2el d, Oneida Co:, T35N, RIE, S17, July 8-29, 1975, Gypsy moth M.T.; 1 2, Dane Co., July 17, 1918, W. S. Marshall. QUE- BEC: 1 2, Wakefield, June 21, 1966, G.S. Walley; 1 2, Foster, August 2, 1929, GS. Waley; 1 2, Foster, August 2, 1929, G. S. Walley. ONTARIO: 1 2, 1 ¢, One Sided Lake, July 19, 1930, S. M. Clark; 1 6, Fin- land, July 21, 1960; S: M. Clark: 4 9, Thartway Island, St. Lawrence Is. Nat. Park, July 22—August 11, 1976, W. Reid, Malaise trap; 2 2, Cumberland, July 16, 305 1975; Ly Ling “CANADA: 2 2 2°6>no locality, collection C. EK Baker. Deposited in USNM, RMSEL, TAMU, CNC, NNML, NCDA, AEI. Biology.—Several specimens in the type series were reared from the noctuid Eupsi- lia devia (Grote). Comments.—This species has been con- fused with terminalis and is very similar but differs by the black second metasomal tergum, the black apices of the fore and middle femora, and the longer ovipositor. Etymology.—From the Greek pseudos meaning false in reference to the confusion of this species with terminalis. Aleiodes terminalis Cresson (Figs. 6-7) Aleiodes terminalis Cresson 1869:379. Diagnosis.—Body bicolored black and orange, head and mesosoma black, first and second metasomal terga orange, remainder of terga black, third tergum rarely all or partly orange, antenna light brown basally to black apically, fore leg, middle leg, hind coxa and trochanters orange, hind femur or- ange with black at extreme apex, hind tibia brown with light yellow on basal third, hind tarsus black, wings hyaline, veins brown, stigma brown, tegula yellow; body length, 5.0—8.0 mm; 55—62 antennomeres; face ru- gulose-coriaceous, frons rugose, vertex and temple coriaceous; length of malar space equal to basal width of mandible and %4 eye height (Fig. 7); ocellocular distance about equal to diameter of lateral ocellus; prono- tum rugose; mesonotum and scutellum co- riaceous, mesopleuron smooth medially, sternaulus and subalar sulcus rugose; pro- podeum rugose, median carina complete; first and second metasomal terga costate, median carina complete, third tergum cos- tate on basal half, median carina absent; fore wing with vein lcu-a beyond 1M by distance more than twice length of Icu-a; hind wing marginal cell gradually widen- ing, vein RS straight, vein m-cu present as 306 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON short stub (Fig. 6); tarsal claws not pecti- nate. Type material examined.—Aleiodes ter- minalis Cresson, holotype female, Eastern, Middle, Southern and Western States [ANSP]. Distribution.—Widely distributed over Canada and the United States south to the Mexican border; very rare below the 30th parallel. Biology.—A common parasitoid of the noctuid Pseudaletia unipuncta (Haworth) but has also been recorded from Nephelo- des minians Guenée, Spodoptera frugiper- da (Smith), S. ornithogalli (Guenée) and Xestia smithii (Snellen). Exit hole on mum- my is near the posterior end. Comments.—This is the most commonly collected species in the genus and the ma- jority of museum specimens are terminalis. It can be distinguished from other bicolored black and orange species by the combina- tion of the gradually widening marginal cell in the hind wing, the position of vein Icu- a in the fore wing, and the non-pectinate tarsal claws. It is also commonly collected at lights. ACKNOWLEDGMENTS We thank the curators of all the collec- tions listed in Methods for the loan of spec- imens for this study. Kent Hampton, Kansas State University, prepared the scanning electron micrographs and Linda Lawrence, USDA Systematic Entomology Laboratory, prepared the wing drawings. This research was supported, in part, by grant DEB-930- 6314 from the National Science Foundation to S. R. Shaw. Additional support was pro- vided by supplemental REU (Research Ex- perience for Undergraduates) grants in 1994, 1995 and 1996. LITERATURE CITED van Achterberg, C. 1991. Revision of the genera of the Afrotropical and W. Palaearctical Rogadinae Foerster (Hymenoptera: Braconidae). Zoologische Verhandlingen 273: 1—120. Ashmead, W. H. 1889(1888). Descriptions of new Bra- conidae in the collection of the U.S. National Mu- seum. Proceedings of the United States National Museum 11: 611—671. . 1902. Papers from the Harriman Alaska Ex- pidetion. 28. Hymenoptera. Proceedings of the Washington Academy of Sciences 4: 117—274. Cresson, E. T. 1869. List of the North American spe- cies of the genus Aleiodes Wesmael. Transactions of the American Entomological Society 2: 377— 382. . 1872. Hymenoptera Texana. Transactions of the American Entomological Society 4: 153-292. Cresson, E. T., Jr. 1916. The Cresson types of Hyme- noptera. Memoirs of the American Entomological Society No. 1, 141 pp. Delfin, G. H. and R. A. Wharton. 2000. Historical re- view of the genera Aleiodes and Rogas in Mexico, with a redescription of Aleiodes cameronii (Hy- menoptera: Braconidae). Pan-Pacific Entomolo- gist 76: 58-70. Enderlein, G. 1920(1918). Zur kenntnis ausureuro- paischer Braconiden. Archiv fiir Naturgeschichte 84A(11): 51-224. Fortier, J. C. and S: R. Shaw. 1999. Cladistics of the Aleiodes lineage of the subfamily Rogadinae (Hy- menoptera: Braconidae). Journal of Hymenoptera Research 8(2): 204—237. Harris, R. A. 1979. A glossary of surface sculpturing. Occasional Papers in Entomology of the Califor- nia Department of Food and Agriculture no. 28, pp. 1-31. Huber, J. T. and M. J. Sharkey. 1993. Chapter 3, Struc- ture, pp. 13-59. /n Goulet, H. and J. T. Huber, eds., Hymenoptera of the World: An Identification Guide to Families. Agriculture Canada Research Branch Publication 1894/E, 668 pp. Marsh, P. M. 1979. Family Braconidae, pp. 144-313. In Krombein, K. V. et. al., eds., Catalog of Hy- menoptera in America North of Mexico, Vol. 1, Symphyta and Apocrita (Parasitica). Smithsonian Institution Press, Washington, DC, 1,198 pp. . 1989. Notes on Braconidae (Hymenoptera) associated with jojoba (Simmondsia chinesis) and descriptions of new species. Pan-Pacific Entomol- ogist 65: 58-67. Marsh, P. M. and S. R. Shaw. 1998. Revision of North American Aleiodes Wesmael (Part 3): The seriatus (Herrich-Schaeffer) species-group (Hymenoptera: Braconidae, Rogadinae). Proceedings of the En- tomological Society of Washington 100(3): 395— 408. . 1999. Revision of North American Aleiodes Wesmael (Part 5): The melanopterus (Erichson) species-group in North America (Hymenoptera: Braconidae, Rogadinae). Journal of Hymenoptera Research 8(1): 98—108. Nason, W. A. 1905. Parasitic Hymenoptera of Algon- quin, Illinois-II. Entomological News 16:293-298. Sharkey, M. J. and R. A. Wharton. 1997. Morphology VOLUME 103, NUMBER 2 and terminology, chapter 2, pp. 19-37. In Whar- ton, R. A., PR M. Marsh and M. J. Sharkey, eds., Manual of New World Genera of the Family Bra- conidae. Special Publication of the International Society of Hymenopterists 1, 439 pp. Shaw, M. R. 1983. On[e] evolution of endoparasitism: The biology of some genera of Rogadinae (Bra- conidae). Contributions of the American Ento- mological Institute 20: 307-328. . 1994. Chapter 7, Parasitoid host ranges, pp. 112-144. In Hawkins, B. A. and W. Sheehan, eds., Parasitoid Community Ecology. Oxford Univer- sity Press, Oxford. Shaw, M. R. and T. Huddleston. 1991. Classification and biology of braconid wasps. Handbooks for the Identification of British Insects 7: 1-126. Shaw, S. R. 1995. Chapter 12.2, Braconidae, pp. 431— 463. In Hanson, P. E. and I. D. Gauld, eds., The Hymenoptera of Costa Rica. Oxford University Press, Oxford. . 1997. Subfamily Rogadinae, pp. 403-414. In Wharton, R. A., P M. Marsh and M. J. Sharkey, eds., Manual of New World Genera of the Family Braconidae. Special Publication of the Interna- tional Society of Hymenopterists 1, 439 pp. Shaw, S. R., P. M. Marsh and J. C. Fortier. 1997. Re- vision of North American Aleiodes Wesmael (Part 1): The pulchripes Wesmael species-group in the New World (Hymenoptera: Braconidae, Rogadi- 307 nae). Journal of Hymenoptera Research 6(1): 10— 39) . 1998a. Revision of North American Aleiodes Wesmael (Part 2): The ductor Thunberg species- group in the New World (Hymenoptera: Bracon- idae, Rogadinae). Journal of Hymenoptera Re- search 7(1): 62-73. . 1998b. Revision of North American Aleiodes Wesmael (Part 4): The albitibia (Herrich-Schaef- fer) and praetor (Reinhard) species-group (Hy- menoptera: Braconidae, Rogadinae). Proceedings of the Entomological Society of Washington 100(3): 553-565. Viereck, H. L. 1903. Hymenoptera of Beulah, New Mexico. Transactions of the American Entomo- logical Society 29: 43—100. . 1905. Notes and descriptions of Hymenoptera from the Western United States, in the collection of the University of Kansas. Transactions of the Kansas Academy of Sciences 19: 264—326. . 1917(1916). Guide to the insects of Connect- icut. Part III. The Hymenoptera, or wasp-like in- sects, of Connecticut. Bulletin of the Connecticut State Geological and Natural History Survey 22: 1-824. Withington, C. H. 1909. Insect types and cotypes. A preliminary list of those in the Francis Hungton Snow Entomological Collections. Transactions of the Kansas Academy of Sciences 22: 327-340. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 308-311 NEW RECORDS OF TWO PLANT BUG GENERA (HETEROPTERA: MIRIDAE: PHYLINAE: PILOPHORINI) FROM JAPAN, WITH DESCRIPTIONS OF TWO NEW SPECIES TOMOHIDE YASUNAGA Biological Laboratory, Hokkaido University of Education, Ainosato 5-3-1, Sapporo 002-8075, Japan (e-mail: yasunaga @atson.sap.hokkyodai.ac.jp) Abstract.—Two genera of the ant-mimetic plant bug tribe Pilophorini, Druthmarus Dis- tant and Hypseloecus Reuter, are reported from Japan for the first time. A new species in each genus, Druthmarus miyamotoi, n. sp., and Hypseloecus takahasii, n. sp., is de- scribed from the Ryukyu Islands in southwestern Japan. Key Words: During our continuing investigations in the Ryukyus, or subtropical islands of south- western Japan, two unique species of the phyline plant bug have been discovered. These bugs were confirmed to be undescribed species of Druthmarus Distant and Hypseloe- cus Reuter, or members of ant-mimetic tribe Pilophorini, not previously recorded from Ja- pan. These two new species are described in the present paper. All measurements in the text are given in millimeters. In the generic synonymic lists, only selected references are cited; see Ker- zhner and Josifov (1999) and Schuh (1995) for detailed lists. The type specimens are de- posited in Hokkaido University of Education, Sapporo. Genus Druthmarus Distant Druthmarus Distant 1909: 452; Schuh 1995: 455; Kerzhner and Josifov 1999: 278. Type species: D. magnicornis Dis- tant 1909. Monotypic. This small genus currently comprises only three species in the Oriental Region and Taiwan, and is easily recognized by the fuscous, oval body with densely distributed, silvery, scale-like setae, and a conspicuous, Heteroptera, Miridae, Druthmarus, Hypseloecus, new species, Japan terete or box-like antennal segment II. De- tailed generic diagnosis and redescription, including male genital structure, were pro- vided by Schuh (1984). The present discov- ery of a member of Druthmarus from Oki- nawa Island represents the northernmost distributional record for the genus. Druthmarus miyamotoi Yasunaga, new species (Figs. 1, 4—6) Description.—Body generally fuscous, suboval, small, with densely distributed, sil- very, scale-like setae that are easily rubbed away; dorsal surface subshining, rather sha- greened, impunctate. Head somewhat sha- greened, with dense, silvery, reclining, scale-like setae; vertex carinate basally; head below eyes, except for tylus, polished. Antenna fuscous; segment II terete but dis- tinctly flattened; basal 4%—% of segment III and basal /; of IV yellowish brown; lengths of segments I-IV (6/2): 0.17/0.17, 0.89/ 0.85, 0.31/0.30, 0.34/0.32. Rostrum shiny dark brown, reaching apex of mesocoxa. Thorax unicolorously fuscous, rather sha- greened. Hemelytra weakly shining, with uniformly distributed, dark, simple setae VOLUME 103, NUMBER 2 309 Figs. 1-3. female. and with patches of silvery, scale-like setae; membrane dark grayish brown. Procoxa creamy yellow, with several, dark spines apically; all femora dark brown; tibiae dark brown, with brown spines; apical half of each tibia yellow except for dark apex; tarsi pale brown excluding dark tarsomeres III; lengths of metafemur, tibia and tarsus (3/ 2): 0.88/0.92, 1.32/1.31, 0.35/0.36. Ventral surface of abdomen with patches of silvery, scale-like setae. Male genitalia as in Figs. 4—6; vesica S-shaped, slender, with a sim- ple median branch and minute, comb-like processes near secondary gonopore. Dimensions: 3/2: Body length 2.45/ 2.62; length of apex of tylus to cuneal frac- ture 1.94/2.14; head width 0.84/0.84; basal vertex width 0.35/0.39; rostral length 0.92/ 1.02; basal pronotal width 0.94/1.00; width across hemelytra 1.16/1.26. 1, Druthmarus miyamotoi, male. 2, Hypseloecus takahashii, holotype male. 3, H. takahashii, Holotype.—d, Ban’na Park, Ishigaki Is., Ryukyus, Japan, 4. iii. 1999, T. Yasunaga. Paratypes.—Okinawa Is.: 1 6, Fukuchi- dam, Kunigami Vil., 1. iv. 1999, M. Takai: 1d; 352, Yona Kunieami Vilk120-25- vy, 1993, light trap, T. Yasunaga; 3 2, same locality and collector, 11. x. 1998. Ishigaki Is.: 2 2, Nosoko, 3. iii. 1999, T. Yasunaga; 1 5, 1 2, same data as for holotype; 1 9°, Omoto, 20. i. 1996, M. Takai; 2 2°, same locality, 25. xi. 1997, T. & M. Yasunaga; 2 3, Omoto-Takeda, 7. iii. 1999, T. Yasunaga; 1 3, 1 &, without further locality, 18. i & 21. 11. 1998, K. Takahashi. Iriomote Is.: 2 3, 1 2, Funaura, 10. v. 1993, T. Yasunaga; 7 3, 1 %, same data except for date, 12. v. 1993; 1 3, same data except for collector, Y. Nakatani:*1 *¢; Takana:*22."xie 1997. LT Yasunaga; | ¢°, Toyohara-Haemida, 6. iii. 1999, T: Yasunaga. 310 eae Ss Figs. 4-9. Etymology.—Named after the well- known Japanese heteropterist, Dr. S. Mi- yamoto, celebrating his 88th birthday (a special age for celebration in Japan). Remarks.—This new species is easily distinguished from other congeners by the significantly small size, pale apical half of each tibia, and a simple, mesial branch of the S-shaped, slender vesica. Poppius (1915) reported Druthmarus sp. from Tai- wan, based on a single nymph that may fit the present new species. Druthmarus miyamotoi has been found on Pipturus arborescens (Urticaceae) and Macaranga tanarius (Euphorbiaceae), to- gether with numerous typhlocybinid leaf- hoppers (e.g., Anufrievia sp., Limassolla sp., Davmata (or Tautoneura) sp. [Typhlo- cybinidae, Homoptera]) that may serve as prey. Genus Hypseloecus Reuter Hypseloecus Reuter 1891: 50; Schuh 1995: 456; Kerzhner and Josifov 1999: 279. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Male genitalia. 4—6, Druthmarus miyamotoi. 7-9, Hypseloecus takahashii. 4, 5, 7, 8, Left para- mere. 6, 9, Vesica. Scale lines = 0.1 mm. Type species: Sthenarus visci Puton 1888. Monotypic. This genus contains thirteen species from the Old World tropics and subtropics, S. Europe, and New Guinea, and is recognized by the short, ovoid body with densely dis- tributed, sericeous, flattened setae. Generic characters were provided by Schuh (1974, 1984) and Wagner (1973). Hypseloecus takahashii Yasunaga, new species (Figs. 2—3, 7-9) Description.—Body fuscous, oval, with densely distributed, reclining, sericeous, flattened setae that are easily rubbed away; dorsal surface weakly shining, somewhat shagreened, impunctate. Head weakly sha- greened, below eyes yellow. Antenna dark brown, not incrassate; basal % of segment II, basal % of II] and extreme base of IV yellowish brown; lengths of segments I-IV (3/2): 0.27/0.28, 0.98/0.99, 0.38/0.46, VOLUME 103, NUMBER 2 0.38/0.44. Rostrum shiny dark brown, reaching apex of mesocoxa. Pronotum and scutellum with uniformly distributed, sim- ple, suberect pubescence in addition to se- riceous setae; pleura widely shagreened; ventral margin of propleuron and ostiolar peritreme yellowish brown. Hemelytra bearing dark, stiff setae; membrane dark grayish brown, with a few pale spots and partly pale veins. Coxae and legs fuscous; extreme apex of profemur reddish brown; tibiae with dark reddish brown annulations at bases of fuscous spines; tarsi pale brown; apical parts of tarsomeres III widely dark brown; lengths of metafemur, tibia and tar- sus (6/2): 1.05/1.07, 1.53/1.67, 0.39/0.41. Male genitalia as in Figs. 7—9; vesica C- shaped, pointed at apex, with two mesial notches, two simple, slender, subapical branches, and apical comb-like processes. Dimensions: 3/2: Body length 3.19/ 3.27; length of apex of tylus to cuneal frac- ture 2.43/2.55; head width 0.98/1.01; basal vertex width 0.47/0.50; rostral length 1.20/ 1.23; basal pronotal width 1.37/1.44; width across hemelytra 1.65/1.74. Holotype.—6, Nosoko, Ishigaki Is., Ryukyus, Japan, 19. iv. 1999, K. Takahashi. Paratypes.—1 2, Takeda, Ishigaki Is., 9. iv. 1999, K. Takahashi; 1 2, Kuura Riv., Iriomote Is., 11. iv. 1998, K. Takahashi. Etymology.—Named after Dr. K. Taka- hashi, who collected all type material. Remarks.—This new species is distin- guished from other congeners by the gen- erally fuscous body, and two mesial notches and two subapical branches of the vesica. No information is available on its biolo- gy. ACKNOWLEDGMENTS Special thanks are due to Dr. S. Miya- moto (Fukuoka, Japan) for continuous ad- St vice and encouragement. I am also much indebted to Mr. M. Takai (Nankoku, Japan), and Drs. K. Takahashi and Y. Nakatani (Tsukuba, Japan) for offering valuable ma- terial, and to Prof. M. Hayashi (Saitama University, Urawa, Japan) for identification of the typhlocybine leafhoppers. Mr. Takai also kindly offered the excellent photo- graphs. Thanks are extended to Dr. T. J. Henry (Systematic Entomology Laboratory, USDA, Washington, DC) and an anony- mous reviewer for improving the manu- script with comments and suggestions. LITERATURE CITED Distant, W. L. 1909. Descriptions of Oriental Capsidae. Annals and Magazine of Natural History (8)4: 440-454. Kerzhner, I. M. and M. Josifov 1999. Miridae Hahn, 1833. In Aukema, B. and C. Rieger, eds., Cata- logue of the Heteroptera of the Palearctic Region, Vol. 3, Cimicomorpha II. The Netherlands Ento- mological Society, 576 pp. Poppius, B. 1915. H. Sauter’s Formosa = Ausbeute: Nabidae, Anthocoridae, Termatophylidae, Miri- dae, Isometopidae und Ceratocombidae (Hemip- tera). Archiv fiir Naturgesichte 80A(8): 1—80 (1914). Reuter, O. M. 1891. Ein falscher und ein echter Sthen- arus (Capsidae). Wiener Entomologische Zeitung 10: 49-51. Schuh, R. T. 1974. The Orthotylinae and Phylinae (Heteroptera: Miridae) of South Africa with a phylogenetic analysis of the ant-mimetic tribes of the two subfamilies for the world. Entomologica Americana 47: 1—332. . 1984. Revision of the Phylinae (Heteroptera, Miridae) of the Indo-Pacific. Bulletin of the Amer- ican Museum of Natural History 177: 1—462. . 1995. Plant bugs of the world (Insecta: Het- eroptera: Miridae). Systematic catalog, distribu- tions, host list and bibliography. The New York Entomological Society. xi1 + 1329 pp. Wagner, E. 1973. Die Miridae Hahn, 1831, des Mit- telmeerraumes und der Makaronesischen Inseln (Hemiptera, Heteroptera). Teil 2. Entomologische Abhandlungen, Dresden 39 supplement: 1—421. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 312-318 A NEW SPECIES OF ADHEMARIUS OITICICA (LEPIDOPTERA: SPHINGIDAE) FROM MEXICO MANUEL A. BALCAZAR-LARA AND CARLOS R. BEUTELSPACHER (MAB-L) Facultad de Ciencias, Universidad de Colima, Apdo. Postal 25, Av. 25 de Julio #965, Col. Villas San Sebastian, C. P. 28000, Colima, Mexico, (e-mail: mab] @cgic.ucol.mx); (CRB) Instituto de Historia Natural, Calz. a Cerro hueco s/n A.P. No. 6, C.P. 29000, Tuxtla Gutiérrez, Chiapas, Mexico Abstract.—Adhemarius mexicanus, new species, is described from the Sierra Madre Oriental mountains, Mexico. A key to males of the A. donysa species group (A. donysa, A. dariensis, A. mexicanus, A. globifer, and A. blanchardorum) is presented, and specific characters are discussed. Key Words: Adhemarius blanchardorum, Adhemarius dariensis, Adhemarius donysa, Adhemarius globifer, Adhemarius mexicanus, distribution, Mexico The taxonomy of the species that are closely related to Adhemarius donysa (Dru- ce) has been problematic. D’Abrera ([1987]), who illustrated A. globifer (Dyar) and the types of A. dariensis (Rothschild and Jordan) and A. donysa, suggested that all three of these taxa might be synony- mous. He also overlooked the most recently described species, A. blanchardorum (Hodges 1985). In contrast, Carcasson and Heppner (1996) treated A. globifer and A. donysa as separate species, with dariensis as a subspecies of the latter. Recently, Ca- diou and Hodges (1998) reviewed the spe- cies of Adhemarius from North and Central America, with special reference to the don- ysa group. They concluded that there were four species, and undertook a thorough re- view of the names and previous taxonomic confusion. For several years, we had rec- ognized four species in Mexico. The iden- tities of three were clear following Cadiou and Hodges’ paper: A. donysa, A. dariensis, and A. globifer (A. blanchardorum has not yet been found in Mexico). However, with regard to A. dariensis, Cadiou and Hodges (1998) wrote “‘Specimens from Veracruz and farther north tend to have more elon- gate forewings, with a narrower apex, than do specimens from Chiapas and _ farther south ... we have not found clear charac- ters to subdivide this taxon.’ We believe that these northern populations correspond to what we considered was the fourth spe- cies found in Mexico. After close exami- nation of the external morphology and gen- italia of a quite large series of specimens, we have concluded that it is indeed a hith- erto undescribed species. Adhemarius mexicanus Balcazar and Beutelspacher, new species (Figs. 1-2, 5—6, 9) Description.—Forewing length: ¢43-—51 mm (x = 47.8, n = 31); 249-52 &@ = 50.3, n = 4). Male (Fig. 1).—Head: Gray; labial palp ventrally orange, especially basal segments (distal one can be greenish); zone from base of antenna to vertex greenish brown. Tho- rax: Dorsally pale yellowish gray; dark ar- eas greenish brown. Legs with coxae and VOLUME 103, NUMBER 2 313 Figs. 1—4. “AN Habitus of Adhemarius species. 1-2, A. mexicanus. 1, Holotype d (forewing length: 48 mm). 2, Paratype 2 (forewing length: 61 mm), Mexico, Veracruz, Las Minas, 15 Oct 1972, C. Beutelspacher. 3, A. dariensis 3 (forewing length: 53 mm), Mexico, Chiapas, Rancho Nuevo, San Cristobal de las Casas, 19—22 Oct 1992, C. Beutelspacher. 4, Measurements used for the PCA analysis (AM = anal margin length; AM—CM = length from the concave area at the anal margin to the interception of the postmedial line with costal margin; AN = width of the apex notch; FWL = forewing length; OM = outer margin length). trochanters yellowish green; femora, tibiae, and tarsi greenish brown. Abdomen: Green- ish brown followed by pale yellowish gray then pale grayish green, with a faint medial line on segments 2—6. Forewing: Narrow and very elongate. Apex produced; very narrow; external margin almost straight (can be slightly wavy, but not crenulate); tornus very acute. Forewing dorsally gray green; extreme base yellowish gray; lunule dark greenish brown; median line clearly oblique; posterior margin just beyond me- 314 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 5-8. 7-8, A. dariensis. 7, Aedeagus. 8, Right valva (scale line = 1 mm). dial line greenish gray (with a lighter patch near costa); discocellular vein well outlined. Dark submarginal area both basally and dis- tally convex, with maximal width at M2, merging smoothly into wing pattern. Fore- wing ventrally pale yellowish green; anal angle with the same color as rest of wing; basal area red; discocellular vein clearly outlined. Hindwing: Base red becoming paler toward outer margin; dark mark in anal area dark greenish brown immediately preceded and followed by pale yellowish gray or green; median line slightlly oblique; antemarginal black pattern well marked and extending distad along veins, almost reach- ing posterior margin. Medial line curved basad when reaching costal margin. Male genitalia (Figs. 5—6): Valva elongate, not broadly rounded to apex; costal margin with a small lobe not followed by a sharp angle. Sacculus very poorly developed; lacking setae. Female (Fig. 2).—Similar to male, but larger. Forewing pattern less contrasting, and with a pinkish cast. Female genitalia (Fig. 9): Base of ductus bursae broad, about two-thirds width of ostium bursae. Male genitalia of Adhemarius species. 5—6. Adhemarius mexicanus. 5, Aedeagus. 6, Right valva. Signum narrowly arrow shaped; inwardly directed cones evenly graduated from short to very short. Types.—Holotype: 6; MEXICO: Oaxa- ca, Santiago Comaltepec, La Esperanza, Si- erra de Judrez, 17°37'45"N, 96°22'5’W, 1600, 28 Mar 1984 (coll. M. Garcia). Par- atypes: MEXICO: same locality as holo- type, 26 Mar 1976 (coll. ND)—d; 27 Mar 1976 (coll. ND)—<6; 30 Mar 1976 (coll. A. Diaz Francés)—¢; 22 Mar 1977 (coll. A. Diaz Francés)—2 3; 28 Mar 1984 (coll. A. Ibarra)—2 <; (coll. H. Delfin)—3 36; (coll. V. Hernandez)—2 ¢; 29 Mar 1984 (coll. A. Ibarra)—2 3; (coll. H. Delfin)—1 ¢; Pueb- la, Xicotepec, Dos Caminos, 20°14'35’N, 97°58'12”W, 29 Jun 1976 (coll. ND)—1 ¢; Villa Juarez, 19°52'0"N, 97°47'0"W, 3 Apr 1954 (coll. G. Pérez H.)—1 6; 1 May 1954 (coll. C. Marquez M.)—1 6; Jul-Aug 1960 (coll. ND)—2 6; Sep—Oct 1960 (coll. ND)—1 @, 1 6; 22 Oct 1960 (coll. ND)— 1 5; 26 Sep 1961 (coll. C. Marquez M.)— 1 6; Xicotepec; 20°16'32"N;)97°57 377W, 1150, 18 Aug 1978 (coll. R. Boue)—1 ¢; 20 Sep 1980 (coll. R. Boue)—1 ¢6 11 Oct 1980 (coll. R. Boue)—1 ¢; 4 Jul 1981 VOLUME 103, NUMBER 2 Fig. 9. Female genitalia of Adhemarius mexicanus (scale line = 1 mm). (coll. C. Beutelspacher B.)—1 6; 10 Apr 1985 (coll. C. Beutelspacher B.)—1 ¢. Other specimens studied.—MEXICO: Oaxaca, San Juan Lachao, Sierra Madre del Sur, +20 mi Norte de San Gabriel Mixte- pec loro 24"N: 97°7 36 W, 10° Jul 1991 (coll. ND)—1 ¢; Querétaro, Landa de Ma- tamoros, Km 8 carr. Agua Zarca -Neblinas, 21°15'14°N, 99°4’/58"W, 1150, 22 Jul 1998 (coll. Oliver & Ibarra)—1 3; Veracruz, Las Minas, Las Minas, 19°41'27"N, 97°8'48"W, Pas0rm, 28 Sep’ 1972 (coll? €. Beutel- spacher B.)—1 6; 15 Oct 1972 (coll. C. Beutelspacher B.)—1 92; Xalapa, Xalapa, 19°32'24"N, 96°55'39"W, 1,460 m, 14 Sep 1984 (coll. J. Pena M.)—1 a. Deposition of types.—The holotype and 25 paratypes are in the Coleccién Nacional de Insectos (CNIN), Universidad Nacional Aut6noma de México. One paratype male will be deposited in each of the following institutions: National Museum of Natural 315 History, Smithsonian Institution, Washing- ton, DC, The Natural History Museum, London, American Museum of Natural His- tory, New York, and Muséum national d’ Histoire naturelle, Paris. Etymology.—This new species is named after the country in which it is found. Distribution.—Fig. 10. Flight period.—A. mexicanus has been collected from March to October. KEY TO THE MALES OF THE ADHEMARIUS DONYSA GROUP Adult males of the Adhemairus donysa group of species can be identified using the following key. Additional characters to sep- arate A. mexicanus and A. dariensis are dis- cussed in the next section. The key can also be used for females, except for A. mexican- us and A. dariensis, as discussed below. 1 Antemarginal black pattern of hindwing upper side well marked and extending dis- tad along veins, almost reaching posterior margin; discocellular vein of forewing up- per side well outlined, forewing external margin almost straight; ............. 2 - Antemarginal black pattern of hindwing upper side not extending distad along veins; discocellular vein of forewing upper side not clearly outlined; forewing external margin crenulate near tornus Forewing broad; forewing apex little pro- duced; median line of forewing upper side little oblique; medial line of hindwing un- der side straight Adhemarius donysa - Forewing narrow; forewing apex produced; median line of forewing upper side clearly oblique; medial line of hindwing under side curved basad when reaching costal margin 2(1). Forewing external margin almost straight, apex very narrow (Fig. 1); dorsally usually gray green; forewing length usually less than 51 mm ~ Forewing external margin outwardly con- vex, apex not very narrow (Fig. 3): dorsally usually gray brown; forewing length usu- ally 52 mm or more .. Adhemarius dariensis Forewing dorsally gray green; dark sub- marginal area of forewing upper side gen- erally narrower, indented distally at M2 and at M3; dark submarginal area of forewing upper side sharply delimited basally Adhemarius blanchardorum Are ae Scie Adhemarius mexicanus 4(1). 316 O 200 400 600 Kilometers =—— —— Fig. 10. - Forewing dorsally gray brown; dark sub- marginal area of forewing upper side glob- ally convex, with maximal width at M2; dark submarginal area of forewing upper side merging smoothly into wing pattern OE Bros Lara te Adhemarius globifer Discussion.—Several characters can be used to separate males of Adhemarius mex- icanus from those of A. dariensis. Adhe- mairus mexicanus is a smaller (in most specimens, the forewing is less than 50 mm), a more slender species, with more elongate wings, an almost straight forewing external margin, a very narrow apex and a very acute tornus (Fig. 1). In contrast, A. dariensis is a larger (forewing is usually more than 55 mm), a more robust species, with broader forewings and apex, an out- wardly convex external margin, and a more rounded tornus (Fig. 3). In A. mexicanus, the forewing pattern is more contrasting, and the ground color is greenish in fresh PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Known distribution of Adhemarius mexicanus (dots) and A. dariensis (squares) in Mexico. specimens, turning yellowish in old ones; while in A. dariensis, the ground color is brownish turning orange in old specimens, with a less contrasting wing pattern. Adhe- mairus mexicanus has a costal light gray patch next to the distal margin of the post- medial line that makes the zigzag postme- dial lines very clear; this patch is missing in A. dariensis. The hindwings are usually uniformly red in A. dariensis, whereas in most specimens of A. mexicanus they turn yellowish toward the outer margin. The val- va of A. dariensis is more rounded, and the sacculus, although poorly developed, is bet- ter developed (Fig. 8) than in A. mexicanus, which in addition has a less rounded valva. The aedeagus is long (longer than the bul- bus ejaculatorius) in A. dariensis (Fig. 7) and short (subequal than the bulbus ejacu- latorius) in A. mexicanus (Fig. 5) (1. Kitch- ing, personal communication, uses the term VOLUME 103, NUMBER 2 Fig. 11. Sil7/ 11 Principal component analysis of 30 specimens of Adhemarius mexicanus and 30 specimens of A. dariensis. Three-dimensional graph with the projection of the specimens onto the first three components. This representation expresses 96% of the variation of five characters studied. cuticular simplex instead of bulbus ejacu- latorius, but both terms are obscure). On the basis of a limited number of fe- males, we suggest the following diagnostic characters. The same characters of forewing apex, tornus, red hindwing color, and ro- Table 1. Summary of the data used in the PCA analysis; 30 male specimens of each species were mea- sured. FWL = forewing length; OM = forewing outer margin length; AM-CM = forewing length from the concave area at the anal margin to the interception of the postmedial line with costal margin; AM = anal margin length; AN = width of the apex notch. A. dariensis A. mexicanus FWL ag Syl 47.8 Min 42 43 Max 3)// 3)1I OM xX 30.2 27 Min 26 24 Max 3)5) 30 AM-CM x IS),3) L/L Min 7 15 Max 22 19 AM x 28.6 27.4 Min 25 25 Max 33 30 AN xX 3.4 Pd Min 3 A) Max 4d 2 bustness that separate the males also serve to separate the females. In A. dariensis, the forewing is divided into a light basal area and a darker distal area, separated by the medial line; in A. mexicanus, the basal and distal areas are less contrasting, and have a pinkish cast. No diagnostic characters were identified in the female genitalia. To analyze the apparent overlap in the external appearance of some specimens of both species, we conducted a _ principal component analysis (PCA). Five measure- ments were used: forewing length (FWL), forewing outer margin (termen) length (OM), forewing length from the concave area at the anal margin to the interception of the postmedial line with costal margin (AM-CM), anal margin length (AM), and width of the apex notch (AN) (Fig. 4). We measured 30 males from Mexico north of the Isthmus of Tehuantepec and 30 males from Chiapas and Costa Rica. The data are summarized in Table | (The data are avail- able on request from the senior author). All measurements were standardized and ana- lyzed using NTSysPc version 2.02c (Rohlf 1989). The first three components extracted account for 96% of the variation observed 318 Table 2. Eigenvectors, corresponding eigenvalues, and percentage of variation explained for 30 male Ad- hemarius mexicanus and 30 A. dariensis studied. Percent Cumulative Eigenvalue 1 3.97187564 79.4375 79.4375 DZ 0.57496201 11.4992 90.9368 3 0.23571324 4.7143 95.6510 4 0.12598810 2.5198 98.1708 5 0.09146101 1.8292 100.0000 (Table 2). The loadings of each variable are shown in Table 3. There are two distinct clouds of specimens on the three-dimen- sional graph (Fig. 11), a very compact group comprising the specimens from North of the Isthmus of Tehuantepec, (A. mexicanus) and a more dispersed group comprising the remaining specimens (A. dariensis). Although the two clouds are very close, there is no overlap. Based on the few character measured, the external differences between A. mexicanus and A. dariensis seem to be largely a matter of scale, with A mexicanus being a smaller species than A. dariensis. The same pattern appears to be true for the females, but we did not have a sufficiently large number for an analysis. We expect that knowledge of immature stages will help clarify status of taxa. This new species seems to be closely as- sociated with cloud forest (“bosque mes6ofi- lo’) sensu Rzedowski (1978) in the states of Veracruz, Hidalgo, Querétaro, Puebla, and Oaxaca, and at altitudes between 1,150 and 1,600 m a.s.l. (Fig. 10). This cloud forest is well known for its relictual and endemic taxa (Llorente & Escalante 1992). We have one specimen from the Sierra Madre del Sur province on the Pacific slope in Oaxaca, again from an area with cloud forest. How- ever, more specimens are needed from that area to confirm its presence there. The few specimens of A. dariensis that we have seen from Mexico were all collected at altitudes between 1,900 and 2,400 m, and never at lower altitudes, in the state of Chiapas. Fur- thermore, the specimens illustrated as A. dar- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 3. Correlations (loadings) between principal components and the five variables analyzed. I 2 3 FWL 0.9348 0.1786 0.1134 OM 0.9615 —0.0747 0.0323 AM-CM 0.9286 —0.0207 0.2981 AM 0.8344 0.4516 —0.2967 AN 0.7843 SV —0.2120 iensis by Cadiou and Hodges (1998: figs. 3— 4) are actually A. mexicanus ACKNOWLEDGMENTS We thank Jorge Corrales (INBio, Costa Rica) for providing information on Costa Rican specimens, and Adolfo Ibarra for his assistance at the National Collection of In- sects (CNIN, Mexico). We thank Ian Kitch- ing (Department of Entomology, The Nat- ural History Museum, London), and Ronald W. Hodges for reviewing the manuscript. This study was possible in part thanks to CONABIO grant FB269/H021/96, and DGAPA, UNAM grant IN201497. LITERATURE CITED Cadiou, J. M. and R. W. Hodges. 1998. Adhemarius donysa (Druce): Identification and notes on close- ly related species (Lepidoptera: Sphingidae). Pro- ceedings of the Entomological Society of Wash- ington 100(2): 202-208. Carcasson, R. H. and J. B. Heppner. 1996. Sphingo- idea. In Heppner, J. B., ed., Atlas of Neotropical Lepidoptera 5B, Checklist (part 4B): i-l + 1-87. d’Abrera, B. [1987]1986. Sphingidae Mundi—Hawk moths of the world. E. W. Classey, Faringdon, x + 226 pp. Hodges, R. W. 1985. A new species of Amplypterus from the Chisos Mountains, Texas (Lepidoptera: Sphingidae). Proceedings of the Entomological Society of Washington 87(2): 323-328. Llorente, J. E. and P. Escalante 1992. Insular Bioge- ography of Submontane Humid Forests in Mexi- co. In Darwin, S.P. and A. Welden, eds., Bioge- ography of Mesoamerica. Supplementary Publi- cation. Tulane Studies in Zoology and Botany 1: 139-146. Rohlf, E J. 1989. NTSYS-pc. Numerical Taxonomy and Multivariate Analysis System. Exeter Publish- ing, LTD, Setauker, New York. (Software) Rzedowski, J. 1978. La Vegetacion de México. Edi- torial Limusa, Mexico, 432 pp. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 319-324 TWO NEW SPECIES OF ZONITIS F. (COLEOPTERA: MELOIDAE) FROM SOUTHWESTERN NORTH AMERICA, WITH COMMENTS ON GENERIC DEFINITIONS IN THE NEMOGNATHINAE JOHN D. PINTO Department of Entomology, University of California, Riverside, CA 92521, U.S.A. (e-mail: john.pinto @ucr.edu) Abstract.—Two new species of Nemognathinae, Zonitis stevewardi and Zonitis min- utissima, are described from southwestern North America. These species do not easily fit current generic definitions, and they tentatively are placed in Zonitis (Neozonitis). Prob- lems associated with current generic concepts in the Nemognathinae are discussed. Key Words: The meloid fauna of the United States is reasonably well known and it no longer is common to encounter taxa not previously recognized at some level. The two closely related species described here both occur in southern Arizona, a popular area for collec- tors, but they apparently have been missed by most because of small size and low numbers. The few specimens in collections were mixed with Gnathium Kirby based on general appearance and size. Although I am confident these species do not belong to Gnathium as currently defined, placing them to genus is not straightforward. The genera of Nemognathinae are impossible to define objectively and are in need of thor- ough revision. Definitions proposed by MacSwain (1951) and followed by Enns (1956) in his revision of several genera of U.S. nemognathines do not entirely accom- modate known diversity. The new species, a case in point, do not clearly fit into any described genus, yet because they are char- acterized by several features assumed to be primitive, a new taxon is not justified. For this paper they are tentatively assigned to Zonitis (Neozonitis) Enns. The justification for this assignment, as well as a brief dis- Meloidae, Nemognathinae, blister beetles, Zonitis cussion of nemognathine generic concepts, follows the descriptions. Both new species can be separated from all known Nemognathinae by their size (body length < 5 mm) which places them among the smallest of Meloidae, the max- illary galeae which are not produced into a sucking tube (Figs. 4—5), and the fringe of setae on the ventral blade of the tarsal claws (Figs. 6—7). The latter trait, known in all Eleticinae (Pinto and Bologna 1999) and some Old World Meloinae (personal obser- vation) has not been previously reported in the Nemognathinae. Zonitis (Neozonitis) stevewardi Pinto, new species (Figs. 2—4, 6, 8) Description.—Body small, relatively slender, surface shiny; length (with head in hypognathous position) = 3.9—4.2 mm. Color primarily fulvous with tibiae, tarsi, antennal segments III—XI, maxillary palpus and mandible darker brown; elytra each with a single longitudinal dark brown vitta (Figs. 2—3), vitta relatively broad, broadest at base and particularly at apex where it oc- cupies entire elytron width, narrowing to 320 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-3. 1, Zonitis minutissima, holotype 2 (23.8). 2—3, Z. stevewardi elytra showing variation in width of vitta (2, from 1 mi. E. Douglas, AZ, paratype 2 [18.1]; 3, from 9 mi. SE Picacho, AZ [17.6X]). slightly less than half width of elytron at middle (but see Variation); eyes black. Pu- bescence dark, relatively sparse; elytral se- tae longest, reclinate, ca. 0.1 mm in length, separated from others by a distance sube- qual to their length; head and pronotum with setae shorter, lighter and sparser; ven- ter with light, sparse, inconspicuous setae. Antenna ca. half as long as body, relatively loosely articulated, with dense procumbent setation and a few short distinctly erect se- tae; segments elongate, subfiliform, apical segments only slightly wider, with relative lengths of I-XI in an exemplar female as follows: 20:17:24:20:20:19:20:19:19:19:28, proportions similar in male. Head capsule (Fig. 4) 0.74 (0.71—0.79) as long as wide, widest at eyes; interocular distance 0.66 (0.64—0.68) greatest head width; surface with relatively small, moderately dense, in- conspicuous punctures, area between dor- sum of eyes largely impunctate. Labrum reaching apex of mandibles. Maxillary ga- leae penicillate, extending well beyond mandibles but not joined to form a sucking tube (Fig. 4). Eyes suboval, slightly emar- ginate behind antennal fossa, relatively small and not extending beyond mandibles on underside of head. Pronotum slightly longer than wide (length to width ratio = 1.10 [1.07—1.14]); widest at apical %, sides very slightly convergent to base and mod- erately convergent to apex; surface shiny, sparsely, shallowly punctate. Elytra smooth, shiny, shallowly and inconspicuously punc- tate. Tarsal claws (Fig. 6) with 7 elongate teeth in inner row; ventral blade of claws distinctly fringed. Fore- and mid-tibial spurs spiniform; hind tibial spurs (Fig. 8) moderately spathulate, similar in shape, uniformly dark reddish brown in color. Ae- deagus of male genitalia with two small sclerotized lobes associated with median tube. Etymology.—I take pleasure in naming this species after Steve Ward, of Port An- geles, Washington, in recognition of his help collecting Meloidae over the past 35 VOLUME 103, NUMBER 2 32) Figs. 4—S. years and his enjoyable company on several field trips in southern California and Baja California. Types.—Holotype ¢. UNITED STATES. Arizona: Douglas, 1 mi. E (Cochise Co.); vuli-31-1971; on Tidestromia lanuginosa (Nutt.); John D. Pinto. Allotype 2? and two paratypes (1 6, 1 2); same data as holo- type. Holotype and allotype deposited in the California Academy of Sciences, San Fran- cisco (CAS). Paratypes reside in the De- partment of Entomology Research Muse- um, University of California, Riverside (UCRC). Additional material examined.—UNIT- ED STATES. Arizona: McNeal, 8 mi. S (Cochise Co.); vili-10-1976; sweeping 7i- destromia lanuginosa; 1 2; John D. Pinto. Picacho, 9 mi. SE (Pinal Co.); viii-25-1976; on Tidestromia lanuginosa; 1 3; John D. Pinto. Variation.—The elytral vitta varies in width. In most specimens it is widest api- cally and basally and narrows only slightly in between where it occupies ca. half the elytral width (Fig. 2). In the specimen from Head capsule of Zonitis spp. 4, Z. stevewardi. 5, Z. minutissima. Arrow indicates maxillary galea. SE of Picacho, however, the vitta is consid- erably narrower. It is of normal width api- cally but narrows anteriorly to less than ¥, the width of the elytron, and is not widened at the elytral base (Fig. 3). Diagnosis.—The only species Z. steve- wardi can be confused with is Z. minutis- sima, described below. Their separation is treated in the description of the latter. Remarks.—All collections of this species are from Tidestromia lanuginosa (Amar- anthaceae). However, feeding has never been observed; the few specimens taken were collected by lightly sweeping the plant with an aerial net. It is not clear if Z. stev- ewardi is extremely uncommon or simply difficult to collect. Several attempts to find additional specimens were unsuccessful. 77- destromia lanuginosa is relatively common in the Southwest but because of its prostrate growth form it is a difficult plant to sample. Zonitis minutissima Pinto, new species (Piss: 1,5... 7) Description.—Differing from Z. steve- wardi as follows: Body length perhaps av- 322 Figs. 6-8. itis spp. 6, Hind tarsal claw, Z. stevewardi (inner view of posterior claw with ventral blade below). 7, Hind tarsal claw, Z. minutissima (as in 6). 8, Hind tibial spurs, Z. stevewardi. Scanning electron micrographs of Zon- eraging smaller, length (with head in hy- pognathous position) = 3.0—4.1 mm. Color darker with legs and venter of metathorax brown, head brown except fulvous at frons; elytral vitta (Fig. 1) considerably broader, encompassing most of disk (0.6—0.7 elytral width at middle) except for a fulvous mar- gin along suture extending to apical 7,, of elytra and a narrow lateral border extending to apical 4%, extreme apex of elytra entirely dark brown. Pubescence denser, that on el- ytra of similar length but setae separated PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON from one another by a distance equal to ca. Y%—% their length; light colored pubescence on venter moderately dense, conspicuous. Head capsule (Fig. 5) wider, 0.64 (0.61— 0.69) as long as wide; galea penicillate but considerably shorter, extending only to apex of mandibles (Fig. 5). Eyes slightly more bulged. Pronotum wider than long (length to width ratio = 0.93 [0.90—0.95]). Elytral surface distinctly rugulopunctate, dull. Tar- sal claws (Fig. 7) with 7 elongate and 1 very short teeth in inner row, teeth slightly longer. Aedeagus of male genitalia with ventral lobes associated with median tube somewhat larger, pointed apically. Etymology.—The specific name refers to the extremely small size of this species. Types.—Holotype 2. UNITED STATES. Arizona: Picacho Pass (Pinal Co.); vili-7- 1940; on Boerhaavia (Nyctaginaceae); P. Timberlake; deposited in CAS. Paratype °, as above except collected on different plant (label illegible) (UCRC). Additional material examined.—MEXI- CO. Baja California Sur: Loreto, 33.8 mi. NNW; at light; 1 ¢6; E Andrews and D. Faulkner; in the collection of the California Department of Food and Agriculture, Sac- ramento. Diagnosis.—Zonitis minutissima 1s most similar to Z. stevewardi. The two are easily separated by the structure of the galea, the dimensions of the head and pronotum, and color. In Z. minutissima the galeae of the maxillae are short and extend only to near the apex of the mandibles; in Z. stevewardi the galeae are distinctly penicillate and elongate, extending well beyond the man- dibles (cf. Figs. 4-5). In Z. minutissima both the head and pronotum are broader than in Z. stevewardi (Figs. 4—5; also see quantitative data above). Finally, color pat- tern, particularly of the elytra, differs in the two species. Zonitis minutissima is darker than Z. stevewardi and its elytral vitta is considerably broader (cf. Figs. 1—3). Both species can be separated from all other Neo- zonitis by their minute size (body length < 5 mm), and the fringed ventral blade of the VOLUME 103, NUMBER 2 tarsal claws (Figs. 6-7). The only species of Neozonitis with a similar color pattern is Z. bilineata Say. In addition to size and claw structure, Z. bilineata is separated by its moderately inflated tempora, a typical trait of most North American Zonitis Fa- bricius, which represent the widest aspect of the head. In the new species, the head is widest at the eyes and the tempora are not inflated. Because of size and general shape, Z. stevewardi and Z. minutissima are most likely to be confused with Gnathium. Un- like Gnathium, however, the galeae are not prolonged into a sucking tube, and the la- brum reaches the apex of the mandibles rather than only approaching their mid- point. Also, the ventral blade of the tarsal claws in Gnathium lack fringe setae, a fea- ture absent in all other nemognathines as well. Although, as in Gnathium, the anten- nae are slightly widened apically, they are less distinctly so. Also, the entire antenna is longer in the new species and the seg- ments are more loosely articulated. The two new species cannot be easily identified to genus using the recent key to New World meloid genera (Pinto and Bo- logna 1999). This is due to the presence of the fringed ventral blade of the tarsal claws. Until discovered in these nemognathines, the only New World blister beetles known with this feature were genera of the primi- tive subfamily Eleticinae, a group not found in North America. This character, along with others distinctive of eleticines, is used in the first couplet of the generic key. Con- sequently, the new species fit neither choice offered by couplet 1. However, the combi- nation of the fringed ventral blade and combed claws, a characteristic of virtually all Nemognathinae, immediately identifies them. Remarks.—Zonitis minutissima appears to be sympatric with Z. stevewardi. Both have been collected in August at or near Picacho Pass in Pinal Co. Arizona, although in different years. The females of Z. minu- 323 tissima from Arizona are very similar to the single male collected in Baja California Sur. DISCUSSION As indicated earlier, the generic assign- ment of Z. stevewardi and Z. minutissima is questionable. MacSwain (1951) placed Zonitis and Gnathium in the Zonitini and separated them from Nemognatha Illiger, Tricrania LeConte and Hornia Riley (Nem- ognathini) by the structure of the aedeagus. Rhyphonemognatha Enns was later added to the Nemognathini by Enns (1956). The Zonitini were characterized by the bilobed, and heavily sclerotized median tube. The Nemognathini included genera with a mem- branous median tube. Unfortunately the dis- tinction drawn by MacSwain is not clear- cut and in certain species of Nemognatha a relatively small but distinctly sclerotized bi- lobed structure also occurs. For example, this is the case in species which Enns (1956) treated as Nemognatha (Pronemog- natha) Enns. The two new species also fit this category. The bilobed structures are not as well developed as in most North Amer- ican Zonitis but they are present. The other character which leads me to place them in Zonitis is hind tibial spur structure. In these species, as in most North American Zonitis, the spurs are similar in shape, spathulate and completely dark reddish brown in color (Fig. 8). The hind tibial spurs of most other nemognathine genera in North America are structured differently (see Enns 1956). The one obvious exception is Pronemognatha, which as indicated by aedeagal structure as well, may be inappropriately placed in Nemognatha. The genera of Nemognathi- nae require considerable study. The generic definitions by MacSwain (1951) and Enns (1956) were based on a limited sampling of North American species and no attempt has been made to apply them to all of our spe- cies or to the considerable nemognathine fauna occurring elsewhere in the world. Within Zonitis, the new species are placed in the subgenus Neozonitis. Most of their features are consistent with this as- 324 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON signment. Neozonitis is separated from Par- azonitis Enns, the only other subgenus rec- ognized in North America, by galea struc- ture. In Neozonitis the galeae are scarcely modified and not produced into a sucking tube as they are in Parazonitis (Enns 1956). Considering that this clearly is a primitive character, the monophyly of Neozonitis has yet to be demonstrated. Also, its relation- ship to Old World Zonitis, most of which similarly lack a sucking tube, requires clar- ification. ACKNOWLEDGMENTS This study was supported in part by a grant from the National Science Foundation (DEB-9508735). Figures 4 and 5 were pre- pared by Marina Planoutene; Gary Platner was primarily responsible for the prepara- tion of all others. LITERATURE CITED Enns, W. R. 1956. A revision of the genera Nemog- natha, Zonitis and Pseudozonitis (Coleoptera: Me- loidae) in America North of Mexico, with a pro- posed new genus. University of Kansas Science Bulletin 37: 685—909. MacSwain, J. W. 1951. New North American species of Nemognatha and Zonitis. The Pan-Pacific En- tomologist 27: 72-80. Pinto, J. D. and M. A. Bologna. 1999. The New World genera of Meloidae (Coleoptera): A key and syn- opsis. Journal of Natural History 33: 569-620. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 325-333 STUDY ON CHINESE MELANAPHIS VAN DER GOOT (HOMOPTERA: APHIDIDAE) WITH DESCRIPTIONS OF THREE NEW SPECIES LIKUN ZHANG, GEXIA QIAO AND GUANGXUE ZHANG Institute of Zoology, Chinese Academy of Sciences, 19 Zhongguancun Lu, Haidian, Beijing 100080, People’s Republic of China Abstract.—The genus Melanaphis van der Goot 1917 includes 10 species in China. Three new species are described, Melanaphis arthraxonophaga, M. grossisiphonellus, and M. zhanhuaensis. Melanaphis siphonella is a new record for China. A key to the ten species is given. Key Words: Melanaphis is a genus of Rhopalosiphi- na, Aphidinae, and it was described in 1917 by van der Goot. Twenty-one species of Melanaphis have been described. This ge- nus is associated with Gramineae and Ro- saceae, and only a few species have proven host alternation (Blackman and Eastop 1994). The species of this genus were re- ported from Europe (Barbagallo and Stroy- an 1982, Stroyan 1984, Heie 1986), Africa, Australia, U.S.A., Uruguay, and Argentina (Blackman and Eastop 2000), and Asia. So- rin (1970) reviewed the Japanese species, Raychaudhuri and Banerjee (1974) reported the species in India, Pashchenko (1988) gave a key to the species of the former So- viet Far East, and Hodjat (1998) listed the species in Iran. Blackman and Eastop (1994, 2000) reviewed this genus on the world’s crops and trees. The reports from China (Zhang and Zhong 1983; Tao 1990; Zhang et al. 1992, 1999) came from several provinces; thus, a review of this genus in China is needed. MATERIALS AND METHODS Structural terminology follows Zhang and Zhong (1983). The unit of measure- ment is in millimeters (mm). Aphididae, Melanaphis, new species, China The studied specimens are deposited in the Institute of Zoology, Chinese Academy of Sciences, Beijing. RESULTS Melanaphis van der Goot 1917 Melanaphis van der Goot 1917:61. Type species: Aphis bambusae Fullaway 1910. Synonyms: Longiunguis van der Goot 1917, Yezabura Matsumura 1917, Yezaphis Matsumura 1917, Geoktapia Mordvilko 1921, Pyraphis Borner 1931, Piraphis Bor- ner 1932, Nevskia Mordvilko 1932, Mas- raphis Soliman 1938, Schizaphidiella Hille Ris Lambers 1939 (Remaudiére and Re- maudiére 1997). Melanaphis differs from the other genera of Rhopalosiphina by the following char- acters: Siphunculus short, cylindrical or subcylindrical and alata frequently with dorsal sclerites or cross bars on abdominal tergites in front of the siphunculi. Seven species have been reported in China. In our study, we did not find M. pyraria (Passer- ini) in China. The specimens collected from Pyrus sp. and described by Zhang et al. (1999) as M. pyraria are not the same as Passerini’s (1861) description, such as the sclerotic dorsal patch on the abdomen, the 326 length of the siphunculus, and the shape of the cauda. The appearance of the aphid dif- fers remarkably on its secondary host plants according to the species and condition of the aphid grass colonized (Blackman and Eastop 2000), but there is no remarkable difference on the same host. Therefore, this species is not included in this paper. Three new species and one new record to China were found. KEY TO CHINESE SPECIES OF MELANAPHIS 1. Dorsal cuticle with reticulation formed by rows OSS PINES ey. ene eae M. zhanhuaensis, n. sp. — Dorsal cuticle without reticulation formed by RONWAS Ol GowMNlas: dress Se Wise ola 5 Hd co stain 2, . Abdomen with 3-9 pairs of marginal hairs on segments I-III, their length 2.0 times as long as or longer than basal diameter of 3rd antennal SCPIMEME Gye titra cert, es ite eee oe Oe ene ate 3 — Abdomen with 1-2 pairs of marginal hairs on segments I-III, their length at most 1.5 times as long as or longer than basal diameter of 3rd antennal segment 3. Genital plate with 2 hairs on its anterior part; abdomen with 5—6 dorsal hairs on segments I- JOU ots chain ot icine M. grossisiphonellus, n. sp. — Genital plate with 5—10 hairs on its anterior part; abdomen with at least 15 dorsal hairs on segments I-III 4. Abdomen with 7—9 pairs of marginal hairs on anterior segments, with 20—38 dorsal hairs on segments I-III, cauda with 8-11 hairs ........ Ry doe rst ge ehovntwctet wank hs Oe Deke ae M. pyrisucta — Abdomen with 3-5 pairs of marginal hairs on anterior segments, with 15—22 dorsal hairs on segments I-III, cauda with 6 hairs ao MARIA EMA RM wee es Meo dy oh M. arundinariae 5. Cauda with 4—6 hairs; siphunculus as long as or longer than cauda (with dark coxae)....... ESR NOTOR OT haley See ome cacao, oo oon M. bambusae — Cauda with at least 8 hairs; siphunculus shorter than cauda™ fir. a. ston ee eee ea eee es 6 6. Abdomen with 4-8 hairs on tergite VIII... Se ofa heoep eta « Moe ese s Breas, + M. graminisucta i) Abdomen with 2 hairs on tergite VII ...... 7 7. Longest hairs on hind tibia as long as its mid- GEICO TAIMELE I ras cae rae ee aie ee ence atc 8 — Longest hairs on hind tibia shorter than its its MINA GLE diameter a. eeens meee eee, ae 9 8. Processus terminalis about 3.7 times as long as base of ultimate antennal segment; ultimate rostral segment shorter than 2nd hind tarsal segment; genital plate with 4 hairs on its an- IOP (ORM 5 oo See oe M. arthraxonophaga, n. sp. — Processus terminalis at most 2.5 times as long PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON as base of ultimate antennal segment; ultimate rostral segment as long as 2nd hind tarsal seg- ment; genital plate with at least 7 hairs on its aNntenlOn Part. 4... c= 5.5 seen M. siphonella 9. Dorsal hairs of head as long as or a little longer than basal diameter of 3rd antennal segment; ultimate rostral segment 0.8 times as long as 2nd hind tarsal segment; genital plate with 6— 13 hairs on its anterior part ....... M. sacchari — Dorsal hairs of head about 0.5 times as long as basal diameter of 3rd antennal segment; ulti- mate rostral segment almost as long as 2nd hind tarsal segment; genital plate with 2-3 hairs on its anterior part M. formosana Melanaphis arthraxonophaga Zhang, Qiao, and Zhang, new species (Figs. 1-5, 14) Apterous viviparous female.—Body 1.088 in length, 0.675 in width. Antenna 0.925, segments I—V: 0.057, 0.051, 0.270, 0.137, 0.086 + 0.324. Ultimate rostral seg- ment 0.062. Hind femur 0.283, hind tibia 0.484, 2nd hind tarsal segment 0.072. Si- phunculus 0.103 in length, 0.118 in basal diameter, 0.046 in tip diameter. Cauda 0.157 in length. Body elliptical. In cleared specimens, head, rostrum, antennal segments I—II and V, basal and apical parts of tibia, tarsi, si- phunculi, cauda, anal plate and genital plate brown; other parts pale. Spiracles circular and open, spiracular plates elliptical, brown. Mesosternal furca without stem. Marginal tubercles on prothorax and ab- dominal tergites I and VII. Vertex slightly convex, antennal tubercles undeveloped. Antennal segments IIJ—VI and 2nd tarsal segments transversely imbricate. Head with 1 pair of frontal hairs, 1 pair of lateral fron- tal hairs and 3 pairs of dorsal hairs. Two dorsal hairs on abdominal tergite VIII. Length of frontal hairs, marginal hairs on abdominal tergite I, and dorsal hairs on ab- dominal tergite VIII 0.60, 0.67 and 1.20 times as long as basal diameter of antennal segment III, respectively. Antenna 5-seg- mented, without secondary rhinaria. Pro- cessus terminalis 3.71 times as long as base of antennal segment V. Primary rhinaria round, ciliated. Rostrum reaching middle VOLUME 103, NUMBER 2 B27] Tei etka ee Figs 1-9. 1-5, Melanaphis arthraxonophaga. 1—4, Apterous viviparous female. 1, Antenna. 2, Ultimate rostral segment. 3, Siphunculus. 4, Cauda. 5, Alate viviparous female, segment III of antenna. 6-9, M. grossi- siphonellus. 6-8, Apterous viviparous female. 6, Ultimate rostral segment. 7, Siphunculus. 8, Cauda. 9, Alate viviparous female, antenna. 328 coxae; ultimate rostral segment short, in length 0.92 times as long as its basal di- ameter, 0.86 times as long as 2nd hind tarsal segment, with | pair of accessory hairs and 3 pairs of primary hairs. Hind femur 1.05 times as long as antennal segment III, hind tibia 0.44 times as long as body. Hairs on hind tibia 0.041, almost as long as the mid- dle diameter of tibia. First tarsal segment chaetotaxy: 3, 3, 2. Siphunculus short, im- bricated, without flange, 0.87 times as long as its basal diameter, 2.22 times as long as its tip diameter, 0.63 times as long as cauda. Cauda distinctly constricted in the middle, 1.66 times as long as its basal diameter, with 16—20 hairs. Anal plate broad, circular, with 21—24 hairs. Genital plate broadly egg- shaped, bearing 13 hairs, among them 4 hairs on its anterior part. Alate viviparous female.—Body 1.238 in length, 0.642 in width. Antenna 5 or 6-seg- mented, 1.143 (or 1.051), 5-segmented, segments I-V: 0.072, 0.052, 0.371, 0.165, 0.103 + 0.381; 6-segmented, segments I— VI: 0.062, 0.052, 0.185, 0.155, 0.160, 0.093 + 0.345. Ultimate rostral segment 0.069. Hind femur 0.326, hind tibia 0.615, 2nd hind tarsal segment 0.076. Siphunculus 0.086 in length, 0.062 in basal diameter, 0.039 in tip diameter. Cauda 0.129 in length. Body elliptical. In cleared specimens, head, rostrum, antenna, legs (except middle part of tibiae), and siphunculus dark brown; cauda, anal plate and genital plate brown; others pale. Spiracles circular and open, spiracular plates elliptical, brown. Abdo- men with dark brown marginal sclerites on tergites I-VI, middle sclerites on tergites III-V, and cross bars on tergites VI—VIII. Two dorsal hairs on abdominal tergite VIII. Vertex flat and straight, antennal tubercles undeveloped. Head with 1 pair of frontal hairs, 1 pair of lateral frontal hairs and 4 pairs of dorsal hairs. Length of frontal hairs, marginal hairs on abdominal tergite I, and dorsal hairs on abdominal tergite VIII 0.67, 1.00 and 1.00 times as long as basal di- ameter of antennal segment III, respective- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ly. Antenna 5 or 6-segmented. Antennal segment III with 6—9 secondary rhinaria, which are round and convex. Processus ter- minalis about 3.75 times as long as base of ultimate antennal segment. Primary rhinaria round; ciliated. Rostrum reaching middle coxae; ultimate rostral segment short, as long as its basal diameter, 0.93 times as long as 2nd hind tarsal segment, with 1 pair of accessory hairs and 3 pairs of primary hairs. Hind tibia 0.50 times as long as body. Siphunculus short, imbricated, without flange, 1.42 times as long as its basal di- ameter, 0.68 times as long as cauda. Cauda distinctly constricted in middle, 1.32 times as long as its basal diameter, with 13 hairs. Anal plate bearing 19-21 hairs. Genital plate bearing 16 hairs, among them 5 hairs on its anterior part. Other characters as for apterous viviparous female. Discussion.—This new species is closely related to M. montana Sorin, but differs from M. montana by the following char- acters: 1) Hairs on hind tibia almost as long as the middle diameter of the tibia [the lat- ter (Sorin 1970): 2.00 times]; 2) dorsal hairs on abdominal tergite VIII 1.20 times as long as basal diameter of antennal segment III (the latter: 3.00 times); 3) cauda bearing 16-20 hairs (the latter: 10—11 hairs); and 4) abdomen without marginal tubercles on ter- gites I-IV (the latter: present). Types.—Holotype: apterous viviparous female, No. Y2898-1-1.4, July 30, 1980, Liaoning Province (Tieling City, 123.8°E, 42.3°N), on Arthraxon hispidus (Thumb.), collected by Liu Lijuan. Paratypes: | apter- ous viviparous female, 3 alate viviparous females, No. Y2898, data same as holotype. Melanaphis arundinariae (Takahashi 1937) Aphis arundinariae Takahashi 1937: 12. Host plant.—Sinarundinaria sp. Distribution in China.—Sichuan and Taiwan. Melanaphis bambusae (Fullaway 1910) Aphis bambusae Fullaway 1910: 35. VOLUME 103, NUMBER 2 Synonyms: Yezabura sasae Matsumura 1917, Yezabura sasicolla Matsumura 1917, Yezabura? photiniae Matsumura 1918, Masraphis phyllostachia Soliman 1938 (Remaudiére and Remaudiére 1997). Host plant.—Bambusa sp. Distribution in China.—Hunan, Zhe- jiang, Sichuan, Yunnan, Guangdong and Taiwan. Melanaphis formosana (Takahashi 1921) Aphis formosana Takahashi 1921: 54. Synonym: Aphis miscanthi Takahashi 1921 (Eastop and Hille Ris Lamberrs 1976, Re- maudiére and Remaudiére 1997). Host plants.—Echinochloa sp., Oryza sp., and Panicum sp., Distribution in China.—Hebei and Tai- wan. Melanaphis graminisucta Zhang 1992 Melanaphis graminisucta Zhang 1992: 150. Host plant.—Species of Gramineae. Distribution in China.—Hunan. Melanaphis grossisiphonellus Zhang, Qiao, and Zhang, new species (Figs. 6—9, 15) Apterous viviparous female.—Body 1.408 in length, 0.947 in width. Antenna 0.947, segments I-VI: 0.045, 0.049, 0.222, 0.129, 0.135, 0.093 + 0.274. Ultimate ros- tral segment 0.076. Hind femur 0.381, hind tibia 0.635, 2nd hind tarsal segment 0.083. Siphunculus 0.088 in length, 0.076 in basal diameter, 0.055 in tip diameter. Cauda 0.108 in length. Body elliptical, dark green and with white wax. In cleared specimens, head dark, prothorax with a dark cross bar and a pair of marginal sclerites; mesothorax and meta- thorax with a pair of dark marginal scler- ites, respectively; rostrum, legs (except the middle parts of tibiae), siphunculus, cauda, anal plate, and genital plate dark brown; an- tennal segments I—II and VI brown; others pale. Spiracles circular and open, spiracular plates elliptical, dark. Intersegmental mus- 329 cle sclerites light brown. Mesosternal furca brown, without stem. Marginal tubercles on prothorax and abdominal tergites 1 and VII, dark and very small, as large as hair-bearing tubercles. Vertex slight convex, antennal tu- bercles slightly developed. Head with 1 pair of frontal hairs, 1 pair of lateral frontal hairs, and 5 pairs of dorsal hairs. Prothorax with 3 pairs of dorsal hairs, 1 pair of mar- ginal hairs. Abdomen with 5—6 dorsal hairs on tergites I-VI, 3 pairs of marginal hairs on tergites I-IV and VI-VII, 5—6 pairs of marginal hairs on tergite V, and 4—6 dorsal hairs on tergite VIII. Length of frontal hairs, marginal hairs on abdominal tergite I, and dorsal hairs on abdominal tergite VIII 2.94, 2.48 and 3.36 times as long as basal diameter of antennal segment III, respec- tively. Antenna 6-segmented, without sec- ondary rhinaria. Processus terminalis 2.93 times as long as base of antennal segment VI. Primary rhinaria round, ciliated. Anten- na slightly imbricated. Rostrum not reach- ing middle coxae; ultimate rostral segment short, as long as or a little longer than its basal diameter, 0.96 times as long as 2nd hind tarsal segment, with 1 pair of acces- sory hairs and 3 pairs of primary hairs. Hind femur 1.72 times as long as antennal segment III, hind tibia 0.45 times as long as body. Length of hairs on hind tibia 0.092, 2.30 times as long as middle diam- eter of tibia. First tarsal segment chaeto- taxy: 3, 3, 2. Second tarsal segment trans- versely imbricated. Siphunculus short, smooth, with a small flange, 1.16 times as long as basal diameter, 1.60 times as long as its tip diameter, 0.82 times as long as cauda. Cauda 1.54 times as long as its basal diameter, with 5—6 hairs. Anal plate broad circular, with 14—17 hairs. Genital plate kidney-shaped, bearing 12—15 hairs, among them 2 hairs on its anterior part. Alate viviparous female.—Body 1.408 in length, 0.640 in width. Antenna 1.132, seg- ments I-VI: 0.049, 0.048, 0.323, 0.160, 0.161, 0.091 + 0.300. Ultimate rostral seg- ment 0.080. Hind femur 0.389, hind tibia 0.720, 2nd hind tarsal segment 0.078. Si- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 13 Figs. 10-13. ment. 12, Siphunculus. 13, Cauda. phunculus 0.099 in length, 0.071 in basal diameter, 0.046 in tip diameter. Cauda 0.097 in length. Body elliptical, dark green and with white wax. In cleared specimens, head and thorax dark brown; antenna and legs brown; abdomen with dark brown cross bars on ter- gites III-VII. Antenna 6-segmented, anten- nal segments HJ—VI with 23-40, 4-18, 6— 12, O—1 secondary rhinaria, respectively, which are round and convex. Processus ter- minalis about 3.32 times as long as base of ultimate antennal segment. Other characters as for apterous viviparous female. Discussion.—This new species is closely related to M. arundinariae, but differs from it by the following characters: 1) Genital plate with 2 hairs on its anterior part (the latter: 7—8 hairs); 2) abdomen with 5—6 dor- sal hairs on segments I-III (the latter: 15— 22 hairs); and 3) alate viviparous female with 23—40, 4—18, 6-12, O—1 convex sec- ondary rhinaria on antennal segments IIJ— VI, respectively (the latter: 12, 5, O—2). Types.—Holotype: apterous viviparous female, No. 7151-2-3-4, May 24, 1980, Yunnan Province (Lijiang, 2400m, 100.2°E, 26.8°N) on Bambusa sp., collected by Zhong Tiesen. Paratypes: 9 apterous vivip- arous females, 7 alate viviparous females, No. 7151, data same as holotype. 0.05mm 11 Melanaphis zhanhuanesis, apterous viviparous female. 10, Antenna. 11, Ultimate rostral seg- Melanaphis pyrisucta Zhang and Qiao ioe Melanaphis pyrisucta Zhang and Qiao 1999, in Zhang et al. 1999: 589. Host plant.—Pyrus sp. Distribution in China.—Fujian. Melanaphis sacchari (Zehntner 1897) Aphis sacchari Zehntner 1897: 551. Synonyms: Aphis pheidolei Theobald 1916, Aphis sorghella Schouteden 1906, Aphis sorghi Theobald 1904 (Eastop and Hille Ris Lamberrs 1976, Remaudiére and Remaudiére 1997). Host plants.—Echinochloa sp., Oryza sp. and Panicum sp. Distribution in China.—Heilongjiang, Jilin, Liaoning, Nei Mongol Auto. Reg., Hebei, Henan, Shandong, Shanxi, Anhui, Zhejiang, Jiangsu, Sichuan, Yunnan, Guangdong, Taiwan. Melanaphis siphonella (Essig and Kuwana 1918) Aphis siphonella Essig and Kuwana 1918: TS: Host plant.—Pyrus betulaefoli. Distribution in China.—Hebei (Beidaihe City). This is a new record for China. VOLUME 103, NUMBER 2 Eo Figs. 14-16. M. zhanhuanesis. Examined specimens.—1 apterous vivpa- rous female, 2 alate vivparous females. Melanaphis zhanhuaensis Zhang, Qiao, and Zhang, new species (Figs. 10-13, 16) Apterous viviparous female.—Body 1.542 in length, 0.960 in width. Antenna 5 or 6-segmented, 0.781 (or 0.839), 5-seg- mented, segments I—V: 0.055, 0.042, 0.244, 15 Melanaphis spp., habitus illustrations. 16 14, M. arthraxonophaga. 15, M. grossisiphonellus. 16, 0.113, 0.076+0.252; 6-segmented, seg- ments I-VI: 0.055, 0.042, 0.155, 0.118, 0.118, 0.080+0.271. Ultimate rostral seg- ment 0.078. Hind femur 0.336, hind tibia 0.546, 2nd hind tarsal segment 0.076. Si- phunculus 0.092 in length, 0.052 in basal diameter, 0.042 in tip diameter. Cauda 0.128 in length. Body elliptical and purplish red. In cleared specimens, cuticle on thorax and 352 abdomen with reticulation formed by rows of spinules; antennal segments IV—V (or VI), rostrum at apex, siphunculus, cauda, anal plate, and genital plate dark brown; legs brown; others pale. Spiracles circular and open, spiracular plates elliptical and brown. Mesosternal furca brown, without stem. Marginal tubercles on prothorax and abdominal tergites I and VII, with height a little longer than their basal diameter. Ver- tex slightly convex, antennal tubercles de- veloped. Body with sparse, fine, short dor- sal hairs. Head with 1 pair of frontal hairs, 1 pair of lateral frontal hairs, and 2—3 pairs of dorsal hairs. Prothorax with 3—4 pairs of dorsal hairs. Abdomen with 2 dorsal hairs on tergite VIII. Length of frontal hairs, mar- ginal hairs on abdominal tergite I, and dor- sal hairs on abdominal tergite VIII, 0.84, 0.64 and 1.29 times as long as basal di- ameter of antennal segment III, respective- ly. Antenna 5 or 6-segmented, without sec- ondary rhinaria. Processus terminalis 3.22— 3.32 times as long as base of ultimate an- tennal segment. Primary rhinaria round, cil- iated. Antenna slightly imbricated. Hairs on segment III 0.008, 0.33 times as long as basal diameter. Rostrum reaching middle coxae; ultimate rostral segment short, 1.16 times as long as its basal diameter, 1.03 times as long as 2nd hind tarsal segment, with 1 pair of accessory hairs and 3 pairs of primary hairs. Hind tibia 0.35 times as long as body. Length of hairs on hind tibia 0.034, 0.94 times as long as middle diam- eter of tibia. First tarsal segment chaeto- taxy: 3, 3, 2. Second tarsal segments trans- versely imbricated. Siphunculus short, slightly imbricated, with a flange, 1.76 times as long as its basal diameter, 2.20 times as long as its tip diameter, 0.72 times as long as cauda. Cauda 1.22 times as long as its basal diameter, with 12 hairs, con- stricted in middle. Anal plate broad, circular at apex, with 36 hairs. Genital plate bearing 21 hairs, among them 8 hairs on its anterior part. Discussion.—This new species is closely related to M. luzulella and M. pyraria on PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON secondary host, but differs from M. luzu- lella by the following characters: 1) Si- phunculus 0.72 time as long as cauda [the latter (Heie 1986): a little longer than cau- da]; 2) cauda with 12 hairs (the latter: 5—7 hairs); and 3) abdomen pale in cleared spec- imens (the latter: with dark cross bar on ter- gite VIII, sometimes also with paler, shorter cross bars on tergites VI and VII). Melan- aphis zhanhuaensis differs from M. pyraria on the secondary host by the following characters: 1) Siphunculi and canda dark brown [the latter (Heie 1986): pale, whit- ish]; and 2) living on the subterranean part of the host (the latter: living above the ground). Types.—Holotype: apterous viviparous female, No. Y2594-1-1-1, June 28, 1982, Shandong Province (Zhanhua County, 118.1°E, 37.3°N), on subterranean part of Imperata cylindrica var. major (Nees) C. E. Hubb., collected by Zhang Tingzhu. Para- type, 1 apterous viviparous female, No. Y2594, data same as holotype. ACKNOWLEDGMENTS We are very grateful to Ms. Liu Lijuan, Mr. Zhong Tiesen, Mr. Zhang Tingzhu, and Mr. Tian Shibo who collected the speci- mens, to Mr. Zhong Tiesen who made the slides, and to Mr. Mai Guoqing who took the photographs of the aphids. We also thank Dr. M. Sorin to allow examination of some named specimens of this genus. This work was supported by a grant from the National Natural Science Foundation of P. R. China (No. 39770119). LITERATURE CITED Barbagallo, S. and H. L. G. Stroyan. 1982. Biological, Ecological and Taxonomic Notes on the Aphid Fauna of Sicily. Frustula Entomologica Nuova Serie Vol. 3 (16): 1—182. Blackman, R. L. and V. E Eastop. 1994. Aphids on the World’s Trees. An Identification and Infor- mation Guide. CAB International. 987 pp. . 1994. Aphids on the World’s Crops, An Iden- tification and Information Guide. Second Edition. John Wiley & Sons, Ltd. 466 pp. Eastop, V. E and D. Hille Ris Lambers. 1976. Survey VOLUME 103, NUMBER 2 of the World’s Aphids. Dr. W. Junk b. v., Publish- ers, The Hague. 573 pp. Heie, O. E. 1986. Fauna Entomologica Scandinavica Vol. 17: The Aphidoidea (Hemiptera) of Fennos- candia and Denmark. III. E. J. Bill/Scandinavian Science Press Ltd. Leiden. Copenhagen, 313 pp. Hodjat, S. H. 1998. A list of aphids and their host plants in Iran. Shahid-Chamran University Print- ing & Publication Center. 148 pp. Pashcheko, N. F 1988. 4. Suborder Aphidinea-aphids, pp. 546—686. /n Ler, P. A., Key to the identifica- tion of insects of the Soviet Far East. Vol. 2. Ho- moptera and Heteroptera. Nauka. Leningrad. Passerini, G. 1861. Additamenta ad indicem Aphidi- narum. Quas hucusque in Italia legit. Atti della Societa Italiana di Scienze Naturali 3: 398—401. Raychaudhuri, D. N. and C. Banerjee. 1974. A study on the genus Melanaphis (Homoptera: Aphididae) with descriptions of new taxa from India. Oriental Insects 8(3): 365-284. Remaudiére, G. and M. Remaudiére. 1997. Catalogue of the World’s Aphididae (Homoptera Aphidoi- dea). Paris, INRA Editions, 475 pp. Sorin, M. 1970. Longiunguis of Japan (Homoptera: 333 Aphididae). Insecta Matsumurana, Supplement 8: 5-17. Stroyan, H. L. G. 1984. Handbooks for the Identifi- cation of British Insects Vol. 2, part 6: Aphids- Pterocommatinae and Aphidinae (Aphidini) Ho- moptera, Aphididae. London, Royal Entomologi- cal Society of London, 232 pp. Tao, C. C. 1990. Aphid-Fauna of Taiwan Province, China. Taiwan Province Museum Press, Taipei, 327 pp. Zhang, G. X. and T. S. Zhong 1983. Economic Insect Fauna of China. Fasc. 25, Homoptera: Aphidinea, Part 1. Science Press, Beijing, 387 pp. Zhang, G. X., T. S. Zhong and W. Y. Zhang. 1992. Homoptera: Aphidoidea, pp. 140—156. Jn Huang, F S., ed., Insects of Wuling Mountains Area, Southwestern China. Science Press, Beijing. Zhang, G. X., G. X. Qiao and T. S. Zhong. 1999. Ho- moptera: Aphicoidea, pp. 512-661. Jn Huang, P. K., ed., Fauna of Insects Fujian Province of China, Vol. 2. Science & Technology Press of Fujian Province, Fuzhou. van der Goot, P. 1917. Zur Kenntnis der Blattlaiise Java’s. Contributions a la Faune des Indes Néer- landaises dirigées par le Dr. J. C. Koningsberger 1(3): 1-301. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 334-336 AMPHIAREUS CONSTRICTUS (STAL) (HEMIPTERA: HETEROPTERA: ANTHOCORIDAE) FROM CALIFORNIA: CLARIFICATION OF PREVIOUS RECORD AND CITATION JOHN D. LATTIN AND TAMERA LEWIS (JDL) Department of Entomology, Oregon State University, Corvallis, OR 97331-2907, U.S.A.; (TL) Agricultural Research Service, U.S. Department of Agriculture, 5230 Kon- nowac Pass Road, Wapato, WA 98951, U.S.A. Abstract.—Clarification is provided on the origin of specimens of Amphiareus constric- tus (Stal) from San Francisco, California. These specimens were interecepted in a ship- ment of orchids from Colombia, South America. Key Words: Amphiareus constrictus (Stal) 1860 was described from Rio de Janeiro, Brazil, in the genus Xylocoris. Subsequently, it has been reported from other locations around the world [e.g. Australia (Gross 1955); Ga- lapagos Islands (Herring 1966); Hawaii (Zimmerman 1948); India (Distant 1906); Japan (Hiura 1958); Mexico (Champion 1900); United States (Blatchley 1926)], of- ten under names now considered synonyms of Xylocoris constrictus. Several species were synonymized with this species includ- ing Xylocoris fulvescens Walker, 1872; Xy- locoris fumipennis Walker, 1872; and Car- diastethas macilentus Hiura, 1958. The reader is referred to Herring (1965) for the review and resolution of the taxonomic problem and Henry (1988) who reviewed selected references. Note that Péricart (1996) did not consider Xylocoris fumipen- nis Walker, 1872, a synonym of A. constric- tus. The only published record of the oc- currence of it in the United States was that of Blatchley (1926) who reported it under the combination Poronotus constrictus from Dunedin, Florida, collected “‘Dec 11 - April 14”’ where it was “‘common beneath cover in dry sandy places near the bay Anthocoridae, introductions, customs interceptions, United States beach; especially so in or beneath piles of dead cabbage palmetto leaves, in decaying fungi and beneath freshly cut pine blocks.” Hiura (1958) described Cardiastethus macilentus from Jinzenji, K6chi-City, Shi- koku, Japan, and provided a detailed de- scription and comparison with C. pygmaeus Poppius, an illustration of the adult, and a measured outline of the head, antenna, and prothorax. Subsequently, Hiura (1960) placed his species into synonymy with Am- phiareus fulvescens (Walker), a species lat- er synonymized under Amphiareus_ con- strictus (Stal) (Herring 1965), after receiv- ing specimens of Poronotellus constrictus (Stal) from R.I. Sailer (then of the U.S. De- partment of Agriculture, Systematic Ento- mology unit at the National Museum of Natural History, Washington, DC) labeled San Francisco, California. Hiura (1960) provided a discussion of the complicated problem of the proper name for the species; the solution resolved ultimately by Herring (1965). Referring to the habits of his own species, Hiura (1960) stated ‘‘Habits: - the bug lives in the piles of harvested stems and leaves (e.g., wheat, sweet potato-vine), fire- wood, and sometimes is attracted to light at VOLUME 103, NUMBER 2 night.”’ Further, he captured a specimen that had flown into the cabin of a passenger boat touching port. In Guam, specimens were obtained from “dead orange twigs.’’ His descriptions of other habitats where the spe- cies was found were similar to those given by Blatchley (1926). Amphiareus constrictus is a small species (2.2—2.8 mm). The second antennal seg- ment is slightly enlarged and segments three and four are slender. The pronotum has a distinct transverse impression that sets off the anterior portion while the posterior portion has a longitudinal impression and a broadly concave posterior margin. The an- terior and posterior angles of the pronotum have a long erect seta and another long seta originates about one-quarter the distance back from each anterior angle. The clavus is distinctly punctate with coarse punctures with a seta in each impression, the balance of the hemielytra have scattered, semi-de- cumbent setae; the membrane has three veins. Hiura (1960) illustrated the male genital capsule with the clasper in place (as Poronotellus constrictus Stal), and Zim- merman (1948) illustrated the adult of the Same species under the name Cardiastethus fulvescens (Walker). T. J. Henry was unable to locate any specimens in the collection of the National Museum of Natural History, Smithsonian Institution, Washington, DC with labels similar to the publication by Hiura (1960) (personal communication with T: Lewis, 1999). The two specimens sent to Japan by Sailer are in the Osaka Museum of Natural History, Osaka, Japan. I. Kangzawa, now responsible for the insects of this museum, very kindly provided us the following in- formation: a specimen with the label ‘‘Co- lombia, wild orchids. D.C. Inspect. House. XII, 26. 39. Poronotellus constrictus Stal’’; and one specimen missing, but the label “Colombia, on orchid. S. Francisco, Cal. 4- 30-41. Poronotellus constrictus’’ present (presumably Hiura used the specimen for dissection for comparison with his own spe- cies). This information clarifies the origin 335 of the specimen(s) collected in San Fran- cisco, California, in 1941 - their intercep- tion in a shipment of orchids from Colom- bia, South America. More recently, Cassis and Gross (1995) reported this species (as Amphiareus con- strictus) from Australia and other localities around the world, suggesting introductions elsewhere. The details of the collection at San Francisco provide at least one mode of introduction, movement in commercial plant materials. This species appears to be native to South America (Stal’s original description from Brazil in 1860), but now seemingly introduced into many other parts of the world. An established population in the San Francisco area has not yet been documented but might be found in Golden Gate Park, San Francisco, where another non-indige- nous species of Anthocoridae, Buchanan- iella continua (White), now established, has been found. Judging from the comments of Blatchley (1926) and Hiura (1958, 1960), the species occurs most commonly beneath piles of dead plant materials and fungi on the ground, while specimens from Guam were beaten from dead orange twigs (Hiura 1960). These habitats resemble those of other species of Anthocoridae found on Pa- cific Islands and elsewhere where many of the bugs have been taken beating clusters of dead leaves and branches. Some of these instances included the occurrence of Pso- coptera living in the same environment (e.g., Lattin 1999). The generalist predatory feeding habits of some Anthocoridae in habitats likely to be found many places around the world might result in wide dis- persal: close attention to specific habitats where future specimens are collected will add to our knowledge. ACKNOWLEDGMENTS We dedicate this paper to the late I. Hiura (Osaka Museum of Natural History), who published many fine papers on the Antho- coridae fauna of Japan and other areas. We 336 thank T. J. Henry (Systematic Entomology Laboratory, U.S. Department of Agricul- ture, Washington, DC) and I. Kanazawa (Osaka Museum of Natural History, Osaka, Japan) for their assistance in providing in- formation on critical specimens, two anon- ymous reviewers for their useful comments, and L. Parks for her careful preparation of the manuscript. LITERATURE CITED Blatchley, W. S. 1926. Heteroptera or true bugs of east- ern North America, with especial reference to the faunas of Indiana and Florida. Nature Publishing Company, Indianapolis. 1,116 pp. Cassis, G. and G. C. Gross. 1995. Hemiptera: Heter- optera (Coleorrhyncha to Cimicomorpha) Antho- coridae, pp. 23—42. Jn Houston, W. W. K. and G. V. Maynarad, eds, Zoological Catalogue of Aus- tralia. Vol. 27.3A. Melbourne. CSIRO, Australia. 506 pp. Champion, G. C. 1897-1901. Insecta: Rhynchota (He- miptera - Heteroptera). Vol. I. Jn Godwin and Sal- vin, eds. Biologia Centrali - Americana, London. XVI + 416 pp., 22 plates. [1897: 1—32; 1898, 33- 192; 1899: 193-304; 1900: 304—344; 1901: i-xvi + 345-416]. Distant, W. L. 1906. Fauna of British India, including Ceylon and Burma. Rhynchota. Vol. HI Heterop- tera and Hemoptera, 503 pp. Gross, E G. 1955. A revision of the flower bugs (Het- eroptera: Anthocoridae) of the Australian and ad- jacent Pacific Regions, part II. Records of the South Australian Museum 11: 409—422. Henry, T. J. 1988. Family Anthocoridae, pp. 12—28. In Henry, T. J. and R. C. Froeschner, eds. Catalog of PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON the Heteroptera, or true bugs, of Canada and the Continental United States. E. J. Brill, Leiden. 958 Pp. Herring, J. L. 1965. The status of Amphiareus Distant, Buchananiella Reuter, and Poronotellus Kirkaldy (Hemiptera: Anthocoridae). Proceedings of the Entomological Society of Washington 67: 202— 203. . 1966. The Anthocoridae of the Galapagos and Cocos Islands (Hemiptera). Proceedings of the Entomological Society of Washington 68: 127— 130. Hiura, I. 1958. Two new species of Cardiastethus from Japan (Hemiptera: Anthocoridae). Entomological Review of Japan 9: 38—40, pl. 7. . 1960. Contribution to the knowledge of An- thocoridae from Japan and its adjacent territories (Hemiptera - Heteroptera). 2. Bulletin of the Osa- ka Museum of Natural History No. 12: 43-55. Lattin, J. D. 1999. Bionomics of the Anthocoridae. An- nual Review of Entomology 44: 207-231. . 1999. Dead leaf clusters as habitats for adult Calliodis temnostethoides and Cardiastethus lur- idellus and other anthocorids (Hemiptera: Heter- optera: Anthocoridae). Great Lakes Entomologist 32(1): 33=38. Pericart, J. 1996. Anthocoridae, pp. 108-140. Jn Au- kema, B. and C. Rieger eds. Catalogue of the Het- eroptera of the Palaearctic Region. Vol. 2., Cimi- comorpha 1. Netherlands Entomological Society, Amsterdam, The Netherlands. 359 p. Stal, C. 1860-1862. Bidrag till Rio Janeiro - traktens Hemipter-fauna. Kongliga Svenska Vetenskaps- Akademien Handlingar 2(7): 184(1860); 3(6): 1— 75 (1862). Zimmerman, E. C. 1948. Insects of Hawaii. Vol. 3. Heteroptera. University of Hawaii Press, Hono- lulu. 255 pp. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 337-348 A REVIEW OF THE FAMILY NANNODASTIIDAE (DIPTERA) LASZLO PAPP AND WAYNE N. MATHIS (LP) Department of Entomology, Hungarian Natural History Museum, Baross utca 13, Budapest, Hungary (e-mail: lpapp@zoo.zoo.nhmus.hu); (WNM) Department of Entomol- ogy, Smithsonian Institution, Washington, D.C. 20560-0169 U.S.A (e-mail: mathis. wayne @nmnh.si.edu) Abstract.—The two genera and five species currently assigned to the family Nanno- dastiidae are reviewed. The genera, Azorastia Frey and Nannodastia Hendel, are very similar and closely related, although each was originally described in a different family, Asteiidae and Ephydridae respectively. Adults of both genera are very small, lengths often less than 1 mm, and exhibit many highly derived, often reduced, structures, making their phylogenetic placement enigmatic. Thus, we remain unsure of the family’s phylogenetic relationships and placement in acalyptrate classification or even of the group’s status as a family. Nannodastia atlantica is newly described (type locality: Belize. Stann Creek District: Glover’s Reef (Long Cay)) and is the first record of this genus in the New World. Key Words: review, Diptera, Nannodastiidae, Azorastia, Nannodastia The higher classification of many taxa in- cluded in the so-called acalyptrate Diptera remains an inscrutable puzzle, frequently lacking resolution at essentially any level (Yeates and Wiegmann 1999). This _per- plexing condition is particularly true of taxa that have highly derived morphological structures, often greatly reduced, as exem- plified by the ‘“‘Nannodastiidae.’’ The Nan- nodastiidae were accorded familial status less than 10 years ago (Carles-Tolra 1994) largely because they did not agree with the characterization of any other known family and because of the unusual morphology, ex- tremely small size, and reduction of struc- tures. The objectives of our review of this so-called family, are to describe a new spe- cies in the genus Nannodastia Hendel, to provide new information on the distribution and morphology of the group, and to high- light this poorly known family in an attempt to foster further research on it. The descriptive terminology follows Papp (1980). Because specimens are ex- tremely small, often less than 1 mm in length, study and illustration of the male and female terminalia required use of a compound microscope. Except for the new species, other species are provided with a diagnosis in the appropriate key to species. Family Nannodastiidae L. Papp Nannodastiinae L. Papp 1980:425 [subfam- ily in Ephydridae]. Type genus: Nanno- dastia Hendel 1930.—Cogan 1984:144 [Palearctic catalog, subfamily in Ephy- dridae].—Pitkin 1989:600 [Australasian and Oceanian catalog, subfamily in Chy- romyidae].—Mathis 1989:6—7 [Nanno- dastia in Chyromyidae].—Colless and McAlpine 1991:766, 773 [subfamily in Chyromyidae]. Nannodastiidae: Carles-Tolra 1994:199 [ac- corded familial status]—L. Papp and Mathis 1998:309-—314 [review of Pale- arctic fauna]. 338 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-2. Azorastia minutissima, paratype male. 1, Posterior portion of preabdomen and genitalia in lateral view. 2, Surstylus in widest view. Scales: 0.2 mm for Fig. 1; 0.1 mm for Fig. 2. Diagnosis.—Minute to very small flies, length 0.70—1.25 mm; body mostly brown, generally dull colored, sparsely to moder- ately microtomentose; wings hyaline; legs lacking long setae. Head: higher than long; 3 reclinate fron- to-orbital setae, anterior seta inserted at lev- el of lunule; inner and outer vertical and ocellar setae distinct, ocellar seta inserted immediately laterad of anterior ocellus; postvertical seta lacking; | pair of minute, moderately divergent and proclinate posto- cellar setulae. Antenna: scape small; pedicel lacking a suture, bearing long setae dorsally and ventrally, with a process (projected into the flagellomere cavity, see Figs. 3, 16) api- cally or ventroapically; flagellomere with a posterior cavity, bearing very long setae apically; arista short, bearing sparse short rays (see Figs. 3, 16). Face small in width and height, weakly sclerotized medially; ventral facial margin straight to very shal- lowly convex; clypeus small; 3 well-devel- oped, slightly upcurved, parafacial setae and some rather long peristomal setae; vi- brissae and a vibrissal angle lacking. Thorax: 1 dorsocentral seta (O+1); 1 postalar seta; 2 scutellar setae, apical seta longer, curved dorsally; 1 row of dorsocen- tral setulae; 1 row of intra-alar setulae; ac- rostichal setulae in 2 mostly regularly ar- ranged rows; postpronotal setae lacking; 2 notopleural setae, posterior seta inserted at much higher level than anterior seta; 4—5 anepisternal setae, 2—3 curved dorsally; 1 katepisternal seta. Legs lacking character- istic setae (see Fig. 19 of L. Papp 1980). Wing hyaline; costal vein broken twice, basal break distad to humeral vein, apical break just before vein R,; subcosta rudi- mentary, blunt apex free; vein M_ thin; crossvein r-m lacking; crossvein dm-cu pre- sent or absent; anal cell and anal vein lack- ing; alula lacking; wing margin bearing long fringe. Halter short, knob compara- tively large. Abdomen: Spiracles apparently lacking. Male abdomen with 6 exposed segments VOLUME 103, NUMBER 2 (Figs. 1, 18-19), tergites 1 and 2 separate, although tergite 1 very short and thin me- dially, Ist segment otherwise not too short; a transverse narrow sclerite in membrane between tergite 6 and epandrium, forming a half ring (here considered a synsternite 7+8). Male postabdomen symmetrical; epandrium with or without a posteroventral process; lateral wall of surstylus fused with epandrium (but not medial wall); cerci weakly sclerotized, bearing very long setae; genitalia comprising a lever structure; hy- pandrium elongate, rather dorsally placed on epandrium, fused anteriorly with lateral extensions of aedeagal apodeme; aedeagal apodeme fused with aedeagus; aedeagus elongate; gonopods or parameres lacking; a sclerotized plate connecting posterior ends of hypandrium and cerci (under the epan- drium) that is here considered an interpar- ameral sclerite (perhaps sternite 10); genital opening bordered by hypandrial arms (L. Papp 1980). Papp (1980) described the function of the genitalia but misinterpreted the aedeagus and some other structures of the terminalia. Female abdomen normal; tergites 7—8 undivided, but more strongly sclerotized laterally; last dorsal sclerites of female (here interpreted as cercit+epiproct) as weakly sclerotized plates, bearing long (Azorastia) or short (Nannodastia) setae; spermathecae and a sclerotized ventral re- ceptacle apparently lacking. Discussion.—The two genera comprising this family remain enigmatic with essen- tially unknown familial affiliations within the Schizophora (Opomyzoidea/Carnoidea). The type genus, Nannodastia Hendel, was first described in the Ephydridae, and Azo- rastia Frey, the only other included genus, in the Asteiidae. Sabrosky (1956) first noted a close relationship between these genera, and he, followed by Frey (1958) and Sa- brosky and Wirth (1959), treated these gen- era as members of the family Ephydridae, tribe Atissini. L. Papp (1980) described Nannodastiinae as a subfamily of Ephydri- dae, based in part on the advice of other acalyptratae experts. L. Papp (1980: 426), B59 however, qualified his classification with the statement that: ““The majority of the above differences [of Nannodastiinae and other ephydrid subfamilies] separate super- families in Griffiths’ (1972) system; they seem sufficient to separate subfamilies here.’ Pitkin (1989), as part of the Austra- lasian/Oceanian catalog, recognized Nan- nodastiinae as a subfamily in the family Chyromyidae. No synapomorphies linking these two families have been elaborated, however (the number and position of the fronto-orbital setae are regarded as a con- vergence, shared by other families, such as the Tethinidae and Acartophthalmidae), and familial affiliations remain unsubstantiated. We have not discovered the sister group of Nannodastiidae nor its placement among the superfamilies of Griffiths (1972: Fig. 14), partially as a consequence of the re- duced and highly modified morphology. For the present, we are uncertain about the relationships of Nannodastiidae except that they lack the synapomorphies that charac- terize Ephydroidea (Zatwarnicki 1992, Mathis and Zatwarnicki 1998), and we are likewise doubtful of Chyromyidae. The two genera presently included in Nannodastiidae, Azorastia and Nannodas- tia, are very similar externally, as are, to lesser extent, the structures of the male ter- minalia. Their close resemblance led Sa- brosky and Wirth (1959) and L. Papp (1980) to question their status as distinct genera. We confirm their similarity, and certainly the two groups are sister taxa, but until their familial placement is resolved, we prefer to recognize them as distinct gen- era. Although similar, the two genera are easily diagnosed by external characters and by structures of the male terminalia. Natural history.—Adults, which are very small (often less than | mm), are usually associated with maritime beaches in tropi- cal and subtropical zones. Specimens were often collected in caves or under cliff over- hangs where the substrate was rich in or- ganic matter from the droppings of bats, small mammals, and birds. Nothing is 340 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON /, : Se I Xi ~~ ~ TEN gw 3 4 Moya (AYN ANANDA Figs. 3-7. Azorastia mediterranea, male. 3, Antenna, medial view. 4, Wing, dorsal view. 5, Genitalia, lateral view (anus shown by a small arrow). 6, Same, ventral view (hypandrium foreshortened, interparameral sclerite hatched). 7, Right surstylus in widest, medial view, i.e., posterior edge at right. Scales: 0.5 mm for Fig. 4; 0.1 mm for Figs. 3, 5—7. terior portion of surstylus bearing 3 long setae; posterior portion with a digitiform process with 3 thornlike setae apically; hypandrium less dor- sally placed on epandrium (Figs. 13, 22), short- er, anterior part robust with ventral dilatation. Female cerci bearing short setulae only (Figs. 1. Crossvein dm-cu lacking (Fig. 17); vein M at- DAO UNCLEAR Nannodastia Hendel known about the immatures, although pre- sumably they are in the richly organic sub- strate. KEY TO GENERA OF NANNODASTIIDAE tenuate apically, not extended to wing margin; — Crossvein dm-cu present (Fig. 4); vein M dis- costal vein continued past apex of vein R,,, but not to vein M. Male syntergosternite 7+8 very much reduced (Figs. 18-19); epandrium with a posteroventral process, bearing a thick thornlike structure apically (Figs. 12, 20); an- tinct to wing margin; costal vein extended to vein M. Male syntergosternite 7+8 distinct (Fig. 1); epandrium lacking a posteroventral process; surstyli longer without any processes or thorns, medial surface with numerous short VOLUME 103, NUMBER 2 341 Figs. 8-11. Azorastia gemmae. 8, Postabdomen of paratype female, lateral view. 9, Same, dorsal view. 10, setae (Figs. 1-2, 5—7); hypandrium rather dor- sally placed on epandrium (Fig. 6); hypan- drium and aedeagus very long and slender. Fe- male cerci bearing at least 1 long seta (Figs. =e). a eielar arena a cramer oe te: Azorastia Frey Genus Azorastia Frey Azorastia Frey 1945:72 [misquoted as *1958:Commentat. biol., 18(4):40,”" by Cogan (1984:144)]. Type species: A. min- utissima Frey 1945, by original designa- tion; 1958:40—41 [discussion, key to gen- era].—Sabrosky 1956:217 [removed from Asteiidae, placed in Ephydridae, subfamily Psilopinae, tribe Atissini, near Atissa Haliday].—Sabrosky and Wirth 1956:18 [placed in Ephydridae, tribe Atissini, compared with Nannodastia}; Same, ventral view (with the right evertible sack in 2X magnification). 11, Genital opening with its sclerites in ventral view, higher magnification, hypoproct dashed. Scales: 0.2 mm for Figs. 8-10; 0.1 mm for Fig. 11. 1959:109—-110 [discussion, list, Hawaii (Oahu)].—L. Papp 1980:427—431 [re- view and diagnosis].—Cogan 1984:144 [Palearctic catalog].—Carles-Tolra 1994: 202 [list]; 1995:258 [list]—L. Papp and Mathis 1998:310—314 [figs., key, discus- sion]. Diagnosis.—Azorastia is distinguished from Nannodastia by the following com- bination of characters: Wing: crossvein dm- cu present (Fig. 4); vein M distinct to wing margin; costal vein extended to vein M. Male terminalia: syntergosternite 7+8 dis- tinct (Fig. 1), normally retracted within ter- gite 6; epandrium lacking a ventroapical process; surstylus relatively long, lacking WwW £ Nw Figs. 12—15. view. 13, Genitalia, lateral view. 14, micropylar end of three eggs, sublateral view. 15, Postabdomen of female, dorsal view. Scales: 0.1 mm for Figs. 12—13; 0.2 mm for Fig. 14; 0.1 mm for Fig. 15. any processes or thornlike setae, medial surface with numerous short setae (Figs. 1— 2, 5—7), bearing setulae on medial surface, lacking contact with interparameral sclerite; lateral wall of surstylus fused with epan- drium (but not medial wall); aedeagus in unflexed position with flat lamellae parallel with dorsum, horizontally turned out (Figs. 1, 5); hypandrium rather dorsally placed on epandrium (Fig. 6); hypandrium and aedea- gus very long and slender. Female cerci bearing at least 1 long seta (Figs. 8—9). KEY TO SPECIES OF THE GENUS AZORASTIA FREY 1. Male surstylus very long, length over twice width, narrow basally, thereafter becoming PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Nannodastia atlantica, holotype male. 12, Epandrium, surstyli, and cerci, oblique lateroventral wider to apical %, then curved posteriorly and pointed (Figs. 1-2, 4 of Carles-Tolra 1994) .. Oe ee Se re eens A. gemmae Carles-Tolra — Male surstylus moderately long to short, length less than twice width, not narrowed basally .. 2 . Male surstylus short, apex more rounded, an- terior edge more or less protuberant in profile (Figs. 5—7). Ratio of intercrossvein section of vein M/terminal section of vein Cu usually IL. OOk* 5 oie was come oars A. mediterranea L. Papp — Male surstylus moderately long, anterior mar- gin nearly straight in lateral view (Figs. 1—2); Ratio of intercrossvein section of vein M 1.20— 1.30 A. minutissima Frey N Azorastia minutissima Frey (Figs. 1—2) Azorastia minutissima Frey 1945:73; 1958: 40-41 [fig. of wing].—Sabrosky and VOLUME 103, NUMBER 2 Wirth 1959:110 [placed in Ephydridae, tribe Atissini].—L. Papp 1980:430 [lec- totype designation].—Cogan 1984:144 [Palearctic catalog].—Carles-Tolra 1994: 202 [list]_—L. Papp and Mathis 1998:310 [figs. of male terminalia]. Material examined.—The lectotype male (designated by L. Papp, 1980: 430) and 2 female paralectotypes; another three speci- mens of the type series not discussed by Papp (1980) are here recognized as para- lectotypes: Ist paralectotype ¢ (ZMH, head and left wing lost): Azor: S. Miguel, Ri- beira Grande, 26.V. Stora/2894/Mus. Zool. Helsinki Loan No. D 4954 [yellow]/ Mus.Zool. Helsinki Loan No. D90-184. 2nd paralectotype ° (ZMH, head, legs and right wing lost, abdomen on a separate minuten): Azores: Flores, Sta Cruz, 1—15.VI. Stora/ 1086 [blue]/Mus.Zool. H:fors, Spec. typ. No. 89.95. Azorastia minutissima Frey’’/ as on previous label but with D 4952/as on previous label but with D90-183 (recog- nized here as a paralectotype; state of pres- ervation poor). 3rd paralectotype ¢ (ZMH, originally double-mounted on a minuten, now abdomen and genitalia preserved in an Andersson’s microvial with glycerol): Azor: Flores, Sta Cruz, VI. Stora/4889 [blue]/ Mus.Zool.H:fors, Spec. typ. No. ‘89-96 Azorastia minutissima Frey’’/Mus.Zool. Helsinki Loan No. D 4953/Mus.Zool. Hel- sinki, No. D 90-185. Measurements of the three paralectotypes noted previously: body length: 1.00—1.03 mm; wing length 0.98—-1.12 mm; wing width 0.43-0.52 mm; distance between crossveins r-m and dm-cu/apical section of vein M 1.17—1.35. A very long seta on cer- cus ventrally (see L. Papp 1980:430). Distribution.—Palearctic: Azores. Remarks.—This species is closely related to A. mediterranea but differs in the shape of the male surstylus. Azorastia mediterranea L. Papp (Figs. 3—7) Azorastia mediterranea L. Papp 1980: 427.— L. Papp and Mathis 1998:309— 343 310 [habitus fig., figs. of male termina- lia]. Material examined.—Type series (see L. Papp, 1980: 430). Newly collected ma- terial: ISRAEL. Migdal ’Afeq (32°05'N, 34°NS57’E; also known as Midgal Zedek), 25 Jul 1976, A. Freidberg (9 5, 1 ¢); ibid., 4 Sep 1977, A. Freidberg (11 3); ibid., 24 Sep 1981, A. Freidberg, EF Kaplan (25 6, 3 2); ibid., 11 Jul 1993, A. Freidberg (17 6, 1 2); ibid., 3 Jun 2000, D. and W. N. Math- is (3 6) (HNHM, TAU, USNM). Distribution.—Palearctic: Croatia (Dal- matia), Greece, Israel (new record), Tunisia. Natural history.—The specimens from Israel were collected by sweeping just above the substrate within the cavelike ru- ins of an Ottoman fortress (Migdal Afeq) where sheep, goats, and other mammals of- ten seek shelter. The fortress is not imme- diately coastal but is on the coastal plain, approximately 17.5 km from the eastern shore of the Mediterranean Sea. Azorastia gemmae Carles-Tolra (Figs. 8-11) Azorastia gemmae Carles-Tolra 1994:199; 1995225 Sq (hist Material examined.—A paratype female is labeled (HNHM through exchange; com- pare with Carles-Tolra 1994:201) ‘‘26—29.6. 88 [26—29 Jun 1988], Ibiza (Baleares), Pun- ta Arabi, penado muerto, M. Carles-Tolra leg/Azorastia gemmae Carles-Tolra, PAR- ATYPO 92.” The paratype was mounted from alcohol on a 0.1 mm minuten. The abdomen is in an attached plastic microvial with glycerine. Distribution.—Palearctic: Spain (Bale- aric Islands: Ibiza). Remarks.—The following remarks are additions or corrections to Carles-Tolra’s original description. The female preabdo- men comprises 6 segments and the 6th is normally developed (Fig. 8). Segment 7 is a syntergosternite (Figs. 9-10) with a small medial subanterior dark spot on the sternal portion (also on Fig. 7 of Carles-Tolra 344 Figs. 16-19. lateral view. 19, Same, ventral view. Scales: 0.1 mm for Fig. 16; 0.5 mm for Fig. 17; 0.2 mm for Figs. 18—19. 1994). No “‘internal sclerites,’’ as in Carles- Tolra’s Fig. 8, were observed in the para- type. We spectulate that these might be sec- tions of tracheae, although abdominal spi- racles are not discernible in nannodastiids. Carles-Tolra’s “‘lateral slightly sclerotized zones’’ were found between sclerites of the 6th and 7th segments, as in Fig. 10. They lack a definite shape, are three-dimensional (seen in Fig. 8), and the microchaetae inside are directed against each other. We suspect the microchaetae are eversible and are pos- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON \ WSS My > NW wopy pH GNAANANHNNNN Nannodastia horni, male. 16, Antenna, medial view. 17, Wing, dorsal view. 18, Abdomen, sibly invaginations of the integument (ever- sible sacks) for pheromone excretion. Ter- gite 8 is a pair of comparatively large lateral sclerites that are rather elongate ventrally. Sternite 8 is a pair of medium-long, narrow sclerites that are positioned on the apical part of a very large genital opening (Figs. 10—11); sternite 8 bears some medium-long setae. The very large genital opening cor- relates with the comparatively large eggs. The epiproct is a minute sclerite at the base of the cerci and bears a pair of asymmet- VOLUME 103, NUMBER 2 345 Figs. 20-23. surstylus, and cercus, lateral view. 23, Surstylus of a 2nd specimen, ventral view. Scales: 0.1 mm for Figs. 21— 22; 0.05 mm for Figs. 20, 23. rically placed, comparatively long setae (Figs. 8-9). The hypoproct (Figs. 8, 11), al- though not very small, is weakly sclerotized and bears a pair of asymmetricaly placed, short setae (Fig. 11). The cerci (Figs. 8—9) are moderate sized and bear two pairs of very long and some shorter setae. Genus Nannodastia Hendel Nannodastia Hendel 1930:68. Type spe- cies: N. horni Hendel 1930:70, by origi- nal designation.—Frey 1958:40—41 [dis- cussion].—Sabrosky 1956:217 [placed in Ephydridae, subfamily Psilopinae, tribe Atissini, near Atissa Haliday].—Sabrosky and Wirth 1956:18 [list, Hawaii, Oahu]; 1959:109-110 [discussion, placed in Ephydridae, tribe Atissini, compared with Azorastia|.—Sabrosky 1999:209—210 [nomenclature]. Nannodastia horni, male. 20, Genitalia, lateral view. 21, Same, ventral view. 22, Epandrium, Diagnosis.—Nannodastia is distin- guished from Azorastia by the following combination of characters: Wing: crossvein dm-cu lacking (Fig. 17); vein M attenuate apically, not extended to wing margin; cos- tal vein continued past apex of vein R,,, but not to vein M. Male terminalia: synter- gosternite 7+8 greatly reduced (Figs. 18— 19), almost lacking, only a thin, half ring in membrane between tergite 6 and epan- drium; epandrium with a posteroventral process, bearing a thick thornlike structure apically (Figs. 13, 20); anterior portion of surstylus bearing 3 long setae; posterior portion with a digitiform process with 3 thornlike setae apically; hypandrium less dorsally placed on epandrium (Fig. 19), shorter, anterior portion robust with ven- trally directed, dilated process. Female cerci bearing short setae only (Figs. 24—27). 346 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ty Thy ply Ly ine Figs. 24-27. Nannodastia horni, female. 24, Abdomen, dorsal view. 25, Postabdomen, ventral view. 26, Posterior end of abdomen, dorsal view. 27, Same with cerci flattened, dorsal view. Scales: 0.2 mm for Fig. 24; 0.1 mm for Figs. 25-27. Egg (from abdomen of N. atlantica par- atype female) comparatively large, up to 0.25 mm (length of female abdomen about 0.55 mm). Micropylar end (Fig. 14) with a pair of small thin and pointed processes. Chorion with curved or meandering longi- tudinal grooves of uneven length (as in some Ephydridae). KEY TO SPECIES OF THE GENUS NANNODASTIA HENDEL 1. Epandrial process much shorter than posterior process of surstylus (Figs. 12-13); thick setae on anterior portion of surstylus on projections (particularly anterior seta); surstylar process somewhat slender and long (Figs. 12—13); hy- pandrium short, not much longer than height Oepandrumiee eee N. atlantica, new species VOLUME 103, NUMBER 2 — Epandrial process nearly as long as posterior process of surstylus (Figs. 18-22); thick setae on anterior part of surstylus sessile (Figs. 20— 23); surstylar process slightly thicker than in N. atlantica; hypandrium nearly twice as long as height of epandrium ....... N. horni Hendel Nannodastia atlantica L. Papp and Mathis, new species (Figs. 12-15) Nannodastia atlantica L. Papp and Mathis 1998:312 [nomen nudum, figs. of female terminalia]. Description.—A minute brown fly: length of body 0.86—-1.05 mm (holotype 0.86 mm); wing length 0.87—0.96 mm (ho- lotype 0.87 mm), wing width 0.38-0.41 mm (holotype 0.38 mm). Very similar to N. horni in having 3 pairs of long parafacial setae and several shorter peristomal setae; 1 pair of minute postocel- lar setulae just behind posterior ocelli; 2 an- terior pairs of fronto-orbitals distinct though very short; acrostichal setulae arranged in 2 mostly regular rows; second/third costal sections 0.28/0.33 mm, ratio 0.84 (holo- type), 0.29/0.38 mm, ratio 0.77 (1 female paratype); halter dark brown. So far as we can determine, the only di- agnostic characters are in the male genita- lia: epandrial process much shorter than posterior process of surstylus (Figs. 12—13); thick setae on anterior portion of surstylus on stalks (particularly the anterior seta); surstylar process somewhat slender and long (Figs. 12-13); hypandrium short, not much longer than height of epandrium (see key). The armature of the female cerci is virtually identical to that of N. horni (Figs. 15, 24-27). Type material—The holotype ¢ is la- beled’ “BELIZE. Stann Creek District: Glover’s Reef, Long Cay[,] 27-28 July 1989[,] Wayne N. Mathis.’’ The holotype is double mounted (minuten in block of plas- tic), is in good condition (eyes collapsed; abdomen removed, dissected, and stored in an attached microvial), and is deposited in the USNM. Four paratypes (3 2 USNM, 1 347 ? HNHM) bear the same label data as the holotype. Other paratype: BARBADOS. Christ Church: Bockley Beach (13°4.4’N, 59°35.3'W), 1 Sep 1997, W. N. Mathis (1 2? USNM). Distribution.—Neotropical: Belize, West Indies (Barbados). Etymology.—This species, occurring in Belize and on Barbados and possibly on other seashores of the Atlantic Ocean, is named atlantica. Nannodastia horni Hendel (Figs. 16—27) Nannodastia horni Hendel 1930:70.—Frey 1958:40—41 [discussion].—Sabrosky and Wirth 1956:18 [list as Nannodastia sp, Hawaii, Oahu]; 1959:109-110 [discus- sion, Ephydridae in tribe Atissini].—L. Papp, 1980:426.—Pitkin 1989:600 [Aus- tralasian and Oceanian catalog, in Chy- romyidae].—Mathis 1998:6—7 [list, Hen- derson Island]. Material examined.—Holotype @ and 1 additional 2 (DEI): for data see L. Papp (1980:426). Newly collected material: AUS- TRALIA. Christmas Island (10°30’S, 105°34’E): near Hosnie Spring, Oct 1983, L. Hill (2. 2; AM). HENDERSON ISLAND (24°04’S, 129°07'W). North Beach, 17—20 May 1987, W. N. Mathis (4 6, 9 2; USNM); Northwest beach, 17-20 May 1987, W. N. Mathis (6 6, 3 2; USNM). PITCAIRN IS- LAND (24°04’S, 130°06’W), Down Rope, 25 May 1987, W. N. Mathis (3 6; USNM). HA- WAII. Oahu: Kailua (at window near beach), 1 Jun 1946, W. W. Wirth (4 2; USNM). PAPUA NEW GUINEA. Central: Hisiu (swept from coastal Pandanus), 13 Jul 1986, J. W. Ismay (2 3; AM). Distribution.—Australasian/Oceanian: Christmas Island (new record), Hawaii (Oahu), Henderson Island, Papua New Guinea (new record), Pitcairn Island (new record). Oriental: Taiwan. This species is probably widespread in the Pacific basin along maritime shores in tropical and sub- tropical zones. 348 Remarks.—The specimens from the Pit- cairn Group of islands were collected in caves and under cliff overhangs above the beach and on the cliff face. ACKNOWLEDGMENTS We are grateful to Dr. Rauno Vaisanen (Division of Zoology, Zoological Muse- um of the University (ZMH), Helsinki, Finland) for the loan of the type series of A. minutissima Frey. Other specimens studied herein are from the Australian Museum, Sydney, Australia (AM; Dr. Da- vid K. McAlpine); Deutsches Entomolo- gisches Institut, Eberswalde, Germany (DEI; Dr. Joachim Ziegler); Hungarian Natural History Museum, Budapest, Hun- gary (HNHM); National Museum of Nat- ural History, Smithsonian Institution, Washington, D.C., USA (USNM); and Tel- Aviv University, Tel-Aviv, Israel (TAU; Dr. Amnon Freidberg). We are grateful to Dr. Carles-Tolra for making the female paraty- pe of A. gemmae available for dissection, study, and illustration. For reviewing a draft of this paper, we thank Drs. Allen L. No- trrbom, Amnon Freidberg, Richard S. Zack, and Stephen D. Gaimari. This is contribution number 593 from the Caribbean Coral Reef Ecosystems (CCRE), Smithsonian Institution, which is_ partly supported by a grant from the Exxon Cor- poration. LITERATURE CITED Carles-Tolra, M. 1994. Azorastia gemmae: a new nan- nodastiid species from the Isle of Ibiza (Spain, Balearic Isles) (Insecta: Diptera: Nannodastiidae). Reichenbachia 30(34): 199-202. . 1995. Nuevas citas de dipteros acalipteros para las Islas Baleares, junto con una lista preli- minar de once familias (Diptera: Acalyptrata). Boletin de la Asociacion Espanola de Entomolo- gia 19(1—2): 253-265. Cogan, B. H. 1984. Family Ephydridae, pp. 126-176. In Soos, A. and L. Papp, eds., Catalogue of Pa- laearctic Diptera, Vol. 10. Elsevier Science Pub- lishers, Amsterdam, and Akadémiai Kiad6, Bu- dapest, 402 pp. Colless, D. H. and D. K. McAlpine. 1991. Diptera (flies), pp. 717-786. In The insects of Australia. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON A textbook for students and research workers, 2nd ed. Vol. 2. Melbourne University Press, Mel- bourne, 1137 pp. Frey, R. 1945. Tiergeographische Studien tiber die Dipteren der Azoren. Societas Scientiarum Fen- nica. Commentationes Biologicae 8(10): 1-114. . 1958. Zur Kenntnis der Diptera brachycera p-p. der Kapverdischen Inseln. Societas Scientia- rum Fennica. Commentationes Biologicae 18(4): 1-61. Griffiths, G. C. D. 1972. The phylogenetic classifica- tion of Diptera Cyclorrhapha, with special refer- ence to the structure of the male postabdomen. The Hague, Junk ed., Series Entomologica 8: 1— 340. Hendel, E 1930. Eine neue interessante Ephydriden- gattung (Dipt.). Konowia 9(1): 66—70. Mathis, W. N. 1989. Diptera (Insecta) or true flies of the Pitcairn Group (Ducie, Henderson, Oeno, and Pit- cairn Islands). Atoll Research Bulletin 327: 1—15. Mathis, W. N. and T. Zatwarnicki. 1998. Chapter 49. Family Ephydridae, pp. 537-570. In Papp, L., and Béla Darvas, eds., Contributions to a Manual of Palaearctic Diptera (with special reference to flies of economic importance), Vol. 3. Higher Brachy- cera. Science Herald, Budapest, 880 pp. Papp, L. 1980. New taxa of the acalyptrate flies (Dip- tera: Tunisimyiidae fam. n., Risidae, Ephydridae: Nannodastiinae subfam. n.). Acta Zoologica Hun- garica 26(4): 415-431. Papp, L. and W. N. Mathis. 1998. Chapter 27. Family Nannodastidae, pp. 309-314. In Papp, L., and Béla Darvas, eds., Contributions to a Manual of Palaearctic Diptera (with special reference to flies of economic importance), Vol. 3. Higher Brachy- cera. Science Herald, Budapest, 880 pp. Pitkin, B. R. 1989. 95. Family Chyromyidae, p. 600. In Evenhuis, N. L., ed., Catalog of the Diptera of the Australasian and Oceanian Regions, Bishop Museum Press and E.J. Brill, 1155 pp. Sabrosky, C. W. 1956. Additions to the knowledge of the Old World Asteiidae (Diptera). Revue Fran- caise d’Entomologie 23: 216—243. . 1999. Family-group names in Diptera. Myia 10: 1-576. Sabrosky, C. W. and W. W. Wirth. 1956. Notes and exhibitions. Proceedings of the Hawaiian Ento- mological Society 16: 18. . 1959. A Formosan ephydrid new to Hawaii (Diptera: Ephydridae). Notulae Entomologicae 38(1958): 109-110. Yeates, D. K. and B. M. Wiegmann. 1999. Congruence and controversy: Toward a higher-level phylogeny of Diptera. Annual Review of Entomology 44: 397-428. Zatwarnicki, T. 1992. A new classification of Ephy- dridae based on phylogenetic reconstruction (Dip- tera: Cyclorrhapha). Genus 3(2): 65-119. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 349-355 TEXAS CAENINAE (EPHEMEROPTERA: CAENIDAE), WITH DESCRIPTION OF A NEW SPECIES W. P. MCCAFFERTY AND J. R. DAVIS (WPM) Department of Entomology, Purdue University, West Lafayette, IN 47907 (e- mail: pat-mccafferty@entm.purdue.edu); (JRD) Texas Natural Resource Conservation Commission, P.O. Pox 13087, MC 165, Austin, TX 78711 (e-mail: jdavis@tnrcc.state. tx.us) Abstract.—The subfamily Caeninae is examined with respect to its distribution within the state of Texas. Texas fauna includes Amercaenis ridens (McDunnough), Caenis amica Hagen, C. arwini, new species, C. bajaensis Allen and Murvosh, C. hilaris (Say), C. diminuta Walker, C. latipennis Banks, and C. punctata McDunnough. The report of C. diminuta represents a new state record. Some 26 new county records are provided for all species except C. bajaensis, and new Texas biotic province records are established for C. amica (Chihuahuan), C. arwini (southwestern Balconian), C. diminuta (Austroriparian and Texan), and C. hilaris (Kansan). Species ranges within and without the state are discussed. The highly distinctive C. arwini is described from a substantial series of larvae that demonstrate structural affinities primarily with C. bajaensis. Key Words: miayflies, Texas, Caeninae The only broad-scale account of the Ephemeroptera fauna of Texas has been that of Lugo-Ortiz and McCafferty (1995), where close to 100 now valid species were treated, including five species of Caeninae (family Caenidae). Four of those five Caen- inae species had been originally recorded from Texas by Provonsha (1990) in his re- vision of the genus Caenis Stephens in North America, and in fact the Caeninae had been entirely unrecorded in Texas prior to 1990. Lugo-Ortiz and McCafferty (1995) first reported a species of Amercaenis Pro- vonsha and McCafferty from Texas, and one additional species of Caenis was sub- sequently reported from Texas by Baum- gardner et al. (1997). Numerous samples of larvae of Caenis and Amercaenis—the only North American members of the subfamily Caeninae of the family Caenidae (see McCafferty and Wang 2000)—were taken from 19 counties in var- ious regions of Texas between 1975 and 1997 by personnel associated with the Tex- as Natural Resource Conservation Commis- sion or its predecessor agencies. Eight spe- cies were represented in the samples, in- cluding a distinctive new species of Caenis described herein, as well as a new state re- cord for C. diminuta. Of previously known Texas species, only C. bajaensis was not represented in the studied samples. In ad- dition to the new species and new state re- cord, 23 new county records were associ- ated with the six species of Caeninae pre- viously known from Texas. Herein we provide distributional data for each of the eight species of Caeninae now known from Texas, including brief refer- ences to the previous records (Provonsha 1990, Lugo-Ortiz and McCafferty 1995, Baumgardner et al. 1997, Wiersema and 350 McCafferty 1999) and new records detailed in full; brief remarks concerning the ranges of each species within and without the state; and the description of the new species as well as a comparison of it with other North American species of Caenis. Material ex- amined of all species are made up of larvae housed in both the J. R. Davis personal col- lection and the Purdue Entomological Re- search Collections. Biotic provinces within the state that we refer to are after Blair (1950) (see also Lugo-Ortiz and Mc- Cafferty 1995). Preliminary species identi- fications were verified by A. V. Provonsha. Amercaenis ridens (McDunnough) Previous records.—Lugo-Ortiz and McCafferty (1995): Austin Co. New records.—Colorado Co., San Ber- nard R. at FM 3013, VII-17-1997, J. R. Da- vis. Remarks.—This mainly north-central and northeastern North American species is known from the Austroriparian and Texan biotic provinces of eastern Texas. It should also occur in the Great Plains region of the Kansan biotic province (panhandle of Tex- as) because it is known typically to inhabit sandy bottom rivers in the plains states (Provonsha and McCafferty 1985). Caenis amica Hagen Previous records.—Provonsha (1990): Austin Co., Jasper Co., and Orange Co.; Lugo-Ortiz and McCafferty (1995): Gua- dalupe Co., Travis Co., and Victoria Co.; Baumgardner et al. (1997): Hardeman Co. and Uvalde Co. New records.—Presidio Co., Rio Grande, 10.5 km upstream from Presidio, XII-30- 1975, J. R. Davis. Remarks.—The new county record of this widespread species also represents a first record for the Chihuahuan biotic prov- ince of far western Texas. The species is now known from all six biotic provinces within the state. The new Rio Grande re- cord, being shared with Mexico, also serves as a new Mexican record, although the re- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON cent report of C. punctata from Nuevo Leon, Mexico by Wiersema and Baumgard- ner (2000), is possibly attributable to C. amica (see also remarks under C. punctata, below). Larvae of the two species may be easily confused (Provonsha, personal com- munication). Caenis arwini McCafferty and Davis, new species (Figs. 1—4) Larva.—Body length 3.0—4.0 mm. Head: Dorsal color pattern often as in Fig. 1; pair of relatively large more or less light spots well defined posterior to lateral ocelli; single large, light, ovate spot bordering pos- terior median stalk of epicranial suture; and single light spot anterior of epicranial su- ture branches between antennal bases. An- tenna generally pale. Thorax: Dorsum with light to medium brown ground color (Fig. 1). Venter pale, cream, in some slight in- dication of light brown on _ prosternum. Pronotum (Fig. 1) with pale lateral borders. Mesonotum (Fig. 1) with conspicuous light spots, most with rounded margins (spots sometimes less apparent on wingpads). All legs pale except for brown coxal bases, and in some slight indication of distal-posterior, light brown dash on femora (Fig. 1). Fore- femur (Figs. 1-2) with transverse row of spatulate setae in distal half (about one third distance from apex); posterior edge with moderately stout, sharp setae intermixed with short, fine, simple, hairlike setae; an- terior edge with short, fine, simple, hairlike setae in basal half. Inner margin of foretibia (Fig. 2) with rows of marginal and sub- marginal stout setae. Hindtarsus (Fig. 3) with row of 8—10 stout, sharp, simple setae and row of 8—10 stout, fimbriate setae along inner (ventral) margin. Hindclaw with 15— 20 minute marginal denticles. Abdomen: Tergum 1 usually light brown throughout; terga 2—9 with broad, pale lateral margins; terga 7—9 with broad, light, medial area; ter- gum 10 pale medially and light brown lat- erally. Sterna pale; sternum 9 (Fig. 4) with broadly notched distal margin (somewhat VOLUME 103, NUMBER 2 Fig. 1. Caenis arwini, larval habitus. 352 ) S x = of Pulee Pe a — Ze Wao pean NX Q a = NS NN ‘N Figs. 2—4. 9 (posterior margin). V-shaped medial emargination). Operculate gills light to medium brown and usually with conspicuous light spotting similar to, or approaching that, shown in Figure 1; se- tae on dorsal Y-ridge restricted to basal half of inner branch. Caudal filaments (Fig. 1) pale, with whorls of short setae at every other segmental joining. Adult.—Unknown. Egg.—Ovate with one or two polar caps restricted to ends of egg; polar caps covered by thick membrane and made up of long coiled threads [see type II eggs in Caenis discussed by Provonsha (1990)]. Material examined.—Holotype: Mature larva, TEXAS, Val Verde Co., San Filipe Cr. at Academy St. in Del Rio (pool), X-8- 1996, J. R. Davis (deposited in PERC, the Purdue Entomological Research Collection, West Lafayette, Indiana). Paratypes: Six mature larvae, same data and deposition as holotype; an additional six mature larvae, Val Verde Co., San Felipe Cr. at US 277 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 3 Caenis arwini, larva. 2, Foreleg (dorsal). 3, Hindtarsus and claw (lateral). 4, Abdominal sternum (pool), X-7-1996, J. R. Davis (deposited in PERC). Additional material: 45 variously mature larvae, same data as types and de- posited in PERC or J. R. Davis personal collection. Etymology.—We are pleased to name this species after Arwin Provonsha, the foremost American authority on the genus Caenis. Remarks.—The overall dramatic color pattern of the larva of C. arwini is unique among North American Caenis. Although C. hilaris may also have a distinctively mottled color pattern [see Fig. 5 in Provon- sha (1990)], it does not have the almost uni- formly rounded light spots that appear on the dorsum of C. arwini. Also C. arwini larvae lack the sublateral dark dashes on the abdominal terga that are found on many C. hilaris larvae. Structurally, the larvae of C. arwini and C. hilaris are highly dissimilar. The short whorls of setae associated with only every other segmental joining on the VOLUME 103, NUMBER 2 caudal filaments of C. arwini are shared only with C. anceps and C. bajaensis. All other North American Caenis larvae have long lateral setae associated with every seg- mental joining on the caudal filaments (Pro- vonsha 1990). The caudal filament setation, leg armature of the larvae, and the charac- terization of the polar caps of the eggs, all of which C. arwini shares with C. bajaen- sis, Would appear to suggest some close re- lationship of the two species, but only if these traits prove to be apomorphic. Caenis arwini larvae, however, are easily distin- guished from those of C. bajaensis by both the color pattern, as elaborated above, and the more broadly notched abdominal ster- num 9 [compare Fig. 4 herein with Fig. 45 of Provonsha (1990)]. If C. arwini larvae were being keyed out with use of the Provonsha (1990: 810) key, they would come out to couplet 3, along with C. anceps and C. bajaensis. The notch of abdominal sternum 9 would tend to place C. arwini with C. anceps, whereas the lon- ger fimbriate setal row of the hindtarsi would place C. arwini with C. bajaensis. Thus, the unique combination of the these two latter characteristics will diagnose C. arwini from the other two species. How- ever, the color pattern differences of the operculate gills alone could be used to im- mediately distinguish the three species. Caenis bajaensis Allen and Murvosh Previous records.—Baumgardner et al. (1997): Jeff Davis Co. and Culberson Co. Remarks.—We doubt that this primarily southwestern USA and northern Mexico spe- cies ranges east of far western Texas (the Chi- huahuan biotic province) based on its present distributional pattern (see McCafferty et al. 1997); however, it has been commonly taken to the north in small streams of far western Nebraska (Provonsha 1990). Caenis diminuta diminuta Walker New records.—Titus Co., Tankersky Cr. at FM 899, W Mt. Pleasant, [X-9-1997, J. R. Davis; Wharton Co., West Bernard Cr. 353 at county road off CR 252, ESE Lissie, VII- 16-1997; Wharton Co., West Mustang Cr. at CR 378, IX-17-1997, J. R. Davis. Remarks.—This species was taken from the Austroriparian biotic province (far east- ern Texas) and the southern part of the Tex- an biotic province (eastern Texas). This dis- tribution is consistent with the fact that C. diminuta diminuta is a northeastern and southeastern North American subspecies. The species is known from Central America (McCafferty and Lugo-Ortiz 1992) as the subspecies C. diminuta latina McCafferty and Lugo-Ortiz, but the species has yet to be taken from Mexico, where we expect it to range at least along the gulf corridor. Caenis hilaris (Say) Previous records.—Provonsha (1990): Austin Co. and Victoria Co.; Lugo-Ortiz and McCafferty (1995): Kimble Co. and Val Verde Co. New records.—Bosque Co., N. Bosque R. at Hwy 22 in Meridian, I[V-25-78, D. Pe- trick; Colorado Co., San Bernard R. at FM 3013, VII-18-1997, J. R. Davis; Jackson Co., Arenosa Cr. at county road off Hwy 59, 3.5 mi N Inez, I[X-6-1988, S. Twidwell: Lipscomb Co., Wolf Cr. at FM 1454 E Lip- scomb, VII-18-1990, S. Twidwell; Mason Co., James R. at private road off FM 2389 SW Mason, VII-23-1987, S. Twidwell; Montgomery Co., Lake Cr. at private road off FM 1488 SE Conroe, VII-20-1988, S. Twidwell; Wheeler Co., Sweetwater Cr. at Oklahoma-—Texas Line Road, [X-9-1987, S. Twidwell; Wilson Co., Cibolo Cr. at FM 537 SW Stockdale, V-8-1980, J. R. Davis. Remarks.—This widespread species is re- ported for the first time from the Kansan bi- otic province in Texas, and thus is now known from all six biotic provinces in the state. Caenis latipennis Banks Previous records.—Provonsha (1990): Austin Co., Jasper Co., La Salle Co., Palo Pinto Co., and Webb Co.; Lugo-Ortiz and McCafferty (1995): Blanco Co., Kerr Co., 354 Kimble Co., and Travis Co.; Baumgardner et al. (1997): Reeves Co.; Wiersema and McCafferty (1999): Victoria Co. New records.—Bosque Co., N. Bosque R. at Hwy 22 in Meridian, IV-25-78, D. Pe- trick; Cameron Co., Rio Grande 0.3 km downstream from El] Jardin intake SE Brownsville, I-14-1993, J. R. Davis; Don- ley Co., Lelia Lake Cr. at FM 2471 NE Le- lia Lake, VIII-17-1989, S. Twidwell; Harris Co., Greens Bayou at Hardy Road in Al- dine, VI-27-1995, J. R. Davis; Lipscomb Co., Wolf Cr. at FM 1454 E Lipscomb, VII- 18-1990, S. Twidwell; Mason Co., James R. at private road off FM 2389 SW Mason, VII-23-1987, S. Twidwell; Newton Co., Big Cypress Cr. at SH 87 NW Deweyville, [X- 12-1989, S. Twidwell; Val Verde Co., San Felipe Cr. at US 277 in Del Rio (pool), X- 7-1996, J. R. Davis; Wheeler Co., Sweet- water Cr. at Oklahoma—Texas Line Road, IX-9-1987, S. Twidwell. Remarks.—This widespread species has been known from all of the biotic provinces in Texas, and the new county records fur- ther demonstrate its somewhat ubiquitous nature. Caenis latipennis is also known from all of the Mexican states bordering Texas (Randolph and McCafferty 2000). Caenis punctata McDunnough Previous records.—Provonsha (1990): Jasper Co.; Lugo-Ortiz and McCafferty (1995); Kimble Co., Lavaca Co., Mason Co., Travis Co., and Victoria Co.; Baum- gardner et al. (1997): Crosby Co. and Tom Green Co. New records.—Bosque Co., N. Bosque R. at Hwy 22 in Meridian, [V-25-78, D. Pe- trick; Galveston Co., Dickinson Bayou near American Canal, SE League City, X-5- 1987, S. Twidwell; Harris Co., Greens Bay- ou at Hardy Road in Aldine, VI-27-1995, J. R. Davis; Harris Co., Buffalo Bayou at Shepherd Dr. in Houston, VII-17-1985, J. R. Davis; Jackson Co., Arenosa Cr. at coun- ty road off U.S. 59, 3.5 mi N Inez, IX-6- 1988, S. Twidwell; Randall Co., Prairie PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Dog Town Fork Red R. below Tanglewood Dam, NE Canyon, V-13-1992, S. Twidwell. Remarks.—This northeastern and south- eastern North American species is known from all biotic provinces in Texas except the Tamaulipan (far south) and Chihuahuan (far west). If it occurs in Mexico, it more than likely dispersed southward via the Gulf of Mexico Maritime Corridor (see Lugo-Ortiz and McCafferty 1995). ACKNOWLEDGMENTS We greatly appreciate assistance from A. V. Provonsha (West Lafayette, Indiana), for confirming and in some cases correcting preliminary identifications of Caeninae lar- vae from Texas, and for the preparation of figures used herein. Research was support- ed in part by NSF grant DEB-9901577 to WPM. This paper has been assigned Purdue University Agricultural Research Program Journal No. 16275. LITERATURE CITED Baumgardner, D. E., J. H. Kennedy, and B. C. Henry, Jr. 1997. New and additional records of Texas mayflies (Insecta: Ephemeroptera). Transactions of the American Entomological Society 123; 55— 69. Blair, W. E 1950. The biotic provinces of Texas. Texas Journal of Science 2: 93-117. Lugo-Ortiz, C. R. and W. P. McCafferty. 1995. The mayflies (Ephemeroptera) of Texas and their bio- geographic affinities, pp. 151-169. Jn Corkum, L. D. and J. J. H. Ciborowski, eds., Current Direc- tions in Research on Ephemeroptera. Canadian Scholars’ Press, Toronto. McCafferty, W. P. and C. R. Lugo-Ortiz. 1992. Regis- tros nuevos y notas los Ephemeroptera de Nica- ragua. Revista Nicaragiiense de Entomologia 19: 1-7. McCafferty, W. P. and T.-Q. Wang. 2000. Phylogenetic systematics of the major lineages of pannote may- flies (Ephemeroptera: Pannota). Transactions of the American Entomological Society 126: 9-101. McCafferty, W. P., C. R. Lugo-Ortiz, and G. Z. Jacobi. 1997. Mayfly fauna of New Mexico. The Great Basin Naturalist 57: 283-314. Provonsha, A. V. 1990. A revision of the genus Caenis in North America (Ephemeroptera: Caenidae). Transactions of the American Entomological So- ciety 116: 801-884. Provonsha, A. V. and W. P. McCafferty. 1985. Amer- VOLUME 103, NUMBER 2 caenis: new Nearctic genus of Caenidae (Ephem- eroptera). International Quarterly of Entomology 1: 1-7. Randolph, R. P. and W. P. McCafferty. 2000. Mexican mayflies: Inventory and additions. Annales de Limnologie 36: 113-121. Wiersema, N. A. and D. E. Baumgardner. 2000. Dis- 855 tribution and taxonomic contributions to the Ephemeroptera fauna of Mexico and Central America. Entomological News 111: 60—66. Wiersema, N. A. and W. P. McCafferty. 1999. Ameri- cabaetis (Ephemeroptera: Baetidae) from Texas: first USA record and adult description of A. ple- turus. Entomological News 110: 36-38. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 356-363 A NEW FAMILY, GENUS, AND SPECIES OF SCALE INSECT (HEMIPTERA: COCCINEA: KUKASPIDIDAE, NEW FAMILY) FROM CRETACEOUS ALASKAN AMBER JAN KOTEJA AND GEORGE O. POINAR, JR. (JK) Department of Zoology and Ecology, Cracow Agricultural University, al. Mick- iewicza 24, 50-059 Krakow, Poland. (e-mail: rzkoteja@cyf-kr.edu.pl); (GOP) Department of Entomology, Oregon State University, 2046 Cordley Hall, Corvallis, Oregon 97331- 2907, U.S.A. (e-mail: poinarg @bcc.orst.edu) Abstract.—A new genus and species of scale insect, Kukaspis usingeri is described from Cretaceous Alaskan amber and placed in a new extinct family, the Kukaspididae. This fossil is a derived member of the superfamily Orthezioidea, with six pairs of uni- corneal eyes forming lateral rows, a scutum with a large subrectangular membrane, a tubular scutellum separated from the mesopostnotum by a large membrane, wings narrow with a clear posterior (claval) flexing line, but a reduced anterior one, narrow parallel- sided halteres; a unique waxy tail consisting of four soft filaments arising from the last abdominal tergite and a penial sheath divided into basal capsule and stylus with a hook- like apex. Relationships of this peculiar Lower Cretaceous form with both extant and extinct forms are discussed. Key Words: ceous Alaskan amber Alaskan amber is perhaps the most in- triguing and least known of the Cretaceous fossiliferous ambers. Its outcrops between the Brooks Range and northern coastline comprising the Arctic Coastal Plain and Foothills were discovered and explored by R. Usinger and R. Smith (see Poinar 1992 for further discussion and early exploration of the Alaskan amber deposits). While there was originally some question whether the amber deposits originated from the Upper or Lower Cretaceous, the most recent study places the Kuk deposits as Late to Middle Albian (97-104 million years) (Scott and Smiley 1979). Chemical analysis of the am- ber indicates that the plant source was a member of the Araucariaceae, probably Agathis (Lambert et al. 1990). The coccid male described in this paper fossil scale, Hemiptera, Orthezioidea, Coccinea, Kukaspis usingeri, Creta- is only the second description of an organ- ism in Alaskan amber, the first being a cer- aphronid wasp, the present whereabouts of which is unknown (Muesebeck 1963). Some insect groups from Alaskan amber were mentioned in the reports of Hurd et al.(1958), namely empidid flies, an eulo- phid wasp and an adelgid, and Langenheim et al. (1960) reported ceratopogonid and empidid flies and an eulophid and cera- phronid wasp (the latter fossil presumably the one later described by Muesebeck 1963). Studies on Cretaceous scale insects were initiated by Beardsley (1969) on Upper Cretaceous Canadian (Cedar Lake) amber. Further Upper Cretaceous coccids have been described from Siberian (Taimyrian) (Koteja 1989a) and New Jersey amber (Ko- VOLUME 103, NUMBER 2 teja 2000). Lower Cretaceous impressions have been found in Siberia and southern England (Koteja 1988, 1989b, 1999). Un- described scales also occur in Cretaceous Lebanese, Canadian, and Burmese amber (Schlee 1972; Poinar 1992, 1998; Pike 1994; Rasnitsyn 1996; D. Azar, D. Grimal- di, A. Ross, personal communications), thus scale insects appear to be quite abundant in Cretaceous deposits. The present study deals with a coccid male found in Alaskan amber, first pub- lished by Poinar (1998), as a “‘male scale insect.”” The specimen appears to be a pe- culiar archeococcid (Orthezioidea), but with some derived features characteristic of neo- coccids (Coccoidea). The combination of characters exhibited by the male does not fit any established group, thus a new extinct family is created to contain it. MATERIAL AND METHODS The present fossil was collected by R. Usinger and R. Smith from Cretaceous out- crops on the West side of Kuk Inlet at Pug- nik Beach, in 1955, which occurs between Brooks Range and the northern Alaskan coastline. The amber piece is flat, ca. 9 mm long by 8 mm wide, transparent, yellowish brown with a dark brown film separating subsequent resin flows which is also char- acteristic of Canadian and Lebanese amber. Except for the coccid, no other organic or inorganic inclusions are in the piece. At the present state of preparation, the inclusion can be observed only from its right side and many structural details are difficult to see because of the thick layer of resin overlying the specimen, however further polishing could damage the specimen. The inclusion is well preserved and not obscured by im- purities or bubbles. The dark color of the specimen makes observation difficult and some details cannot be seen despite the use of high light intensity. Sclerotized and membranous body parts can clearly be dis- tinguished, the former being black and the latter silver in direct light. Ordinary light Si5)// microscopes were used to examine and draw the inclusion. The thick layer of resin limited the use of objectives higher than 16X (ca. 160 combined magnification). Descriptive terms for the males as well as other morphological details were taken from Theron (1958). Kukaspididae Koteja and Poinar, new family Type genus.—Kukaspis, new genus. Etymology.—The family name refers to the Kuk Inlet in which the amber was found. Description.—Archeococcids (Orthezioi- dea) with dinintegrated compound eyes; single ommatidia forming oblique lateral rows; spherical pedicel; cylindrical flagellar segments bearing short setae; scutum with large subrectangular membrane; scutellum short, tubular; wings membranous, subcos- tal and cubital ridges weak, anterior flexing line reduced, posterior one present; halters narrow; all trochanters with numerous se- tae; abdomen with four soft waxy filaments arising from last tergite; penial sheath formed of basal capsule and stylus with hooked apex; aedaegus sickle-like, acute; structure of tail organs unique in scale in- sects; with an apically hooked penial sheath which is shared with the Putoidae. Kukaspis Koteja and Poinar, new genus Type species.—Kukaspis usingeri, new species. Description.—Archeococcid (Orthezioi- dea) with a circle of 10—12 unicorneal eyes; pedicel short, cylindrical flagellar segments, antennal setae short, capitate setae absent; scutum oval, large rectangular scutal mem- brane very short, tubular scutellum with an oval foramen; mesopostnotum separated from scutellum by large membrane; wings elongate-oval, finely sculptured; microtri- chia absent; subcostal and cubital ridges weak with only posterior (claval) flexing membrane developed; halters narrow, with 2 setae; legs short, all trochanters with nu- merous setae, tarsal and ungual digitules 358 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON capitate; tail organs consisting of four soft waxy filaments arising from last abdominal tergite; penial sheath with a globose basal capsule and stylus, latter with a hook-like extension dorsally; aedaegus acute, extend- ing beyond apex of penial sheath. Etymology.—The generic name refers to The Kuk river, Alaska. Kukaspis usingeri Koteja and Poinar, new species (Figs. 1—9) Holotype.—Alate male Cocc-876 inclu- sion in Alaskan amber, deposited in the col- lection of G. Poinar (HO-4-30) maintained at Oregon State University, Lower Creta- ceous, Kuk Inlet at Pugnik Beach between Brooks Range and the northern Alaskan coastline. Etymology.—The species is named in honor of Dr. Robert Usinger, one of the first to realize the scientific value of Alaskan amber. Description.—Only holotype available; accessible from right side, but some dorsal and ventral structures also can be examined. Sclerotized body parts dark brown. Body 1.32 mm long; wing expanse 3.20 mm; head 120 pm long, 180 pm high, with strong ocular sclerites bearing 6 (possibly only 5) pairs of simple (unicorneal) eyes in oblique lateral rows and a pair of ocelli; dorsum of head seems completely membra- nous; a few short setae recognizable at out- line of head; antenna filiform, 10-segment- ed, 1,170 wm long, slightly shorter than body; scape difficult to define; pedicel glo- bose, 65 jm in diameter; flagellar segments cylindrical, ca 35 wm wide (some flattened after preservation), terminal segment with acute apex; first flagellar segment 90 pm, others 140-160 pm long, apical segment longest; all segments bearing numerous se- tae about as long as segment width, includ- ing hair-like and thick setae; antennal bris- tles recognized at terminal segments but number and position difficult to ascertain; capitate setae apparently absent; other types of sensilla not recognized. Thorax 510 wm long, 360 wm deep. Prothorax with large bulged post-tergites; scutum perhaps oval, medial membrane very large, subrectangu- lar; scutellum cylindrical, subrectangular in dorsal view, very short (40 wm), with an oval foramen; mesopostnotum well devel- oped, separated from scutellum by a large membrane, overlapped with a metathoracic fold; mesosternum strongly bulging; a few short setae noted on various parts of thorax. Legs (in oblique positions on drawing) rel- atively short, 890 wm long, slender; coxae 110 wm; trochanters+femora 300 pm, tib- iae 280 wm, tarsi 110 wm, claws 30 pm; all segments (including trochanter) with nu- merous short straight setae, some spine- like; tibiae with few apical spurs; claw nar- row but with broad base, without denticles; tarsal and ungual digitules knobbed, 35 wm and 25 pm respectively. Wings 1,500 pm long, 600 wm wide, with fine granulate sculpture; presence of microtrichia prob- lematical, not recognized on wing margin where usually best visible; subcostal and cubital ridges weakly sclerotized; anterior flexing line absent, posterior (claval) one distinct and represented by a membranous patch which does not reach wing margin (an associated trachea visible on right wing); pouch for holding halteral setae elongate, strongly sclerotized; basal alar se- tae and cupolae along subcostal ridge pre- sent, difficult to count. Haltere narrow, par- allel sided, 120 ym long, bearing 2 setae. Abdomen cylindrical (compressed as shown on Fig. 9), 695 ym long, with well developed tergal and sternal sclerites, short setae and wax glands (secretions forming a granulate coating). Tail organs peculiar, consisting of four flexible filaments (tubes?) protruding from last abdominal tergite; each apparently secreted by a single multilocular gland; particular threads arising from single loculi intertwined and forming a rope, per- haps as long as body. Penial sheath (in lat- eral view) consisting of distinct capsule and stylus, both 120 wm long, capsule 35 pm high; stylus with peculiar hooked apex (as VOLUME 103, NUMBER 2 359 Sy CSRS we Be a ieee ‘ Figs 1-5. Kukaspis usingeri, holotype. 1, Outline of body in laterodorsal view (note one of the waxy tails above the left wing). 2, Right ocular sclerite and antenna (segments 4 and 5 compressed). 3, Mid tibia and tarsus. 4, Foreleg. 5, Apex of tarsus. 360 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs 6-9. mesothorax (dorsal; note the metathoracic fold above mesopostnotum). 8, Abdomen (lateral); note four waxy tails bent headward. 9, Optical cross-section of abdomen showing dorsoventral compression. in Puto); a group of setae on ventral face of capsule, sensilla on stylus not noted. Ae- daegus acute, with a slit-like dorsal gono- pore projecting beyond apex of penial sheath. DISCUSSION Scale insects are believed to have split into two groups at an early stage of their radiation: the archeococcids (Orthezioidea) and neococcids (Coccoidea). The former Kukaspis usingeri, holotype. 6, Head (lateral) and thorax (laterodorsal). 7, Reconstruction of bear several plesiomorphic features, such as abdominal spiracles and compound eyes, but developed also specialized structures and behaviors sometimes paralleling those of the neococcids, both in extant and fossil faunas (Koteja 2000). The neococcids are descendants of the Putoidae (Koteja 1996) and are monophyletic; the ancestor of the archeococcids is unknown and their mono- phyly is not evident. Studying the phylog- eny of scale insects and relationships of VOLUME 103, NUMBER 2 particular groups based on fossil material presents two problems: some essential mi- croscopic structures are not discernable (e.g., abdominal spiracles) and some spe- cialized convergent features occur in both archeo- and neococcids, thus the assign- ment of fossil forms to one of these branch- es can be difficult. At first glance, the Alas- kan amber male resembles a neococcid, with tubular (rectangular in dorsal view) scutellum, large rectangular medial scutal membrane, well developed capitate tarsal and ungual digitules, reduced “venation” of forewing, disintegrated compound eyes, four waxy tails and other characters. How- ever, a detailed analysis of these features indicates that this species (family) must be a derived member of the archeococcids. In scale insects, the prothorax is largely membranous, with minute paired pronotal and post-tergal sclerites in some groups, the largest in the digging Margarodidae. In in- clusions, the prothorax is significantly shrunken, almost non-existent, and the me- sothoracic prescutum seems to be attached to the head. In K. usingeri, there are con- spicuous bulged sclerites on each side of the prothorax comparable in size with those in the margarodids, however, the anterior legs are of normal walking (not digging) structure. A short broad tubular scutellum, often with a foramen, is characteristic of ad- vanced neococcids (Coccidae, Diaspididae and others); however, it has been found also in the Steingeliidae and Electrococcidae among the aberrant archeococcids (Theron 1958, Koteja 2000). In neococcids, the tro- chanter bears a few setae only (perhaps a maximum of 10 in Puto), usually with a distinct long apical seta. In archeococcids there are numerous subequal setae on this joint. The conditions in K. usingeri corre- spond with those in archeococcids. The wing skeleton (subcostal and cubital ridges) is simple and weak in K. usingeri, as in the neococcids and some specialized archeococcids, but there is a distinct claval flexing line, never encountered in the neo- 361 coccids, but present in the archeococcids. Neococcid wings bear microtrichia, devel- oped also to some extent in fossil archeo- coccids (Koteja 1996). Although microtri- chia have not been observed on the wing margin (usually best visible there) of K. usingeri, there are “‘spots’”? on the mem- brane that could be interpreted as microtri- chia. However, such a pattern with micro- trichia absent is also characteristic of the Steingeliidae. The halteres are very narrow and slightly S-shaped, as in Ortheziidae (Koteja 1986). The structure of the penial sheath in Puto is characterized by its apical hook-like ex- tension, a feature unique in scale insects so far (Reyne 1954). Yet the same hook or thorn occurs in K. usingeri. However, the aedaegus in some species of Puto is bifur- cate apically but is acute in K. usingeri. Owing to some deformation of the ab- domen it is difficult to comment about the four conspicuous waxy tails in K. usingeri. They do not correspond with the dorsal glassy straight waxy tubes forming a tuft characteristic of the Ortheziidae, Matsucoc- cidae, Margarodidae and some other ar- cheococcids (well preserved in amber in- clusions and rock impressions), neither with the lateral paired waxy rods in the neococ- cids. In the latter, the rods arise from a pocket of the wall that produces the rods and are supported by two or more long se- tae inserted at the bottom of the pouch. Two pairs of rods, on two terminal abdominal segments, are produced in some Phenacoc- cinae (Pseudococcidae) and a species from New Jersey amber (Koteja 2000), one pair in all other neococcids. Among archeococ- cids, one pair of tail organs has been de- scribed in the Steingeliidae and Phenaco- leachiidae (for more details, see Koteja 1996). In K. usingeri, the four tails arise dorsally from one (the last) segment and are “soft”? and flexible; they have been bent forward in the resin. Each of the tail fila- ments seems to have been produced by a single multilocular pore or tube (see de- scription of species). They must be consid- 362 ered in the scale insects as unique tail de- vices which possibly developed through a modification of the archeococcid waxy tuft. Most important in this discussion is the structure of the head. Advanced neococcid males bear four simple (unicorneal) eyes: one dorsal and one ventral pair. Some neo- coccids have more than two pairs of eyes (Putoidae, Kermesidae, some Coccidae), the dorsal and ventral eyes being the larg- est. Archeococcids bear, as a rule, com- pound eyes, but some of them have also simple eyes (perhaps 5 to 8 pairs) forming a circle or lateral rows, but never two pairs as the neococcids. It is obvious that simple eyes are the result of the degeneration of compound eyes, and that a circle or rows of simple eyes is an intermediate degener- ative stage with eventual reductions result- ing in two pairs of disintegrated ommatidia only. It is also evident that this process of degeneration and reduction occurred many times in various groups. For instance, some Coccidae have several pairs of eyes, others only two pairs which means that further re- duction occurred within this family (Gili- omee 1967). The Kermesidae bear several pairs of simple eyes, but their close rela- tives, the Eriococcidae, possess only two pairs which indicates that the eriococcid- kermesid ancestor had many eyes, and that reduction of their number occurred in the eriococcid branch, but not in the kermesid clade (Afifi 1968, Koteja and Zak-Ogaza 1972): Among the archeococcids, reduction of eyes occurred in the following groups: Pi- tyococcidae, Steingeliidae, Phenacoleachi- idae (all extant), Electrococcidae and at least three other Cretaceous groups (Koteja, unpublished). With the degeneration of compound eyes and reduction of the num- ber of ommatidia the head capsule and es- pecially the ocular sclerites undergo signif- icant structural changes which are more striking than the reduction of the number of ommatidia. Unfortunately, it is impossible to examine all the details of the head struc- ture in fossils. In general, the head in K. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON usingeri is definitely different from that in the Putoidae, Kermesidae and Coccidae, but shows some similarity with the head of the above mentioned archeococcids. In all of them, the ocular sclerites are narrow and small compared with the large and project- ing ommatidia which, in lateral view, form oblique rows. However, considering other features, it must be assumed that the reduc- tion of eyes to simple ommatidia occurred independently in various archeococcids. In conclusion, the Kukaspididae are ar- cheococcids that developed many special- ized and some unique features and at this time it is not possible to affiliate them with any fossil or extant group. LITERATURE CITED Afifi, S. A. 1968. Morphology and taxonomy of adult males of the families Pseudococcidae and Erio- coccidae (Homoptera: Coccoidea). Bulletin of the British Museum of Natural History (Entomology) Suppl. 13, 210 pp. Beardsley, J. W. 1969. A new fossil scale insect (Ho- moptera: Coccoidea) from Canadian amber. Psy- che 76: 270-279. Giliomee, J. H. 1967. Morphology and taxonomy of adult males of the family Coccidae (Homoptera: Coccoidea). Bulletin of the British Museum of Natural History (Entomology) Suppl. 7, 168 pp. Hurd, Jr., PR. D., R. E Smith, and R. L. Usinger. 1958. Cretaceous and Tertiary insects in arctic and Mex- ican amber. Proceedings of the Tenth International Congress of Entomology (1956) 1: 851. Koteja J. 1974. On the phylogeny and classification of the scale insects (Homoptera, Coccinea) (discus- sion based on the morphology of the mouthparts). Acta Zoologica Cracoviensia 19: 267-325. . 1986. Morphology and taxonomy of male Or- theziidae (Homoptera, Coccinea). Polskie Pismo Entomologiczne 56: 323-374. 1988. Eomatsucoccus gen. n. (Homoptera, Coccinea) from Siberian Lower Cretaceous de- posits. Annales Zoologici 42: 141-163. . 1989a. Inka minuta gen. et sp. n. (Homoptera, Coccinea) from Upper Cretaceous Taymyrian am- ber. Annales Zoologici 43: 77-101. . 1989b. Baisococcus victoriae gen. et sp. n.— A Lower Cretaceous coccid (Homoptera, Cocci- nea). Acta Zoologica Cracoviensia 32: 93-106. . 1996. The scale insects (Homoptera: Cocci- nea) a day after, pp. 65-88. Jn Schaefer, C. W., ed., Studies on Hemiptera Phylogeny. Proceedings of Thomas Say Publications in Entomology; En- VOLUME 103, NUMBER 2 tomological Society of America, Lanham (Mary- land) 244 pp. . 1999. Eomatsucoccus andrewi sp. nov. (He- miptera: Sternorrhyncha: Coccinea) from the Lower Cretaceous of southern England. Creta- ceous Research 20: 863-866. . 2000. Scale insects (Homoptera: Coccinea) from Upper Cretaceous New Jersey amber, pp. 147-229. In Grimaldi, D., ed., Studies on Fossils in Amber, with Particular Reference to the Cre- taceous of New Jersey. Backhuys Publishers, Lei- den, The Netherlands, 498 pp. Koteja, J. and B. Zak-Ogaza. 1972. Morphology and taxonomy of the male Kermes quercus (L.) (Ho- moptera, Coccoidea). Acta Zoologica Cracovien- sia 17: 193-216. Lambert, J. B., J. S. Frye, and G. O. Poinar, Jr. 1990. Analysis of North American amber by Carbon-13 NMR Spectroscopy. Geoarchaeology 5: 43-52. Langenheim, R. L., Jr., C. J. Smiley, and J. Gray. 1960. Cretaceous amber from the arctic coastal plain of Alaska. Bulletin of the Geological Society of America 71: 1345-1356. Muesebeck, C. FE E. 1963. A new ceraphronid from Cretaceous amber (Hymenoptera: Proctotrupidae). Journal of Paleontology 37: 129-13. 363 Pike, E. M. 1994. Historical changes in insect com- munity structure as indicated by hexapods of Up- per Cretaceous Alberta (Grassy Lake) amber. Ca- nadian Entomologist 126: 695-702. Poinar, G. O. Jr. 1992. Life in amber. Stanford Uni- versity Press, Stanford, 350 pp. . 1998. Palaeontology of amber. Geology To- day 14: 154-160. Rasnitsyn, A. P. 1996. Burmese amber at the Natural History Museum, London. Inclusion-Wrostek 23: 19-21. Reyne, A. 1954. A redescription of Puto antennatus Sign. (Homoptera, Coccidea), with notes on Cer- oputo pilosellae Sule and Macrocerococcus su- perbus Leon. Zoologische Medelingen 32: 291— 324. Schlee, D. 1972. Bernstein aus dem Lebanon. Kosmos (Stuttgart) 68: 460—463. Scott, R. and C. J. Smiley. 1979. Some Cretaceous plant megafossils and microfossils from the Nan- ushuk Group, Northwest Alaska: a preliminary re- port. United States Geological Survey Circular 794: 89-111. Theron, J. G. 1958. Comparative studies on the mor- phology of male scale insects (Homoptera: Coc- coidea). Annals of the University of Stellenbosch 34(sec. A): 1-71. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 364-366 DISTRIBUTION OF THE WESTERN CONIFER SEED BUG, LEPTOGLOSSUS OCCIDENTALIS HEIDEMANN (HETEROPTERA: COREIDAE) IN CONNECTICUT AND PARASITISM BY A TACHINID FLY, TRICHOPODA PENNIPES (F.) (DIPTERA: TACHINIDAE) GALE E. RIDGE-O’ CONNOR Department of Entomology, Connecticut Agricultural Experiment Station, PO. Box 1106, New Haven, CT 06504-1106 U.S.A. (e-mail: gale.ridgeoconnor @ po.stae.ct.us) Abstract.—The western conifer seed bug, Leptoglossus occidentalis Heidemann, was first collected in Connecticut on December 27, 1985 in Fairfield County. Subsequently, it dispersed throughout the state with further geographic expansion into Massachusetts, Rhode Island, New Hampshire and Maine. In 1997, five adults of L. occidentalis were found parasitized by Trichopoda pennipes (EF) (Tachinidae). Adults of L. occidentalis invade homes to overwinter and, thus, are mainly considered a nuisance species. Key Words: forest pest, Connecticut, Leptoglossus occidentalis, Trichopoda pennipes, Diptera, Tachinidae, Heteroptera, Coreidae, biological control The western conifer seed bug, Leptoglos- sus occidentalis Heidemann, is a household nuisance in New England. I observed that they can tolerate freezing temperatures but are unable to survive if exposed to wet con- ditions in combination with cold. In the wild, L. occidentalis seeks shelter under dry bark or in rodent or bird nests. Because buildings offer the same protection, they frequently enter homes and commercial fa- cilities in late fall and early winter (Wheeler 1992): Connelly and Schowalter (1991) consider L. occidentalis to be a forest pest. It feeds on over 30 known species of trees, primar- ily Abies, Picea, Pinus and Tsuga species (Koerber 1963, Krugman 1969, Rice 1985). It gains nourishment on the endosperm of the maturing seeds in warm weather months and on the needles during the cold months. Leptoglossus occidentalis feeds at the base of host conifer needles, probably ingesting sugars produced by over wintering trees. However, there is no evidence that this ac- tivity is damaging. Leptoglossus occiden- talis is also a pest in commercial plantations growing nursery seed stock (Schowalter and Sexton 1990). Leptoglossus occidentalis was first re- corded from Utah, California, Colorado and British Columbia by Heidemann (1910). Originally, it was confined to the western third of the United States and Canada. Gall (1992) presented strong evidence that the eastern range expansion during the last 70 years was mediated by humans. Adults are active fliers and can fly over 200 feet in a single flight. This aids dispersal once the insect is introduced to an area. The insect probably reached Connecticut in the early to mid-1980’s. The first specimen of L. oc- cidentalis collected in Connecticut was from a Douglas-fir on 27 December 1985 in Fairfield, Fairfield Co. Dr. R. J. Pack- auskas (then at the University of Connect- icut) identified the insect in 1990. The spec- VOLUME 103, NUMBER 2 365 rom le imen is in the insect collection at Yale Uni- versity. In 1996, the public information office at The Connecticut Agricultural Experiment Station (CAES) in New Haven began re- ceiving calls about this insect from citizens across the state. In response, a survey of pest control operators, Christmas tree grow- ers, state agencies, and citizens was con- ducted to determine the geographic distri- bution of L. occidentalis in Connecticut. The survey was facilitated by telephone in- terviews and questionnaires. Specimens of L. occidentalis were collected from all eight counties in Connecticut (Fig. 1). Most spec- imens were collected in urban areas in the southern part of the state. Additional adult specimens were collected from widely sep- arated sites in Massachusetts, Rhode Island, New Hampshire, and Maine and were brought to CAES for identification. Dr. Ri- chard Dearborn (personal communication) Distribution of Leptoglossus occidentalis in Connecticut. collected the first adult from Kennebec City, Kennebec County, Maine, on 8 Oc- tober 1994. In most instances, adults were collected from homes and other buildings and were considered a nuisance. There are no previously known parasites or pathogens of L. occidentalis. During Oc- tober 1997, two adult insects, each with an egg cemented to the dorsal surface of the head were collected. These insects were held in the laboratory, and, after death, brown puparia were found near each insect. From these, two adult females of Tricho- poda pennipes (FE) (Diptera: Tachinidae) emerged 14 days later. Once the maggots matured, the host died, and the maggots ex- ited through the anus of the host and pu- pated. Trichopoda pennipes, a parasitoid of in- sect species of Pentatomidae Largidae, and Coreidae (Beard 1939, Arnaud 1973), is a conspicuous fly with bright dorsal yellow 366 stripes on the thorax and feather-like setae on the posterior hind tibia. Trichopoda pen- nipes has no official common name, al- though Bratley (1933) referred to it as the *‘feather-legged fly.”’” Although T. pennipes usually laid its eggs on the head of L. oc- cidentalis, eggs were sometimes deposited on the dorsal or ventral surface of the host’s thorax. According to Beard (1939), the maggots burrowed into the head capsule of Anasa tristis (DeGeer) after hatching, and then attached to the tracheal system to feed. In the present study, four adults of L. oc- cidentalis had multiple 7. pennipes eggs, but ten had single eggs. Of the 438 L. oc- cidentalis collected during the winter of 1997-1998, five (1.5%) had eggs on them, while during the winter of 1998-1999, nine (16%) of 57 L. occidentalis had eggs on them. Trichopoda pennipes may contribute to some biological control of L. occidental- is, especially in nursery seed groves in the western United States and Canada. Lepto- glossus occidentalis and T. pennipes are widely distributed in the United States with the geographic range of L. occidentalis con- tinuing to expand. Specimens of T. penni- pes and L. occidentalis from this study are in the Connecticut Agricultural Experiment Station insect collection. ACKNOWLEDGMENTS I thank the countless members of the general public, pest control operators, Christmas tree growers, and state agencies that helped me gather information about L. occidentalis in Connecticut and neighbor- ing states. I especially thank Patty Carr O’Shea, Mr. and Mrs. Louis Bernardini, and Barbara Listro for supplying me with many live specimens. Other scientists, scholars, and private cit- izens I thank for their generous assistance are Dr. Alan Tomlin, Dr. William Krinsky, Dr Leland Humble, Dr. Richard Dearborn, Dr. Richard Packauskas, Dr. Steven Katov- ich, Ms. Sarah Bates, Dr. John Bordon, Mr. and Mrs. Comboni, Dr. Wayne Gall, Dr. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Andy Hamilton, Mr. Jeff Fengler, Dr. Ken Harrison, Dr. Ronald Weseloh, Dr. Louis Magnarelli, Dr. Sharon Douglas, Dr. Chris Maier, Dr. Alan Wheeler, Mr. Kenneth A. Welch, Mr. Jay Donahue, Dr. John Ander- son, Dr. Paul Johnson, Dr. Steven Jacob, Dr. Vernon Nelson, Dr. Richard Zack, Dr. John Bouseman, Dr. Neil McHale, and Dr. Steve Burian. LITERATURE CITED Arnaud, Jr, P H. 1973. A host-parasite catalog of North American Tachinidae (Diptera). Science Education Administration., U.S. Department of Agriculture. Miscellaneous Publication 1319, 860 Pp. Beard, R. 1939. The biology of Anasa tristis DeGeer, with particular reference to the tachinid parasite, Trichopoda_ pennipes Fabr. Ph.D. dissertation, Yale University, New Haven, Connecticut. Bratley, H. EF 1933. Annual report of the Florida Ag- ricultural Experiment Station. 1932. Entomology, pp. 85-86. Connelly, B. C. and T. D. Schowalter. 1991. Seed loss- es to feeding by Leptoglossus occidentalis (Het- eroptera: Coreidae) during two periods of second- year cone development in western white pine. Journal of Economic Entomology 84: 215-217. Gall, W. K. 1992. Further eastern range extension and host records for Leptoglossus occidentalis (Het- eroptera: Coridae) well documented dispersal of a household nuisance. The Great Lakes Entomolo- gist 25: 159-171. Heidemann, O. 1910. New species of Leptoglossus from North America. Proceedings of the Ento- mological Society of Washington 12: 191—197. Koerber, T. W. 1963. Leptoglossus occidentalis (He- miptera: Coreidae), a newly discovered pest of co- niferous seed. Annals of the Entomological Soci- ety of America 56: 229-234. Krugman, S. L. 1969. Effect of cone feeding by Lep- toglossus occidentalis on Ponderosa Pine seed de- velopment. Forest Science 15: 104-111. Rice, R. E. et. al. 1985. New findings on pistachio problems. California Agriculture Department Bul- letin 39: 15-18. Schowalter, T. D. and J. M. Sexton, 1990. Effect of Leptoglossus occidentalis (Heteroptera: Coridae) on seed development of douglas fir at different times during the growing season in western Oregon. Journal of Economic Entomology 83: 1485-1486. Wheeler, A. G. 1992. Leptoglossus occidentalis, a new conifer pest and household nuisance in Pennsyl- vania. Pennsylvania Department of Agriculture Bulletin 18: 29—30. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 367-372 A NEW SPECIES OF EULITHIS HUBNER (LEPIDOPTERA: GEOMETRIDAE) FROM CALIFORNIA DOUGLAS C. FERGUSON AND SEI-WOONG CHOI (DCF) Systematic Entomology Laboratory, PSI, Agricultural Research Service, USDA, % National Museum of Natural History, Washington, DC 20560-0168, U.S.A.; (S-WC) Dept. of Environmental Education, College of Engineering, Mokpo National University, 61 Dorim-ri, Chunggye-myon, Muan-gun, Chunnam 534-729, Republic of Korea Abstract.—A new geometrid moth, Eulithis powellata, of the subfamily Larentiinae, is described from Monterey County, California. It is most similar to the Eurasian Eulithis mellinata (Fabricius) and E. pyraliata (Denis and Schiffermiiller), and its larva feeds on Ribes menziesii (Pursh) (Saxifragaceae). Although one of its Old World relatives, E. mel- linata, has been introduced into eastern Canada, E. powellata is clearly distinct and ap- parently endemic to a small area of coastal California. Sixteen species of Eulithis are now known from North America, including two Eurasian introductions. Key Words: This distinct and easily recognized moth is related to an Old World species group and closely resembles the Palearctic Eulithis mel- linata (Fabricius) and E. pyraliata (Denis and Schiffermiiller), both of which range across Eurasia from Britain and Norway to eastern Siberia. Eulithis mellinata was accidentally introduced into eastern Canada (Sheppard 1975, as Lygris associata Borkhausen; Neil 1978), but the Californian species is clearly different. It is larger, with the antemedial and postmedial lines of the forewing somewhat differently shaped and farther apart, the un- dersurfaces of the wings much more boldly marked (Figs. 1—3), and with conspicuous differences in the genitalia (Figs. 4-8). Eu- lithis propulsata (Walker), E. luteolata Hulst), a yellowish form of E. destinata (Moschler), and possibly E. gracilineata (Guenée) are the only other light-yellowish species of Eulithis that have been collected in California, and the new species is easily distinguished from all of them, as indicated in the description. At the time of writing, the generic placement of the new Californian species remains uncertain. According to one Larentiinae, Eulithis powellata, Ribes, introduced species phylogenetic hypothesis, this species, togeth- er with the Eurasian FE. pyraliata and the North American Ecliptopera atricolorata (Grote and Robinson), should be removed from Eulithis and referred to the otherwise Asian genus Gandaritis Moore 1868. The ju- nior auther is addressing this problem in an- other paper (Choi, in press). Sixteen species of Eulithis (sens. lat.) are now known from North America, including two Palearctic introductions—E. mellinata, mentioned above, and E. prunata (L.), re- cently reported also as being established in southern Quebec (Handfield 1997: 59, 93; Handfield 1999: 355). This new species was investigated and described in manuscript by the two authors working independently, each without knowledge of what the other was doing un- til the accidental overlap came to light; thus the co-authorship. Eulithis powellata Ferguson and Choi, new species (Figs. 1—6) Description.—Appendages of head and thorax structurally similar to those of other 368 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-3. Eulithis powellata, adults. 1, Holotype male, dorsal view. 2, Holotype male, ventral view. 3, Paratype female, University of California Big Creek Reserve, Devil’s Creek, 60—80 m, Monterey County, Cal- ifornia, 3 August 1992, J. Powell, collector. yellow species of genus; antenna of both sexes filiform, but that of male thicker and somewhat compressed; frons nearly flat but with a low, conical, mid-frontal crest of scales in fresh specimens; labial palp long in male, equal to 2% times width of frons, equal to twice width of frons in female. Frons and palpi with mixture of yellowish and light reddish-brown scales; body yel- lowish; legs similarly yellowish but partly shaded with brown, particulary on inner sides of forefemur and tibia and on outer sides toward distal ends of all tibiae and femora, especially of males. Forewing light ochreous yellow with three main transverse lines thin but distinct, brown, approximately right-angled as fol- lows: antemedial line regular or finely cren- ulate, nearly right angled at longitudinal fold in discal cell; medial line nearly par- allel to antemedial line, regular to slightly waved, similarly angled at longitudinal fold in discal cell and sometimes dentate at first anal fold; postmedial line usually regular, angled at M,, with sections before and after that point each nearly straight. Subterminal band diffuse, pale, crenulate in middle of outer third, becoming faintly dark shaded at costa. Basal space shaded with brown near costa; space between antemedial and medial faintly brown-shaded toward middle; me- dial line with slight dark shading distad near costa, and postmedial line immediately preceded by similar brown shading between its angle and costa. Apical angle bisected by interface line separating a paler area cos- tad from a darker area on side toward outer margin. Discal spot minute and inconspic- uous, or absent. Veins thinly outlined with brown scales, especially between postme- dial band and outer margin, more distinctly so than in any closely related Nearctic spe- cies. Fringes reddish brown on both wings, not or hardly variegated. Hindwing pale ochreous yellow, with two gray or blackish, diffuse, wavy trans- verse bands, subparallel to each other and to outer margin, and with some reddish- brown marginal shading between outermost band and somewhat crenulate outer margin. These bands evident only toward inner mar- gin and vary from faint to very distinct; they may represent medial and postmedial bands. Discal spot weak or absent. Undersurfaces of both wings a slightly deeper buff yellow; forewing with anteme- dial and postmedial lines strongly empha- sized with dark brown on anterior half of wing, fading out posteriorly; subterminal band indicated by little more than a weak brown patch at costa; outer margin with large crescent-shaped brown patch just pos- terior to apex; discal spot obsolete. Under- surface of hindwing with same transverse bands as upperside, but complete from in- ner margin to costa, dark brown; medial band wide, double; postmedial band mostly broken into series of intervenular spots. Forewing length: holotype, 18 mm; other males, 15—18 mm; females, 17—19 mm. Male genitalia (Figs. 4—5) with valve simple, lacking dentate process on outer VOLUME 103, NUMBER 2 369 Figs. 4-6. Eulithis powellata, genitalia. 4, Main part of male genitalia (paratype, USNM Slide No. 58872). 5, Aedeagus of same specimen. 6, Female genitalia (paratype, USNM Slide No. 58910). margin characteristic of mellinata (Figs. 7— 8), propulsata, and luteolata (that of pyr- aliata also without dentate process but dif- ferently shaped, with pointed, incurved apex). Vesica also simple, not scobinate, and without cornuti, in contrast to patches of cornuti present in other species, one in propulsata, two in mellinata and luteolata, and two or more in pyraliata (appearing as bundles of cornuti if vesica not everted). Also, setose, clavate, basocostal processes twice as long as those of mellinata, and even larger relative to those of some other species. Saccus convex and rounded in powellata as in most species, not emargin- ate as in both mellinata and pyraliata. Male genitalia closest to those of Eulithis diver- silineata (Hiibner) and E. gracilineata with respect to simple valve and vesica without cornuti. However, vesica of powellata not scobinate as in those two species. (This is a case in which the external appearance of 370 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 7-8. imen. the adult moth, plus food plant and female genitalia, may be more revealing with re- spect to relationship than are the male gen- italia). Female genitalia (Fig. 6) much as in E. pyraliata except corpus bursae essentially membranous, thickened only slightly to- ward posterior end. Also, signum of pow- ellata a scobinate, elongate, conical plate with point directed toward ostium (i.e., pos- teriorly); that of pyraliata similar but small- er and more rounded. Ostium simple and membranous in powellata and most others examined, but scobinate in mellinata. Bursa copulatrix of diversilineata and gracilinea- Eulithis mellinata (Europe), male genitalia. 7, Main part of genitalia. 8, Aedeagus of same spec- ta differs from those of powellata and oth- ers examined in having a_ longitudinal, wrinkled, bandlike signum running most of length of corpus bursae. Early stages and host plants.—Several adults were reared from larvae found in May on Ribes menziesii Pursh (Saxifraga- ceae) by J. A. Powell and B. Scaccia. Data from field-collected adults show that the species is double brooded, flying in late May and June and again in August and ear- ly September. The bivoltine life history is consistent with that of the two common eastern yellowish species, E. diversilineata and E. gracilineata, although more northern VOLUME 103, NUMBER 2 or montane species of Eulithis appear al- ways to be univoltine and to overwinter in the egg stage. This is true of E. mellinata in northern Europe, where it also feeds on species of Ribes, and of E. pyraliata, which feeds on species of Galium (Rubiaceae) (Skou 1986: 92-95). Host information for other American species of Eulithis is frag- mentary, although E. propulsata is thought to feed on Ribes; E. diversilineata and E. gracilineata feed on Vitaceae (especially on Virginia creeper, Parthenocissus quinque- folia (L.) Planch.); and other species have been variously reported on Salix, Populus, Alnus, Betula, Physocarpus, and Vaccin- ium. Types.—Holotype 6: California: Big Creek Reserve, U[niversity] C[alifornia] ‘““NLWRS,”’? Monterey County, California, 22 May 1992, J. Powell, collector; So. Ridge Rd., mixed hardwoods, 350 m. Par- atypes (18): 1 d, same data and collector but taken at 500 m on 31 May 1997. 1 6, same data and collector but dated 3/5 Sept. 1991. 2 5, 1 2, same data but collected by R. Zunigo and J. Powell, 5 June 1992 [col- lectors’ names not given on @]. 2 2, same locality, larvae found 17/18 March 1994, emegd. 22 April, 11 May 1994, reared from Ribes menziesi, J. Powell No. 94C26, J.A. Powell and M. McIntosh, collectors. 1 6, same data as holotype but collected at Bru- nette Creek, 60—180 m elev., 3 Aug. 1992; redwood-hardwood. | 2, same data as for holotype but collected at Devil’s Creek, 60— 80 m, 3 Aug. 1992, J. Powell; redwood, hardwoods. 3 6, 3 2, 1 sex unrecorded, same locality as holotype (Big Creek Re- serve), reared from larvae collected by B. Scaccia, as follows: from larva found 12 May 1992 on Ribes menziesii, emgd. 13 June 1992, J. Powell No. 92G74 (¢); from larva found 13 May 1992 on same host, emgd. 9 June 1992, J. Powell No. 92E209 (2); from larva found 23 June 1992 on same host, emgd. 22 Aug. 1992, J. Powell No. 92F133 (@); from larva found 13 May 1992 on same host, emgd. 3 June 1992, J. Powell No. 92E155 (¢); from larva found 371 13/15 March 1993 on same host, emgd. 30 April 1993, J. Powell No. 93C110 (3d); from larva found 13/15 March 1993 on same host, emgd. 1 May 1993, J. Powell No. 93C112 (sex not recorded); from larva found 24/26 April 1993 on same host, emgd. 14 May 1993, J. Powell No. 93D175 (3). 1 6, Eden Vale, Monterey Co[unty), Cal[ifornia] [no date], collection W.H. Broadwell, Acc. No. 36961 (AMNH). 1 6, Bixby Canyon, Monterey Co[unty], Ca- liffornia], 30 July 1949. J.W. Tilden, col- lector). Two additional specimens were tak- en on ‘the residential hill’ in Carmel, Monterey County on 25 June 1969 by R.H. Leuschner; these are not included among the paratypes because we have not seen them. Holotype and nine paratypes depos- ited in Essig Museum of Entomology, Uni- versity of California at Berkeley; eight par- atypes in National Museum of Natural His- tory, Smithsonian Institution, Washington, DC; and the last two paratypes listed are in American Museum of Natural History, New York. Remarks.—This species is named after Jerry A. Powell, who collected it during a survey of the University of California Big Creek Reserve and sent specimens to the senior author for identification. This survey was the source of all known specimens ex- cept the two older ones found by the junior author in the American Museum of Natural History and the two reported by Leuschner. Larvae were first collected and reared and the foodplant established by Brian Scaccia, who assisted with the survey in 1992. The Big Creek Reserve is situated within the northern section of the Los Padres National Forest, near the Ventana Wilderness Area, and toward the southern end of the Santa Lucia Mountains, at about 36°05’N, 121°35'W. It appears that Eulithis powellata may exist only as this localized, relict pop- ulation. ACKNOWLEDGMENTS We thank Jerry A. Powell for supplying most of the material of this species, and we BZ PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON acknowledge the contribution of Brian Scaccia in finding and rearing larvae, thus establishing the host plant. We also thank James Adams of Dalton, GA, Charles V. Covell, Jr. of Louisville, KY, and Ron Leus- chner of Manhattan Beach, CA, for review- ing the paper. The genitalia drawings are by Lisa Roberts, who worked as a contract il- lustrator in the Systematic Entomology Laboratory, USDA. LITERATURE CITED Choi, S.-W. In press. Phylogeny of Eulithis Hiibner and related genera (Lepidoptera: Geometridae), with an implication of Miillerian mimicry. Sys- tematic Entomology. Handfield, L., avec la collaboration de Jean-Francois Landry, Bernard Landry et J. Donald Lafontaine. 1997. Liste des Lépidopteres du Québec et du Labrador. Association des entomologistes ama- teurs du Québec. Fabreries, Suppl. 7. 155 pp. Handfield, L., avec la collaboration de J. Donald La- fontaine, Bernard Landry, Fran¢ois Lessard, Denis Rancourt, Eric Rassart, et Patrice Thibault. 1999. Le Guide de Papillons du Québec, version scien- tifique. Broquet Inc., Boucherville, Québec, 982 Pps Ele 3rcole pls: Neil, K. 1978. A second locality for Eulithis mellinata (Geometridae) in North America. Journal of the Lepidopterists’ Society 32: 224-225. Sheppard, A.C. 1975. Lygris associata Bork., a new record for North America (Lepidoptera: Geome- tridae). Annales de la Société Entomologique de Québéc 20: 7. Skou, P. 1986. The geometrid moths of North Europe. Entomonograph, Vol. 6. E.J Brill/Scandinavian Science Press, Leiden, Copenhagen. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 373-375 A NEW SPECIES OF THE GENUS ATRICHOPOGON KIEFFER (DIPTERA: CERATOPOGONIDAE) FROM MEXICO HERON HUERTA Departamento de Entomologia, InDRE, Carpio 470, Col. Santo Tomas, 11340, México, D.E, México (e-mail: cerato_2000 @ yahoo.com) Abstract.—A new species, Atrichopogon mexicanus, is described from Chiapas, Mex- ico. This species is included in the subgenus Atrichopogon. Key Words: The 75 Neotropical species of Atricho- pogon are poorly understood, with no mod- ern keys and with many species undescri- bed in the genus. Six named species are currently recognized from Mexico (Borkent and Spinelli 2000), and Huerta (1996) de- scribed an additional five, but unnamed spe- cies from the state of Chiapas. One of these is described and named in this paper. The specimens were slide mounted using the technique of Wirth and Marston (1968) but were stained with pink lignin to en- hance observation of some features. Terms for taxonomic characters follow those by Downes and Wirth (1981), Wirth (1994), and Szadziewski et al. (1996). All type ma- terial, is deposited in the Collection of Ar- thropods with Medical Importance of the Institute of the Diagnostic y Epidemiology Referencia (InDRE). Atrichopogon (Atrichopogon) mexicanus Huerta, new species (Figs. 1—9) Diagnosis.—A large brown species; fe- male with one long spermatheca; male gen- italia with pilose lobule on sternite IX. Description.—Female: Head (Fig. 1): Brown. Eyes bare. Antenna (Fig. 2) with lengths of flagellar segments in the dimen- sions (wm): 71.7-41-41-41-41-41-43-43- Diptera, Ceratopogonidae, Atrichopogon, Mexico, new species 133.3-128.2-143.5-143.5-174.3; antennal ratio (AR): 1.99; distal flagellomeres brown; proximal pale. Length of flagellum about 1.08 mm. Palpus (Fig. 3) brown, with segment lengths (wm): 51.2-51.2-92.3-41- 51.2; third segment with deep sensory pit opening located on distal half. Palpal ratio (PR): 2.1. Length of last two segments of palpus equal to length of third segment. Palpus length: 0.28 mm. Mouth parts sim- ple, without teeth. Thorax: Brown, scutellum pale brown with 12 bristles (Fig. 5), postscutellum brown. Paratergite with 1 seta. Wing length 1.76 mm (1.74-1.79; n = 2); width 0.72 mm (n = 2) (Fig. 6), with radial veins pale brown, membrane with dense microtrichia. Macrotrichia present on all veins, cells, more dense on distal half. Cells r,, r, well- developed. Second radial cell about 3 times longer than first. Costal ratio (CR): 0.75. Halter pale. Legs pale yellowish, with basal and subapical pale band of all femora; basal and distal third pale band of all tibiae. Tar- someres pale. Tibial comb with 10 spines. TR) 2.4, TRAD 2.8, TRUID 2.5. Scutum with dense vestiture of setae (Fig. 7), stripes not visible. Abdomen: Pale with moderately dense vestiture of brownish setae. Genital sclero- tization as in Fig. 8, with sternite VIII well 374 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 0.50 Figs. 1-9. Atrichopogon mexicanus. 1-3, 5-8, Female. 4, 9, Male. 1, Head. 2, Antennal segments. 3, Palpus. 4, Palpus. 5, Scutellum. 6, Wing. 7, Scutal vestiture. 8, Genitalia segments and spermatheca. 9, Ventral aspect of genitalia. Scales in millimeters. VOLUME 103, NUMBER 2 sclerotized with dense hairs. Spermatheca (Fig. 8), 0.28 mm by 0.14 mm, without neck. Male: Similar to female with usual sex- ual differences. Palpus (Fig. 4) brown, with segment lengths (wm): 41-51.2-92.3-46.1- 61.5; third segment with sensory pit located on distal half. Palpal ratio (PR): 2.9; palpus length: 0.29 mm. Thorax: Scutellum with 12 bristles, short setae. Tibial comb with nine spines, and long spur. TRI) 2.5, TRAD) 2.1. Wing length 1.79 mm, width 0.56 mm. Second radial cell 3.4 times longer than first. Abdomen: Genitalia (Fig. 9) yellow brown. Sternite [X short with prominent pi- lose lobule on posterior margin; tergite IX round, caudomedian process bearing setae; gonocoxite wide, pilose; gonostylus short, wide basally, with apical extreme round, bearing | apical tooth. Aedeagus membra- nous medially, short and broad, with basal arch high 0.6 of total length; basal arms markedly sclerotized; caudosubmedian pro- jections enlarged at apex, without lateral teeth. Etymology.—tThe specific epithet “mexi- canus’’ refers to the country of origin. Distribution.—Mexico (Chiapas). Types.—Holotype: 3d, Mexico, Chiapas, El Vergel, 5 June 1935, light trap, Col. Dampf (ME 4268). Paratypes: 2 2, same date as for holotype. Deposited in the Col- lection of Arthropods with Medical Impor- tance of the InDRE. Discussion.—This species is included in the subgenus Atrichopon by the presence of a single spermatheca, female sternite 8 lacking bifurcating projection, more or less uniformly colored, gonostylus with single apex, sensory pit not at apex of segment three of maxillary palpus, fifth palpal seg- ment rounded apically, and second radial 35) cell much longer than first. The male of this species can be associated with the female by the body coloration and legs, vestiture of the abdomen and scutum, and the form of the palpus and clypeus. Atrichopogon mexicanus differs from all other Neotropical members of the genus by having a male genitalia with a distinctive setose lobe on the posterior margin of ster- nite 9. The female is similar to those of a number of other species of Atrichopogon, but the length of the spermatheca is unusu- al. ACKNOWLEDGMENTS I am deeply indebted to Dr. Art Borkent for suggestions and changes in the manu- script. I thank an anonymous reviewer for improving the manuscript. LITERATURE CITED Borkent A. and G. R. Spinelli. 2000. Catalog of the new world biting midges south of the United States of America (Diptera: Ceratopogonidae). Contributions on Entomology, International 4, 107 Pp. Downes, J. A. and W. W. Wirth. 1981. cap. 28. Cera- topogonidae, pp. 393-421. In McAlpine J. F, B. V. Peterson, G. E. Shewell, H. J. Teskey, R. J. Vockeroth & Wodd D. M. eds. Manual of Nearctic Diptera. Vol. 1. Agriculture Canada Research Branch Monograph No. 27. 674 pp. Huerta, H. 1996. Los ceratopogonidos colectados por Alfonso Dampf en Chiapas, México (Diptera: Ceratopogonidae). Tesis Facultad de Ciencias, Universidad Nacional Aut6noma de México, México, D.E 199 pp. Szadziewski, R., E. Kaczorowska and J. Krzywinski. 1996. Redescriptions of some European species of Atrichopogon (Diptera: Ceratopogonidae). Polskie Pismo Entomologiczne 65: 297-318. Wirth, W. W. 1994. The subgenus Atrichopogon (Lo- phomyidium) with a revision of the Nearctic spe- cies (Diptera: Ceratopogonidae). Insecta Mundi 8: 17-36. Wirth, W. W. and N. Marston. 1968. A method for mounting small insects on microscope slides in Canada balsam. Annals of the Entomological So- ciety of America 61: 783-784. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 376-385 DIFFERENTIAL PERFORMANCE OF A CONTARINIA GALL MIDGE (DIPTERA: CECIDOMYIIDAE) ON ANT-DEFENDED ACACIA CORNIGERA (FABACEAE) ZHIWEI LIU AND HORACIO BONFIL (ZL) Department of Entomology, Swedish University of Agricultural Sciences, Box 7044, S-750 07 Uppsala, Sweden; Division of Invertebrate Zoology, American Museum of Natural History, New York, NY 10024, U.S.A.; current address: Division of Insects, Field Museum of Natural History, Chicago, IL 60605, U.S.A. (e-mail: zliu@fmnh.org); (HB) Center of Ecology, National University of Mexico, P.O. Box 70-275, Ciudad Univ- ersitaria, 0410, D.E, Mexico Abstract.—Galls induced by an unidentified species of Contarinia gall midge were found on leaves of Acacia cornigera L. in Los Tuxtlas, Veracruz, Mexico. The host plant is protected against other herbivores by the mutualistic ant species Pseudomyrmex fer- rugineus (Smith). We found that the density of the galls was significantly different among A. cornigera trees with different functional categories of ant colonies, as determined by the colonies’ development stages. The galls were also unevenly distributed among posi- tions within leaves. More leaves were galled on the rachis and pinna than at other posi- tions, but galls at the leaf base and extrafloral nectary were larger than those on the rachis or pinna. No difference was found for parasitism of mature galls by parasitic wasps between the trees with different functional categories of ant colonies. We conclude that ant protection of acacia trees may be effective against specialized herbivores like gall- makers. A trade-off may exist for the gall midge in that gall size is largest at leaf positions where risk of predation is greatest, creating the uneven distribution of galls among leaf positions. The hypothesis that the gall midges of Contarinia sp. may benefit by living in ‘enemy-free space’ created by the ant is not supported by the present study. Key Words: Acacia cornigera, ant protection, Contarinia, Cecidomyiidae, enemy-free space, gall makers, mutualism, parasitism, Pseudomyrmex ferrugineus The relationship between Pseudomyrmex spp. and Acacia spp. has been widely cited as obligate mutualism (Futuyma 1986, Huxley and Cutler 1991, H6lldobler and Wilson 1990, Krebs 1994). The ant-acacia tree provides the ant with hollow thorns as nesting sites, and Beltian bodies and ex- trafloral nectar as food. The ant, in return, provides the tree with protection against herbivores. Several ant-acacias have lost the chemical traits that protect other Acacia species from herbivores (Rehr et al. 1973, Seigler et al. 1978, Seigler and Ebinger 1995). Field experiments have shown that acacia trees grow less and are frequently killed when deprived of ant protection (Jan- zen 1966, 1967). Similar associations have also been reported for other myrmecophytic plants associated with ants (Longino 1991, Fowler 1993, Fonseca 1994). The insect fauna associated with myr- mecophytic plants is usually less diverse and less abundant, and consists of more specialized species compared to non-myr- VOLUME 103, NUMBER 2 mecophytic plants (Jolivet 1991). Some specialists are able to circumvent ant attack through fast escape, building shelters or making galls (Heads and Lawton 1985, Koptur 1992, Loeffler 1996, Eubanks et al. 1997). Such insects may even take advan- tage of the fact that their natural enemies may be chased off, thereby living in ‘ene- my-free space’ created by the ants (Koptur 1991, 1992). For shelter-builders and gall- makers, however, the ant threat still exists for egg-laying females and the life stages not yet concealed, such as eggs and young larvae. Ants may actively remove those ex- posed insects from their host plant surface (Stephenson 1982, Fiala et al. 1991). In Los Tuxtlas, Mexico, two major spe- cialist herbivores were found on Acacia cornigera L., which harbored the obligate mutualistic ant Pseudomyrmex ferrugineus (Smith). The larva of Polyhymno sp. (Lep- idoptera: Gelechiidae), a shelter builder, can defeat the ant-defense of A. cornigera and inflict mortality-level defoliation on the host plant (Eubanks et al. 1997). The other, Contarinia sp., is a cecidomyiid fly that makes galls on the plant, mostly on the leaves. According to our preliminary observa- tions, the leaf galls at extrafloral nectaries appeared to be larger than those at other positions of the leaves. In many systems, large galls not only produce large adults with high fecundity (Weis et al. 1988), but also provide better protection against natu- ral enemies such as parasitoids (Price and Clancy 1986, Weis et al. 1988). Thus, the gall inducer may prefer to lay more eggs at extrafloral nectaries. However, these posi- tions are also resources for, and frequently visited by, both the ants and generalist par- asitoids (Koptur 1991), and therefore rep- resent a risk for the gall-maker as well. The present study was designed to ex- amine the Acacia-ant cecidomyiid interac- tions. Specifically, the following questions were asked: (1) Does the presence of ants significantly reduce herbivory inflicted on Acacia trees by specialized herbivore in- S/T sects such as gall-making cecidomyliids? (2) Are galls evenly distributed among leaf po- sitions and what role, if any, do the ants and parasitic wasps play in determining such a pattern? (3) Do the gall midges gain pro- tection from the ants against parasitic wasps, thus living in ‘enemy-free space’? NATURAL HISTORY Host plant.—Acacia cornigera L. (Le- guminosae: Mimosaceae) is naturally dis- tributed in wet to relatively dry, mostly dis- turbed habitats at lower elevations from southern Mexico to Costa Rica (Seigler and Ebinger 1995). At the base of each com- pound leaf is a pair of thorns, which are modified stipules. The thorns are excavated and occupied by the ants when the thorns are still very young (Liu, personal obser- vation). The compound leaf consists of pet- iole, rachis, pinna, and pinnules or leaflets. The Beltian bodies are located at the tips of the pinnules of young leaves (Fig. 1), but are rarely seen because they are usually har- vested by the ants as soon as the young leaves develop (Seigler and Ebinger 1995). Extrafloral nectar is produced on a contin- uous basis by canoe-shaped nectaries on the dorsal side of the petiole and, rarely, also on the rachis (Janzen 1966, 1967, 1974; Seigler and Ebinger 1995; Liu, personal ob- servation). Associated ant species.—Pseudomyrmex ferrugineus (Smith) (Hymenoptera: Formi- cidae), often erroneously cited as P. ferru- ginea (e.g., Janzen, 1966, Mintzer et al., 1987, but see Ward, 1989, 1993), has be- come well known because of Janzen’s stud- ies (1966, 1967), which provided strong ex- perimental evidence for the mutualism be- tween the ant and Acacia cornigera. It has a distribution ranging from eastern and southern Mexico to El Salvador and Hon- duras. It is a common species and uses as domatium hosts all swollen-thorn acacia species growing within its range, including Acacia chiapensis, A. collinsi, A. cookii, A. cornigera, A. gentlei, A. globulifera, A. hindsii, A. janzenii, A. mayana, and A. 378 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Rigale nectary; c = gall induced by a Contarinia gall midge; d = rachis; e = pinna; f = pinnule; g = Beltian body; h = ant exit. sphaerocephala (Ward 1993). Pseudomyr- mex ferrugineus resides in the thorns of the acacias. Usually only one colony exists in an acacia tree. Workers of young colonies leave the thorns only long enough to collect nectar and Beltian bodies, but they begin patrolling the plant surface when the colony has reached a size of 50-100. When ant numbers reach 200—400, the workers be- come more aggressive and start attacking competing ant colonies, as well as warding off other insects that attempt to land in the vicinity. It takes about 10 months for an ant colony to reach this stage (Janzen 1967). Janzen (1967) classifies the ant colonies into three functional categories on the basis of their developmental stages: Establishing (< 50 worker ants per tree), transient (SO— 200), established (> 200) (summarized in Holldobler and Wilson 1990; terms of the functional categories coined by the authors of the current article). Herbivores.—An unidentified Contarinia Diagram of a branch of ant acacia, Acacia cornigera, with Contarinia sp. gall (a = thorn; b. = sp. (Diptera: Cecidomyiidae) induces pin- nule-shaped galls in A. cornigera at the leaf base, rachis plus petiole, pinna or extraflor- al nectaries. Another type of gall is also found on the stems of the acacia trees, but the inducer was not identified. No infor- mation is available in the literature about either of the gall-inducers and their biology. METHODS Study sites.—The study was carried out about one kilometer northwest of the Los Tuxtlas Biological Station of the National University of Mexico, Veracruz, Mexico (95°04'W and 18°30'N). The Los Tuxtlas region constitutes the northernmost limit of tropical rain forest in the New World (Dirzo and Miranda 1991). The average annual rainfall is 1,966 mm, of which 80% occurs from June to October. December through May is relatively dry (Soto 1976). The study site was an extensively managed pas- tureland by the forest edge with an area of VOLUME 103, NUMBER 2 approximately 8 hectares. Acacia trees were abundant in the pasture, and had been pe- riodically cut. New sprouts growing from the stumps were common. A few larger trees were also present. Sampling was mainly done within a 50 by 30 m plot. Sev- eral additional trees along a main road by the studied pasture and along forest trails around the field station were included in the study. Estimation of infestation rate of host trees with gall-makers.—In total, 85 trees were surveyed to examine the infestation rates from the gall-makers. Both the leaf galls and the stem galls per tree were count- ed. To verify the presence of ant colony on each tree, thorns of all trees were opened until ants were found or all thorns had been opened and no ant was found. Examining the relationship between gall density and ant abundance.—Twelve trees from the pasture plus one tree along the road were selected to determine whether gall density and ant density were correlated. Measurements, for each tree, included 1) the number of thorns, 2) the number of thorns occupied by ants, 3) the number of leaf galls, and 4) the number of stem galls. Because old thorns that have been aban- doned by ants do not have a leaf at their base, we used the total ant-occupied thorns as an estimate of the number of leaves per tree. Between two to four (mostly three) av- erage-sized thorn pairs were sampled from each tree and the number of ant workers was counted. The average number of ants per thorn was then multiplied by the num- ber of ant-occupied thorns of the tree to cal- culate the ant colony size. The ant colonies of acacia trees were grouped according to Janzen’s classification (1967) into three functional categories: Establishing (< 50 ants per tree), transient (S0—200), estab- lished (> 200) (summarized in H6lldobler and Wilson 1990). Examining the distribution pattern of leaf galls and parasitism.—Leaves were sam- pled from a set of 17 trees to document the distribution of leaf galls on the trees and 379 incidence of parasitism of these galls by parasitic wasps. Two of these trees were growing along the road, one was close to the research station, and the others were from the pasture. On each tree, all leaves with mature galls were collected, bagged separately, labeled, and brought back to the laboratory for dissection. Gall size (width across the middle), the number of galls, and the positions of galls were recorded for each leaf. Galled leaves were classified as being galled at leaf base, rachis plus petiole, pinna, or extrafloral nectary (hereafter BAS, RACH, PINN, and NECT respective- ly). Only galls with a direct connection with a nectary will be classified as being galled at extrafloral nectary. The few leaves (n = 2) that were galled at more than one posi- tion were not included in the analysis. Oc- currences of parasitism and predation were also recorded. Parasitism was determined by counting the number of galls with exit holes by parasitic wasps and/or the number of galls with parasitic wasp larvae or pupae still remaining. Predation was estimated by counting the number of galls that had been broken open. Voucher specimens of the leaf gall-in- ducer are deposited in the National Muse- um of Natural History, Smithsonian Insti- tution, Washington, DC (Dr. Gagné) and in the senior author’s collection, and of the parasitoids in the National Museum of Nat- ural History, Smithsonian Institution, Washington, DC (Dr. Schauff). RESULTS The leaf gall induced by Contarinia sp. consists of a basal larval chamber in the stem, petiole, rachis, or pinna, and an upper part that resembles a pinnule in shape and color. A single developing larva feeds with- in the basal larval chamber, moves up into the pinnule-like part as a mature larva, and pupates there. Galls crack at the tip when mature and the adult midges emerge. The exit holes of the parasitoids were round and made at the lateral side of the pinnule-like upper part of the gall. 380 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 10 > c 8 N Z. ca a 6 = = 5 bor < = = jo e 2 oe 0 a . ESTABLISHING ESTABLISHED FUNCTIONAL CATEGORY OF ANT COLONY Fig. 2. The per leaf gall density of galls caused by Contarinia sp. is different between trees with establishing (n = 6) and established (n = 7) ant colonies (Mann-Whitney U-test, P = 0.01, U = 2.00). Shown is sample mean (inner rectangle) £1.00 standard error (outer rectangle), and +1.96 standard error (parallel horizontal lines). The stem gall is an inconspicuous swell- ing with larvae living in larval chambers in the woody part of the stem. The larvae dis- sected out of the stem galls could not be identified to genus, although it is clear that it does not belong to Contarinia. Two and one species of parasitoid wasps were reared from the leaf galls and the stem galls, respectively. The two species reared from the leaf galls are Closterocerus sp. (Eulophidae) and Eurytoma sp. (Eurytomi- dae). The parasitic wasps reared from the stem galls belong to a unidentified genus of Tanaostigmatidae (Chalcidoidea), a family whose members are generally gall-inducers with one species known as parasitoid (Gib- son 1993). More than half of all A. cornigera trees studied were found to have leaf galls (61.2%). Overall leaf gall density per tree was 9.56 (n = 85, Xx = 95, SD = 16.52). Slightly more than half of all host trees had stem galls (51.8%). Stem gall density per tree was 2.18 (n 85, Xnax = 18, SD = 3.53). All trees checked for ant presence in the study (n 85) were found to host P. ferrugineus colonies. Only two functional categories of ant colony, Establishing and established, were found on the studied trees. The average number of galls per leaf of each tree was not significantly related to the size of the ant colony in terms of the num- ber of worker ants inhabiting the tree (P = 0.10, r° = 0.22), but differed significantly between the trees that host establishing ant colonies and those that host established ones (Mann-Whitney U-test, P = 0.01, U = 2.00) (Fig. 2). The number of stem galls per tree was not related to ant colony size (P = 0.25, r? = 0.12), nor to the functional categories of ant colony (Mann-Whitney U- test, P = 0.22, U = 12.5). Galls were unevenly distributed among galling positions within a leaf. Galls were more often found on the rachis and pinna VOLUME 103, NUMBER 2 28 = nN nN > —) & NUMBER OF GALLED LEAVES Ss 381 PINN NECT GALL POSITION OF LEAVES Ric 3: Observed proportion of leaves galled by Contarinia sp. on different positions. The galled leaves were classified into four types according to gall forming positions of a leaf: at leaf base (BASE), at rachis (RACH), at pinna (PINN) and at extrafloral nectary (NECT). x?-test showed that galls were unevenly distributed (P < 0.05, x? = 8.70, df = 3). than on extrafloral nectaries or at the leaf base (x? = 8.70, P < 0.05, df = 3) (Fig. 3). Leaf gall size differed significantly among positions of gall formation (P = 0.01). Galls formed on extrafloral nectaries or at the leaf base were larger than those formed at the rachis or pinna (Fig. 4). The overall parasitism rate was 12.3% (n = 382). No difference in parasitism of leaf galls was found among gall-forming posi- tions (One-way ANOVA, F(2, 68) = 0.695, P = 0.55), or between A. cornigera trees hosting ant colonies of different functional categories (Mann-Whitney U-test, P = 0.77, U = 7.0). Data on parasitism of stem galls were not sufficient for statistical anal- ysis. Predation on leaf galls was generally low: only 3.9% of the 382 sampled leaf galls were observed to have been bitten open by predators, apparently by other in- sects. No predation was observed on stem galls. DISCUSSION Even though all A. cornigera trees sur- veyed were colonized by P. ferrugineus, gall midge infestation rates were rather high, suggesting that the gall midges can somehow circumvent the ant defense. The per leaf density of Contarinia galls of each tree was not related to the ant col- ony size in terms of the number of worker ants. There does appear, however, to be a significant relationship between per leaf gall density of each tree and the functional categories of ant colonies found on the trees. This indicates that ant protection does reduce gall density. It also suggests that the degree of biotic protection for the trees is likely to be affected by the socially initiated behavior of the protecting ant colony. Dur- ing the Establishing stage, the ant workers are less active and the protection is weak. When the ant colony becomes large and the workers are more aggressive, ant protection becomes more intensive. We did not find 382 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 2.8 similar results for the per tree density of stem galls between trees hosting different functional categories of ant colonies. It is likely that the stems are less important to the ants as a food resource and less fre- quently visited. Therefore the activity of the stem gall midge is less likely to be affected by ant activities in terms of the size and development stage of the ant colonies. Galls suffered very little predation. Heads and Lawton (1985) observed that ants on bracken (Pteridium aquilinum) were attracted to the extrafloral nectaries and pro- tected the plant from caterpillars by attack- ing them. The ants, however, appeared to have no effect on the two species of gall midges that induce galls on bracken pinna because the midges were concealed inside the galls. The fact that more leaves were found with galls at the rachis and pinna than at the leaf base or the extrafloral nectaries may indicate that the egg-laying female of Con- 2.4 =e = S 2.0 a = s < S 1.6 1.2 BASE RACH NECT GALL FORMING LOCATION ON LEAVES Fig. 4. Comparison of average size of galls induced by Cnotarinia sp. on ant acacia leaves at different gall- forming positions. Samples with different letters at upper left are significantly different using the Newman-Keuls Test (P = 0.01). The galled leaves were classified into four types according to gall forming positions of a leaf: at leaf base (BASE), at rachis (RACH), at pinna (PINN) and at extrafloral nectary (NECT). PINN tarinia sp. prefers the former sites. How- ever, it does not seem to be beneficial for her offspring to develop at such sites be- cause those galls were significantly smaller than those at the nectaries and leaf base. In general, large galls result in high fecundity (Weis et al. 1988) and better protection from natural enemies (Price and Clancy 1986, Weis et al. 1988). Thus it does not seem to be an adaptive behavior for the ovi- positing female midge to prefer rachis and pinna for egg-laying, unless they are at- tacked by ants at the other sites. One alternative explanation for the un- even distribution pattern of galls on leaves is that egg-laying females, eggs and young larvae at leaf bases and extrafloral nectaries are subject to more intense predation than those at the rachises. The acacia tree is pro- tected by the ant because it provides nesting thorns, Beltian bodies and extrafloral nec- tars. The exclusion of herbivores from ant- acacia tree by the ants is in fact the result VOLUME 103, NUMBER 2 of resource competition, which can become more intense if it is a direct one. For ex- ample, when an extrafloral nectary is galled by the leaf gall midge, it can no longer function as a nectar provider. Therefore, the ant workers should chase away the egg-lay- ing female gall midge from extrafloral nec- taries, and actively remove eggs and unen- capsulated larvae. Acacia thorns, as we ob- served, were excavated at very young stag- es by the ant, and are thus under ant protection from the very beginning. The proximity of the leaf base to the thorns will expose an egg-laying female of Contarinia sp. and her offspring to a higher risk com- pared to the other positions. Therefore, the gall midge female may prefer to lay her eggs on the rachis and pinna rather than at the leaf base or the extrafloral nectary, thus creating the uneven distribution pattern of galls. Alternatively, she may not demon- strate a preference for a particular oviposi- tion site, but the different degrees of ant predation create the pattern. It is also pos- sible that a combination of both takes place. We found no difference in the rate of par- asitism of galls on trees with different de- grees of ant protection derived from func- tional categories of ant colony, or among positions within leaves. Other studies have shown herbivores on plants defended by ants to be less frequently parasitized than those on plants without ants, and concealed feeders to be parasitized less frequently in areas of high ant activity (reviewed in Kop- tur 1992). In our study, the acacia trees with established ant colonies were taller and had more branches, and thus had more available extrafloral nectaries. It is possible that the many nectaries attract more parasitoids (cf. Koptur 1991). Thus, a larger number of parasitoids may circumvent the ant protec- tion for the leaf galls. An alternative expla- nation is that the parasitoids are highly spe- cialized and are not affected by the ants. Regardless of the explanation, the “enemy- free space’ for the Contarinia gall midge in ant protected acacia trees does not seem to 383 exist in terms of parasitic wasps in the pre- sent study. In conclusion, because galls were found on so many ant-colonized acacia trees, data from this study supports the theory that concealed herbivores such as gall-making insects can circumvent ant attack. Because the incidence of galls was much lower on trees hosting established colonies of ag- gressive ants, this study also indicates that ant protection for the ant-acacia could be effective even against highly specialized herbivores like the Contarinia leaf gall midge in the present system. A trade-off may exist for the gall midge in that gall size is largest at leaf positions where risk of pre- dation is greatest. Finally, based on evi- dence of parasitic wasps in this system, the Contarinia leaf gall midge does not appears to enjoy ‘enemy-free space’ on ant-protect- ed acacia trees. ACKNOWLEDGMENTS We thank Drs. Betty Benrey, Robert Denno, Rodolfo Dirzo, William Lamp, Stig Larsson and our classmates at the Tropical Insect Ecology course conducted at Los Tuxtlas for encouragement and friendship. The course was a collaborative project of the Center of Ecology, National University of Mexico, the Department of Entomology, University of Maryland, and the Depart- ment of Entomology, Swedish University of Agricultural Sciences. Parts of the writ- ing were carried out by ZL during a Kalb- fleisch Postdoctoral Research Fellowship at the American Museum of Natural History, New York, and a Boyd Postdoctoral Re- search Fellowship at the Field Museum of Natural History, Chicago. Our particular thanks are due to Dr. Larsson for criticism on our initial ideas and the manuscript and to Drs. Raymond Gagné and Michael Schauff, Systematic Entomology Labora- tory, USDA, Washington, DC, for identifi- cation of the gall midges and the parasitic wasps, respectively. Comments from Drs. Amy Berkov, Christer Bj6rkman, Raymond Gagné, Christine Johnson, Géran Nordlan- 384 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON der, and an anonymous reviewer have helped to improve the manuscript signifi- cantly. LITERATURE CITED Dirzo, R. and A. Miranda. 1991. Altered patterns of herbivory and diversity in the forest understory: A case study of the possible consequences of con- temporary defaunation, pp. 273—287. In Price, P. W., T. W. Lewinsohn, G. W. Fernandes, and W. W. Benson, eds. Plant—Animal Interactions: Evo- lutionary Ecology in Tropical and Temperate Re- gions. John Wiley & Sons, Inc. New York, NY. Eubanks, M. D., K. A. Nesci, M. K. Petersen, Z. Liu, and H. Bonfil. 1997. The exploitation of an ant- defended host plant by a shelter-building herbi- vore. Oecologia 109: 454—460. Fiala, B., U. Maschwitz, and T. Y Pong. 1991. The association between Macaranga trees and ants in South-east Asia, pp. 263—270. Jn Huxley, C. and D. Cutler, eds. Ant—Plant Interactions. Oxford Univ. Press, New York. Fonseca, C. R. 1994. Herbivory and the long-lived leaves of an Amazonian ant-tree. Journal of Ecol- ogy 82: 833-842. Fowler, H.G. 1993. Herbivory and assemblage struc- ture of myrmecophytous understory plants and their associated ants in the central Amazon. In- sectes Sociaux 40(2): 137-145. Futuyma, D. J. 1986. Evolutionary Biology, 2nd edi- tion. Sinauer Associates, Sunderland, Massachu- setts. Gagné, R. J. 1989. The plant-feeding gall midges of North America. Cornell University Press, Ithaca, NY. Gibson, G. A. P. 1993. Superfamilies Mymaromma- toidea and Chalcidoidea, pp. 570—655. Jn Goulet, H. and J. T. Huber, eds. Hymenoptera of the World: An Identification Guide to Families. Re- search Branch Agriculture Canada Publication 1894/E. Heads, P. A. and J. H. Lawton. 1985. Bracken, ants and extrafloral nectaries. II]. How insect herbi- vores avoid ant predation. Ecological Entomology 10: 29-42. H6lldobler, B. and E. O. Wilson. 1990. The Ants. Har- vard University Press, Cambridge, Massachusetts. Huxley, C. and D. Cutler. 1991. Ant—Plant Interac- tions. Oxford Univ. Press, New York, NY. Janzen, D.H. 1966. Coevolution of mutualism between ants and acacias in Central America. Evolution 20: 249-75. . 1967. Interaction of the Bull’s-Horn acacia (Acacia cornigera L.) with one of its ant inhabi- tants (Pseudomyrmex ferruginea F Smith) in east- ern Mexico. Kansas University Science Bulletin 47: 315-558. 1974. Swollen-thorn Acacias of Central America. Smithsonian Contributions to Botany 13: 1-131. Jolivet, P. 1991. Ants, plants and beetles: A triangular relationship, pp. 397—406. In Huxley, C. and D. Cutler, eds. Ant—Plant Interactions. Oxford Univ. Press, New York, NY. Koptur, S. 1991. Extrafloral nectaries of herbs and trees: Modeling the interaction with ants and par- asitoids, pp. 213—230. /n Huxley, C. and D. Cutler, eds. Ant—Plant Interactions. Oxford Univ. Press, New York. 1992. Extrafloral nectary-mediated interac- tions between Insects and plants, pp. 81-129. In Bernays, E., ed. Insect—Plant Interactions IV, CRC Press, Boca Raton, Florida. Krebs, C. J. 1994. Ecology: The experimental analysis of distribution and abundance, 4th edition. HarperCollins, New York. Loeffler, C. C. 1996. Adaptive trade-offs of leaf fold- ing in Dichomeris caterpillars on goldenrods. Eco- logical Entomology 21: 34—40. Longino, J. T. 1991. Azteca ants in Cecropia trees: Taxonomy, colony structure, and behaviour, pp. 271-288. In Huxley, C. and D. Cutler, eds. Ant— Plant Interactions. Oxford Univ. Press, New York. Mintzer, A. C., H. J. Williams, and S. B. Vinson. 1987. Identity and variation of hexane soluble cuticular components produced by the acacia ant Pseudo- myrmex ferruginea. Comparative Biochemistry and Physiology 86B: 27-30. Price, W. P. and K. M. Clancy. 1986. Interactions among three trophic levels: Gall size and parasit- oid attack. Ecology 67: 1593-1600. Rehr, S. S., P. P. Feeny, and D. H. Janzen. 1973. Chem- ical defense in Central American non-ant-acacias. Journal of Animal Ecology 42: 405-416. Seigler D. S., J. E. Dunn, E. E. Conn, and G. L. Hol- stein. 1978. Acacipetalin from six species of Aca- cia of Mexico and Texas. Phytochemistry 17: 445-446. Seigler D. S. and J. E. Ebinger. 1995. Taxonomic re- vision of the ant-acacias (Fabaceae, Mimosaceae, Acacia, Series Gummiferae) of the new world. Annals of the Missouri Botanical Garden 82: 117— 138. Soto, E. M. 1976. Algunos aspectos climaticos de la region de los Tuxtlas. Jn Gomez-Pompa, A. et al., eds. Regeneracion de Selvas. Compania Editorial Continental, Mexico City, Mexico. Stephenson, A. G. 1982. The role of the extrafloral nectaries of Catalpa speciosa in limiting herbiy- ory and increasing fruit production. Ecology 63: 663-669. Ward, P. S. 1989. Systematic studies on Pseudomyr- mecine ants: revision of the Pseudomyrmex ocu- latus and P. subtilissimus species groups, with VOLUME 103, NUMBER 2 385 taxonomic comments on the other species. Quaes- Weis, A. E., R. Walton, and C.L. Crego. 1988. Reac- tiones Entomologicae 25: 393-468. tive plant tissue sites and the population biology . 1993. Systematic studies on Pseudomyrmex of gall makers. Annual Review of Entomology 33: acacia-ants (Hymenoptera: Formicidae: Pseudo- 467-86. myrmecinae). Journal of Hymenoptera Research Zar, J. H. 1984. Biostatistical Analysis, 2nd edition. 2: 117-168. Prentice-Hall, Englewood Cliffs, New Jersey. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 386-388 NOTES ON THREE SPECIES OF ANTHOCORIDAE (HEMIPTERA: HETEROPTERA) FROM HAWATI, INCLUDING THE FIRST RECORD OF BUCHANANIELLA CONTINUA (WHITE) GREGORY J. BRENNER AND JOHN D. LATTIN (GJB) 1520 NW 14th Place, Corvallis, OR 97330, U.S.A.; (JDL) Department of En- tomology, Oregon State University, Corvallis, OR 97331-2907, U.S.A. Abstract.—Three species of Anthocoridae (Hemiptera: Heteroptera) are reported from a study of the palila bird Loxioides bailleui (Oustalet) on the island of Hawaii: Amphiareus constrictus (Stal), Buchananiella continua (White), and Orius tristicolor (White). All are non-indigenous and B. continua is a species not previously reported from the Hawaiian Islands, making a total of 21 species known there, of which 15 are non-indigenous. Key Words: The insect fauna of the Hawaiian islands is large and diverse. Eldredge and Miller (1997) reported 7,979 species, including 2,582 that were non-indigenous (32%), likely the highest percentage we know in the world. Taxa within the Insecta display different percentages of introduced species as evidenced by the minute pirate bugs. The family Anthocoridae, or minute pirate bugs, contains 20 reported species in Hawaii that includes 14 introduced species - 70% (As- quith and Messing 1992; Nishida 1994; Eldredge and Miller 1995; Miller and Eld- redge 1996; Eldredge and Miller 1997; Lat- tin 1999, 2000). The family contains small, usually predaceous, insects (~2.5 mm), that are found in a variety of habitats in asso- ciation with living and dead plants and litter (Péricart 1972; Lattin 1999, 2000). Nishida (1994) reported 19 species from the Ha- waliian Islands. Montandoniola moraguesi (Puton) should be added to that list. It was introduced in 1964 as a biological control agent against the newly introduced Cuban laurel thrips and quickly spread over sev- eral islands (Davis and Krauss 1965). In this paper, we report Orius tristicolor Anthocoridae, Hawaiian Islands, non-indigenous species, introductions (White), Amphiareus constrictus (Stal) (previously reported under the name Car- diastethus fulvescens (Walker) by Zimmer- man 1948; Herring 1966), and Buchanan- iella continua (White) from the Hawaiian Islands. MATERIALS AND METHODS The specimens presented in this paper are from a long-term survey by the Biolog- ical Resources Division of the United States Geological Survey that is investigating the diet and prey items of the endangered palila bird, Loxioides bailleui (Oustalet) Part of the effort includes a series of studies of in- sects and their relatives found in the dry, montane woodland forest on the western slopes of Mauna Kea between 1,700 m and 2,835 m in elevation (5,580 ft—9,300 ft). Part of the study involves rearing insects from pods of the mamane tree, Sophora chrysophylla (Salisbury) Seeman, a major food item of palila (Scott et al. 1986). Ma- mane pods were collected monthly from October 1995 through April 1997 along an elevation gradient and placed in rearing cages covered with a fine mesh screen. As- VOLUME 103, NUMBER 2 sociated insects were removed from the cages as they emerged from the pods, and placed in alcohol with appropriate docu- mentation and labels. The specimens re- ported here were among those collected during that study. All Anthocoridae speci- mens were examined and identified by JDL. Specimens will be deposited with the B. P. Bishop Museum in Honolulu, Hawaii. Orius tristicolor (White) Orius tristicolor, was introduced from Arizona into Oahu (Davis and Krauss 1965), where, according to Clausen (1978), it failed to become established. Our speci- mens from the collections made in 1996 on Hawaii reconfirm the establishment of O. tristicolor in Hawaii. Described from Cali- fornia by White (1879), O. tristicolor is widely distributed in western North Amer- ica and has a range that extends to South America (Herring 1966b). The male geni- talia of O. tristicolor are distinct from those of O. insidious (Say), another species of Orius also introduced into the Hawaiian Is- lands (Kauai and Oahu) (Weber 1953). It, too, was considered not to have become es- tablished (Oatman 1978), but Takara and Nishida (1981) have confirmed its estab- lishment on the island of Oahu. The status of the endemic Orius persequens (White) and the introduced O. pumilio (Champion), the latter reported by Kirkaldy (1910) from Oahu, remains uncertain. Amphiareus constrictus (Stal) There is a single specimen of what ap- pears to be Amphiareus constrictus, but it lacks the terminal segments of the abdo- men. This species was previously reported by Zimmerman (1948) as Cardiastethus fulvescens (Walker), but see Herring (1966a) for clarification. Additional speci- mens are required for final documentation. Buchananiella continua (White) Originally described from the island of Madeira by White (1879), it has been intro- duced into many countries around the 387 world. The female of Buchananiella contin- ua (White) represents the first record for the Hawaiian Islands. While a male would be useful, the distinctive appearance of this species, including the short, semideclining setae on the dorsum; the pronotum with dis- tinct calli having a transverse sulcus behind and the posterior half with a median, lon- gitudinal impression; the two large, well- defined punctures on the scutellum; the in- distinct punctures on the clavus and balance of the hemelytra; the shape of the ostiolar canal; and the characteristic omphalophore on the middle of the venter of abdominal segment VII (Péricart 1972) combine to provide positive identification. The discovery of Buchananiella continua brings the total of species of Anthoridae to 21. Fifteen of these are introduced (71.4%), either accidentally or intentionally. These numbers indicate the vulnerability of the Hawaiian Islands to non-indigenous spe- cies. More intensive collecting in all parts of Hawaii likely will disclose additional ex- otic species. Much more work is needed to document the insect fauna of these unique, isolated, and vulnerable islands. ACKNOWLEDGMENTS We thank our colleagues working on the Hawaiian Islands: A. Asquith, L. G. Eld- redge, E G. Howarth, S. E. Miller, and G. M. Nishida for their current work on the insect fauna of the Hawaiian Islands and their interest in the occurrence and impact of non-indigenous species; P. Banko, USGS/BRD Kilauea Field Station, Hawaii, for his inspired leadership of the palila res- toration project; P. Oboyski and the many volunteers at the USGS/BRD Kilauea Field Station for their assistance in field collect- ing and laboratory rearing; two reviewers for their helpful comments; and to L. Parks for her work on the manuscript. LITERATURE CITED Asquith, A. and R. Messing. 1992. Annotated insect distribution records for the Island of Kauai. Pro- 388 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ceedings of the Hawaiian Entomological Society 31: 151-158. Clausen, C. P. 1978. Thysanoptera. Phlaeothripidae, pp. 18-19. Jn Clausen, C. P. ed. Introduced para- sites and predators of arthropod pests and weeds: a world review. United States Department of Ag- riculture., Agricultural Research Service, Agricul- ture Handbook No. 480. 545 pp. Davis, C. J. and N. L. Krauss, 1965. Recent introduc- tions for biological control in Hawaii. Proceedings of the Hawaiian Entomological Society 19: 88— 90. Eldredge, L. G. and S. E. Miller. 1995. How many species are there in Hawaii? Bishop Museum Oc- casional Papers No. 41: 3-18. . 1997. Numbers of Hawaiian species: Supple- ment 2, including a review of freshwater inverte- brates. Bishop Museum Occasional Papers No. 48: 3-22. Herring, J. L. 1966a. The correct name for an antho- coric predator of the Cuban laurel thrips. Proceed- ings of the Entomological Society of Washington 68: 93-94. . 1966b. The genus Orius of the Western Hemi- sphere (Hemiptera: Anthocoridae). Annals of the Entomological Society of America 59: 1093— 1109. 1979. Keys to genera of Anthocoridae of America north of Mexico, with description of a new genus (Hemiptera: Heteroptera). Florida En- tomologist 59: 143-150. Kirkaldy, G. W. 1910. Further notes on Hemiptera, chiefly Hawaiian. Proceedings of the Hawaiian Entomological Society 2: 118-123. Lattin, J. D. 1999. Bionomics of the Anthocoridae. An- nual Review of Entomology 44: 207-231. . 2000. Chapter 26. Economic Importance of minute pirate bugs (Anthocoridae), pp. 607—637. In Schaefer, C. W. and A. R. Panzini, eds. Heter- optera of Economic Importance. CRC Press, New York. 828 pp. Miller, S. E. and L. G. Eldredge. 1996. Numbers of Hawaiian species: Supplement 1. Bishop Museum Occasional Papers No. 45: 8-17. Nishida, G. M., ed. 1994. Hawaiian terrestrial arthro- pod checklist. 2nd ed. Bishop Museum Technical Report No. 4. 287 pp. Oatman, E. R. 1978. Noctuidae, pp. 205-210. In Clau- sen, C. P., ed. Introduced parasites and predators of arthropod pests and weeds: A world review. United States Department of Agriculture, Agri- cultural Research Service, Agricultural Handbook No. 480. 545 pp. Péricart, J. 1972. Faune de 1’ Europe et du Bassin Med- iterranéen. No. 7. Hémiptéres Anthocoridae, Ci- micidae et Microphysidae de 1’ Ouest-Paléartique. Masson et Cie, Paris. 402 pp. Scott, J. M., S. Mountainspring, EK L. Ramsey, and C. B. Kepler. 1986. Forest bird communities of the Hawaiian Islands: Their dynamics, ecology and conservation. Studies in Avian Biology 9: 1—431. Takara, J. and T. Nishida. 1981. Eggs of the Oriental fruit fly for rearing the predacious anthocorid, Or- ius insidious (Say). Proceedings of the Hawaiian Entomological Society 23: 441-445. Weber, P. W. 1953. Recent liberations of beneficial in- sects in Hawaii. 2. Proceedings of the Hawaiian Entomological Society 15: 127-130. White, FE B. 1879. Descriptions of new Anthocoridae. Entomologist’s Monthly Magazine 16: 142-148. Zimmerman, E. G. 1948. Insects of Hawaii. Vol. 3 Heteroptera. University of Hawaii Press, Hono- lulu. 255 pp. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 389-395 A NEW GENUS AND SPECIES OF PSEPHENINAE (COLEOPTERA: DRYOPOIDEA: PSEPHENIDAE) FROM MEXICO ROBERTO ARCE-PEREZ AND RODOLFO NOVELO-GUTIERREZ Instituto de Ecologia, A.C. Departamento de Entomologia, Apartado Postal 63, 91000 Xalapa, Veracruz, Mexico. (e-mail: entomol@ecologia.edu.mx; novelor@ecologia. edu.mx) Abstract.—Psephenotarsis triangularis, n. gen. and n. sp., is described and illustrated from Veracruz State, Mexico. Psephenotarsis triangularis can be separated from other Psepheninae by the shape of the body and length of the antenna (Pheneps), the size proportion between the phallobase and the parameres (Pheneps), the pronotum and tarsi (Psephenops), and the tarsi and the median lobe of male genitalia (Psephenus). Psephen- otarsis appears to be more closely related to Psephenops than other Psepheninae. A key for the New World genera of Psepheninae is provided. Resumen.—Se describe e ilustra a Psephenotarsis triangularis, n. gen. y n. sp., con base en especimenes del estado de Veracruz, México. Psephenotarsis triangularis se puede diferenciar de otros Psepheninae por la forma del cuerpo y la longitud de la antena (Pheneps), la proporcion entre la falobase y los parameros (Pheneps), el pronoto y los tarsos (Psephenops), y los tarsos y el l6bulo medio de los genitales masculinos (Pse- phenus). Al parecer Psephenotarsis esta mas cercanamente relacionado a Psephenops que al resto de los Psepheninae. Se proporciona una clave para la identificaci6n de los géneros de Psepheninae. Key Words: Psephenidae, Psepheninae, new genus, description, key, Veracruz State, Mexico The subfamily Psepheninae is represent- ed in the New World by the genera Pse- phenus Haldeman 1853, Psephenops Grou- velle 1898 and Pheneps Darlington 1936 (Brown 1981, Spangler 1982). Psephenus contains 15 species (seven in North Amer- ica, two of them shared with Mexico; four species in Mesoamerica; three species in Brazil and one species in Peru); Psephen- ops has seven species (Arce-Pérez and Novelo-Gutiérrez 2000) with tropical dis- tribution (Mexico, Guatemala, Costa Rica, Colombia, Argentina, Haiti, and the Antil- les); Pheneps has four species also with tropical distribution (Venezuela, Surinam, Haiti, and Cuba). In this paper, we describe a new genus and a new species based on two males and one female recently collect- ed in a mountain cloud forest at 1800 m altitude, in the Municipality of Xico, Ve- racruz State, Mexico. Psephenotarsis Arce-Pérez, new genus (Figs. 1—3) Description.—Head short, transverse, concave dorsally between compound eyes, quite visible from above (Fig. 1); eyes spherical, very prominent, reddish yellow; antenna short (e.g., hardly reaching the apex of scutellum), moniliform, 11-seg- 390 Figs. 1-5. mented, scape robust, larger than remainder of antennomeres, scape and pedicel yellow- ish brown, flagellomeres dark brown to black. Pronotum trapezoidal, its posterior mar- gin quite smooth, never crenulate nor ru- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1-3, Psephenotarsis triangularis. 1, Habitus, dorsal view. 2, Prosternum, ventral view. 3, Protarsal segments. 4, Habitus of Pheneps cursitatus, dorsal view. 5, Protarsal segments of Pheneps antennalis. (Fig. 4 from Spangler 1987; Fig. 5 from Spangler and Steiner 1983). gose, lateral margins expanded, without dorsal carinae; prosternum short, neither raised nor carinate at its mediobasal por- tion, prosternal process long, parallel-sided at middle then gradually diverging to form a lanceolate apex, with a weak longitudinal VOLUME 103, NUMBER 2 carina on its apex (Fig. 2); mesosternum clearly grooved and widely bifurcate api- cally, making both procoxa and mesocoxa appear more separate from each other than in other Psepheninae. Basal four tarsomeres notably triangular (Fig. 3), flattened and widely emarginate at apex, beset with long black setae, tarsomeres 1 and 2 with dense ventral vestiture of long setae, setae on first tarsomere reaching basal half of second one, setae on second tarsomere extending to base of fifth one, covering tarsomeres 3 and 4 ventrally; fifth tarsomere longest, as long as preceding ones together, stout, sub- cylindrical; tarsal claws entire, never bifid or split, widely separate, downcurved in apical half, widened at base and without teeth or accessory membranes. Male genitalia subrectangular, trilobate; phallobase shorter than parameres; para- meres shorter than median lobe, internal margin of apical half sinuate, ending in a more or less sharply-pointed tip, the later- oapical portion membranous; median lobe long, wide, in lateral view moderately con- vex ventrally, sclerotized. Remarks.—Psephenotarsis can be sepa- rated from other genera of Psepheninae by the following: body more slender in Phe- neps, antenna longer, reaching 0.75 the length of elytra, pronotum and elytra with- out expanded margins (Fig. 4), tarsomeres 1—2 clearly stouter than any of apical three tarsomeres (Fig. 5), phallobase notably larger than parameres (Fig. 9). The basal half of pronotum in Psephen- ops has a longitudinal crest (Fig. 6); tarso- meres | and 2, and occasionally 3, bearing long, wide, ventral pads which totally cover the apical tarsomeres (Fig. 7); median lobe of male genitalia mostly membranous (Fig. 10). The basal four tarsomeres in Psephenus are slender and subcylindrical, slightly emarginate at apex, with delicate, minute setae or papillae ventrally (Fig. 8). The me- dian lobe of male genitalia mostly membra- nous and shorter or as long as parameres (Fig. 11). 391 Psephenotarsis appears to be more close- ly related to Psephenops than other Pse- pheninae, mainly by the short antennae, lat- eral margins of pronotum and elytra ridged, posterior margins of sternites | and 2 slight- ly sinuate or emarginate, the basal two tar- someres with long vestiture of setae, and in general appearance. Etymology.—From the Greek Psepheno = hidden, and tarsis = tarsomeres, making reference to this hitherto unknown genus with a characteristic tarsal shape which dis- tinguishes this genus from other Psepheni- nes. Psephenotarsis triangularis Arce-Pérez, new species (Figs. 1-3, 12—13) Description.—Holotype male: Body oval and depressed, total length 4.20 mm, max- imum humeral width 2.05 mm. Head, pron- otum and scutellum black, covered with in- termingled short and long dark-reddish se- tae; elytra dark reddish-brown, densely cov- ered with short, brilliant yellow setae arranged in four longitudinal bands, and long, dark reddish setae on entire surface. Venter reddish-brown, with short and long reddish-yellow setae. Coxae, tibiae and tarsi reddish-brown, trochanters and femora yel- lowish. Head: short, transverse (Fig. 1), concave dorsally between compound eyes; eyes spherical, very prominent; clypeus subrect- angular, wider than long, slanting, forming an angle less than 90 degrees in relation to frons, its apex widely emarginate; labrum as clypeus but shorter; antenna short, hardly reaching apex of scutellum, moniliform, 11- segmented, covered with long reddish-black setae, scape larger than remainder of anten- nomeres, slightly curved and yellowish, pedicel half as long as scape; maxillary palp 4-segmented (Fig. 12), slightly longer than first two antennomeres combined, bas- al three palpomeres subcylindric, first one shortest, second one 3 times longer than first one and wider, third palpomere half as long as second one but wider, fourth one 892 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 6-10. 10 6-7 and 10, Psephenops mexicanus. 6, Pronotum, dorsal view. 7, Protarsal segments. 8, Protarsal segments of Psephenus sp. 9, Male genitalia of Pheneps antennalis, dorsal view. 10, Male genitalia of P. mexicanus, (a) ventral view, (b) dorsal view. (Figs. 6—7 and 10 from Arce-Pérez and Novelo-Gutiérrez 2000; Fig. 9 from Spangler and Steiner 1983). slightly longer than 2nd one but stouter and acuminate, laterally compressed and trun- cate at apex, all palpomeres reddish brown and covered with black setae; labial palp small (Fig. 12), 3-segmented, not extending beyond basal margin of second maxillary palpomere, basal two labial palpomeres cy- lindric, larger than third one which is sub- spherical, all of them reddish brown with black setae. Thorax: (basal Pronotum trapezoidal width 1.75 mm, apical width | mm), wider than long (width 1.75 mm, length 0.85 mm), its anterior margin arcuate, posterior margin bisinuate and slightly narrower than elytral bases, lateral margins straight and convergent at their anterior half, then sud- denly expanded at posterior half, postero- lateral angles obtuse. Scutellum short, tri- angular, its apex rounded. Elytra 3.2 mm length (from base to apex of elytral suture), 2.01 mm width (measured across calla hu- VOLUME 103, NUMBER 2 393 Figs. 11-13. otarsis triangularis. 12, Maxillary and labial palpi. 13, Male genitalia, (a) ventral view, (b) dorsal view, (c) lateral view. meralis), gradually widening caudad, reach- ing its maximum width at middle then nar- rowing to apex entirely covering abdomen, beset with four longitudinal bands of yel- low, dense, short setae intermingled with four longitudinal bands of reddish brown, sparse, long setae, these two types of bands 11, Male genitalia of Psephenus palpalis, (a) ventral view, (b) lateral view. 12—13, Psephen- extending along entire length of elytra. Prosternum large, as described for genus; procoxa globose; femur ovate and thick, its internal margin and apex dark; tibia slender and straight, slightly shorter than femur, with small ventral tooth on apex; tarsi as described for genus but first tarsomere 394 widely concave at its apical margin, second to fourth tarsomeres with their apical mar- gins V-shaped, sharply-pointed lateral ex- tensions on second tarsomere totally cov- ering third one (Fig. 3). Mesosternum short, mesosternal groove deep for reception of prosternal process, clearly bifurcate caudad; mesocoxa prominent, slightly transverse, other features of midleg as described above except tibia without apical tooth. Metaster- num large, slightly convex, with longitudi- nal, shallow, wide groove at middle which ends in a semi-triangular cavity; metacoxa transverse and wide, trochanter long and tri- angular, other features as described above. Abdomen: Convex ventrally but slightly flattened on midline, with seven sterna, sec- ond sternum largest, sixth sternum shortest and slightly emarginate at middle, posterior margin of sterna | and 2 slightly sinuate at middle; abdominal segment 7 strongly re- duced to an oval sclerite; pygidium oval, convex, with long yellowish black setae. Genitalia: Short, wide, subrectangular, trilobate; paramere shorter than median lobe (Figs. 13a, b), robust, in dorsal view (Fig. 13b) its basal half subrectangular with internal margin in close contact with that of other paramere, its apical half abruptly and strongly narrowed at 0.75 of paramere’s length ending in a sharp point, its internal margin sigmoid and widely separate from that of other paramere by a drop-shaped space; a translucid membrane at lateroapi- cal part (Fig. 13). Median lobe long, sub- cylindric, sclerotized (Figs. 13a, c), in lat- eral view (Fig. 13c) moderately convex ventrally. Total length (from basal margin of phallobase to apex of median lobe) 0.56 mm; length of phallobase 0.26 mm, para- meres 0.30 mm, median lobe 0.33 mm. Allotype female: Slightly larger and stouter than male, total length 4.2 mm, maximum width (at level of humeri) 2.2 mm. In other features similar to male ex- cept: abdomen wider than that of male, with six sterna, first two sterna with their pos- terior margins slightly sinuate. Type material——Holotype d labeled: PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON “MEXICO, VERACRUZ, XICO, Xico Viejo, 1,800 m, bosque mes6dfilo de mon- tana 16-VIII-98, Roberto Arce col.” Allo- type ¢: same data as holotype, except: 3- VIII-97. Paratype d: same data as holotype. All material deposited at Colecci6én Ento- moldgica del Instituto de Ecologia, A.C., Xalapa, Veracruz, México. Etymology.—The name triangularis re- feres to the triangular shape of the basal four tarsomeres of the legs. Type locality.—xXico Viejo, Municipality of Xico, Veracruz State, México. Habitat.—Specimens of Psephenotarsis triangularis were captured in a stream with rapid water flow, on overhanging riparian vegetation and on big rocks not exposed di- rectly to the water current but in the spray zone. KEY TO THE GENERA OF PSEPHENINAE OF THE NEw WORLD [Modified from Brown (1976) and White & Brigham (1996)] 1. Antenna long, filiform, extending notably be- yond the elytral humeri (Fig. 4); pro- and me- sotarsomeres | and 2 markedly stouter than the following three and slightly emarginate at apex (Fig. 5); phallobase much longer and wider than) parameres| (E19) een eee eee Pheneps — Antenna short, moniliform or subserrate, scarcely or not extending the elytral humeri; pro- and mesotarsomeres | and 2 of variable length but never stouter than following three (Figs. 3,7—8); phallobase subequal in length to parameres . Tarsomeres 1 and 2, and, occasionally tarso- mere 3, with long, spongy-like laminar expan- sions almost completely covering the remain- ing tarsomeres (Fig. 7); pronotum with a well developed, medio-basal, longitudinal carina (Fig. 6) Psephenops — Tarsomeres | and 2 without such laminar ex- pansions; pronotum without longitudinal carina i) 3. Tarsomeres 1—4 short, triangular, flattened, deeply emarginate for the reception of the fol- lowing tarsomere; tarsomeres 1—2 with long ventral setae or papillae, those on 2 reaching the base of the fifth tarsomere (Fig. 3) ite 1S aha Nas aregs tight eda ela Psephenotarsis, n. gen. — Tarsomeres 1—4 subconic and slender, scarcely emarginate at apex; tarsomeres 1—2 usually VOLUME 103, NUMBER 2 with ventral setae or papillae that never cover the contiguous tarsomere (Fig. 8) .... Psephenus ACKNOWLEDGMENTS We extended our thanks to Dr. Harley P. Brown (University of Oklahoma) for his in- valuable criticism on the final manuscript as well as for the English corrections. We also thank Biol. Leonardo Delgado (Xalapa) for his comments on a preliminary manuscript. Biol. Cesar V. Rojas (Xalapa) edited the il- lustrations. LITERATURE CITED Arce-Pérez, R. and R. Novelo-Gutiérrez. 2000. First record of the genus Psephenops (Coleoptera: Pse- phenidae) from Mexico, with a description of a new species. Entomological News 111(3): 196— 200. Brown, H. P. 1976. Aquatic dryopoid beetles (Cole- optera) of the United States. United States Envi- ronmental Protection Agency. Water Pollution 395 Control Research Series 18050/72 (Second Print- ing). Aquatic Biology Section. Cincinnati, Ohio, 82 pp. . 1981. A distributional survey of the world genera of riffle beetles (Coleoptera: Dryopidae, Elmidae and Psephenidae). The Pan-Pacific En- tomologist 57(1): 133-148. Spangler, P. J. 1987. A new species of water penny beetle, Pheneps cursitatus, from Cerro de la Neb- lina, Venezuela (Coleoptera: Dryopoidea: Pse- phenidae). Proceedings of the Entomological So- ciety of Washington 89(2): 219-225. . 1982. Coleoptera, pp. 323-395. Jn Hurlbert, S. H. and A. Villalobos-Figueroa, eds. Aquatic Bi- ota of Mexico, Central America and West Indies. San Diego, University of San Diego, California. Spangler, P. J. and W. E. Steiner. 1983. New species of beetles of the genera E/moparnus and Pheneps from Suriname (Coleoptera: Dryopidae: Psephen- idae). Proceedings of the Entomological Society of Washington 85(4): 826-839. White, D. S. and W. V. Brigham. 1996. Aquatic Co- leoptera, pp. 399—473. In Merritt, R. W. and K. W. Cummins, eds. An Introduction to the Aquatic Insects of North America. Third Edition, Kendall/ Hunt Publishing Company, Dubuque, Iowa. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 396-402 REVIEW AND FIRST NEW WORLD ENDEMIC OF THE STREPSIPTERAN GENUS CORIOXENOS BLAIR (STREPSIPTERA: CORIOXENIDAE: CORIOXENINAE) JERRY L. COOK Department of Biological Sciences, Sam Houston State University, Huntsville, TX 77341-2116, U.S.A. (e-mail: bio_jlc@shsu.edu) Abstract.—Corioxenos acucyrtophallus, n. sp., described from two males collected in Chiapas, Mexico, represents the first endemic new world species of the genus. A female Corioxenos is reported from its host in Vera Cruz, Mexico, but is left undescribed because it is not possible to determine if it is the female of C. acucyrtophallus n. sp. or if it represents a new species. A key to the Corioxenos species is given and wing venation terminology of Corioxenos is discussed. Key Words: The genus Corioxenos Blair is a small group of Strepsiptera characterized by males having seven antennal segments, with lateral flabella on the third and fourth segments; four segmented tarsi, with the apical segment short, bilobed, and without claws; mandibles absent; aedeagus without an apical hook; and hind wings with de- tached veins. However, all of these char- acters were not correctly diagnosed in the original generic description (Blair 1936). Blair considered the antennae to be “‘five- jointed,’ but Kinzelbach (1972) correctly diagnosed the antennae as seven-segment- ed. Furthermore, Baliga (1967) mistakenly described Corioxenos raoi as having three segmented tarsi, although the habitus draw- ing clearly shows four tarsal segments. Bal- iga also misdiagnosed the number of anten- nal segments as being five instead of seven. Females are characterized as having an elongated cephalothorax oriented such that its ventral side is towards the host; narrow brood canal opening in the head region of the cephalothorax; rudiments of ommatidia on the cephalothorax; and abdomen with taxonomy, key, wing morphology, Corioxenos, Pentatomidae eight dorsal papillae (Kathirithamby 1989). All known hosts are in the heteropteran family Pentatomidae. Previously, only two species of Coriox- enos have been described. Corioxenos an- testiae Blair was described from male, fe- male, and triungulin specimens associated with Antestia lineaticollis Stal from the slopes of Kilimanjaro, Tanganyika (Blair 1936). Kinzelbach (1971) listed Antestia faeceta Germar as another host of C. an- testiae, and Kinzelbach (1971) reported C. antestiae from the Dominican Republic, but inferred that it was not native to that area. Kathirihamby (1992) reported the same dis- tribution, with no further explanation. A second species, C. raoi Baliga, was de- scribed from male, female, and triungulin specimens associated with Antestiopsis cru- ciata (Fabricius) from around Bangalore, India (Baliga 1967). Here I describe the first Corioxenos spe- cies endemic to the Western Hemisphere from two males collected in Chiapas, Mex- ico. I also report on a female Corioxenos that may be the female of this newly de- VOLUME 103, NUMBER 2 scribed species. However, there is not enough evidence at this time to consider it conspecific. Along with the new species de- scription, a key to all species of Corioxenos males is provided. KEY TO SPECIES OF CORIOXENOS MALES 1. R, and R, subequal in length (Fig. 9); aedeagus almost twice as long as first tarsal segment; INTEXI CON aatancc sheen cian C. acucyrtophallus, n. sp. — R, much shorter than R,;; aedeagus length about equal to first tarsal segment ........ 2, . Aedeagus begins tapering at about % its length from base; maxillary palp % as long as 5th an- tennal segment; flagella of antennal segments 3 and 4 subequal to 5 + 6 + 7; Tanganyika, DomimicansRepublicga ee C. antestiae — Aedeagus begins tapering at over % its length from base; maxillary palp % as long as Sth an- tennal segment; antennal segment 3 shorter that 4, both 3 and 4 shorter than 5 + 6 + 7; India C. raoi i) Corioxenos acucyrtophallus Cook, new species (Figs. 1-4, 9) Male description.—Body coloration: heavily sclerotized regions of head and tho- rax golden-brown to brown, other regions tan. Total body length 2.2 to 2.4 mm. Head rectangular, strongly transverse, enlarged somewhat at eyes, sclerites as in Fig. 1. Head width including eyes 0.56 mm; dis- tance between eyes 0.45 mm. Antenna with flabellum on segments IV and V; segments I and II with pubescent hairs; segments IJ-— VII with pubescent hairs and sensory cups. Length of antennal segments, including fla- bellum where present (mm.): I = 0.05, Il = 0.05, Il = 0.62, IV = 0.58, V = 0.38, VI = 0.07, VII = 0.25. Mouthparts: only max- illary palp present, 2-segmented, second segment tapering to sharp point, I = 0.10 mm, II = 0.20 mm (Fig. 2). Thorax sclerites as in Fig. 1; pronotum narrow, transverse; mesonotum irregular-shaped; metascutum- prescutum rounded anteriorly; prealar tri- angular; scutellum triangular, much longer than wide; postscutellum elongate, rounded posteriorly; trapezoidal membrane between scutellum and postscutellum. Fore wing 397 club-shaped with R half way to apex. Hind wing (Fig. 9) smoky brown, no veins ex- tending to wing margin; Sc extending half way to wing margin or less; R, extending % to wing margin; R, beginning posterior to and near apex of R,; extending to near wing margin; R, with distinct bend, sube- qual to R,; R, extending about %4 to wing margin; R, beginning posterior and just pri- or to end of R,; M extending to near wing margin; CuA, extending about %4 to wing margin; CuA, extending about % to wing margin; CuP near wing posterior, extending to near 2 wing margin. Leg shapes and rel- ative lengths as in Fig. 4; prothoracic and mesothoracic femora grooved laterally such that tibiae will be held in grooves when folded; prothoracic and mesothoracic fem- ora subequal in length to corresponding tib- iae; metathoracic femur L-shaped, shorter than metathoracic tibia; tarsi 4-segmented on all legs; Ist tarsal segment without cir- cular sensory spot; 2nd tarsal segment with sensory spot near base; 3rd and 4th tarsal segments with sensory spot on directed op- posite of sensory spot on 2nd tarsal seg- ment, located in middle of segment; 4th tar- sal segment bilobed and without claws. Ab- domen as in Fig. 1. Aedeagus thin and gradually tapering from base to sharp apex; curved convexly in lateral view (Fig. 3). Female.—Unknown. Larva.—Unknown. Diagnosis.—Corioxenos acucyrtophallus can be easily distinguished from both C. antestiae and C. raoi by the length rela- tionship of the R, and R, wing veins (Fig. 9), shape of the aedeagus (Fig. 3), and gen- eral shape of the thoracic tergites. In C. ac- ucyrtophallus, R, and R,; are subequal in length (Fig. 9), while in C. antestiae and C. raoi R, is less than half as long as R, (Figs. 5-8). The R, vein of C. acucyrtophallus is distinctly bent, while this vein is straight in both C. antestiae and C. raoi. The aedeagus of C. acucyrtophallus is long and needle- like, tapering gradually from the base to the apex (Fig. 3). The aedeagus of C. antestiae and C. raoi begin tapering some distance 398 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Corioxenos acucyrtophallus, adult male, habitus. Scale bar = 1.0 mm. from the base, at about % its length in C. antestiae and at about % its length in C. raoi. The differences in general shapes of the thoracic tergites between the Coriox- enos species is most apparent in the scutel- lum and postscutellum. In C. acucyrtophal- lus, these tergites are relatively longer than those of C. antestiae and C. raoi. In gen- eral, the two Old World species have char- acters that are much more similar to each other than to C. acucyrtophallus. Etymology.—The specific epithet is de- VOLUME 103, NUMBER 2 399 Fig. 2-4. scriptive of the aedeagus using the Greek roots ‘“‘acu’’ = needle, “‘cyrto”” = curved, convex, and “‘phallus’’ = the penis. Holotype.—¢d, Mexico, Chiapas, Mpio. San Cristobal, Reserva Huitepec, 7,860’, 16°46’06"N; 92°41'04”"W, 2-14 VIII 1997, Woolley, Gonzalez, and Galdamez, Malaise trap 97/068. Deposited in Colecci6n Ento- molégica, El Colegio de la Frontera Sur (ECOSUR). Paratype.—1 36, Mexico, Chiapas, Mpio. Corioxenos acucyrtophallus. 2, Maxillary palps. 3, Aedeagus and genital capsule. 4, From top to bottom, prothoracic leg, mesothoracic leg, and metathoracic leg. San Cristobal, Reserva Huitepec, 8,160’, 16°46'06"N; 92°41'04”"W, 2-14 VIII 1997, Woolley, Galdamez, & Gonzalez, Malaise trap 97/070. Deposited in the Museum of Entomology, Texas A&M University. Corioxenos sp. A female of a Corioxenos species was collected by A. R. Gillogly in Veracruz, Mexico, in 1997. This specimen was col- lected in a species of Edessa (Pentatomi- 400 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. 5-8. raoi redrawn from Beliga (1967), 7, C. antestiae redrawn from Cooper (1938). 8, C. antestiae redrawn from Kinzelbach (1971). dae), 8 km south east of Monte Pio and can be identified as Corioxenos using the key published by Miyamoto and Kifune (1984). However, the only evidence that this is the female to C. acucyrtophallus is that they both were collected in southern Mexico, and no other species of Corioxenos is known from that area of the New World. Due to these circumstances, I choose not to describe this specimen as a new species at this time, thus eliminating the possibility of Corioxenos hindwing veins, as originally labeled. 5, C. antestiae redrawn from Blair (1936). 6, C. erecting a possible synonym, or incorrectly associating it with the males described here. The specimen is deposited in the collection of Jerry Cook. CORIOXENOS WING VEINATION There is considerable discrepancy in the labeling of veins in Corioxenos wings. Blair (1936) described C. antestiae as having ‘“‘wings with median vein forked distally, the branch detached.”’ From the figure that VOLUME 103, NUMBER 2 eno: Hindwing of Corioxenos acucyrtophallus. accompanies his description (redrawn in Fig. 5), it is clear that he was referring to the radial vein, specifically R, and R,. Blair also described a third anal vein “‘lying close to the inner margin,” which should be des- ignated as CuP. Baliga (1967) also consid- ered part of the radial vein, R,, to be the medial vein (redrawn in Fig. 6). Baliga did not designate the detached radial veins as being radial or medial, and named R, as simply another detached vein. Baliga la- beled the vein now considered to be M as Cu and followed Blair’s naming of the re- maining posterior veins. Cooper (1938) gave a different diagnosis of the wing veins (redrawn in Fig. 7) and designated what I now consider R, as the radial sector. He joined veins R, and R, and called this vein M,,,. The detached vein was designated as M,,,. The other posterior veins were then given an improper designation of Cu, 2nd A and 3rd A, respectively. Cooper’s diag- nosis was made using a wing “‘about half way through the pupal stadium.”’ Kinzel- bach (1971) attempted to straighten out this terminology, labeling the pupal wing draw- ing of C. antestiae as drawn by Cooper 401 (1938) for his diagnosis (redrawn in Fig. 8). This pupal wing did not show the devel- opment of R,, the differentiation of CuA, and CuA,, or the presence of CuP. This pu- pal wing also did not show any of the veins as yet detached, but aids in showing that the radial veins that are detached in the adult wing are truly radial veins. Part of the confusion of wing vein des- ignation may have arisen from the difficulty in seeing the origin of the wing veins. Blair (1936) and Beliga (1967) did not clearly show the origin of veins, and Cooper (1938) drew veins with a combined origin in places where they are clearly separated. The specimens used to describe C. acucyr- tophallus being in pristine condition facili- tated the correct designation of wing veins that I propose. The origins of all veins are clearly seen in both the holotype and par- atype. Therefore, species of Corioxenos have five radial veins, three of which are detached in the adult wing. The next vein, posteriorly, is designated as M because it has a separate origin. Kinzelbach (1971) la- beled this vein as MA, but since there is no evidence of any other medial veins, it seems inappropriate to designate it as the anterior medial. The next two veins have a common origin and are designated CuA, and CuA, respectively. The posterior most vein has a separate origin and is thus des- ignated as postcubitus, CuP. This new di- agnosis is shown in the wing of C. acucyr- tophallus shown in Fig. 9. ACKNOWLEDGMENTS I thank Edward Riley for the loan of specimens from the Texas A&M Entomol- ogy Museum and James Woolley for allow- ing me to describe the new species from his specimens. I also thank Ver6nica Galdamez Estrada and Lorena Riuz Montoya, both of El Colegio de la Frontera Sur, San Cristobal de Las Casas, Chiapas, Mexico, for their part in making these specimens available and Alan Gillogly, Texas A&M University, for the parasitized Edessa specimen. I also 402 thank the anonymous reviewers for critical- ly reviewing the manuscript. LITERATURE CITED Baliga, H. 1967. A new species of Corioxenos (Sty- lopoidea) parasitizing Antestiopsis cruciata (FE) (Homoptera, Pentatomidae) in India. Bulletin of Entomological Research 57: 387-393. Blair, K. G. 1936. A new genus of Strepsiptera. Pro- ceedings of the Royal Entomological Society of London, Series B, 5: 113-117. Cooper, B. 1938. The internal anatomy of Corioxenos antestiae Blair (Strepsiptera). Proceedings of the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Royal Entomological Society of London, Series A, 13: 31-54. Kathirithamby, J. 1989. Review of the order Strepsip- tera. Systematic Entomology 14: 41—92. . 1992. Strepsiptera of Panama and Mesoam- erica, pp. 421—431. /n Quintero, D. and A. Aiello, eds. Insects of Panama and Mesoamerica, Oxford University Press, Oxford, England. Kinzelbach, R. K. 1971. Morphologische Befunde an Facherfiiglern und ihre phylogenetische Bedeu- tung (Insecta: Strepsiptera). Zoologica 41: 1—256. Miyamoto, S. and T. Kifune. 1984. Descriptions of a new genus and two new species of the Strepsiptera parasitic on Japanese Heteroptera (Strepsiptera, Corioxenidae). Kontya 52: 137-149. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 403-408 INCISALIA HENRICI (GROTE AND ROBINSON) (LEPIDOPTERA: LYCAENIDAE) REARED ON REPRODUCTIVE AND NON-REPRODUCTIVE TISSUES OF THREE DIFFERENT PLANT SPECIES GORDON FE PRATT AND CECILIA L. PIERCE Department of Entomology, University of California, Riverside, CA 92521, U.S.A. Abstract.—Larvae of the small hairstreak butterfly, Henry’s Elfin (/ncisalia henrici), feed on a wide variety of plants and plant parts. The larvae of this Elfin along the eastern coastal plain feed on young leaves and flowers of hollies (//ex sp.). Throughout much of the interior southeast the major food plant is believed to be flowers, flower buds, and young leaves of redbud (Cercis canadensis). While in the northern part of its range larvae have been found feeding on the fruits of a variety of plants including blueberries (Vac- cinium sp.) and cherries (Prunus sp.). The effects of food plant species (hollies, redbud, and cherry) and plant parts (leaves vs. flowers) were studied on percent survival, pupal weights, and development times. Larvae survived well on the different test host plants and host plant parts. The //ex reared pupae, despite being reared on the natal host, were significantly lower in weight than pupae of larvae reared on the other two plant species. Larvae reared on Cercis flowers and flower buds performed best in all categories. We conclude the subspecies utilizing //ex as a food plant did not become adapted to this plant because larvae perform better than on other plants, but due to other variables such as perhaps the availability or abundance of plants in the habitat. Key Words: It has been suggested that some lycaen- ids, particularly eumaeine hairstreaks, which feed on a broad variety of plants, are able to do so because they specialize on the flowers and fruits of their hosts (Pratt and Ballmer 1991). Presumably these reproduc- tive tissues exhibit a higher proportion of amino acids and protein to that of alkaloids and other secondary plant compounds (Chew and Robbins 1984, Pratt and Ball- mer 1991). Specializations to multiple food plants can be either as separate species or by local adaptations of a single species that is capable of feeding on a broad variety of plants (Fox and Morrow 1981). Lycaenid butterflies fit both types from species with restricted host ranges to generalists that have become locally specialized (Ballmer Lycaenidae, hairstreaks, host races, Theclinae, foodplant effects and Pratt 1989). For instance some lycaenid genera, such as Celastrina that feed on a broad variety of plants, exhibit both mo- nophagous species and generalists with lo- cal specialization in food plants as geo- graphic races (Pratt et al. 1994). In contrast some species, such as Strymon melinus Hubner, not only feed on a broad variety of plants locally, but on a variety of plant parts as well (Ballmer and Pratt 1989). Henry’s Elfin, Incisalia henrici (Grote and Robinson), is an interesting model spe- cies to test larval adaptations, since it feeds on the flowers, fruits, and leaves of a broad variety of plants and exhibits a number of geographic races (Pavulaan 1998, Gatrelle 1999). It uses six host families in nature: Aquifoliaceae [Jlex cassine L., I. opaca 404 Ait., I. vomitoria Ait., Nemopanthus mu- cronatus (L.) Trel.]; Caprifoliaceae [Vibur- num acerifolium (L.)]; Ebenaceae [Diospy- ros texana Scheele]; Ericaceae [Vaccinium pallidum Ait., Vaccinium corymbosum L., Leucothoe racemosa (L.) A. Gray, Gaylus- sacia baccata (Wangenn.)]; Fabaceae [Cer- cis canadensis L., Lupinus texensis Hook., Sophora secundiflora (Ort.) Lag ex. DC.]; Rhamnaceae [Rhamnus frangula L.]; and Rosaceae [Prunus virginiana L., Prunus domestica L., Prunus pennsylvanica L. f., Prunus americana Marsh., Prunus serotina J. E Ehrh.] (Gifford and Opler 1983, Opler and Krizek 1984, Scott 1986, Pratt, person- al observations, Pavulaan and Wright, in litt.). Larvae can be collected from the leaves and flowers of /lex opaca (Pratt, pers. obs.), leaves of Cercis canadensis (Wright, pers. comm.), or fruits of V. cor- ymbosum and P. virginiana (Pratt pers. obs.). Henry’s Elfin exhibits an interesting pat- tern of host utilization throughout its range. From the southern Appalachians of North America west to the Prairies it seems to specialize on Cercis, while along the coast- al plain it specializes on the evergreen /lex species. To the north of the range of these hosts in eastern Canada it specializes on a different variety of hosts particularly Pru- nus and Vaccinium species. Therefore this butterfly could exhibit local adaptations to the different plant species or may be adapt- ed to a broad variety of plants and using specific plants locally because of food plant availability (Fox and Morrow 1981). Gatrelle (1999) separates the subspecies of Henry’s Elfin into two groups, those that are Ilex feeding and those that are non-/lex feeding. He suggested that these groups could have already separated into sibling species, since larvae of one race of the /lex feeding group had poor survival when fed Cercis (redbud). Redbud is a major food plant of the non-/lex feeding group. Pavu- laan (1998) supported Gatrelle’s conclu- sions since he found holly-associated fe- males from Prince George’s County, Mary- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON land, when set up on Cercis would not ovi- posit, yet immediately did so when transferred to /lex opaca. Pavulaan also found that larvae of Incisalia henrici viri- dissima Pavulaan, a race that belongs to the Ilex feeding group, survived poorly on Cer- cls. Host race adaptations are believed to have played an important role in the spe- ciation of many insects, particularly in sym- patry (Tauber and Tauber 1989). These ad- aptations involve traits that relate to fitness within the different groups of the Henry’s Elfin as suggested by Gatrelle (1999). If the different races have made adaptations to their different food plants, then their sur- vival and fecundity on their natural food plants should be higher than on food plants of the other races. The objectives of this paper are to determine how the coastal race of I. henrici adapted to Ilex performs as to survival, weight, and development time on its natal verses host plants of its other races. MATERIALS AND METHODS Ova (n > 350) of /. henrici were field collected on April 20 1992 from Jlex opaca at Nanticoke Wildlife Refuge, 10 km west of Laurel, Sussex Co., Delaware. Ova were easily found by scanning the upper surface of older leaves (often near the midrib). First instar larvae which eclosed from these ova two days later were separated into six groups of 50 each and reared either on flowers or leaves of the following plants: Cercis canadensis (Fabaceae), Ilex opaca (Aquifoliaceae), and Prunus serotina (Ro- saceae). The newly eclosed larvae were transferred prior to any larval feeding via a camel hairbrush to their respective test host and reared in groups of 50 within square plastic storage containers (10 < 10 X 6 cm) at room temperature (23°C). Branches of Prunus and Ilex were ob- tained from Nanticoke Wildlife Refuge, and Cercis were collected from a woodlot at the University of Delaware, Newark, DE, and maintained in refrigerated plastic Ziploc® bags until needed. Larvae that were fed VOLUME 103, NUMBER 2 flowers (from male plants for /lex) were given branches stripped of leaves, while lar- vae fed leaves were given branches stripped of flowers. Due to the season of the year, at which the butterfly occurs, most of the leaves and flowers larvae were reared on were fresh young growth of all three plant species. Fresh and old plant parts were add- ed and removed daily, and the containers were cleaned twice daily at 12-hour inter- vals. When larvae reached fourth instar, to reduce the quantity of frass buildup in each container, they were divided in two equal groups and reared on the same host and plant part in two storage containers. Development times in days and number that survived to pupation and to adult were recorded for each of the six treatments. Two weeks after the last larva pupated, all pupae were weighed to the nearest milligram. On October 15, 1992, all pupae were placed on moist sterilized sand at 4°C for a four- month period. The pupae were removed in mid February 1993, and placed at 20—22°C in an incubator with 24 h of darkness until all live pupae eclosed. The data were analyzed by the SAS gen- eral linear models procedure (SAS Institute 1990). Data were grouped by treatments (Cercis leaves, Cercis flowers, Ilex leaves, Ilex flowers, Prunus leaves, and Prunus flowers), by host species, and by plant part (leaves and flowers). Means and standard errors were calculated for each group. Sig- nificance between groups for larval devel- opment time to pupation, pupal weights, and adult eclosion time and adult percent survival were determined by Tukey’s Stu- dentized Range (RSD) Test. RESULTS The lack of significant differences be- tween development times of larvae on leaves and flowers suggests that the plant parts did not consistently affect the rate of development in the same way across spe- cies, yet there were significant differences of larvae reared on flowers and leaves with- in some species (Cercis and Prunus) (Table 405 Table 1. Jncisalia henrici \arval development and proportion survival to pupation on different hosts and plant parts. Mean Survival Host Part N Development* (percent) Cercis Flowers 50 15.98 (0.80) A 100 Cercis Leaves 43 17.91 (0.97) C 86 Ilex Flowers 42 16.36 (1.78) AB 84 Ilex Leaves 46 16.68 (1.14) AB 92 Prunus Flowers 48 18.85 (1.54) D 96 Prunus Leaves 31 17.00 (1.48) B 62 * Means that are followed by a different letter are significantly different (P < 0.05). The standard devi- ations are in parentheses beside the mean development times. 1). Larvae reared on flowers and leaves of the same food plant were not significant (P > 0.05). The mean development times of larvae reared on Prunus were significantly slower than that of the other hosts (P < 0.05). The differences between species were in large part due to the long devel- opment times of larvae reared on Prunus flowers (Table 1). Most of the variations in development times amongst the six rearing treatments were between flowers of the dif- ferent hosts, with larvae reared on Cercis flowers taking the shortest time to pupate and those on Prunus flowers taking the lon- gest. The pupae of larvae reared on the natal host [lex opaca weighed significantly less from those reared on the other hosts (P < 0.0001). Pupal weights of larvae reared on flowers were not significantly different from those reared on leaves (P > 0.05). The heaviest mean pupal weights were of larvae reared on Cercis flowers and Prunus leaves, which were significantly different from the other four treatments but not from each oth- er (Table 2). The highest percent survival to adult was on Cercis flowers (98%), which was signif- icant from all other treatments (Table 3). The percent survivals of the other five treat- ments were not significant from each other. Adult eclosion times of /. henrici pupae from larvae reared on Prunus were signif- 406 Table 2. Mean pupal weights of /. henrici reared on different hosts and plant parts. Host Part N Mean Weight* Cercis Flowers 50 0.099 (0.01) A Cercis Leaves 37 0.089 (0.01) B Tlex Flowers 38 0.085 (0.01) B Ilex Leaves 4] 0.082 (0.01) B Prunus Flowers 36 0.088 (0.01) B Prunus Leaves 29 0.104 (0.02) A * Means that are followed by a different letter are significantly different (P < 0.05). The standard devi- ations are in parentheses beside the mean weights. icantly shorter than those reared on the oth- er two hosts (P > 0.0005). Also the eclo- sion times of pupae of larvae reared on flowers of a respective host were signifi- cantly different from pupae of larvae reared on leaves of the same host (P < 0.05). DISCUSSION In our model system, Jncisalia henrici thrived best on Cercis flowers. This treat- ment yielded the quickest larval develop- ment time (Table 1), a very high mean pu- pal weight (Table 2), and the greatest sur- vival to adult (Table 3). It has been sug- gested that flowers may provide advantages to lycaenid larvae due to their higher ratio of nutritional resources (amino acids and proteins) to non-nutritional resources (al- kaloids and other secondary chemicals) (Chew and Robbins 1984, Pratt and Ball- mer 1991). The family Fabaceae (which in- cludes Cercis) is known to have some of the highest ratios of amino acids and pro- teins to carbohydrates, most likely due to their association of nitrogen-fixing bacteria. Members of the Fabaceae are thought to be great nutritional resources to lycaenid lar- vae (Pratt and Ballmer 1991). In nature redbud (Cercis), is not the nat- urally available host for this model popu- lation of I. henrici. Cercis canadensis and Ilex opaca are utilized by I. henrici almost everywhere they occur. Yet these hosts do not exhibit much distributional overlap in nature (Gatrelle 1999). Prunus virginiana is PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 3. J/. henrici mean adult eclosion times from pupae and proportion survival from first instars to adults on different hosts and plant parts. Mean Survival Host Part N Eclosion* (percent) Cercis Flowers 49 9.67 (2.15) A 98* Cercis Leaves 37 6:81 G02) Bie 74: Ilex Flowers 35 6.91 (1.29) B 70 Ilex Leaves 37 9.51 (1.84) A 74 Prunus Flowers 30 7.73 (1.48) B 60 Prunus Leaves Diy 6.67 (0.78) B 54 * Means followed by an asterisk or a different letter are significant at P < 0.05. The standard deviations are in parentheses beside the mean eclosion times. common in many localities of both races, but does not appear to be used as a food plant, except in the northern part of the range where Cercis canadensis and Ilex species are both absent. Euphydryas editha, a butterfly that has had extensive research for over 30 years, has this type of host uti- lization, since when its natural host is ab- sent the butterfly will switch to other avail- able food plants that are not normally used (Singer et al. 1992, Singer and Thomas 1996). Interestingly, Incisalia henrici is quite abundant in areas where it utilizes hollies as food plants, while the butterfly is gen- erally uncommon in areas where the species is believed to be using redbud (Pratt, per- sonal observation, Pavulaan, personal com- munication). This observation suggests there could be advantages of feeding on holly over that of redbud. Also, Cercis flowers are not likely used to a great extent even in populations specifically adapted to Cercis. Cercis blooms early in nature be- fore the leaves (probable larval food in the field) break bud and well before most adult butterflies eclose from over-wintering pu- pae. The success of natural /lex-feeding /. henrici larvae on Cercis in the laboratory may be explained by 1) the ancestral food plant is redbud and holly is a recent adap- tation, 2) the ratio of nutritive to non-nutri- tive resources in redbud flowers is higher VOLUME 103, NUMBER 2 than its own food plant, 3) redbud flowers are devoid of growth toxins present in other hosts, and/or 4) unknown factors unmasked by laboratory conditions. In nature the abil- ity to feed and thrive on a specific plant is an obvious adaptive trait, but it may not be the only variable that makes one food plant better than another. Cercis may harbor large numbers of predators and parasites, espe- cially around young budding leaves where most eggs on Cercis are laid. In /lex pop- ulations, Trichogramma (Hymenoptera: Trichogrammatidae) parasitized eggs may be far less frequent on the old /lex leaves and have a distinct advantage over eggs Oviposited upon flowers, flower buds, or leaf buds (Pratt, personal observation). Old Ilex leaves on the other hand can’t feed first instar larvae, so newly hatched larvae must travel great distances in search of develop- ing leaves. If there were no advantage to Ovipositing on these old leaves, this adap- tation would therefore be counterproductive to the species. These sort of adaptations made by insects to escape parasitization have been observed in other host/parasite systems (Moore 1989, Schreiber et al. 2000). It would seem from the data in this study that Incisalia henrici has not formed sibling species based on host adaptations as sug- gested by Gatrelle (1999). If the Incisalia henrici races adapted to these different plants had formed species, the model pop- ulation adapted to /lex would be expected to do better on //ex than on either Cercis or Prunus. Instead this population adapted to Ilex did best on Cercis flowers, rather than on the natal host, /lex opaca. Instead Incis- alia henrici appears to fit more the pattern of a generalist that is able to feed on a broad variety of plants and has made local adap- tations to local food plants (Fox and Mor- row 1981). As expected in this model there was actually little differences in perfor- mance of larvae on the different food plants. Our J. henrici model system showed no significant differences in the developmental 407 parameters between larvae reared on flow- ers versus those reared on leaves. This elfin is adapted to a variety of food plants and plant resources. It does not appear to be adapted to any specific plant or plant part. The adaptations made by this species are broad in that it is able to feed on the flow- ers, fruits, and leaves of many plant species and it is not particularly specialized. That a butterfly with this capacity has formed dis- tinct localized races suggests that it has made localized adaptations independent of its ability to feed and grow on locally avail- able food plants. ACKNOWLEDGEMENTS We thank David M. Wright and Harry Pavulaan for reading the manuscript and all of their thoughtful suggestions provided throughout this study. Two reviewers, Rob- ert K. Robbins and anonymous reviewer, also provided much helpful editing and sug- gestions. We also thank John Pesek at the University of Delaware for his help with the statistics. We also acknowledge the late lep- idopterist Ben Zeigler for his deep interest in the food plant utilization of the Henry’s Elfin and other butterflies found in North America. LITERATURE CITED Ballmer, G. R. and G. EF Pratt. 1989. A Survey of the Last Instar Larvae of the Lycaenidae (Lepidop- tera) of California. Journal Research on the Lep- idoptera 27: 1—80. Chew, EF S. and R. K. Robbins. 1984. Egg-Laying in Butterflies, pp. 65-79. Jn Vane-Wright, R. I. and P. R. Ackery, eds. The Biology of Butterflies. Symposium of the Royal Entomological Society of London Number 11, Academic Press, Harcourt Brace Jovanovich, Publishers, 429 pp. Fox, L. R. and P. A. Morrow. 1981. Specialization: Species Property or Local Phenomenon? Science 211: 887-892. Gatrelle, R. R. 1999. An evolutionary subspecific as- sessment of Deciduphagus henrici (Lycaenidae) based on its utilization of //ex and non-//ex hosts: description of a third //ex associated subspecies. Designation of a neotype and type locality for De- ciduphagus irus. The Taxonomic Report of the In- ternational Lepidoptera Survey 1(6): 1—10. Gifford, S. M. and P. A. Opler. 1983. Natural history 408 of seven hairstreaks in coastal North Carolina. Journal of the Lepidopterists Society 37(2): 97— 105. Moore, S. D. 1989. Patterns of juvenile mortality with- in an oligophagous insect population. Ecology 70: 1726-1737. Opler, P. A. and G. O. Krizek. 1984. Butterflies East of the Great Plains. John Hopkins University Press, Baltimore MD, 294 pp. Pavulaan, H. 1998. A new subspecies of /ncisalia hen- rici (Grote & Robinson) (Lepidoptera: Lycaeni- dae) from the Outer Banks of North Carolina. Maryland Entomologist 4(2): 1-16. Pratt, G. EK and G. R. Ballmer. 1991. Acceptance of Lotus scoparius (Fabaceae) by larvae of Lycaenidae. Journal of the Lepidopterists Society 45: 12-28. Pratt, G. F, D. M. Wright, and H. Pavulaan. 1994. The various taxa and hosts of the North American Ce- lastrina (Lepidoptera: Lycaenidae). Proceedings of the Entomological Society of Washington 96: 566-578. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON SAS Institute. 1990. SAS/STAT user’s guide, version 6 4th ed, Vol. 2. SAS Institute, Cary, North Car- olina. Schreiber, S. J., L. R. Fox, and W. M. Getz. 2000. Coevolution of contrary choices in host-parasitoid systems. The American Naturalist 155: 637-648. Scott, J. A. 1986. The butterflies of North America. Stanford University Press, Stanford, CA, 583 pp. Singer, M. C., D. Ng, D. Vasco, and C. D. Thomas. 1992. Rapidly evolving associations among ovi- position preferences fail to constrain evolution of insect diet. The American Naturalist 139: 9-20. Singer, M. C. and C. D. Thomas. 1996. Evolutionary responses of a butterfly metapopulation to human- and climate-caused environmental variation. The American Naturalist 148: S9—S39. Tauber, C. A. and M. J. Tauber. 1989. Sympatric spe- ciation in insects: perception and perspective. /n Otte, D. and J. A. Endler, eds. Speciation and its Consequences, Sinauer Associates, Inc., Sunder- land, Massachusetts, 679 pp. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 409-427 A REVIEW OF THE NORTH AMERICAN SPECIES OF MEGARCYS KLAPALEK (PLECOPTERA: PERLODIDAE) BRIAN J. VANWIEREN, Boris C. KONDRATIEFF, AND BILL P. STARK (BJVW, BCK) Department of Bioagricultural Science and Pest Management, Colorado State University, Fort Collins, CO 80522, U.S.A. (e-mail: bkondrat@ceres.agsci. colostate.edu); (BPS) Department of Biology, Mississippi College, Clinton, MS 39058, WES; A: Abstract.—The arcynopterygine genus Megarcys in North America is reviewed. Five species, M. irregularis (Banks), M. signata (Hagen), M. subtruncata. Hanson, M. water- toni (Ricker) and M. yosemite (Needham and Claassen) are recognized. New illustrations for the males and females and a key to species are presented. Scanning electron photo- micrographs of the eggs are given. Key Words: The arcynopterygine genus Megarcys in North America (Stark and Szczytko 1988) has had a complex taxonomic history since the original description of Dictyopteryx sig- nata by Hagen in 1874. Currently, five spe- cies are recognized from North America (Stark 1998), following Ricker’s (1952) evaluation of the group. However, as Ricker pointed out “‘they could be treated almost equally as species, or as subspecies of Ha- gen’s signata.”’ Hanson’s (1942) review of Megarcys recognized only two species, M. signata and his new species, M. subtrun- cata. A current survey of the stoneflies of Mt. Rainier National Park, Washington, the type locality of two species, M. irregularis (Banks) and M. subtruncata, and the oc- currence of M. yosemite (Needham and Claassen), has provided material for a re- view of the genus in North America, and comparative descriptions and illustrations of the five North American species. MATERIALS AND METHODS Specimens examined in this study are housed in the collections of the Monte L. Plecoptera, Perlodidae, Megarcys, North America Bean Life Science Museum, Brigham Young University (BYU), C.P. Gillette Mu- seum of Arthropod Biodiversity, Colorado State University (CSU); Cornell University (CU); Llinois Natural History Survey (INHS); and the B. P. Stark Collection (BPSC). Illustrations were made of the male ter- minalia, head and prothorax, and female subgenital plates using a camera lucida. Techniques of Stark and Szczytko (1981) were used to prepare photomicrographs of eggs. Terminology used for describing adult morphological characters and eggs follow Stark and Szczytko (1988). We have used the term ‘“‘windsock-like”’ for the descrip- tion of the posterior process (ventral epi- proct surface) of the aedeagus. This term was used by Stark et al. (1998). The aedea- gus of three of the species, M. irregularis, M. signata and M. yosemite are illustrated for future comparisons, but are not used to distinguish taxa. The holotypes of M. irre- gularis, M. signata, and M. subtruncata were not examined, since the identity of these species were not in doubt. 410 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fe A N E | r gaps" tly t TT AAU tly eel CF : IY gree WHA 3 7 Figs. 1-7. Megarcys signata. 1, Male head and prothorax, dorsal. 2, Male terminalia, oblique view. 3, 9" tergum, dorsal. 4, Hemitergal lobe, lateral. 5, Lateral stylet, apex. 6, Aedeagus, dorsal. 7, Aedeagus, lateral. VOLUME 103, NUMBER 2 Megarcys signata (Hagen) (Figs. 1-7, 32, 37—40) Dictyopteryx signata Hagen 1874: 576. Type locality, Colorado. Holotype ¢, MCZ, not examined. Megarcys signata: Hanson (in part), 1941: 398. Arcynopteryx (Megarcys) signata: Ricker 525: Megarcys signata: Illies 1966: 371. Diagnosis.—The male of M. signata can be distinguished from the cognate species M. irregularis by presence of numerous spi- nule in the median cleft of the 9" tergum (Fig. 3). The female of M. signata is also very similar to M. irregularis, but can be usually separated by the subgenital plate produced % or more length of 9" sternum (Fig. 32). General color brown. Head with dark rugosities medially and posterior to the compound eyes, darker V-shaped pattern encompassing anterior ocellus; pale mid- dorsal stripe on prothorax (Fig. 1). Male.—Length of forewing 10-16 mm, body 15-18 mm. Anterior (dorsal) surface of epiproct sclerotized, posterior (ventral) surface windsock-like with narrowly round- ed anterodorsal hump, apex acutely tapered, not decurved at tip (Fig. 2). Hemitergal lobes U-shaped, greater than 14% as long as wide (Figs. 2, 4), bluntly rounded with spinules covering entire tip; lateral stylet with forward directed spine (Fig. 4). Ninth tergum swollen submedially, divided by median cleft, spinules covering entire cleft (Figs. 2-3). Aedeagus socklike, constricted about % distally (Figs. 6—7); apical lobe with basal setal patch dorsally, apically with deep evagination, lateral tubular processes extend proximally from basal lobe (Fig. 6). Female.—Length of forewing 18-21 mm, body 20—21 mm. Subgenital plate pro- duced to % or more of the 9" sternum, rounded with median cleft usually widest posteriorly (Fig. 32). Egg.—Collar stalked, rim usually deeply and irregularly incised, posterior surface of shoulder with ornate processes (Figs. 37— 411 38). Chorion covered with irregular deep pits without hexagonal pattern; surface ir- regularly indented including posterior pole (Figs. 38—40); micropylar row subequato- rial, ecolosion line absent (Fig 37). Distribution.—Megarcys signata, a widespread western North American spe- cies, can be common in small to medium sized high elevation streams of AK, BC, CO, ID, MT, NM, NV, UT, and WY (Stark 1998). In Colorado, this large predatory stonefly species is often found with Kogo- tus modestus (Banks) above 3,300 m. Remarks.—Stark and Szczytko (1988) provided good descriptions of the male ter- minalia and eggs of this species. All the known biological information for North American Megarcys refers to M. signata (see references cited by Stewart and Stark 1988). Recently, Taylor et al. (1999) re- ported that this species is semivoltine with delayed egg hatch. The nymph has been de- scribed in detail by Stewart and Stark (1988). Material examined.—COLORADO: Boulder Co., Middle St. Vrain, FR. 114 W. of Peaceful Valley, 17 July 1993, B. Kon- dratieff and W. Painter, 1 ¢ (CSU); Boulder Co., Trib. to Jasper Cr., Indian Peaks Wild, 12 September 1998, B. Kondratieff, 1 @ (CSU)z Eagles Go> Deep Cr Ra 7 June 1997, B. Kondratieff, 2 6,7 2 (CSU); Gilpin Co., South Boulder Cr., Rollinsville, 19 June 1986, B. Kondratieff, 7 6, 2 2 (CSU); Gilpin Co., South Boulder Cr., 2 mi. W. Tolland, 7 July 1991, B. Kondratieff and Re Durkee. 12.686) 2(GSU); Grand Co, Fraser R., above Robbers Roost Camp., 5 August 1973, R.W. Baumann, B.P. Stark, 11 6.03) Sox BY): Graad: Co.) Timbers Cr; Backcountry campsite, RMNP, 26 July 1989, C. Barker and M. Harris, 5 2 (CSU); Grand Co., W. St. Louis Cr, Fraser Exp. Forest, 24 July 1999, H. Rhodes, 1 ¢ (CSU); Gunnison Co., East R., near Avery campground, 13 August 1986, B. Armitage, 1 6, 1 2 (CSU); Gunnison Co., tribs. to Oh-Be Joyful Cr., Rt. 754, 14 August 1996, B. Kondratieff and N. Lorenzon, 1 ¢, 1 @ 412 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 8-12. Megarcys subtruncata. 8, Male head 10, 9 tergum, dorsal. 11, Hemitergal lobe, lateral, 12 (CSU); Hinsdale Co., Silver Cr., Redcloud Pk., 1 August 1992, B. Kondratieff, 1 2 (CSU); Hinsdale Co., Silver Cr., Silver Cr. Tr. 2 mi. from trailhead, 12,000 ft., 5 Sep- tember 1993, B. Kondratieff and R. Durfee, 12 and prothorax, dorsal. 9, Male terminalia, oblique view. , Lateral stylet, apex. 1 3, 1 2 (CSU); Hinsdale Co., Silver Cr. 12,000 ft., Redcloud Peak, 10 September 1993, S. Fitzgerald and T. Eckberg, 1 6, 3 2 (CSU); Hinsdale Co., Silver Cr. 12,000 ft., Redcloud Peak, 13 August 1995, S. VOLUME 103, NUMBER 2 Fitzgerald and A. Foley, 1 6 (CSU); Hins- dale Co., Nellie Cr. 12,000 ft., 1 August 1997, A. Ellingson, 1 2 (CSU); Hinsdale Co., Cataract Gulch 9,700 ft., 10 July 1998, A. Ellingson, 1 2 (CSU); Jackson Co., Michigan R., off Rt. 14 Lake Agnes, 2 Au- gust 1986, B. Kondratieff, 1 2 (CSU); Jackson Co., Michigan R., by Lake Agnes Rd., 17 July 1987, B. Kondratieff, 1 @ (CSU); Larimer Co., Spruce Cr. 9,700 ft., RMNP, 29 July 1989, M. Harris and M. Scott, 1 6 (CSU); Larimer Co., Skin Gulch, Stove Prairie Rd. 1 mi. S. of Hwy. 14, 8 May 1993, B. Kondratieff and R. Durfee, 1 2 (CSU); Larimer Co., Rawah Wilderness outlet Cr. Twin Crater Lakes 10,800—11,000 ft., 17 September 1995, S. Fitzgerald and A. Foley, 3 6,5 2 (CSU); Park Co., Four- mile Cr, 17 km W Pairplay, 13 July 2000, Haves Evans, 4.46, 2° 2° (CSU); Pitkin ‘Co., Independence Pass, Mt. Boy Park., 6 Au- gust 1943, J.A.R. and H.H. Ross, 1 ¢, 3 2 (BYU); Summit Co., Clear Cr., near Love- land Pass., 8, August 1973, R.W. Baumann and B. P. Stark, | d6 (BYU); Summit Co., Cr. in Avalanche Chute, W. side Peak 5, 2 September 1995, S. Fitzgerald and A. Fo- ley, | female (CSU); Summit Co., Union Cr., Copper Mt., 6 July 1997, B. Kondra- tieff, 1 2 (CSU); MONTANA: Gallatin Go. Pine Cr; 325i. off Traik’Cr. Rd. 10 June 1987, B. Kondratieff and E Kirchner, 556,42 (CSU); Glacier Co., Piegan Cr, Glacier N.P., 8 May 1965, A.R. Gaufin, 1 3 (BYU); Granite Co., Ranch Cr. at Grizzly Campground, 19 June 1967, M.L. Miner, 1 $6 (BYU); UTAH: Salt Lake Co., Millcreek Canyon Cr. at Log Haven, 12 May 1966, R.W. Baumann, 3 6, 2 2 (BYU); Utah Co., Stewarts Cr. Sundance Ski Area, 10 July 1986, R.W. Baumann and R. Nelson, 2 3 (BYU); Utah Co., Summit Cr. Tinney Flat Campground, Santaquin Canyon., 25 June 1998, S.M. Clark, 1 36, 1 2 (BYU); WY- OMING: Sheridan Co., Whedon Spring at Hwy 14A, 25 July 1998, D. Ruiter, 1 (CSU); Sublette Co., Bull Cr., Rt. 191/189 mile marker 153, 11 June 1987, B. Kon- dratieff, 1 ¢ (CSU); Teton Co., Granite Cr. 413 Falls, Bridger Teton N.E, 18 July 1999, P. Opler, 3 2 (CSU); CANADA: ALBERTA, Lake Louise, W.W. Baniff N.P., 26 July 1987,, J. Rabold? 1) 2.(ESU): Megarcys subtruncata Hanson (Figs. 8-12, 36, 41-44) Megarcys subtruncata Hanson 1942: 400. Type locality: Paradise Valley, Mt. Rain- ier National Park, Washington. Holotype 3, USNM, not examined. Arcynopteryx (Megarcys) subtruncata: Ricker 1952275. Megarcys subtruncata: [lies 1966: 371. Diagnosis.—The combination of the acute apex of the hemitergal lobe and trun- cate apex of the lateral stylet easily distin- guish the male of M. subtruncata from M. watertoni and M. yosemite. The female of M. subtruncata is very similar to M. water- toni. The subgenital plate of M. subtruncata usually covers less than % of sternum 9, whereas the subgenital plate of M. water- toni usually covers % or more of sternum 9. General color dark brown. Head with dis- tinct wide V-shaped pattern posterior to lat- eral ocelli, an inverted U-shaped pattern connecting ocelli; pale prothorax middorsal stripe (Fig. 8). Male.—Length of fore wing 13-16 mm, body 17—21 mm. Posterior surface of epi- proct windsock-like with broadly rounded anterodorsal hump; apex tapered and de- curved at tip (Fig. 9). Ninth tergum swol- len, submedially divided by median cleft, spinules sparse in cleft, dense on swollen areas (Fig. 10). Hemitergal lobes U-shaped, distal arm 2X longer than wide, apex of distal arm acute, twisted medially, spinules covering entire arm (Figs. 9, 11). Lateral apex of stylets blunt, subparallel, subacute process on ventral half (Figs. 9, 12). Female.—Length of fore wing 19-20 mm, body 19-25 mm. Subgenital plate ex- tends to % or less of 9" sternum, rounded with small median cleft, cleft sometimes absent, if present, not reaching anteriorly to the posterior edge of 8" sternum (Fig. 36). 414 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Vee pL Gin, ha RT ee 8 V4 Nt 19 15 Figs. 13-19. Megarcys irregularis. 13, Male head and prothorax, dorsal. 14, Male terminalia, oblique view. 15, 9" tergum, dorsal. 16, Hemitergal lobe, lateral. 17, Lateral stylet, apex. 18, Aedeagus, dorsal. 19, Aedeagus, lateral. VOLUME 103, NUMBER 2 Egg.—Collar stalked, rim usually deeply and irregularly incised; posterior surface of shoulder with ornate processes (Figs. 41, 42). Chorion with irregular deep pits with- out hexagonal pattern; surface irregularly indented; posterior pole rounded (Figs. 42— 44); micropylar row subequatorial; eclosion line absent (Fig. 41). Distribution.—Cascade Mountains and Northern Rocky Mountains of BC, ID, MT, OR and WN (Stark 1998). Remarks.—This species is common in most streams of Mt. Rainier National Park, often sympatric with M. irregularis. Material examined.mOREGON: Hood River Co., Upper Elk Cr. Near Lolo Pass, 12 Mune 1983, Gas Fiala, 1 ¢ (BYU); WASHINGTON: Lewis Co., Skate Cr, 4150 Skate Cr. Rd., G. Pinchot N.F, 28 May, 1997), B.. Kondratieff, 11 ¢,°2 (CSU); Lewis Co., Ohanapecosh R., Grove of the Patriarchs, Mt. Rainier N.P., 28 May 1997, B. Kondratieff, 1 ¢6 (CSU); Pierce Co., Deer Cr., Rt. 123, Mt. Rainier N.P., 27 May 1997, B. Kondratieff, 2 3d (CSU); Pierce Co., Panther Cr., Rt. 123, Mt. Rainier N.P., 29 May 1997, B. Kondratieff, 1 3 (CSU); Bierce Co., Goat Cr, Rt. 410, 29 May 1997, B. Kondratieff, 4 d, 1 2 (CSU); Pierce Co., Glacier Basin, Mt. Rainier N.P., 20 July 1997, R. Lechleitner, 4 2 (CSU); Pierce Co., Glacier Basin, Mt. Rainier N.P, 2 auly 19972 RR. jeechicitner, 1°¢,. 11 2 (CSU); Pierce Co., Frying Pan Cr., Sunrise Rd. Mt. Rainier N.P., 6 July 1999, B. Kon- dratict, IG, I2, (CSU); Pierce-Co., Par- adise R. above Narada Falls, Mt. Rainier N.P., 16 August 1999, B. Kondratieff, R. Kechileiner, 2 2°(ESU); Pierce Go. Frying Pan Cr., Sunrise Rd. Mt. Rainier N.P., 16 August 1999, B. Kondratieff, 1 2 (CSU); Skamania Co., Cr. near June Lake, Mt. St. Helens, 12 July 1997, S. Fitzgerald, A. Fo- ley, 8 2 (CSU); Whatcom Co., Wells Cr. Falls jct. of Bar Cr. and Wells Cr., 14 July 1988, R.W. Baumann, S. Wells and M. Whiting, 1 ¢ (BYU); Yakima Co., Union Cr. 20 miles W. Cliffdell, 27 June 1971, D. Loreth, 1 6 (BYU). 415 Megarcys irregularis (Banks) (Figs. 13-19, 35, 45-48) Dictyopteryx irregularis Banks 1900: 243. Type locality: Mt. Rainier National Park, Washington. Holotype 2, MCZ, not ex- amined. Arcynopteryx (Megarcys) irregularis: Rick- er 1952-375. Megarcys irregularis: Ulies 1966:370. Diagnosis.—The male of M. irregularis is very similar to M. signata but can be distinguished by the lack of spinule in the median cleft of the 9" tergum. Females of both species are similar, but the characters given used in the key, the length of the su- bgenital plate and chorion pattern of the eggs usually allow separation. General col- or brown. Head with an inverted U-shaped mark between compound eyes; pale mid- dorsal line on prothorax (Fig. 13). Male.—Length of fore wing 15—16 mm, body 18-20 mm. Posterior surface of epi- proct windsock-like with narrowly rounded anterodorsal hump, apex tapered to tip (Fig. 14). Ninth tergum swollen submedially, di- vided by median cleft; spinule patch distal to median cleft, spinules sparse around edg- es of cleft and swollen areas (Fig. 15). Hemitergal lobes U-shaped, with distal arm 1%xX longer than wide, distal arm blunt, twisted medially, spinules covering entire apex, sparsely covering membranous por- tion of lobe (Figs. 14, 16). Lateral stylets expanded twards tip, with a terminal pro- cess pointing posteriorly (Figs. 14, 17). Ae- deagus sock-like constricted about % dis- tally, setal patch basal and dorsally on api- cal lobe, evagination on proximal end of apical lobe (Figs. 18, 19); lateral tubular processes directed ventral and distal, mar- gin of basal lobe (Fig. 19). Female.—Length of fore wing 20-21 mm, body 21—23 mm. Subgenital plate pro- longed to cover ¥% or less of the 9" sternum and rounded posteriorly; medial cleft reach- ing anteriorly to the posterior margin of 8" sternum, cleft narrow posteriorly, expand- ed, oval-shaped anteriorly (Fig. 35). 416 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 24 Figs. 20-24. Megarcys watertoni. 20, Male head and prothorax, dorsal. 21, Male terminalia, oblique view. 22, 9" tergum, dorsal. 23, Hemitergal lobe, lateral. 24, Lateral stylet, apex. Egg.—Collar stalked, rim usually deeply and irregularly incised; posterior surface of shoulder with ornate processes (Figs. 45, 46). Chorion covered with irregular hex- agonal follicle cell impressions, chorionic folds absent or obscure (Figs. 46—48); mi- cropylar row subequatorial; eclosion line absent (Fig. 45). Distribution.—Coast and Cascade Moun- tains of BC and WN (Stark 1998). Remarks.—Other than M. yosemite, this species seems to have the most restricted VOLUME 103, NUMBER 2 417 aes ee i 29%, 40,3, a 31 raf Figs. 25-31. Megarcys yosemite. 25, Male head and prothorax, dorsal. 26, Male terminalia, oblique view. 27, 9" tergum, dorsal. 28, Hemitergal lobe, lateral. 29, Lateral stylet, apex. 30, Aedeagus, dorsal. 31, Aedeagus, lateral. 418 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON fa Layer an ( ' (tune, + Nee . {i Te tet pat mith nage 32 ene on i UT Fe ie Wane TUT My rt eg Figs. 32-36. Female subgenital plates, ventral. 32, Megarcys signata. 33, M. yosemite. 34, M. watertoni. 35, M. irregularis. 36, M. subtruncata. VOLUME 103, NUMBER 2 Figs. 37-40. Megarcys signata. 37, Egg, lateral. 38, Egg, lateral, anterior pole. 39, Egg posterior pole. 40, Chorionic detail, lateral. 420 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 41-44. Megarcys subtruncata. 41, Egg, lateral. 42, Egg, lateral, anterior pole. 43, Egg posterior pole. 44, Chorionic detail, lateral. VOLUME 103, NUMBER 2 48 Figs. 45—48. Megarcys irregularis. 45, Egg, lateral. 46, Egg, lateral, anterior pole. 47, Egg posterior pole. 48, Chorionic detail, lateral. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 49-52. Megarcys watertoni. 49, Egg, lateral. 50, Egg, lateral, anterior pole. 51, Egg posterior pole. 52, Chorionic detail, lateral. VOLUME 103, NUMBER 2 Figs. 53-56. Megarcys yosemite. 53, Egg, lateral. 54, Egg, lateral, anterior pole. 55, Egg posterior pole. 56, Chorionic detail, lateral. 424 range of all North American Megarcys. Me- garcys irregularis is very similar to M. sig- nata, indicating a recent species divergence from a common ancestor. Material examined.—WASHINGTON: Pierce Co., Carbon R., at entrance Mt. Rainier N.P., 27 May 1997, B. Kondratieff, 8 5, 3 (CSU); Pierce Co., Nisqually R., Mt. Rainier N.P., 29 May 1997, B. Kondra- tieff, 2 5d, 1 2 (CSU); Pierce Co., White R., Mt. Rainier N.P., 29 May 1997, B. Kon- dratieff, $ 6, 1 2 (CSU); Pierce Co., Ta- homa Cr., Paradise Rd., Mt. Rainier N.P, 30 May 1997, B. Kondratieff, 2 2 (CSU); Pierce Co., Frying Pan Cr, Mt. Rainier N.P, 6 July 1999, B. Kondratieff, 1 @ (CSU); Pierce Co., Frying Pan Cr., Sunrise rd., Mt. Rainier N.P., 16 August 1999, B. Kondratieff, 1 2 (CSU). Megarcys watertoni (Ricker) (Figs. 20-24, 34, 49-52) Arcynopteryx (Megarcys) watertoni Ricker 1952: 77. Type locality: Waterton Lake National Park, Alberta. Holotype <4, CNC examined. Megarcys watertoni: Illies 1966: 372. Diagnosis.—The male of M. watertoni is very similar to M. yosemite, but can be best distinguished by the narrower and longer hemitergal lobes (Fig. 23) and the spinule pattern of the 9th tergum (Fig. 22). The fe- male of M. watertoni is similar to M. sub- truncata, but can usually be distinguished by longer plate length, at least % or more of 9" sternum (Fig. 34) and by the eggs, which are regularly folded and have hex- agonal follicle cell impressions (Figs. 50— 52). General color brown. Head with three light circular patterns, others between ocel- li, near base of antennae; pale middorsal line on the prothorax (Fig. 20). Male.—Length of fore wing 13-16 mm, body 17—21 mm. Posterior surface of epi- proct windsock-like with broadly rounded anterodorsal hump; apex tapered and de- curved at tip (Fig. 21). Ninth tergum swol- len submedially, divided by median cleft, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON spinules uniformly dense in cleft and on swollen areas (Fig. 22). Hemitergal lobes U-shaped with the posterior arm greater 2X longer than wide; posterior arm acute, twisted medially with spinules covering the anterior edge and sparsely covering the membrane of lobe (Figs. 21, 23). Lateral stylets blunt at apex, subequal in width with subacute process (Figs. 21, 24). Female.—Length of fore wing 15-18 mm, body 16—21 mm. Subgenital plate pro- longed to cover % or more of the 9" ster- num, posteriorly rounded with small medi- an notch not reaching the posterior margin of 8" sternum (Fig. 34). Egg.—Collar stalked, rim usually deeply and irregularly incised; posterior surface of shoulder with ornate processes (Figs. 49, 50). Chorion covered with irregular hex- agonal follicle cell impressions; walls of follicle cell impressions thick and constrict- ed irregularly; surface with eight wide stri- ae separated by sulci (Figs. 50—52); micro- pylar row subequatorial; eclosion line ab- sent (Fig 49). Distribution.—Northern Rocky Moun- tains of AB, BC, ID, and MT (Stark 1998). Remarks.—This species is often associ- ated with small high elevation Northern Rocky Mountain streams. Material examined.—CANADA: AL- BERTA: Banff N.P., Red Earth Cr., Hwy 1, 20 miles N Banff., 2 July 1965, A.V. Ne- beker, 10 ¢ (BYU); Banff N.P. Moraine Cr. Hwy 17 mi. S. of Lake Louise, 2 July 1965, A.V. Nebeker, 7 ¢6 (BYU); BRITISH CO- LUMBIA: Kootenay N. P. Bridge on stream 3 mi. east of Radium jct. Hwy 93, 2 July 1965, A.V. Nebeker, 1 ¢ (BYU); IDAHO: Bonner Co., Granite Cr., Lake Pend Oreille., 15 May 1959, S.G. Jewett Jr., 4 d, 2 2 (BYU); MONTANA: Cascade Co., Belt Cr. 19 mi. S.E. of Monarch, 7 July 1966, J.R. Grievson, 1 6 (BYU); Flathead Co., Bear Cr. 5 mi. above jct. with M.F of Flathead R., 24 June 1965, A.V. Nebeker, 3 36,1 2 (BYU); Flathead Co., Cr. % mi. N. Essex Post Office, Hwy. 2., 6 July 1965, A.V. Nebeker, 6 d (BYU); Flathead Co., VOLUME 103, NUMBER 2 Cr. 20 mi. E of W. Glacier, 6 July 1965, A.V. Nebeker, 3 6, 2 2 (BYU); Flathead Co., Doris Cr. 4 mi. S. of Hungry Horse Dam, 18 July 1965, A. V. Nebeker, 1 d, 1 2 (BYU); Flathead Co., Bear Cr. Hwy 2 Morias Pass, 18 June 1970, R.A. Haick, 1 3, 1 2 (BYU); Flathead Co., Glacier N.P. Fish Cr. above Fish Cr. Campground above Lake McDonald, 5 June 1996, R.S. Hansen and B. Ward, 2 2 (BYU); Glacier Co., Ice- berg Cr., Glacier N.P, 30 August 1965, A.V. Nebeker, 1 ¢ (BYU); Glacier Co., Ice- berg Cr., Many Glacier Campground., Gla- cier N.P, 29 July 1969, A.R. Gaufin, 3 6, 3 2 (BYU); Glacier Co., Waterton R., 23 July 1970, C. Tarmoloy, 1 6 (BYU); Gla- cier Co., Iceberg Cr., Glacier N.P., 28 July 1970, A.R. Gaufin, 1 ¢@ (BYU); Glacier Co., Baring Cr. Sunrise Gorge, Glacier N.P., 22 July 1979, B. Stark, K. Stewart and R. Baumann, 5 6, 4 2, (BPSC); Granite Co., Ranch Cr. bridge, Rock Cr., 25 June 1966, M.L. Miner, 1 male, 1 2 (BYU); Granite Co., Ranch Cr. at Grizzly Campground., 19 June 1967, M.L. Miner, 1 ¢ (BYU); Mis- soula Co., N. fork of Elk Cr. at middle flume, 6 July 1970, M.L. Miner, 1 & (BYU). Megarcys yosemite (Needham and Claassen) (Rigs. 25-315 33;,53—56) Perlodes yosemite Needham and Claassen 1925: 56. Type locality: Mt. Lyell, Yo- semite National Park, California. Holo- type 6, CU, examined. Arcynoptergx (Megarcys) yosemite: Ricker 1952775: Megarcys yosemite: Ilies 1966: 371. Diagnosis.—The male and female are very similar to M. watertoni and characters for separation are given in the dignosis for M. watertoni. General color brown. Head with posterior dark patch to the anterior margin of the compound eyes, enclosing posterior ocellus, dark W-shaped pattern be- tween compound eyes encompassing ante- rior ocelli (Fig. 25). 425 Male.—Length of fore wing 16 mm, body 21 mm. Posterior (ventral) surface of epiproct windsock-like, with a high, very broadly rounded anterodorsal hump, nar- rowing gradually distally with decurved apex (Fig. 26). Ninth tergum swollen sub- medially, divided by median cleft; spinules dense in cleft and along posterior edges of swellings (Fig. 27). Hemitergal lobes U- shaped, posterior arm 1%X longer than wide, apex of distal arm acute, twisted me- dially, spinules covering anterior arm, sparsely on membrane (Figs. 26, 28). Lat- eral stylets widest distally, with acute spine (Figs. 26, 29). Aedeagus with rounded pos- terior end with small medial cleft, lateral filaments extended laterally, sclerotized patches laterally from the median dorsal lobe (Figs. 30, 31). Female.—Length of fore wing 21—22 mm, body 17—20 mm. Subgenital plate tri- angulate, produced % or more length of 9" sternum, lobes narrowly rounded at apex, with medial emargination extending to ’2 or more of plate (Fig. 33). Egg.—Collar stalked, rim usually deeply and irregularly incised; posterior surface of shoulder with ornate processes (Figs. 53, 54). Chorion covered with hexagonal folli- cle cell impressions; having smooth thick walls; surface with shallow irregular inden- tations; posterior pole rounded (Figs. 54— 56); micropylar row subequatorial; eclosion line absent (Fig. 53). Distribution.—Cascade Mountains and Sierra Nevada Mountains of CA and WA (Stark 1998). Remarks.—The holotype male consists of dissected terminalia and wings mounted on a single microscope slide. The male from Mt. Rainier agrees completely with the type in details of the epiproct, hemiter- gal lobes and lateral stylets. Megarcys yo- semite is apparently rare, and is associated with high elevation small glacier fed- streams (Ricker 1952). Material examined.—WASHINGTON: Pierce Co., Frying Pan Cr., Sunrise rd. Mt. Rainier N.P, 16 August 1999, B. Kondra- 426 tieff, 1 d, (CSU); Creek at high elevation from interglacier and tribs. near White R., Mt. Rainier N.P., 26 July 1953, W. E. Rick- er and A. Ricker, 2 2, (INHS); Whatcom Co., Coleman Glacier., Mt. Baker, 23 July 1983 gRe Pollocks 1) Jonesy lade, 42 (BYU). KEYS TO NORTH AMERICAN SPECIES OF MEGARCYS Males 1. Hemitergal lobe apices acute (Figs. 11, 23, 28) Rate cree Can tes or nat ane, cee eee M ore SPEED, = 2 Hemitergal lobe apices blunt (Figs. 4, 16)... 4 2. Spinules covering most of distal portion of the hemitergal lobe; lateral stylet truncate apically (Figs. 9, 12); Coast and Cascade Mts ...... Fx ONS RE Oe eC ee M. subtruncata — Spinules only at anterior portion of distal hem- itergal lobe; lateral stylet tapered apically (Figs. 23, 24, 28, 29) 3. Hemitergal lobe narrow, 2 longer than wide; epiproct sack arising near apex (Figs. 21, 23), INorthermpRockys Mts ee M. watertoni — Hemitergal lobe wide, 1% as long as wide; epiproct sack arising at mid-length (Figs. 26, 28); Cascade and Sierra Nevada Mts Sint CUBREAC bot ous ec Se aEO or CON CA ee aE M. yosemite 4. Spinules absent from median cleft of 9" ter- gum, sometimes a few scattered spinules (Fig. 15) ; Coast and Cascade Mts .... M. irregularis — Spinules in the median cleft of 9" tergum (Fig. 3) Rockys Mts: ety 5.5 COR M. signata Females and Eggs 1. Subgenital plate with deep median notch, dis- tinctly separating lobes to, or beyond, posterior margin of 8" sternum (Figs. 32, 35) ....... 2 — Subgenital plate with shallow median notch present or absent, if present not distinctly sep- arating lobes to the posterior margin of 8" ster- num (Figs. 33, 34, 36) 3. Subgenital plate produced % or more length of 9" sternum (Fig. 32); posterior pole and egg surface irregularly indented, chorion covered with irregular deep pits without obvious hex- agonal pattern (Figs. 37—40); widespread along the Rocky Mts eee cea es ee M. signata — Subgenital plate produced % or less length of 9' sternum (Fig. 35); chorion covered with shallow pits surrounded by hexagonal follicle cell impressions (Figs. 45—48); Coast and Cas- cadet Mtsy re. hei cee cu cee ae 4. Subgenital plate triangulate, produced % or more length of 9 sternum, lobes narrowly M. irregularis PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON rounded at apex, medial notch narrow, extend- ing % or more of plate (Fig. 33); egg surface indentations superficial, chorion covered with rather uniform, smooth walled hexagonal fol- licle cell impressions surrounding shallow bowl shaped pits (Figs. 53-56); Cascade and Sista INGWAGEVIMNG, 55 aceon acco aec M. yosemite — Subgenital plate rounded, medial notch, ex- tending % or less of plate (Figs. 34, 36) .... 4 5. Subgenital plate covers less than % of 9" ster- num, (Fig. 36); egg surface indentations prom- inent, chorion covered with irregular deep pits without obvious hexagonal pattern (Figs 41— 44); Coast and Cascade Mts. ... M. subtruncata — Subgenital plate covers % or more than half of 9" sternum (Fig. 34); egg surface indentations prominent and arranged in regular striae and sulci, chorion covered with hexagonal follicle cell impressions (Figs. 49-52); Northern Rocky Mts. M. watertoni ACKNOWLEDGMENTS We thank Richard W. Baumann, Bringh- am Young University, E. R. Hoebeke, Cor- nell University, and Kathleen R. Zeiders, II- linois Natural History Survey, for providing specimens. Special thanks to Richard Lech- leitner, Mt Rainier National Park, for mak- ing collecting possible, and joining the sec- ond author on collecting trips. Jeff Martz provided the illustrations. Partial funding provided by Colorado State University, De- partment of Biology through a biology un- dergraduate research fund. LITERATURE CITED Banks, N. 1900. New genera and species of Nearctic neuropteroid Insects. Transactions of the Ameri- can Entomological Society 25: 239-259. Hagen, H. A. 1874. Report on the Pseudo-neuroptera and Neuroptera collected by Lieut. W. L. Carpen- ter in 1873 in Colorado. Annual Report of the U.S. Geological Survey, Territories, Embracing Colorado 7: 571-577. Hanson, J. EK 1942. Records and descriptions of North American Plecoptera, Il. Notes on North Ameri- can Perlodidae. American Midland Naturalist 28: 389-407. Illes, J. 1966. Katalog der rezenten Plecoptera. Das Tierreich, 82 Walter de Gruyter, Berlin. Needham, J. G. and P W. Claassen. 1925. A mono- graph of the Plecoptera or stoneflies of America north of Mexico. Thomas Say Foundation, Ento- mological Society of America 2, 397 pp. Ricker, W. E. 1952. Systematic studies in Plecoptera. VOLUME 103, NUMBER 2 Indiana University, Publications in Science, Series 18, 200 pp. Stark, B. P. 1998. North American stonefly list. www.mce.edu/~stark/stonefly.html. Stark, B. P. and S. W. Szczytko. 1981. Contributions to the systematics of Paragnetina (Plecoptera: Perlidae). Journal of the Kansas Entomological Society 54: 625-648. . 1988. Egg morphology and phylogeny in Ar- cynopterygini (Plecoptera: Perlodidae). Journal of the Kansas Entomological Society 61: 143—160. 427 Stark, B. PB. S. W. Szczytko, and C. R. Nelson. 1998. American stoneflies: A photographic guide to the Plecoptera. Caddis Press, Columbus, Ohio. Stewart, K. W. and B. P. Stark. 1988. Nymphs of North American stonefly genera (Plecoptera). Thomas Say Foundation, Entomological Society of Amer- ica 12, 460 pp. Taylor, B. W., C. R. Anderson, and B. L. Peckarsky. 1999. Delayed egg hatching and semivoltinism in the Nearctic stonefly Megarcys signata (Plecop- tera: Perlodidae). Aquatic Insects 21: 179-185. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 428-432 DESCRIPTION OF A NEW SPECIES OF UROCERUS GEOFFROY (HYMENOPTERA: SIRICIDAE) FROM JAPAN ICHIJI TOGASHI 1-chome, Honmachi, Tsurugi-machi, Ishikawa Prefecture 920-2121, Japan Abstract.—Urocerus tsurugianus, n. sp., from Japan is described and illustrated. Char- acters are given to distinguish it from U. japonicus Smith. Key Words: Four species of woodwasps belonging to the genus Urocerus are recorded from Ja- pan, U. antennatus Marlatt, U. japonicus Smith, U. multifasciatus Takeuchi, and U. yasushii Yano (Takeuchi 1955, 1962; To- gashi 1972). Recently, I found one speci- men of Urocerus which is closely allied to U. japonicus Smith. After comparing this specimen with about 50 female specimens of U. japonicus collected in Honshu, Shi- koku, and Kyushu, I found it is easily sep- arable by the coloration of the 3rd to 7th and 8th tergites, the shape of the fore inner tibial spur, the small cenchri, the shape of the precornal basin, and the structure of the lancet. Therefore, I concluded that my spec- imen is new to science. In this paper, I de- scribe and illustrate this new species. Urocerus japonicus Smith (Figs. 7, 8, 10-11, 13-14, 17—20, 22-23) I examined about 50 female specimens and about 30 male specimens of U. japon- icus Smith collected in Honshu (Iwate, To- kyo, Saitama, Kanagawa, Nagano, Ishika- wa, and Okayama), Miyake Is., Shikoku (Ehime and Kochi), and Kyushu (Fukuoka). Supplementary note. Antenna 26-seg- mented. Antenno-ocular distance shorter than distance between antennal sockets (ra- tio about 1.0:1.5—1.7). Distance between cenchri longer than breadth of each one as Siricidae, Urocerus, new species, Japan 1.2—1.5:1.0. Wings: length of Ist cubital cell (1R1) 1.4—1.7 times as long as 2nd one (1RS). Front inner tibial spur as in Fig. 8. Cornus longer than length of precornal ba- sin (ratio about 2.3—2.7:1.0); cornus, in dor- sal view, as in Fig. 10; basal portion of cor- nus with or without process; apical portion of the lance as in Figs. 17 and 19; 3rd an- nulus of lancet slightly curved (Figs. 18 and 20). Coloration of abdomen as follows: Ist and 2nd tergites yellow; 3rd to 7th tergites dark brown to black (Fig. 23); 8th tergite yellow but apical % black (Fig. 23); 9th ter- gite black or with yellow macula or entirely yellow (Figs. 22—23); cornus yellow. Variation in coloration. Head yellowish brown, sometimes frontal area and ocellar basin dark brown; Ist, 2nd and 8th tergites yellow but apical % to apical margin of 8th tergite dark brown to black; 3rd to 7th ter- gites dark brown (Fig. 23) (about 95% of examined specimens); sometimes 3rd to 4th tergites yellow (Fig. 22) (about 5% of ex- amined specimens). Urocerus tsurugianus Togashi, new species (Figs. 1-6, 9, 12, 15-16, 21) Female.—Length including cornus 24 mm. Body pale reddish yellow with follow- ing parts dark brown: apex of mandible, malar space, frons, ocellar basin, postocel- VOLUME 103, NUMBER 2 429 aoe ; Figs. 1-8. 1-6, Urocerus tsurugianus, holotype. 1, Head, dorsal view. 2, Basal 5 antennal segments, lateral view. 3, Pronotum, dorsal view. 4, Forewing showing cells 1R1 and 1RS. 5, Front inner tibial spur, lateral view. 6, Claw, lateral view. 7—8, U. japonicus, Ehime Prefecture. 7, Forewing showing cells 1R1 and IRS. 8, Front inner tibial spur, lateral view. lar area, lateral sides of pronotum, tegula, mesonotum, mesoscutellum, under thorax, most of Ist tergite, posterior %4 of 8th ter- gite, 7th to 9th sternites, and sheath; basal % of 8th tergite dirty yellow. Antenna pale reddish yellow. Wings yellowish hyaline, stigma and veins pale reddish yellow. Legs pale reddish yellow with following parts dark brown to black: all coxae and trochan- ters, hind femur, apical % of hind tibia, and apical 2 segments of all tarsi. Head: Dilated behind eyes (Fig. 1); in- terocellar, postocellar, and lateral furrows indistinct; postocellar area rectangular, with distinct median longitudinal furrow (Fig. 1); OOL:POL:OCL = 1.0:1.0:3.6; antenno- ocular distance shorter than distance be- tween antennal sockets (ratio about 1.0: 1.6); malar space long, twice as long as pedicel. Antenna 25-segmented; relative lengths of basal 5 segments about 5.1:1.0:2.8:2.7: 2.6; pedicel rather subquadrate (Fig. 2). Thorax: Normal; midlength of pronotum short, as long as POL or OOL (Fig. 3); cen- chrus rather small, distance between them longer than breadth of each one (ratio about 1.6:1.0). Wings: Ist cubital cell (1R1) longer than 2nd one (1RS) (ratio about 2.1:1.0). Legs: hind basitarsus slightly shorter than fol- 430 lowing 4 segments combined; front inner tib- ial spur as in Fig. 5; claw as in Fig. 6. Abdomen: Normal; precornal basin broad- er than long (ratio between breadth and length about 1.5:1.0) (Fig. 9); length of pre- cornal basin % as long as length of cornus; cornus as in Figs. 9 and 12; basal portion without process (Fig. 12); sheath longer than basal plate (ratio about 1.3—1.4:1.0); apical portion of lance as in Fig. 15; 3rd annulus of lancet strongly curved (Fig. 16). Punctation: Head and thorax distinctly, densely, and rather reticulately punctured, matt; hind orbits nearly impunctate, shining (Fig. 1); abdominal tergites shagreened. Food plant—Unknown. Distribution.—Japan (Honshu). Holotype.—Female, 13. IX. 1980, San- nomiya, Tsurugi-machi, Ishikawa Prefec- ture, I. Togashi leg., deposited in the col- lection of the National Science Museum (Nat. Hist.), Tokyo. ly resembles Urocerus japonicus Smith, but PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 7 ee = = >> — —- = =e rv Figs. 9-14. 9-11, Precornal basin and cornus, dorsal view. 9, Urocerus tsurugianus, holotype. 10, U. ja- ponicus, Ehime Prefecurue. 11, U. japonicus, Kanagawa Prefecture. 12—14, Cornus, lateral view. 12, U. tsuru- gianus, holotype. 13, U. japonicus, Ehime Prefecture. 14, U. japonicus, Kanagawa Prefecture. it is easily separable from the latter by the shape of the front inner tibial spur (see Figs. 5, 8), by the black or mostly black 8th ab- dominal tergite (in japonicus, the anterior *2 or most of the 8th abdominal tergite is yellow, see Figs. 22—23), by the small cen- chri with the ratio between the distance be- tween the cenchri and the breadth of one as 1.6:1.0 (in japonicus, the ratio between the distance between the cenchri and the breadth of each one is 1.2—1.5:1.0), by the long precornal basin with the ratio between the length of the precornal basin and the length of the cornus as 1.5:1.0 (in japoni- cus, the cornus is 2.3—2.7 times as long as the length of the precornal basin, see Figs. 9-11), and by the strongly curved 3rd an- nulus of the lancet (in japonicus, the 3rd annulus of the lancet is slightly curved, see Figs. 16, 18, and 20). ACKNOWLEDGMENTS I cordially thank Dr. David R. Smith, Systematic Entomology Laboratory, USDA, Washington, DC, for his kind ad- VOLUME 103, NUMBER 2 431 20 Figs. 15-20. 15-16, Urocerus tsurugianus, holotype. 17-18, U. japonicus, Tokyo Prefecture. 19-20, U. japonicus, Saitama Prefecture. 15, 17, 19, Apical portion of lance. 16, 18, 20, Apical portion of lancet. Arrow points to 3rd annulus. 432 Figs. 21-23. view. vice and review of this manuscript. I am indebted to Dr. A. Shinohara, National Sci- ence Museum (Nat. Hist.), Tokyo, Mr. H. Makihara, Tohoku Research Center, Forest- ry and Forest Product Research Institute, Morioka, and Dr. K. Maeto, Shikoku Re- search Center, Forestry and Forest Product Research Institute, Kochi, for lending me the specimens of U. japonicus Smith. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 22 23 21, Urocerus tsurugianus, holotype, dorsal view. 22—23, U. japonicus, Tokyo Prefecture, dorsal LITERATURE CITED Takeuchi, K. 1955. Siricidae of Japan (Hymenoptera). Akitu 4:1—9. (In Japanese with English summary.) . 1962. Hymenoptera: Siricidae. Insecta Japon- ica, series 2, part 4, 12 pp. Hokuryukan, Tokyo. (In Japanese with English summary.) Togashi, I. 1972. A note on the seasonal occurrence of sawflies (Hym., Symphyta) on Mt. Hiko, Kyushu. Mushi 46: 129-135. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 433-436 NEW CRANE FLIES (DIPTERA: LIMONIIDAE) FROM LEBANESE AMBER SIGITAS PODENAS, GEORGE O. POINAR, JR., AND RAIF MILKI (SP) Department of Zoology, Vilnius University, Ciurlionio 21/27, Vilnius LT-2009, Lithuania. (e-mail: sigitas.podenas @ gf.vu.lt); (GOP) Department of Entomology, Oregon State University, Corvallis, OR 97331-2907, U.S.A. (e-mail: poinarg@bcc.orst.edu); (RM) Department of Health Science, American University of Beirut, Beirut, Lebanon. (e-mail: rm01 @aub.edu.lb) Abstract.—The new genus Lebania Podenas and Poinar including L. levantia Po- denas and Poinar, n. sp., and L. longaeva Podenas and Poinar, n. sp., is described from Lebanese amber (Lower Cretaceous). These are the first crane flies (Diptera, Li- moniidae) described from these deposits. Key Words: Lebania gen. n., Lebania levantia sp. n., Lebania longaeva sp. n., Limon- iidae, Lebanese amber, fossil, Lower Cretaceous Crane flies of family Limoniidae were previously unknown from Lebanese amber (Lower Cretaceous) (Evenhuis 1994) al- though they have been described from other Cretaceous amber deposits (Poinar 1992). In this study we describe two species (three specimens) in a new genus of the family Limoniidae from the Milki collection of Lebanese amber maintained at the Ameri- can University of Beirut. MATERIALS AND METHODS The specimens originated from Early Cretaceous amber beds in the district of Jezzine, Lebanon. At that site, the amber occurs in primary deposits of the Neocom- ian division of the Early Cretaceous as well as in secondary deposits of the Aptian stage, thus ranging in age from 120—135 million years (Schlee and Dietrich 1970). In the following descriptions, terminolo- gy of genitalia and wing venation follows that presented in the Manual of Nearctic Diptera (McAlpine 1981). Thus m-cu and CuA, are considered separate veins. Acces- sion numbers pertaining to specimens are presented under the section of examined material. Abbreviations used in the drawings are: a—aedeagus; A,—first anal vein; A,—sec- ond anal vein; dm—discal medial cell; gon—gonocoxite; 1 g—inner gonostylus; M—medial; M,,,—combined first and sec- ond medial vein; m-cu—medial cubital cross vein; o g—outer gonostylus; ped— pedicel; pm—paramere; R,—first branch of radius vein; R,—third radial vein; R,— fourth radial vein; R,,,—combined third and fourth radial veins; Rs—radial sector; scp—scape; Sc,—first subcostal vein; Sc,—second subcostal vein; st 9—ninth sternite. Lebania Podenas and Poinar, new genus Type species.—Lebania levantia, n. sp. Description.—Head dark. Vertex forming wide crest. Antenna short; if bent back- wards, reaching wing base; both basal seg- ments thick, flagellum very thin, rod-like; scape cylindrical, pedicel pear-shaped, about 1.7 as long as scape; first flagellom- ere with widened central part, other flagel- 434 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-7. holotype. 4, Wing, holotype. 5, Male genitalia, ventral view, holotype. 6-7. Lebania longaeva, holotype. 6, Wing. 7, Male genitalia, ventral view. lomeres elongated, nearly cylindrical with somewhat flattened basal parts; verticils long, especially on middle segments, ex- ceeding length of respective segment some 4 or 5 times; apical segment large, nearly as long as preceding segment. Wings com- paratively wide, without any dark spots ex- cept faint stigma ; veins brown. Venation: Sc, ends just before end of Rs; Sc, close to tip of Sc,; Rs slightly arcuated at base; R, ends opposite fork of R; and R,; r absent; R, and R, short; discal cell closed; M, en- tirely fused with M,; cross-vein m-cu far removed distally from fork of M, close to distal end of discal cell; anal veins diver- gent, A, comparatively short; anterior ar- culus (vein connecting R with Cu) missing. Meron small. Front leg with one tibial spur; tibiae of middle and posterior legs with two spurs. Abdomen brown, covered with long yellowish hairs. Male genitalia not inverted, with elongated, nearly cylindrical gonocox- ites; two simple gonostyles. Female—un- known. Etymology.—The genus name is based 1-5, Lebania levantia. 1, Maxillary palpus, paratype. 2, Antenna, holotype. 3, Tip of antenna, on Lebanon, the country where the Lower Cretaceous amber deposits are located. Lebania levantia Podenas and Poinar, new species (Figs. 1—5) Diagnosis.—General coloration dark brown due to oxidation in amber. Wings brownish, clear, without any darker spots except very slightly darkened stigma. Male genitalia with elongated gonocoxites, long, slender and simple gonostylus; inner gon- ostyle bearing few strong setae near apicies. Male.—Body length 2.4—2.5 mm, wing length 2.1 mm. Head blackened. Vertex forming wide crest. Rostrum, mouth parts and palpi black. Palpus 0.2 mm long. Basal segment of palpus elongated (Fig. 1); sec- ond and third segments somewhat shorter, both nearly equal in length; apical segment elongated, 1.6 as long as preceding seg- ment. Antenna short (Fig. 2), 15- (or 14-) segmented (limits of apical segments not clear), about 0.7 mm long, brown; if bent backwards, reaching base of wing. Both VOLUME 103, NUMBER 2 basal segments thick, flagellum very thin, rod-like; scape cylindrical, pedicel pear- shaped, 1.7X as long as scape and very wide; flagellomeres elongated, nearly cylin- drical with somewhat flattened basal part; verticils long, especially on middle seg- ments exceeding length of respective seg- ment about five times; apical segment (Fig. 3) long, nearly as long as preceding seg- ment. Dorsum of thorax dark brown, pleura blackened. Wing (Fig. 4) very wide, brown- ish, without dark spots except for faint stig- ma. Veins brown. Venation; Sc, ends oppo- site end of Rs; Sc, close to tip of Sc,; Rs long and nearly straight, just slightly arcuated at base; R, ends just before fork of R,, 4; R3,4 about twice as long as R,; r absent; R,; very short. Discal medial cell wide, just slightly longer than wide, with narrow base and strongly widened distal part; cross-vein m- cu far removed distally from fork of M, close to distal end of discal medial cell; anal veins strongly divergent, A, comparatively short; anterior arculus missing; anal angle wide. Halter brownish, 0.3 mm long. Cox- ae, trochanters, femora and tibiae dark brown, tarsi brown; femur II: 0.8—1.0 mm, III: 0.8—1.5 mm, tibia II: 0.8—1.0 mm, tarsus II: 0.8—1.1 mm; legs covered with short yel- lowish hairs. Tibiae of middle and posterior legs with two strong spurs. Claws narrow and elongated without tooth, arolium nee- dle-like. Abdomen brown, covered with long yel- lowish hairs. Genitalia (Fig. 5) dark brown; ninth tergite could be viewed only laterally, posterior border slightly lifted; gonocoxite elongated, nearly cylindrical; outer gonos- tylus long and slender; inner gonostylus about two-thirds length of outer gonostylus, with somewhat narrowed apex, bearing few strong setae; parameres awl-shaprd; aedea- gus simple, elongated with widened base. Female.—Unknown. Material examined.—Holotype: male, Milki collection at the American University of Beirut: (Acc. # 194—32), Lebanese am- 435 ber. Paratype male, same as holotype (Acc. # 194-15). Etymology.—tThe species name is based on the region (Levant) where the amber originated. Lebania longaeva Podenas and Poinar, new species (Figs. 6—7) Diagnosis.—General coloration dark brown due to oxidation in amber. Wings brownish, clear, without dark spots except for slightly darkened stigma. Male termin- alia with elongated gonocoxites, long slen- der outer gonostylus and shorter inner gon- ostylus. Lebania longaeva n. sp. differs from the foregoing species by the structure of the male genitalia (Figs. 5, 7), position of M vein, and longer setae on dorsal sur- face of posterior femora and tibia. Male.—Body length 2.2 mm, wing length 2.0 mm. Head blackened. Vertex forming wide crest. Rostrum, mouth parts and palpi black. Antenna short, brown; if bent backwards, reaching base of wing. Both basal segments thick, flagellum very thin, rod-like; scape cylindrical, pedicel pear-shaped, about twice as long as scape; flagellomeres elongated, nearly cylindrical with somewhat flattened basal part; verticils long, especially on middle segments ex- ceeding length of respective segment about three times; apical segment nearly as long as preceding segment. Dorsum of thorax dark brown, pleura blackened. Wing (Fig. 6) not widened, brownish, without dark spots, except for nearly indistinct stigma. Veins brown. Ve- nation: Sc, ends opposite the end of Rs; Sc, close to Sc, tip; Rs slightly arcuated at base; R, ends just before fork of R;,4,; R;,, 1.6 longer than R,; r missing; R; comparatively short. Discal medial cell twice as long as wide, with narrow base and widening dis- tally; cross-vein m-cu far removed distally from fork of M, close to distal end of discal medial cell; anal veins divergent, A, com- paratively short; anterior arculus missing; anal angle wide. Halter brownish, 0.3 mm 436 long. Coxae, trochanters, femora and tibiae dark brown, tarsi brown; femur I: 0.9 mm, Ill: 0.9 mm, tibia I: 1.2 mm, III: 0.9 mm, tarsus III: 0.8 mm; legs covered with short, yellowish hairs, posterior femur and tibia with long, yellowish, erect setae, about five times longer than diameter of leg. Front leg with one tibial spur; tibia of posterior leg with two strong spurs. Abdomen brown, covered. with long yel- lowish hairs. Genitalia (Fig. 7) dark brown; gonocoxite elongated, nearly cylindrical, with long setae; outer gonostylus long and slender; inner gonostylus also slender, about two-thirds length of outer gonostylus. Female.—Unknown. Material examined.—Holotype: male, Milki collection at the American University of Beirut, (Acc. # 194-110), Lebanese am- ber. Etymology.—The species name longae- vus is Latin for ancient. DISCUSSION The new genus Lebania resembles cer- tain genera of the Eriopterinae, such as Rhabdomastix Skuse or Gonomyia Meigen, but because of the well developed tibial spurs, it falls into Hexatominae (or Hexa- tomini of American authors) (Alexander and Byers 1981, Savchenko 1989). The new genus is characterized by very primi- tive male genitalia, such as nearly cylindri- cal gonocoxites which are elongated with- out additional structures and two pairs of simple gonostyles. The wing venation is similar to that of Rhabdomastix, but the an- terior arculus or that part of the arculus from R to M is missing. A very specific feature of Lebania is the extremely en- larged pedicel, which is usually small in PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON other crane flies. Differences between both newly described species occur in the wing venation: cell R; is smaller, M vein longer, the discal cell is wider and the anal veins are more divergent in L. levantia (Figs. 4, 6). With the male genitalia, the outer gon- ostylus of L. levantia is shorter and inner gonostylus clearly longer than in L. longae- va and the gonocoxites are comparatively longer and more narrow in L. levantia. (Figs. 5, 7). Also the posterior legs of L. longaeva have very long hairs on the dorsal surface of the femorae and tibiae, while these hairs are not distinguished from oth- ers on the legs of L. levantia. ACKNOWLEDGMENTS We are deeply indebted to Dr. Jon K. Gelhaus (Academy of Natural Sciences, Philadelphia) for arranging the senoir au- thor’s stay in Philadelphia and Pittsburgh, Pennsylvania, and making material avail- able for study. LITERATURE CITED Alexander, C. P. and G. W. Byers. 1981. Tipulidae, pp. 153-190. In McAlpine, J. F et al., eds. Manual of Nearctic Diptera. Ottawa, Agriculture Canada, Re- search Branch, Monograph 27, Vol. 1. Evenhuis, N. L. 1994. Catalogue of the Fossil Flies of the World (Insecta: Diptera). Backhuys Pub., Lei- den, 570 pp. McAlpine, J. E 1981. Morphology and Terminology— Adults, pp. 9-63. In McAlpine, J. E et al., eds. Manual of Nearctic Diptera. Ottawa, Agriculture Canada, Research Branch, Monograph 27, Vol. 1. Poinar, Jr, G. O. 1992. Life in Amber. Stanford Uni- versity Press, Stanford, California, 350 pp. Savchenko, E. N. 1989. Limoniid Crane Flies. Fauna of the USSR. Academy of Sciences, Kiev, 378 pp. (In Russian.) Schlee, D. and H. G. Dietrich. 1970. Insektenfuhrender Bernstein aus der Unterkreide des Lebanon. Neues Jahrbuch der Geologie und Palaéontologie Monat- sheft (Stuttgart) 1: 40—50. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 437-443 A COLLECTION OF FLEAS (SIPHONAPTERA) FROM THE SAN MARTIN RESERVE, VALDIVIA PROVINCE, CHILE MICHAEL W. HASTRITER, MAURICIO E. ALARCON, AND MICHAEL EK WHITING (MWH) Monte L. Bean Life Science Museum, Brigham Young University, 290 MLBM, P.O. Box 20200, Provo, UT 84602-0200, U.S.A. (e-mail: hastritermw @sprintmail.com); (MEA) Departamento de Zoologia, facultad de Ciencias Naturales y Oceanograficas, Universidad de Concepcion, Casilla 160-C, Concepcién, Chile; (MFW) Department of Zoology, 574 Widtsoe Building, Brigham Young University, Provo, UT 84602, U.S.A. (e-mail: Michael_Whiting @ byu.edu) Abstract.—During September 1999, seven species of fleas were collected from small mammals in the San Martin Reserve, Valdivia Province, Chile. The rare species Barrer- opsylla excelsa, Jordon, reported only once in Chile from Aisén Province, is reported from Valdivia Province, Chile. A summary of the known distribution of each of the seven species is provided. Key Words: Beaucournu and Gallardo (1991, 1992) list 91 species of fleas in Chile. Although this is a significant number of species, little is known about the host specificity, season- al, and geographic distribution for most of them. This report provides an account of flea species collected during the early Chi- lean spring within the San Martin Reserve, Valdivia Province, Chile, and summarizes the known host and locality data for each species. MATERIALS AND METHODS Fleas were collected from small mam- mals in the San Martin Reserve, Valdivia Province, Chile (39°38’S, 73°07’W) on September 14—17, 1999. The 72 hectare re- serve, located 30 km northeast of Valdivia in the cool temperate rain forest of the pre- cordillera of southern Chile, receives an- nual rainfall of ~2,500 mm (75% between April and September) (Murua et al. 1987). The dominant climax vegetation includes Nothofagus obliqua (Blume) and Aextoxi- Barreropsylla, fleas, Siphonaptera, Valdivia, Chile con punctatum Ruiz and Pav6n, the bro- meliad Gregia sphacelata Regel, understo- ry schrubs of Rubus constrictus Lefévre and PJ. Mueller, and thick stands of Chus- quea quila Kunth (bamboo, quila) and Rosa moschata Herrmann. The latter two pre- dominate in more open and unshaded areas. Much of the ground surface in densely shaded areas is covered with bryophytes, ferns and fungi. Standard aluminum collapsible Sher- man® traps baited with oatmeal were set on four consecutive nights in linear transects at five locations within the reserve. The density of under-story vegetation necessi- tated trapping only areas adjacent to exist- ing roads and trails. Under-story plant com- munities of C. guila and R. constrictus were dominant trapping habitats. Three weeks of continuous rainy weather preceded trapping and persisted throughout the trapping peri- od. Trapped mammals were shaken from traps into cloth or plastic bags at each cap- ture site, sacrificed by cervical dislocation 438 and transported to the field station. Fleas were obtained by brushing the animals and subsequently washing each thoroughly (Hastriter 2000). They were prepared by conventional techniques and mounted on glass slides in Canada balsam using, 12 mm cover-slips (#1 thickness). Unless otherwise noted, all specimens examined were from the San Martin Reserve. Mammals were preserved in 10% formalin and returned to the Instituto de Ecologia y Evolucion, Universidad Austral de Chile, Valdivia, Chile. Recognition of mammalian taxa fol- lows the regime of Wilson and Reeder (1993) and Smith and Patton (1993). Fifty-five rodents were trapped during 471 trap nights (11.6% capture rate) repre- senting five species [Oligoryzomys longi- caudatus (Bennett) (n = 29, 21 6, 8 @), Akodon olivaceus (Waterhouse) (n = 22, 8 6, 14 2), Rattus rattus (L.) (n = 2 Q), Abrothrix longipilis (Waterhouse) (n = 1 3), and Pearsonomys annectus Patterson (n = | 2)]. Fleas were not recovered from the latter three species. The overall flea index (average number of fleas per host) was 0.87. Forty-one percent of Ol. longicauda- tus (9 56,3 2) and 54% of Ak. olivaceus (4 3, 8 2) harbored one or more fleas. Ctenophthalmidae Chiliopsylla allophyla allophyla (Rothschild 1908) Material examined.—1 6, ex Ak. oliva- ceus, San Martin Reserve; 1 ¢, ex either Ak. olivaceus or Ol. longicaudatus, Castro National Park, Chiloé Island, Chiloé Prov- ince, Chile, April 1999, Patricio Chandia; 3 3, 4 2, ex Dromiciops gliroides Thomas, Cerro Caracol (37°00’S, 72°30’ W), Concep- cidn, Concepcion Province, Chile, 15 April 1981, Artigas; 1 ¢, ex D. gliroides, Cerro Caracol (37°00'S, 72°30'W), Concepcion, Concepcion Province, Chile 1968, Chiang. Other records.—ARGENTINA: Smit (1955): 1 2, ex Oryzomys longicaudatus = Ol. longicaudatus, San Pedro, south shore of Lake Nahuel Huapi, Rio Negro Province, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Argentina, February 1954. CHILE: Roths- child (1908): 1 3, ex Dromiciops australis = D. gliroides, Temuco, Cautin Province, November 1906; Hopkins and Rothschild (1966): 5 36, 5 ¢, plus many females in alcohol, ex D. gliroides, Valdivia, Valdivia Province, 1924; and Beaucornu and Kelt (1990): 1 3, ex Irenomys tarsalis (Philip- pi), 7 km E SE of Puerto Octay, Osorno Province, September 1985. Remarks.—Dr. Roberto Schlatter, Insti- tuto de Zoologia, Universidad Austral de Chile, Valdivia, kindly provided a series of 5 males and 11 females of this uncommon flea to the senior author just prior to our field collection trip. They were collected from a single specimen of D. gliroides in the ‘“‘Valdivia city environs” during July 1981 by Dr. Schlatter. The natural host for C. a. allophyla is the tiny arboreal marsu- pial D. gliroides. Dromiciops may hibernate during the cold Chilean winter months (June through August/September) accord- ing to Rageot (1979). We were unsuccessful in collecting Dromiciops, or locating active nests. A number of older nest boxes (mounted three to four meters high on the trunks of Nothofagus trees) contained de- teriorated nesting materials of Dromiciops from previous summer seasons. Their nests are often constructed in the thickets of quila 60 centimeters or more above the ground, in addition to building nests in the hollows of trees and logs (especially during the win- ter months). Chiliopsylla a. allophyla would likely be found in large numbers ei- ther on the animal, or in its nest during the winter months. It should be noted that the distribution of D. gliroides includes only south-central Chile from Concepcion south to Chiloé Island and east just beyond the Argentina border. This flea has been col- lected throughout the range of Dromiciops except for areas north of Cautin Province, Chile. Hystrichopsyllidae Ctenoparia inopinata Rothschild 1909 Material examined.—1 6, 2 2, ex Ak. olivaceus; 2 5, 1 2, ex Ol. longicaudatus. VOLUME 103, NUMBER 2 Other records.—ARGENTINA: Beau- cournu and Alcover (1989): 49 3,52 @, ex 99 specimens from Ak. longipilis, one from Geoxus valdivianus (Philippi), and one from Or. longicaudatus, adjacent to a series of lakes in Neuquén Province, December 1987—May 1988. CHILE: Rothschild (1909): 1 2, ex Ak. olivaceus, Valparaiso Province; Rothschild (1911b): unspecified number of females, ex unknown, Santiago; Jameson and Fulk (1977): 4 2, ex Ak. lon- gipilis, E of Molina, Santiago Province, April 1973; Beaucournu, et al. (1986): 1 6, 12, ex Ak. olivaceus, Puerto Carmen, Chi- loé Island, Chiloé Province, January 1985, 1 2, ex G. valdivianus, San Martin, Valdi- via Province, June 1985; Beaucornu and Kelt (1990): 1 2, ex Ak. longipilis, 22 km W NW of Puerto Octay, Osorno Province, September 1985, 2 6, 6 2, ex Ak. oliva- ceus, Coyhaique National Reserve, Aisén Province, February 1987. Remarks.—This flea is common in tem- perate rain forests of Neuquén Province, Argentina among Nothofagus trees, but is collected only infrequently in similar Chi- lean temperate rain forests dominated by species of Nothofagus and thick stands of quila. Perhaps more intensive collection ef- forts during winter months would demon- strate more abundant populations in Chilean temperate rain forest, although lower ele- vations of coastal Chile may be limiting for this species. Abrothrix longipilis is the pre- ferred host of C. inopinata. Ctenoparia topali Smit 1963 Material examined.—1 ¢, ex Ak. oliva- ceus. Other records.—ARGENTINA: Smit (1963): 1 3, ex (rodent nest), El Bolson (41°59’S, 71°35’W), Rio Negro Province, June 1961, elevation 700 m; and Beau- cournu and Alcover (1989): 31 6, 36 @, ex Ak. longipilis, Verde Lagoon, Ak. longipilis, Ak. olivaceus, Or. longicaudatus, and I. tar- salis, Lake Curruhue, Auliscomys micropus (Waterhouse,) and Or. longicaudatus, Lake Norquinco, Ak. longipilis, Or. longicauda- 439 tus and I. tarsalis, Ruca Malen, and Akodon sp., Lake Quillen, Neuquén Province, De- cember 1987—May 1988. CHILE: Beaucor- nu, et al. (1986): 1 3, ex Or. longicaudatus, Puerto Carmen, Chiloé Island, Chiloé Prov- ince, July 1985; Beaucornu, et al. (1988): 1 2, ex Ak. olivaceus brachiotis = Ak. oli- vaceus, San Martin, Valdivia Province, May 1986, 1 2, ex Ak. o. brachiotis, Quilan Is- land, Chiloé Province, January 1987, 1 @, ex Akodon sanborni = Abrothrix sanborni (Osgood), Quilan Island, Chiloé Province, January 1987; and Beaucournu and Kelt (1990): 1 2, ex Au. micropus, Coyhaique National Reserve, Aisén Province, Septem- ber 1986, 2 6,3 @, ex Ak. olivaceus and 3 3,5 2, ex Or. longicaudatus, Coyhaique National Reserve, Aisén Province, March 1987. Remarks.—Ctenoparia topali is essen- tially sympatric with C. inopinata, although C. topali does not occur as far north. Its recorded northern limit occurs just north of Concepcié6n, approximately 36°S latitude. This flea displays less host specificity than C. inopinata, occurring on most of the com- mon small mammals (species of Abrothrix, Akodon, Auliscomys, and Oligoryzomys) of the coastal temperate rain forest. Collec- tions in Argentina generally have been made at higher elevations than those in Chile. Populations of both C. topali and C. inopinata appear to increase with elevation as each is more abundant in Argentina than in similar habitats at lower elevations in coastal Chile. Rhopalopsyllidae Tetrapsyllus rhombus Smit 1955 Material examined.—2 6, 4 @, ex Ak. olivaceus; 2 3, 1 ¢, ex Ol. longicaudatus. Other records.—ARGENTINA: Smit (1963): 2 6, 2 2 , ex Or. longicaudatus Philippi = Ol. longicaudatus, August 1961, 3 36, ex Ak. o. brachiotis, April 1961, 3 6, 5 ¢, ex Ak. longipilis suffusus = Ab. lon- gipilis, August 1961, 1 6, 2 @, ex Or. lL. Philippi or Ak. o. brachiotis, January 1961, 440 1 2, ex Or. I. philippi or Ak. o. brachiotis, August 1961, 3 2, ex Ak. o. brachiotis or Ak: slusuffusus, June 19611 S, ex Or. «1. philippi or Ak. o. brachiotis or Ak. 1. suf- fusus, April 1961, 4 3, 6 &, (nest), June 1961, 4 6,3 2, (mest), April 1961, 1 ¢, 1 2, (nest), August 1961, 1 6, forest litter, October 1961, El Bolsén (41°59’S, 71°35’W), Rio Negro Province; Beaucornu and Gallardo (1988): number of specimens, host data, and collection dates omitted, ~ 40 km E NE of Bariloche, Rio Negro Prov- ince; and Beaucournu and Alcover (1989): 31 3, 41 &, [ex Rattus norvegicus (Ber- kenhout) and Ak. longipilis, Lake Huechu- lafquen, ex Ak. longipilis, Verde Lagoon, ex Octodon bridgesi Waterhouse, Ak. longipi- lis, Ak. olivaceus, and Aconaemys sp., Lake Curruhue, ex Ak. longipilis, Hua Hum, ex Ak. longipilis, Ruca Malen, ex Ak. longi- pilis and Or. longicaudatus, Pampa de Hui Hui], Neuquén Province, December 1987— May 1988. CHILE: Beaucournu and Kelt (1990): 1 36, 1 2, ex Ak. olivaceus, 7 km E SE Puerto Octay, Osorno Province, Sep- tember 1985, 1 2, ex Au. micropus, No- vember 1986, 2 d, 2 2, ex Ak. olivaceus, March 1987, 1 2°, ex G. valdivianus, March 1987, Coyhaique National Reserve, Aisén Province, 1 6, 1 @, ex Ak. longipilis De- cember 1986, 2 2, ex Reithrodon physodes = Reithrodon auritus (Fischer), March 1987, 4.5 km E Coyhaique Alto, Aisén Province, 2 6, ex Ak. longipilis, February 1987, Puerto Ibafiez, El Salto, Aisén Prov- ince. Remarks.—The specificity of hosts can not be determined by existing records. The only large series available was collected by Beaucornu and Alcover (1989) and unfor- tunately they did not address the number of T. rhombus that they collected with their associated hosts. Existing host associations would suggest they parasitize the dominant small rodents in their range. Akodon oliva- ceus, Ab. longipilis, and Ol. longicaudatus yield most of the resulting records, but this may only reflect those rodents most fre- quently captured. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Tetrapsyllus tantillus (Jordan and Rothschild 1923) Material examined.—1 ¢, ex Ak. oliva- ceus (collected in an empty lot within the city limits of Valdivia, Valdivia Province, Chile, 12 September 1999, M.W. Hastriter). Other records.—ARGENTINA: Jordan and Rothschild (1923): 4 2, ex Ctenomys haigi Thomas, Cholila, Chubut Province, 700 m, January 1920; Smit (1955): 4 6, 4 2, ex Or. longicaudatus, Irenomys sp., No- tiomys valdivianus fossor = G. valdivianus (nest), Rattus sp., and R. norvegicus, San Pedro, south shore of Lake Nahuel Huapi, Rio Negro Province; Beaucournu and Al- cover (1989): number of specimens, hosts, and date of collections omitted, ex Or. lon- gicaudatus, Ak. longipilis, Au. micropus, and Ak. olivaceus, Neuquén Province; and Smit (1963): 1 6, 1 2, ex Ak. o. brachiotis, 1.X, ex Or. |. philippi, 16,1. 23-exgAieae suffusus, 5 3, 2 °, ex (nest), El Bolsén (41°59’S, 71°35'W), Rio Negro Province. CHILE: Smit (1968): 5 6, 3 2, (from for- malin pit traps), Farellones, Santiago Prov- ince, elevation 2,000—2,400 m; Smit and Rosicky (1972): 1 6, ex Akodon sp., Lake Pehue, Magalanes Province; Jameson and Fulk (1977): 1 2, ex Spalacopus cyanus (Molina), 1 36, 2 2, ex Ak. olivaceus, 2 ¢, ex. Ak. longipilis, 1 3, ex Phyllotis darwini (Waterhouse), Coquimbo Province, 1 6, 4 2, ex Ak. longipilis, Santiago Province; and Beaucournu and Kelt (1990): 10 3, 10 2, ex R. physodes, 6 3,9 &, ex Akodon xan- thorhinus (Waterhouse), 10 6, 10 @, ex Phyllotis xanthopygus (Waterhouse), 12 ¢ 12 2, ex Ak. longipilis, 3 2, ex Euneomys sp., 5 2, ex Au. micropus, Aisén Province. Remarks.—A widely dispersed flea, re- ported from Coquimbo Province in the north through much of Magallanes Prov- ince in the south and east into adjacent re- gions of Argentina. Phyllotis darwini ex- tends only into the northern fringe of the range of 7. tantillus and is probably an ac- cidental association. Abrothrix longipilis is the most commonly infested host of this VOLUME 103, NUMBER 2 flea, although other associated mammals are infested readily. The significant number of specimens (eight!) collected in formalin pit traps (Smit 1968) would suggest that this flea may display a peculiar host seeking behavior (unassociated with the nest) as in some pulicid fleas, e.g., species of Pulex L. and Ctenocephalides Stiles and Collins. Stephanocircidae Barreropsylla excelsa Jordan 1953 Material examined.—1 2°, ex: Ol. longi- caudatus 2, 14 September 1999; and 1 @, ex: Ak. olivaceus 2, 16 September 1999. Other records.—ARGENTINA: Jordan (1953): 1 3d, 1 2, ex Akodon sp., Bariloche, Lake Nahuel Huapi, Rio Negro Province, 7sOrmn:, July, 1952: Smit) (1955)> dic 5, 1.09), ex Euneomys dabbenei = Euneomys peter- soni J.A. Allen, July 1953, 1 2, ex Akodon neocensus Thomas, July 1953, San Pedro, south shore of Lake Nahuel Huapi, Rio Ne- gro Province, elevation 810 m; Smit (1963): 1 2, ex Or. lL. philippi, June 1961, 1 36,2 2, ex Or. I. philippi, August 1961, 1 3, ex Ak. o. brachiotis, April 1961, 1 6, 1 2, ex Ak. o. brachiotis, August 1961, 1 2, ex Ak. Ll. suffusus, June 1961, 1 6,2 °, ex Ak. |. suffusus, August 1961, 1 3d, ex Ak. o. brachiotis or Ak. 1. suffusus, June 1961, iG. lee exrOr il! philippisor Ak. obra: chiotis, August 1961, 1 3, ex Or. lL. philippi or Ak. o. brachiotis or Ak. |. suffusus, April 1961, El Bolsé6n (41°59’S, 71°35'W), Rio Negro Province, elevations 300—480 m; Beaucournu and Gallardo (1988): number of specimens, host data, and collection dates omitted, ~ 40 km E NE of Bariloche (type locality), Rio Negro Province; and Beaucournu and Alcover (1989): 6 6, 14 2, (ex Ak. longipilis and Or. longicaudatus, Lake Curruhue, ex Or. longicaudatus, Lake Norquinco, ex Ak. longipilis, Ruca Malen, and ex Akodon sp., Lake Quillen), Neuquén Province, December 1987 till May 1988. CHILE: Beaucornu and Kelt (1990): 2 6, 1 2, ex Ak. longipilis, Puerto Ibafiez, Aisén Province, February 1987. 441 Remarks.—Barreropsylla excelsa has been collected only in small numbers (one to three) on individual hosts throughout its range. Beaucournu and Kelt (1990) suggest it is primarily a nest flea. This flea appears to prefer Ol. longicaudatus, Ab. longipilis and Ak. olivaceus throughout its limited geographic range. Although more intensive collecting will undoubtedly increase their known range, they are currently found only between a narrow band approximately 39°— 44°S latitude and 71°—73°W longitude. EI- evations range between 300 m and 810 m. Recorded collection dates would suggest that B. excelsa is most abundant during the cold rainy winter months of June through August, although Beaucournu and Kelt (1990) noted three specimens collected dur- ing February. Sphinctopsylla ares (Rothschild 1911) Material examined.—2 6, 3 @, ex Ak. olivaceus (one flea with eggs); 3 6, 15 @, ex Ol. longicaudatus (six fleas with eggs); and 1 6, ex Ak. olivaceus or Ol. longicau- datus (found in bag). Other records.—ARGENTINA: Smit (1955): 12, ex Abrothrix hirta modestior = Ab. longipilis, July 1952, 1 3, 2 2, ex Ab. h. modestior, July 1953, 1 2, ex Akodon sp., July 1952, 3 2, ex Akodon nucus = Akodon iniscatus Thomas, July 1953, 3 @, ex Or. longicaudatus, July 1952, 1 3, ex Or. longicaudatus July 1953, 3 6,5 @, ex Or. longicaudatus, February 1954, 2 2, ex N. v. fossor, July 1953, and 2 9, ex Rattus sp., July 1952, all from San Pedro, South Shore of Lake Nahuel Huapi, Rio Negro Province: Smit (1963): 1 6, 8 2, ex Or. lL. Philippii, 350—480 m, 2 3, 1 @, ex Ak. o. brachiotis, 480 m, 1 3, 2 2, ex Ak. L. suf- fusus, 300 m, 2 3, 4 2, ex Or. 1. philippii, or Ak. 0. brachiotis, 350-360 m, 2 2, ex Ak. o. brachiotis, or Ak. 1. suffusus, 360 m, 4 2, ex Or. l. philippii, or Ak. 0. brachiotis, or Ak. |. suffusus, 480 m, and 1 6,1 2, ex R. norvegicus, 350-380 m,, El Bolsoén (41°59’S, 71°35’W), Rio Negro Province, January—October 1961; Beaucournu and 442 Alcover (1989): 53 6, 105 2, ex R. norn- egicus, Ak. longipilis, G. valdivianus, Lake Huechulafquen, ex Ak. longipilis, Cerro Chapelco, ex Ak. longipilis, Verde Lagoon, ex R. norvegicus, AK. olivaceus, Ak. lon- gipilis, Or. longicaudatus, I. tarsalis, Lake Curruhue, ex Ak. longipilis, Lake Lolog, ex Ak. longipilis, I. tarsalis, Quilanlahue, ex Or. longicaudatus, Ak. longipilis, Lake Norquinco, ex Ak. longipilis, Hua Hum, ex Ak. longipilis, Or. longicaudatus, I. tarsalis, Ruca Malen, ex Ak. longipilis, Volcan Huanqui Hue, Neuquén Province, Decem- ber 1987—May 1988. CHILE: Rothschild (191la): 1 2, ex Ak. olivaceus, Temuco, Cautin Province, February 1908; Smit and Rosicky (1972): 1 2, ex Akodon sp., Estan- cia Pudeto (51°05’S, 73°00'W), Lake Pe- hue, Magallanes Province, February 1969; Jameson and Fulk (1977): 1 @, ex P. dar- wini, Fray Jorge National Park, Coquimbo Province, June 1972, and 1 2, ex Or. lon- gicaudatus, 5 km N of La Serena, Coquim- bo Province, May 1972; Beaucornu and Kelt (1990): 1 2, ex Or. longicaudatus 14.5 km N NW Puerto Octay, Osorno Province, August 1986, 1 66, ex Chelemys macro- nyx (Thomas) September 1986, 1dex C. macronyx, March 1987, Coyhaique Nation- al Reserve, Aisén Province, 1 @, ex Au. micropus, 2 3, 10 2, ex Ak. longipilis, July 1986, 1 2, ex Reithrodon physodes, | 3, 1 2, ex Ak. xanthorhinus, 6 °, ex Ak. lon- gipilis, 2 3,5 2, ex Au. micropus, 4.5 km E Coyhaique Alto, Aisén Province, March 1987, 7 36, 25 2, ex Or. longicaudatus, 18 3, 34 ¢, ex Ak. olivaceus, 1 3, ex G. val- divianus, Coyhaique National Reserve, Aisén Province, March 1987, 4 36,8 @, ex Ak. longipilis, Puerto Ibanez El Salto, Aisén Province, March 1987, and 1 2, ex P. xan- thopygus, Puerto Ibanez, Rocky Bluff, Aisén Province, April 1987. Remarks.—Sphinctopsylla ares is a com- mon flea ranging from the Chilean province of Elqui in the north into the southern prov- ince of Magallanes and east into Neuguén and Rio Negro Provinces, Argentina. The species demonstrates little host specificity, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON occurring on most genera within the sub- family Sigmodontinae (Muridae) that are present within the flea’s range. It has also been reported on species in the genera Aconaemys Ameghino and Spalacopus Wagler (Octodontidae) and on species of the marsupial genus Marmosa Gray (Di- delphidae). Sphinctopsylla ares has been collected during all months of the year ex- cept for May and December [it may have been collected during these months also by Beaucournu and Alcover (1989), since they did not specify dates of collection, other than ‘“‘December 1987 till May 1988’’]. Multiple large eggs were present in seven of the 18 females that we collected during September. These enormous eggs have an extremely thick and well-defined chorion whose gross morphology resembles eggs of hystrichopsyllid species of Stenoponia Jor- dan and Rothschild and Aystrichopsylla Taschenberg. ACKNOWLEDGMENTS The authors express their sincere appre- ciation to Milton H. Gallardo and Fredy Mondaca, Instituto de Ecologia y Evolu- cion, Universidad Austral de Chile, Valdi- via, for the technical and logistic support necessary to accomplish the survey work and for confirmation of mammal identifi- cations, and to Pedro Mujfioz for his kind guidance and support during our stay at the San Martin Reserve facility. Funding for this research was partially provided by a grant from the National Science Founda- tion, Award Number DEB-9983195. LITERATURE CITED Beaucournu, J. C. and J. A. Alcover. 1989. Puces ré- coltées dans la provinces de Neuquén (Argentine); description de 4 nouveaux taxa (Insecta, Siphon- aptera). Annales de Parasitologie Humaine de Comparée 64(6): 489-505. Beaucournu, J. C. and M. H. Gallardo. 1988. Puces nouvelles d’ Argentine (Insecta, Siphonaptera). Revue suisse de Zoologie 95(1): 99-112. . 1991. Catalogue provisoire des puces du Chili (Insecta; Siphonaptera). Bulletin de la Société Frangaise de Parasitologie 9(2): 237—70. . 1992. Catalogue provisoire des puces du Chili VOLUME 103, NUMBER 2 (Insecta; Siphonaptera). Bulletin de la Société Frangaise de Parasitologie 10(1): 93-130. Beaucournu, J. C., M. H. Gallardo, and H. Launay. 1986. Puces (Siphonaptera) nouvelles ou peu con- nues du Chile: Description de Plocopsylla diana n. sp. (Stephanocircidae). Annales de Parasitolo- gie Humaine de Comparee 61(3): 359-366. Beaucournu, J. C. and D. A. Kelt. 1990. Contibution a la faune du Chili: Puces nouvelles ou peu con- nues de la partie and sud (Insecta, Siphonaptera). Revue suisse de Zoologie 97: 647—668. Beaucournu, J. C., J. C. Torres Mura, and M. H. Gal- lardo. 1988. Description de la femelle de Creno- paria topali Smit, 1963 et clef dichotomique du genre Crenoparia Rothschild, 1909 (Siphonaptera, Hystrichopsyllidae). Annales de Parasitologie Hu- maine de Comparée 63(5): 380-383. Hastriter, M. W. 2000. Jordanopsylla becki (Siphon- aptera: Ctenophthalmidae), a new species of flea from the Nevada Test Site. Proceedings of the En- tomological Society of Washington 102(1): 135— 141. Hopkins, G. H. E. and M. Rothschild. 1966. An illus- trated catalogue of the Rothschild collection of fleas (Siphonaptera) in the British Museum. Vol. IV. Hystrichopsyllidae, The British Museum (Nat- ural History), London, 549 pp. Jameson, E. W., Jr. and G. W. Fulk. 1977. Notes on some fleas (Siphonaptera) from Chile. Journal of Medical Entomology 14(4): 401—406. Jordan, H. E. K. 1953. On a remarkable flea from Ar- gentina collected by Dr. J.M. de la Barrera. Bul- letin of the British Museum (Natural History) En- tomology, London 3(5): 179-186. Jordan, K. and N. C. Rothschild. 1923. On the genera Rhopalopsyllus and Parapsyllus. Ectoparasites 1: 320-370. Murua, R., P. L. Meserve, L. A. Gonzalez, and C. Jof- ré. 1987. The small mammal community of a Chi- lean temperate rain forest: Lack of evidence of 443 competition between dominant species. Journal of Mammalogy 68(4): 729-738. Rageot, R. 1979. Notas sobre la biologia del monito del monte. Anales del Instituto de Punta de Betin 12: 83-88. Rothschild, N. C. 1908. New Siphonaptera. Proceed- ings of the Zoological Society of London, pp. 617-629. . 1909. On some American, Australian, and Pa- laearctic Siphonaptera. Novitates Zoologicae 16: 61-68. . 191la. Some new genera and species of Si- phonaptera. Novitates Zoologicae 18: 117-122. 1911b. Liste de Siphonaptera du Museum d’ Histoire Naturelle de Paris, accompagnée de de- scriptions des especes nouvelles. Annales des Sci- ences Naturelles (Zoologie), (Series 9) 12: 203-— 216. Smit, F G. A. M. 1955. Siphonaptera from Bariloche, Argentina, collected by Dr. J.M. de la Barrera in 1952-1954. The Transactions of the Royal Ento- mological Society of London 107: 319-339. . 1963. The zoological results of Gy. Topdal’s collectings in South Argentina. Annales Historico- Naturales Musei Nationalis Hungarici par Zoolo- gica 55: 421—433. . 1968. Siphonaptera taken from formalin-traps in Chile. Zoologischer Anzeiger 180(3/4): 220— 228. Smit, EF G. A. M. and B. Rosicky. 1972. Some Si- phonaptera from Chile. Folia Parasitologica (Pra- ha) 19: 365-368. Smith, M. E and J. L. Patton. 1993. The diversification of South American Murid rodents: evidence from mitochondrial DNA sequence data for the Ako- dontine tribe. Biological Journal of the Linnaean Society 50(3): 149-177. Wilson, D. E. and D. M. Reeder. 1993. Mammal Spe- cies of the World, a Taxonomic and Geographic Reference. 24 edition, Smithsonian Institution Press, Washington, D.C., 1,206 pp. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 444—451 A NEW SPECIES OF CLADOCHAETA COQUILLETT (DIPTERA: DROSOPHILIDAE) AND A NEW RECORD FOR CLADOCHAETA STURTEVANTI WHEELER AND TAKADA IN ARIZONA, WITH NOTES ON NATURAL HISTORY TAM C. NGUYEN Division of Invertebrate Zoology (Entomology), American Museum of Natural History, Central Park West at 79'" Street, New York, NY 10024, U.S.A. (e-mail: nguyen@amnh. org) Abstract.—Cladochaeta johnsonae, new species, is described and figured, including description of larvae and their association with spittlebug (Cercopidae) nymphs. A new record for Cladochaeta sturtevanti Wheeler and Takada is reported from Arizona, previ- ously known only from California, Washington, and southwestern Utah. Key Words: copidae The genus Cladochaeta was erected by the dipterist, D. W. Coquillett (1856-1911) in 1900 for Cladochaeta nebulosa. The ge- neric name refers to the single-branched arista of this species. In 1924, J. R. Malloch described Clastopteromyia for the eastern North American species Clastopteromyia inversa, naming the genus in reference to its association with the cercopid Clastop- tera. These two generic names were sub- sequently used synonymously until Wheel- er and Takada (1971) recognized Clastop- teromyia as a junior synonym of Clado- chaeta. Vilela and Bachli (1990), in a study of type specimens, provided a diagnosis for a monophyletic genus Cladochaeta, in which they included 13 described species. Grimaldi and Nguyen (1999) reported over 100 additional new species to the genus and transferred two species from the genus Dia- thoneura to Cladochaeta. In the present pa- per, I describe the adults and immature stag- es of a new species of Cladochaeta and re- port a new record for C. sturtevanti Wheel- er and Takada. Cladochaeta, Drosophilidae, taxomony, Arizona, larvae, spittlebugs, Cer- MATERIAL AND METHODS During August 22-25, 1998, and Sep- tember 12-17, 1999, spitthkemasses were surveyed at and in the vicinity of the South- western Research Station (SWRS). The sta- tion is located near Portal, Arizona, in the Chiricahua Mountains at elevations of roughly 4,500—5,500 ft. (1,372—1,677 m). Vegetation of the area is dominated by oaks (Quercus spp.), pinyon pine (Pinus edulis Engelmann), juniper (Juniperus spp.), and agave and creosote at slightly lower eleva- tions. Spittlemasses with fly larvae or pupae were collected and kept in plastic vials with branch cuttings from the respective host plants. Some fly larvae and spittlebug nymphs were immediately preserved in 70% ethanol and subsequently critical point dried. The remaining immatures were al- lowed to eclose, were collected, critical point dried, and mounted for study. Diagnostic measurements of the adult head of Cladochaeta were taken as de- scribed in Grimaldi and Nguyen (1999) (see Table 1). Overall body length was also mea- VOLUME 103, NUMBER 2 445 Table 1. Measurements of Cladochaeta johnsonae adults (mm). ThL HW cD ED FW CD/ED FW/HW 3 0.971 0.750 0.086 0.361 0.250 0.238 0.333 0.911 0.742 0.074 OS72Z 0.225 0.198 0.303 0.959 0.750 0.090 0.396 0.237 0.227 0.316 i125) 0.843 0.084 0.407 O278 0.206 0.323 0.980 0.770 0.077 0.369 0.249 0.208 0.323 1.100 0.810 0.099 0.400 0.287 0.247 0.354 O23 0.819 0.082 0.408 0.282 0.200 0.344 1.020 0.706 0.074 O57 0.215 0.207 0.304 0.857 0.687 0.071 0.35 0.251 0.203 0.365 0.858 0.678 0.073 0.338 0.228 0.216 0.336 0.994 0.813 0.083 0.39 0.273 0.213 0.336 0.905 0.716 0.077 0.361 0.249 0.213 0.348 0.791 0.662 0.064 0.349 0.239 0.183 0.361 0.834 0.679 0.070 0.344 0.254 0.203 0.374 0.953 0.792 0.074 0.372 0.271 0.199 0.342 Average 0.952 0.748 0.079 0.372 0.252 0.211 0.337 ? 0.912 0.729 0.074 0.349 0.233 0.212 0.319 0.890 OF/25 0.083 0.345 0.243 0.240 0.335 0.904 0.722 0.070 0.338 0.244 0.207 0.338 0.809 0.681 0.065 0.323 0.226 0.201 0.332 0.923 0.739 0.072 0.373 0.232 0.193 0.314 0.746 0.650 0.068 0.313 0.218 0.217 0.335 0.839 0.677 0.074 0.338 0.192 0.219 0.284 0.906 0.765 0.082 0.361 0.250 0.227 0.327 0.812 0.640 0.070 0.336 0.203 0.208 0.317 0.914 0.696 0.076 0.358 0.223 0.212 0.320 0.784 0.616 0.062 0.318 0.208 0.195 0.338 0.703 0.613 0.062 0.321 0.203 0.193 0.331 Average 0.845 0.688 0.0715 0.339 0.223 0.210 0.324 ThL = Thorax Length, HW = Head Width, CD = Cheek Depth, ED = Eye Depth, FW = Face Width. sured for adult cercopids. To study the mat- ing behaviors of these flies, some flies were maintained in culture using standard instant Drosophila medium (Carolina Biological Supply Company). Virgin females were kept separate from males. Several days after eclosing, a single virgin female was intro- duced to a single male in a plastic vial and observed up to one hour. Genitalia of males and females were dissected and mounted on microscope slides using techniques de- scribed by Grimaldi (1987). Mouthparts of larvae were obtained through dissection of specimens as well as from puparia. Select fly larvae and spittlebug nymphs were sput- ter coated with gold/palladium for study by scanning electron microscopy (SEM). Elec- tromicrographs were also taken of heads of male and female Cladochaeta adults. All SEM’s were taken with a Hitachi S4700 Field Emission SEM. Morphological ter- minology follows Grimaldi and Nguyen (1999). All specimens are deposited in the American Museum of Natural History (AMNH),. RESULTS Individuals of Cladochaeta associated with spittlebugs were found on golden rab- bitbrush (Chrysothamnus nauseosus Brit- ton). Based on dissected male genitalia, the flies associated with cercopids on this plant were identified as C. sturtevanti, which had not previously been reported in Arizona. Previous collections of this species have mostly been from California, with a few in- 446 dividuals from Washington and southern Utah. Eleven male and ten female flies were collected during the first collection period. The Chrysothanmus cercopids were identi- fied as Clastoptera lineatocollis Stal (Doer- ing, 1928). The spittkemass produced by this species of cercopid was very frothy and usually contained only a single nymph. Ap- proximately 30% of the spittlemasses con- tained either a Cladochaeta larva or pupa. Occasionally, two pupae were found within the same spittlemass. Fly pupae were found in locations varying from within the spittle- mass, generally at the edge, to distant of the spittlemass by as much as a centimeter, gen- erally lodged in the node of a small branch apical to the spittlemass. The cercopids were rather small (3.97 mm body length). In several cases, apparently moribund nymphs of this species were found close to a fly pupa, which has never been observed for other Cladochaeta, even C. inversa (Grimaldi and Nguyen 1999). Immature stages of a new species of Cla- dochaeta also were associated with spittle- bugs on wild grape (Vitis arizonica Engel- mann). Three females and nine males of this new Cladochaeta species were reared to adults and collected during the first visit to SWRS. Eleven females and 11 males were collected during the secong collecting period. The wild grape spittlebugs were small (3.66 mm average body length) and keyed out to a species of Clastoptera be- tween Jawsoni Doering and arizonana Doering (Doering 1928). These spittlebugs were found only on wild grape and never on Chrysothamnus. They produced a very mucilaginous spittlemass, which almost al- ways contained fly larvae. The Cladochaeta larvae were observed crawling through the jelly-like portion of the grape spittlemasses more often than those found in Chryso- thamnus spittlemasses. However, many lar- vae were also found attached to nymphs (Fig. 1). These spittlemasses harbored ag- gregations of up to 50 nymphs, many of which were early instars. Attempts at maintaining Cladochaeta in PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON culture were largely unsuccessful. Males survived up to two weeks but females usu- ally died after a few days. It appeared that these females did not eat any of the instant Drosophila medium since their abdomens were shrunken. No results were recorded for the mating behavior of this Cladochae- ta. When introduced into the same vial, males and females ignored one another and usually remained apart and groomed them- selves. Individuals of both sexes, however, were observed to move their wings in a fashion similiar to that reported by Spieth (1952: 454-455) for Chymomyza. No movement of the forelegs was observed. SYSTEMATICS Cladochaeta johnsonae Nguyen, new species (Figs. 1-17) Diagnosis.—Thorax dark yellow to brown with darker areas laterally. Abdomen with light yellow pattern in a V-shape an- teriorly; brown to black posteriorly. Male genitalia similar to Cladochaeta mystaca Grimaldi and Nguyen (see Discussion). Male with long and convergent vibrissae having tips that intertwine to form one full spiral. Male antenna with pedicel having a medially projecting, long, fingerlike lobe with numerous setulae at the apex. Differs from C. mystaca, which has a brush of lon- ger setae on smaller pedicel lobes. Description.—Head: Eyes with very sparse, short pilosity. In male, antenna with pedicel with fingerlike lobe having numer- ous, long setulae; setulae very thick at apex; lobe almost equal in length to flagellomere I (Figs. 2, 3). Female without modified ped- icel (Fig. 4). Flagellomere I dark brown to black in both sexes. Arista with 3 dorsal branches (Fig. 5); one ventral branch be- tween d-3 and apical fork; ventral branch equal in length to d-3; branches of apical fork about 0.5 length of d-3. Frons brown- ish yellow; interfrontal setulae of about equal length to anterior reclinate orbital se- tae but thinner. Proclinate setae about same VOLUME 103, NUMBER 2 a tae lar eth UG i 1.00mm | Figs. 1-6. 447 et ee Seco el, Ua bel 2 73um 2 MeN ts 231um 4 Cladochaeta johnsonae. 1, Cercopid nymph with attached larva. 2, Head of male. 3, Detail of male antennae, showing modified pedicels. 4, Head of female. 5, Arista. 6, Wing. length as posterior reclinate orbital setae; anterior reclinate setae about 0.5 length of proclinates. Postocellar setae medium to small and convergent. Face medium tan and flat; cheeks yellow and of moderate depth. In male, vibrissae long, very fine, and con- vergent, forming spiral; in female, thicker and without spiral. Proboscis and palps yel- low. Thorax: Scutellum yellow; scutum gen- erally yellowish with darker areas in pos- terior and lateral portions. Anterior scutellar setae large and convergent; apical scutellar setae cruciate and slightly shorter than an- 448 terior scutellars. Anterior dorsocentral setae about 0.5 length of posterior dorsocentrals. Legs light yellow. Forefemur with row of 4 stiff dorsolateral setae and row of 3 ventro- lateral setae; in female dorsolateral setae of same thickness as in male but slightly short- er; hindfemur of both sexes without prom- inent setae. Wing hyaline with diffuse in- fuscation along costal edge down to past R,,;; infuscation darkest around costal edge towards but not reaching apex of wing (Fig. 6); very slight clouding of x-vein dm-cu; venation and shape as in C. mystaca. Hal- tere light yellow. Abdomen: Tergites I-III yellow medial- ly, brown laterally; tergites IV—VII uni- formly brown. Sternites yellow; lighter yel- low in female. Female terminalia (Fig. 7): Apical tergite narrow and without setae; simpe inverted U-shaped. Apical sternite sclerotized; base flared into 2 lateral arms curving anteriorly and downwards; tip of sternite undivided, projecting posteriorly, with 12 small setae. Male genitalia (Fig. 8) very similar to C. mystaca (Grimaldi and Nguyen 1999, fig. 64) with following dif- ferences: Cercus larger and triangular shaped; epandrium height about 1.5 width vs. being equal; tapering in distal ends of epandrium more pronounced; each half of epandrium with row of 6—7 setae vs. having 5 setae; distal third of paraphyses much broader; apices of paraphyses without not- icable knob. Type material—Holotype ¢, ARIZO- NA: Cochise Co., vicinity of Portal. 5|000— 5500 ft. VIII/20—25/98. Reared from cer- copids on wild grape. Nguyen and Grimal- di, colls. (AMNH). Paratypes, same data as holotype (2 2, 2 6) (AMNH). Other specimens examined.—ARIZO- NA: Cochise Co., near Portal, 5,000—5,500 ft. [X/13—21/99. Reared from cercopids on wild grape. T. Nguyen, coll. (11 2, 11 3) (AMNH). Same data as holotype (1 2, 6 3) (AMNH). Etymology.—Named for Christine A. Johnson, a colleague at AMNH and veteran researcher at SWRS. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Third instar larva (Figs. 9—17).—Amphi- neustic; anterior spiracle with 3 filaments. Head small (Fig. 10); antenna and maxil- lary palp complex as in other drosophilids. Oral cavity small and round. Oral lamellae in 10—12 transverse rows with long, thin fringes; 3 rows anterior to oral cavity; 1 vertical row of lamellae flanking either side of oral cavity (Fig. 11). Mandible entirely sclerotized, small and hooked with | prom- inent tooth and 3 smaller ones posterior to apical hook on both ventral edges (Fig. 12); teeth form a groove on ventral surface of each mandible (Fig. 13). Anterior portions of cephalopharyngeal skeleton, including hypopharynx, heavily sclerotized; hypopha- ryngeal sclerite moderately sclerotized; dor- sal and ventral cornu not sclerotized (Fig. 14). Six pairs of prolegs, each with crochet of 25-30 sharp, hooked spinules in 3—4 rows; spinules of first row pointing anteri- orly, those of remaining rows pointing pos- teriorly (Fig. 15). Pair of fingerlike and ap- parently eversible structures lateral to anus (Fig. 16) (probably same structures found in C. inversa by Grimaldi and Nguyen 1999, fig. 167c). Posterior spiracles on tele- scoping trunk about 1.5 length of everted spiracular tubes; spiracular tubes separated and eversible (Fig. 17). Spiracular plate with 3 spiracular openings. Spiracular hairs reduced as in other Cladochaeta larvae ex- amined (Grimaldi and Nguyen 1999: 292). DISCUSSION Cladochaeta johnsonae is placed in the sororia species group (Grimaldi and Nguy- en 1999: 92) and is closely related to C. mystaca based on remarkably similar struc- tures in the male genitalia, particularly of the paraphyses. Both species also have modified pedicels and unusual oral vibris- sae (for flies). Despite these similarities, the differences in the modified pedicels of C. jJohnsonae and C. mystaca are striking enough to justify separate species. This is also consistent with the subtle differences in the male genitalia of these two species. McAlpine (1976) suggested that spiral vi- VOLUME 103, NUMBER 2 x50 7/28/99 600um Figs. 7-12. 449 66.0um 10 PUP Pa 8 10.0ur% { 2 Cladochaeta johnsonae. 7, Female terminalia, lateral view. 8, Male genitalia, posterior view. 9, Habitus of third-instar larva. 10, Head of larva. 11, Oral cavity of larva, showing oral lamellae. 12, Detail of larval mouth hook. brissae reported in some male clusiid flies may be used in combat. It is unclear wheth- er this is the case in these two Cladochaeta species. During the second collecting period, no C. sturtevanti larvae nor their associated cercopids were found. This is probably due to the later collecting date during this pe- riod. Both C. sturtevanti and C. johnsonae were restricted to their respective host cer- copids and plants. In only one instance dur- ing the second collecting period, four fly larvae were collected from cercopids on an unidentified vine (possibly an introduced 450 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON af } Bt, is Bly, Jair e ecw anertananer 3.3um 15 | rf? 84.0um 16 10 Ve 1 x600 7/28/99 50.0um 17 Figs. 13-16. Cladochaeta johnsonae. 13, Ventral view of larval mandibles, showing teeth and ventral grooves. 14, Larval cephalopharyngeal skeleton. 15, Detail of proleg, showing crochet of hooks. 16, Everted para-anal organs. 17, Posterior spiracles. species) next to a large stand of wild grape. single female eclosed and was indentified This vine had many spittlemasses of cer- as C. johnsonae. The remaining larvae died copids of apparently the same species as _ and were lost. The presence of Cladochaeta those on wild grape. Only one of these spit- on this different plant may be a chance oc- tlemasses contained any fly larvae. Of the currence due to the proximity of its usual fly larvae collected from this plant, only a host plant. A more thorough examination of VOLUME 103, NUMBER 2 the plants in this area will determine if C. Johnsonae is restricted to cercopids on wild grape. The morphology of C. johnsonae third- instar larvae suggests an intermediate state between parasitic and saprophagous life- styles. The oral lamellae of C. johnsonae are more numerous and developed than in other described Cladochaeta larvae and are closer to those of known saprophagous dro- sophilids (Grimaldi and Nguyen 1999, fig. 170). These lamellae, however, do not con- verge as strongly into the oral cavity as in other sparophagous flies. The mandibles of C. johnsonae are rather blunt and bear teeth. These teeth and ventral grooves on the mandibles are strikingly similar to those of a saprophagous ephydrid reported to live in spittlemasses of Tomaspis inca Guérin- Méneville in Costa Rica (Grimaldi and Nguyen 1999: 293). However, C. johnsonae does possess only a partially sclerotized cornu of the cephalopharyngeal skeleton. In saprophagous species the cornu is heavily sclerotized. The common observation of C. johnsonae larvae attached to spittlebug nymphs also indicates a parasitic lifestyle. These aspects of the morphology of C. jJohnsonae larvae may suggest a facultative- ly spaprophagous lifestyle and possible switching between true parasitism and sap- rophagy in this species. This probably ex- plains why C. johnsonae larvae were found often crawling freely within spittlemasses, a habit which was not seen frequently in C. sturtevanti larvae. ACKNOWLEDGMENTS I am indebted to David A. Grimaldi for his guidance and help with this work. In- 451 deed, my interest in flies, specifically Cla- dochaeta, began through working with Dave. I also thank Michael S. Engel, Molly G. Rightmyer, and Michelle L. Lenzi for their comments and review of the manu- script. For help in identifying the spittle- bugs, I thank K. G. A. Hamilton for his kindness. LITERATURE CITED Coquillett, D. W. 1900. Report on a collection of dip- terous insects from Puerto Rico. Proceedings of the United States National Museum 22: 249-270. Doering, K. C. 1928. The genus Clastoptera in Amer- ica north of Mexico. University of Kansas Science Bulletin 18(1): 5-153. Grimaldi, D. A. 1987. Phylogenetics and taxonomy of Zygothrica (Diptera: Drosophilidae). Bulletin of the American Museum of Natural History 186: 103-268. Grimaldi, D. A. and T. Nguyen. 1999. Monograph on the spittlebug flies, genus Cladochaeta (Diptera: Drosophilidae: Cladochaetini). Bulletin of the American Museum of Natural History 241: 1— 326. Malloch, J. R. 1924. Descriptions of Neotropical two- winged flies of the family Drosophilidae. Pro- ceedings of the United States National Museum 66: 1-11. McAlpine, D. K. 1976. Spiral vibrissae in some clusiid flies (Diptera: Schizophora). Australian Entomo- logical Magazine 3(4): 75—78. Spieth, H. T. 1952. Mating behavior within the genus Drosophila (Diptera). Bulletin of the American Museum of Natural History 99(7): 395-474. Vilela, C. R. and G. Bachli. 1990. Taxonomic studies on neotropical species of seven genera of Droso- philidae (Diptera). Mitteilungen der Schweizer- ischen Entomologischen Gesellschaft 63(suppl.): 1-332. Wheeler, M. R. and H. Takada. 1971. Male genitalia of some representative genera of American Dro- sophilidae. University of Texas Publication 7103: 225-240. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 452-456 A NEW SPECIES OF PROTOTHEORA FROM MALAWI, WITH ADDITIONAL NOTES ON THE DISTRIBUTION AND MORPHOLOGY OF THE GENUS (LEPIDOPTERA: PROTOTHEORIDAE) DONALD R. DAVIS Department of Entomology, National Museum of Natural History, Smithsonian Insti- tution, Washington, DC 20560-127, U.S.A. (e-mail: davis.don@nmnh.si.edu) Abstract.—A new species, Prototheora malawiensis, of the endemic southern African family Prototheoridae is described and illustrated from the Mulanje Massif of Malawi. This constitutes only the second species to be collected outside of South Africa, the first being Prototheora angolae Davis from Angola. Also noted are additional collecting re- cords for Prototheora cooperi Janse, P. corvifera (Meyrick), P. petrosema Meyrick, P. quadricornis Meyrick, and the first record from Cape Province, South Africa, of Proto- theora parachlora Meyrick. Key Words: Prior to my recent revision of the hepi- aloid family Prototheoridae (Davis 1996), this family was known only from South Af- rica, with all but Prototheora parachlora (Meyrick) restricted to Cape Province. In that report I described P. angolae from the central plateau region of Angola, as well as a second species, P. drackensbergae Davis, from Natal Province, South Africa. In Oc- tober 1996, Woffram Mey of Humboldt Universitat, Berlin, collected (by sweeping) an undescribed Prototheora on the Mulanje Massif of southern Malawi. The moth was observed flying rapidly in the afternoon Over a grassy meadow at an elevation of 2,000 meters. The discovery of this species further extends the range of this family in southwestern Africa and brings the total number of described species to 12. Descrip- tion of this species is necessary at this time in order to include it in a synoptic catalogue of the family soon to be published (Davis, in press). Also noted herein are additional records of several previously described species Africa, biogeography, Hepialoidea, South Africa from Cape Province, including the first Cape record of P. parachlora, a species previously known only from from the ho- lotype, collected in Natal. Prototheora malawiensis Davis, new species (Figs. 1—4) Male (Fig. 1).—Wing expanse 20 mm. Head: Vestiture consisting of broad (over frons and vertex) to narrow (between an- tennae), gray, white-tipped scales with mi- nutely serrated apices; scales over occipital area longer, piliform. Antenna ~ % the length of forewing, 34-segmented; scales of scape slender, reduced in size, and same color as head; flagellum dark brown, naked except for dense pubescence of pale sensil- la. Labial palpus same color as head, long and porrect; length ~2.5X eye diameter. Thorax: Scales of dorsum similar in col- or to head, mostly with slender bases; me- tascutum with scattering of long, pale, pil- iform scales. Venter mostly naked, with long, white, piliform scales. Forewing light went-103-02-27-f01 VOLUME 103, NUMBER 2 453 Fig. 1. Adult, P. malawiensis, wing expanse 20 mm. gray, with scattered irroration of fuscous to black scales; a prominent longitudinal, bro- ken, white band from wing base to apex, widest at distal fourth beyond apex of discal cell; another linear concentration of dark fuscous scales along lower border of distal fourth of white band; fringe mostly white, with scattered, longer, gray scales. Hind- wing uniformly light brownish gray. Fore- leg gray dorsally, finely irrorated with slen- der, white-tipped scales; with less irroration ventrally, more brownish over venter of foreleg and most of midleg. Hindleg uni- formly light brown. Abdomen: Predominantly gray, with an irregular, narrow, longitudinal band of white along midventer. Genitalia (Figs. 2—4): Tergal plate a rel- atively small, triangular tergite devoid of processes. Pseudoteguminal plates moder- ately broad at base, articulating aterally with lateral arms of vinculum; pseudote- guminal arms slender, elongate, complex with secondary, spinose branching (Fig. 4) including a short basal spur, and an inverted T-shaped, subapical spinose branch from venter of main arm. Suspensorium weakly developed, with slender lateral arms ex- tending to pseudoteguminal plate. Copula- tory pouch well developed, approximately equal in length to basal third of valva in- cluding saccular lobe. Trulleum with deep ventral groove, the lateral edges of which equal in length to saccular lobe. Valvae moderately broad at base, with a prominent, setose saccular lobe protruding caudally and a greatly extended, spinelike cucullar arm projecting dorsally at right angle to basal third of valva. Vinculum broadly rounded, short, approximately equal to sac- cular lobe in length. Female.—Unknown. Holotype.—¢; Malawi: Mulanje Mts., Chambe Hut — Lichenye Hut, montane meadows, 2000 m, 21 Oct 1996, leg. Mey and Nuss, slide DRD 4114 (MNHU). Distribution.—Known only from the type locality, a montane site in extreme southern Malawi, 16°03”S/35°31"E. Remarks.—The male genital morphology of this species is among the most distinctive of the genus. It is the only species of Pro- totheora discovered thus far to lack tergal processes from the tenth tergum. The pseu- doteguminal arms are the most complex of any species in possessing secondary and tertiary branches of spinose processes. The valvae, although highly modified, bear some resemblance to those of P. drackens- bergae in the development of a setose sac- 454 cular lobe. In addition to adding to our knowledge of the biogeography of this fam- ily, the discovery of this species further in- dicates the great morphological diversity that exists within this group, as well as the great need for more fieldwork. Prototheora cooperi Janse Prototheora cooperi Janse 1942: 69.—Da- vis 1996: 418. New records.—SOUTH AFRICA: Cape Province: Du Toits Kloof, 1 ¢, 1 2, 6 Apr 1989, H. Geertsema, slide USNM 32309 (USEC, USNM). Stellenbosch, Jonkes- shoek FR: 1 odo, H. Geertsema, (USEC). Voélvlei Dam: Gouda: 2 6, 23 May 1987, H. Geertsema, slide USNM 31933 (USEC, USNM). Prototheora corvifera (Meyrick) (Figs. 5—6) Metatheora corvifera Meyrick 1920: 315.—Philpott 1926: 726; 1928: 94.— Janse 1942: 74.—Vari 1958: 75. Prototheora corvifera (Meyrick): Davis, 1996: 425. Because the female of this species was previously unknown, a description of that sex can now be provided. Female.—Wing expanse 19-21 mm. Generally paler, more gray in color than male. Head with vestiture of moderately broad to slender gray scales tipped with white. Antenna 28—29-segmented, ~ % the length of forewing. Labial palpus porrect as in male, similar in color to head; scales rough, slender at base of palpus, becoming more piliform apically. Thorax similar to head in color. Forewings mostly denuded in all four females examined, but generally pale gray with densely scattered dark gray scales. Hindwing pale gray, nearly white. Abdomen pale grayish white. Genitalia (Figs. 5—6): Dorsal plate with a broad, shallow median notch. Sternum IX with lateral lobes well developed, ~ half the length of conjugal process; apex broad- ly rounded, subtruncate (Fig. 5). Median conjugal process broadly triangular; length PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON about 0.75 its basal width. Antrum moder- ately enlarged, membranous. New records.—SOUTH AFRICA: Cape Province: Cape Town, Table Mountain: 2 6, 25 dan, 19952 sed Gr beb 1995-"Gadees o» 7 Heb. 1995, slides “USNM: 31932 32310; 5 6, 11-12 Feb 1995, H. Geertsema (USEC, USNM); vic. Cape Town, Table Mountain, 1,000 m: 1 2, 2 Feb 1995, slide DRD 4060; 14 6, H. Geertsema, slide USNM 32025 (USEC, USNM). Remarks.—The female genitalia of this species is most similar to that of P. serru- ligera, particularly in the triangular form of the conjugal process, with that of corvifera being broader at the base. In serruligera the length of the process is approximately 2.5 x its basal width, compared to 0.75 in corvi- fera. The lateral lobes of serruligera are also more reduced and possess more acute apices. Prototheora parachlora (Meyrick) Metatheora parachlora Meyrick 1919: 229.—Janse 1942: 73.—Vari 1958: 75. Metatheora paraglossa Janse 1942: pl. 62, fig. 2 (misspelling). Prototheora parachlora (Meyrick): Davis, 1996: 411. New records.—SOUTH AFRICA: Cape Province: Tsitsikama State Forest, Goesa- bos, 3 6, 1-13 Mar 1983, Scoble and Kroon, slide USNM 32026 (TMP, USNM). Remarks.—Specimens from the Trans- vaal Museum previously unavailable for study included 3 males of this species col- lected in the Tsitsikama State Forest, a coastal, predominantly temperate, ever- green forest located in southeastern Cape Province. Prototheora parachlora was known only from the holotype, collected 27 January 1917 in the Karkloof Forest of Na- tal. Prototheora petrosema Meyrick Prototheora petrosema Meyrick 1917: 19; 1919: 229; 1920: 314.—Philpott 1928: 94.—Janse 1942: 67.—Vari 1958: 75.— Davis 1996: 416. VOLUME 103, NUMBER 2 455 Fig. 2—6. Genitalia. 2-4, Male P. malawiensis. 2, Ventral view. 3, Lateral view. 4, Lateral detail of pseudo- teguminal arm in Fig. 3. CPo, conjugal pouch; PA, pseudoteguminal arm; PP, pseudoteguminal plate; S, sus- pensorium; SL, saccular lobe of valva; Tr, trulleum; TX, tergum of tenth abdominal segment; V, vinculum; VG, ventral groove of trulleum. Scale = 0.5 mm. 5—6, Female, P. corvifera. 5, Lateral view. 6, Caudal view. CPr, copulatory process; DP, dorsal plate (tergum IX + X); LL, lateral lobe; SP, subanal plate; Ve, vesicle of ductus spermathecae. Scale = 0.5 mm. 456 Prototheora monoglossa Meyrick 1924: 80.—Davis 1996: 416. New records.—SOUTH AFRICA: Cape Province: Citrusdal, Agterland: 1 36, Mar 1995, H. Geertsema (USEC). Hermanus, 1 36,1 2, 12 Mar 1988, H. Geertsema, slide USNM 31934 (USEC, USNM). Stellen- bosch: 1 ¢d, 9 Apr 1996, 2 3, 18 Apr 1991, 1 3d, 10 May 1994, H. Geertsema (USEC); 1 3, Feb 1955, H. Rossouw (USEC). Stel- lenbosch, Jonkershoek FR: 5 3, 20 Apr 1990, slide USNM 32024 (USEC, USNM); 2 6, 20 Apr 1993 (USEC); 1 2, 20 Apr 1993, died 26 Apr 1993, coll. at light, laid eggs 21-26 Apr 1993, ref. egg 001; 1 @, 20 Apr 1993, died 27 Apr 1993, coll. at light, laid eggs 21-26 Apr 1993, ref. egg 002, H. Geertsema (USEC). Prototheora quadricornis Meyrick Prototheora quadricornis Meyrick 1920: 315.—Janse 1942: 72.—Vari 1958: 75.—Davis 1996: 430. New record.—SOUTH AFRICA: Cape Province: Citrusdal, Agterland: 1 6, Mar 1995, H. Geertsema, slide DRD 4115 (USEC). ACKNOWLEDGMENTS I thank Wolfram Mey, Museum fiir Na- turkunde, Humboldt-Universitat, Berlin, Germany, for the generous loan of Proth- eora malawiensis and also for bringing this insect to my attention. I also thank Henk Geertsema, University of Stellenbosch, Stellenbosch, South Africa and Martin Kriiger, Transvaal Museum, Pretoria, South Africa for loans of specimens under their care. As always, I am indebted to Vichai Malikul, Department of Entomology, Smithsonian Institution, for the artwork and John Steiner of the Smithsonian Photo- graphic Services for the photograph. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Finally I acknowledge the cooperation of the institutions listed below and list their acronyms as used in this study. MNHU Museum fiir Naturkunde, Hum- boldt-Universitat, Berlin, Germa- ny. TMP Transvaal Museum, Pretoria, South Africa. USEC University of Stellenbosch, En- tomological Collections, Stellen- bosch, South Africa. USNM_ Collections of the former United States National Museum, now de- posited in the National Museum of Natural History, Smithsonian Institution, Washington D.C., USA. LITERATURE CITED Davis, D. R. 1966. A revision of the South African family Prototheoridae (Lepidoptera: Hepialoidea). Entomologica scandinavica 27: 393-439, 122 figs. . In press. Prototheoridae, Fasicle 11. In Heppner, J. B., ed. Lepidopterorum Catalogus. Janse, A. J. T. 1942. The moths of South Africa AC): 1=xxv, 1—=78: Meyrick, E. 1917. Descriptions of South African Mi- crolepidoptera. Annals of the South African Mu- seum 17: 1-21. . 1919. Exotic Microlepidoptera 2(8—9): 225— 288. . 1920. Descriptions of South African Micro- lepidoptera. Annals of the South African Museum 17: 273-318. . 1924. Exotic Microlepidoptera 3(3—4): 65— 128. Philpott, A. 1926. The maxillae in the Lepidoptera. Transactions of the New Zealand Institute 57: 721-746. . 1928. On the systematic position of Anomoses (Lepidoptera Homoneura). Transactions of the En- tomological Society of London 76: 93-96. Vari, L. 1958. Prototheoridae, p. 75. Jn A list of zoo- logical and botanical types preserved in collec- tions in southern and east Africa 1(2): i-vu, 1— 119. Pretoria. PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 457-466 BOoK REVIEW Fruit Flies (Tephritidae): Phylogeny and Evolution of Behavior. Martin Aluja and Allen L. Norrbom, editors. 1999. CRC Press, Boca Raton, FL. ISBN 0-8493- 1275-2. [xvii +] 944 pp. + 4 color plates and 1 p. of legends. Hard bound. $149.95. The editors of this magnificent and mas- sive volume set forth an ambitious goal of providing a cladistic analysis of relation- ships within this large family and with its near relatives and of using this framework to understand the evolution of basic behav- iors. The book arose as planned from a symposium held in Xalapa, Mexico during February 1998 and attended by some 40 participants. The symposium itself was am- bitious, as the editors invited and attracted most of the world’s top talent in the field for 5 intense days sequestered at the Insti- tuto Ecologia, A. C. Numerous books have been devoted entirely to detailed treatments of individual fly families, but I am aware of none that has attempted so explicitly to combine knowledge of phylogenetic rela- tionships with understanding the evolution of a broad array of behaviors. The family Tephritidae is well-chosen in this regard. It is large with over 4,200 described species, worldwide, diverse in biology and ecology, and includes many species whose biologies and behaviors are very well known because of their considerable economic importance. One of the primary aims of the editors was to provide new information, and not simply a summary of published literature in the field. Thus, for example, most of the tax- onomy chapters undertook a first attempt at formal cladistic analysis of a given group. The editors had high expectations from the start, and the resulting book exceeded those expectations. I can see why. The pool of contributing authors has experience exceed- ing half a millennium in career years of working with tephritid flies. The editors did a fine job of coordinating chapters and channeling the expertise of individual au- thors into a coherent whole. Manuscripts were obviously shared before the book was finalized, thus various chapters draw upon the ideas presented in others. This volume was made possible to a large extent by the culmination of another multi-author, multi- year endeavor that produced the Fruit Fly Expert Identification System and System- atic Information Database (Thompson 1998). It is quite unlike some other prede- cessor books on Tephritidae, such as those resulting from the quadrennial International Symposia on Fruit flies of Economic Im- portance (e.g., McPheron and Steck 1996), which, although organized into general top- ic areas, were largely sets of uncoordinated Proceedings papers. It is also very unlike Robinson & Hooper (1989) with its empha- sis On crop pests, or the very practical, working-man’s White and Elson-Harris i992): The book includes 33 chapters, written by 39 authors, arranged in eight sections (I— VIII). Phylogeny chapters more or less fol- low a common organizational thread that includes discussion of characters, ground- plans, diagnoses of included taxa, and phy- logeny based on cladistic analysis. An as- sortment of species and genera are newly described here. Similarly, behavior chapters include information on larval behavior, host associations, and adult behavior categories such as feeding, dispersal, mating and male-male interactions, oviposition, use of pheromones, and numerous subcategories of these. Each chapter concludes with a set of recommendations for future research di- rections and its own set of literature cita- tions, most very extensive. The following brief comments are intended simply to give readers a feeling for the vast amount of ter- ritory covered in this book and a few of its major conclusions. 458 I. General framework (3 chapters): Korneyev attempts to place the Tephriti- dae within the context of other families of the Tephritoidea and to relate the subfami- lies and tribes of Tephritidae among them- selves. He revisits the character analyses of Hennig, Griffiths, and McAlpine and adds new ones. Two hypotheses that he supports are that Pyrgotidae is the sister group of the Tephritidae, and the currently recognized family Tachiniscidae is actually a subfamily of Tephritidae. Sivinski follows with a sum- mary of breeding habits and sex in eight other tephritoid families. Among the aca- lyptrate flies this superfamily has an unusu- al concentration of elaborate male orna- ments and mating behaviors. Most mem- bers of these families lack a tight relation- ship with one or more host plants as exhibited by the Tephritidae, thus resource relationships are very different. He discuss- es how this may relate to mating systems (e.g., swarming, lekking) and sexual selec- tion. Diaz-Fleisher & Aluja provide a his- torical perspective of 100 years of tephritid behavior studies. Early period studies were primarily observational and descriptive, whereas contemporary work (1970s to the present), heavily influenced by Prokopy, has tended to be much more experimental. Much current research is applied to prob- lems of eradication and control of pest spe- cies using the sterile insect technique. The authors sort behaviors into various major categories, then summarize the develop- ment of theory and a great deal of literature. II. Higher relationships of Tephritidae (2 chapters) Korneyev discusses 111 characters, their states and polarities for analysis of relation- ships among tephritid groups. He recogniz- es six subfamilies, three of them “‘lower Te- phritidae”’ (Tachiniscinae, Blepharoneuri- nae, Phytalmiinae) and three “‘higher”’ (Trypetinae, Dacinae, Tephritinae), and concludes that subfamilies and tribes, as currently defined, appear to be well-sup- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ported monophyletic groups, with the likely exception of Trypetinae. However, the re- lationships among subfamilies and tribes is far from resolved. He freely admits to low confidence in some of the cladistic results due to large gaps in data on genital and lar- val morphology, homoplasy, and computa- tional limitations. Han and McPheron use mitochondrial (mt) ribosomal DNA to test relationships within the family. They pro- vide a basic but helpful explanation of mt DNA characteristics and their phylogenetic constraints, plus tree building methods and Statistical tests for sequence data. Their phylogeny is generally congruent with that based on current analysis of morphological characters, and additionally suggests sev- eral previously unconsidered placements for genera within tribes. III. Subfamilies Blepharoneurinae and Phytalmiinae (3 chapters) These old lineages within the Tephritidae are particularly interesting and challenging groups, as they are relatively rare with re- stricted distributions, often off the beaten path. The five genera of Blepharoneurinae (treated by Norrbom and Condon) are small (one is newly described from a specimen in Dominican amber) and far-flung, each ba- sically restricted to a single tropical region. Blepharoneura, however, may rival Anas- trepha in number of species (200+) once it is fully known. Those few with known bi- ologies are unusual for the family. Unique adult feeding, larvae dispersed by bats, flies restricted to one or the other sex of cucurbit hosts, and very fine niche partitioning among species of Blepharoneura are all highly intriguing. Similarly, the Phytalmi- inae, with scores of mono- or nearly mono- typic genera, are primarily Australasian or Oriental in distribution. Dodson describes the evolution of antler-like facial projec- tions in the Phytalmiinae (first noted by Al- fred Russell Wallace as ‘‘deerflies’’). All antlered species studied to date exhibit a re- source defense mating system. Antlers may be spectacular in color and size; they occur VOLUME 103, NUMBER 2 on males only; they’re used in physical con- tests to control female oviposition sites; and, as antler size is tightly correlated with body size, they probably serve to signal likelihood of success in territorial defense. IV. Subfamily Trypetinae (7 chapters) Roughly 1,000 species belong to this subfamily. Because of the economic impor- tance of some of its members, deep “‘ver- tical’’ studies have been conducted here, that is, in-depth research into many aspects of taxonomy, ecology, behavior, physiolo- gy, etc. of a select few genera and species. These include, for example, Rhagoletis, with notable pests such as the apple mag- got. This genus has been subject to more intensive phylogenetic work than any other single group of tephritid flies, and the les- sons learned here are particularly instruc- tive to studies throughout the remainder of the family. Smith & Bush discuss phylog- eny of Rhagoletis and nine related genera. The authors emphasize biogeography more than authors of other chapters, as these taxa are widespread (Holarctic, Neotropical, and Oriental), and different mechanisms (sym- patric vs. allopatric) seem to have driven speciation within different groups. The in- terplay between analyses based on morpho- logical features and those based on mole- cules show that divergences in both mor- phology and sequences probably occur at different rates in different lineages, possibly due to action of different processes such as natural selection and genetic drift. Despite the considerable effort devoted to Rhago- letis to date, most of it has been directed at Nearctic taxa, leaving many aspects of its phylogeny on a global basis unresolved. As taxa from other regions are being more ful- ly studied, it does indeed appear that Rha- goletis is not the monophyletic group it was once thought to be. Prokopy and Papaj rich- ly describe and discuss what is known of behavior in this group. Much of the discus- sion is in terms of foraging, that is, the el- ements involved in searching for and ac- quiring food, mates, and oviposition sites, 459 and how these may be tempered by numer- ous internal and external factors, such as voltinism, breadth of host range, dispersal and learning ability, competitors and pred- ators. A great deal is known about many of these factors among Rhagoletis species, much of it through direct and focused ex- perimentation. Unlike members of other major fruit fly pest genera, Rhagoletis spe- cies do not engage in lekking behavior or complex courtship signaling. Host marking pheromones (HMP), and their associated behaviors and evolutionary ecologies are especially well known among Rhagoletis species. The authors touch on the potential importance of fly chemosensory capabili- ties with respect to plant volatile profiles in determining likelihood of host plant shifts. Lastly, the authors expound upon the im- portance of knowing and testing the proxi- mal conditions (e.g., environmental, physi- ological, and informational states) that elicit various behaviors; otherwise, a behavior may be phylogenetically miscoded as ab- sent even though a genetically-based capa- bility is present but goes unobserved due to lack of necessary proximal stimuli. This is a very well-written chapter that explicitly addresses behavioral data and phylogeny of Rhagoletis. Han’s chapter on phylogeny of flies in the tribe Trypetini is a major contribution to the internal classification of Tephritidae. Prior to his study, this tribe was a dumping ground for numerous genera that could not easily be placed into other groups based on good derived characters. Han initially pos- tulated that leaf-mining evolved only once in the Tephritidae, and so chose for study the 20 known leaf-mining species, their pre- sumed congeners, and presumed related genera. In all, 250 species were examined and reclassified. Indeed, all of the leaf-min- ing tephritids are included in a single sub- tribe, the Trypetina, where they occur across several genera, which also include non-leaf-miners. Norrbom, Zucchi and Hernandez-Ortiz provide a phylogenetic analysis of the gen- 460 era Anastrepha, Toxotrypana, and Hexa- chaeta, Neotropical groups that include im- portant fruit pests. Most of the approxi- mately 200 described Anastrepha species and 13+ Toxotrypana species can be con- fidently placed into one of 18 species groups, but very little is yet possible in de- termining deeper relationships among them. Readers may be surprised to learn that the plesiomorphic state of host feeding in An- astrepha may be seed-feeding rather than pulp-feeding, contrary to our usual concep- tion of tephritids as “‘fruit fly’’ pests. In a sister chapter, McPheron, Han, Silva and Norrbom use 16S rRNA mitochondrial DNA sequences to generate possible phy- logenies. This analysis corroborates some major groupings of the preceding chapter based on morphology, including monophy- ly of Anastrepha+ Toxotrypana, and con- tradicts other groupings. In general, se- quence data are far more productive than morphology in suggesting relationships within and among members of the various proposed species groups. Landolt describes in detail the behavior of 7. curvicauda, the papaya fruit fly, physically as spectacular a fly as are the antlered phytalmiines. Direct comparison of its behaviors with those of Anastrepha species is provided in a chapter by Aluja, Pinero, Jacome, Diaz-Fleischer and Sivinski. Host marking could be a de- rived behavior in Anastrepha, as it is lack- ing in some of the putative basal groups and T. curvicauda; perhaps it derives from post- Oviposition aculeus cleaning behavior by females attacking latex-producing fruits. Resource defense has been reported for males of 7. curvicauda and one Anastrepha species, but otherwise lek polygyny pre- dominates as a mating strategy in Anastre- pha. The daily pattern of male “calling” rhythms appears to be one of the most hard- wired behaviors in Anastrepha. These seem to be controlled proximally by light inten- sity, and may ultimately derive from selec- tion pressure to exclude interaction with sympatric Anastrepha species. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON V. Subfamily Dacinae (6 chapters) This is another very large and economi- cally important group of approximately 1,000 species. De Meyer addresses phylog- eny of the genus Ceratitis, which includes Mediterranean fruit fly (medfly), an infa- mous agricultural scourge, among its 78 species. De Meyer makes the first attempt at cladistic analysis of the genus, first using 17 characters from which earlier authors postulated the current 6 subgenera; this analysis failed to support most of the orig- inal groupings. De Meyer adds 19 charac- ters and re-analyzes the mix. So far, only a single subgenus, Pterandrus, is supported as monophyletic. Host plant relationships and lure responses, once known for more species, likely will be useful characters to add to the analysis. Yuval and Hendrichs examine the behavior of flies in the genus Ceratitis, for which substantial information is available for the medfly, quite limited in- formation for the Natal fruit fly and none other than host plants for other species. Among tephritids, lekking behavior has been studied in most detail for medfly; par- ticipation in leks is not a strict requirement for male mating success. Medfly’s great plasticity in oviposition behavior likely is an important factor in its high polyphagy and colonization ability. Eberhard provides further intimate details on medfly sexual behavior, specifically on what determines the success or failure of a given courtship. He discusses the possible avenues of sexual selection including cryptic, post-intromis- sion, female choice of different males with whom they have mated. Some of the details and ramifications of such “‘internal behav- ior’ are stunning. Despite considerable re- search effort, our understanding of many aspects of the chemical ecology of lures and pheromones acting between and within the sexes remains elusive. Drew and Hancock use morphological, distributional, and biological (especially host associations and male lure response) data to suggest a phylogeny of the tribe Da- VOLUME 103, NUMBER 2 cini (=subfamily Dacinae, by some reck- onings). Though only four genera are in- cluded, this is a particularly difficult group because of the large number of species in- volved, nearly 700 in all, and convergences due to reduced chaetotaxy and wasp mim- icry. All but three species fall into two gen- era, Bactrocera, a rain forest-adapted group centered in southeast Asia, Australia, and the South Pacific, and Dacus centered in and more adapted to savannah habitats of the Afrotropics. Any group of this size in- vites subdivision, and the authors recognize 30 subgenera of Bactrocera, which they cluster into four sets. The authors eschew a cladistic analysis of the tribe, owing to the limited knowledge of the complicated Af- rotropical fauna. Instead, they anecdotally walk us through a postulated reconstruction of the evolution of the tribe and its geo- graphical dispersion in an old-school man- ner. Thus, the reader is spared Henning&86 and its nitty-gritty of weightings, additivi- ties, indices, and overflows; and the authors are spared a rigorous defense of their many subgenera of Bactrocera. White provides a model approach to the specific aims of the book in attempting to correlate evolution of form, function, and behavior in the Dacini. He elaborates on male pheromone/lure re- sponse, acoustics, wasp mimicry, microtri- chia patterns, aculeus length and possible resource partitioning, and aculeus tip mor- phology. As noted in other chapters, the pattern of speciation, i.e. sympatric vs. al- lopatric, may be important in differentially driving evolution of characters and behav- iors. White picks up where Drew and Han- cock left off, and provides a formal cladis- tic analysis of 51 pest species. The data and taxa are a subset of the very large number of both species and characters included in White and Hancock’s (1997) expert system. White questions the validity of the large number of Bactrocera and Dacus subgen- era, based as they are on nearly all possible permutations of a few binary characters; rather, he would reduce the number of ma- jor monophyletic groups to about five in 461 Dacus and three to five in Bactrocera. Drew and Romig provide further commen- tary on the biology and behavior of Dacini. They elaborate on the concept of host plant as a “‘center of activity for a species pop- ulation,’ as hosts are so tightly tied to all aspects of the flies’ biology, including courtship and mating, adult and larval nu- trition, dispersal, etc. Bacteria seem to play a larger role in the biology of dacines than in other tephritid groups, and may mediate many host-fly interactions by serving as nu- trients for adult females and possibly lar- vae, olfactory cues to attract flies to hosts, and defense against pathogens. The authors usefully note a number of conflicting inter- pretations of behavior, biology, and mor- phology, for example, the relevance of methyl eugenol in nature and the signifi- cance of aculeus tip shapes. VI. Subfamily Tephritinae (4 chapters) Korneyev analyzes relationships among tribes and subtribes of the Tephritinae, the “most specialized subfamily” of Tephriti- dae. Nearly half of the family belongs here, some 1800+ species that primarily infest seed heads of Asteraceae. A mosaic distri- bution of characters makes it difficult to de- fine the subfamily and analyze relationships within it. He presents a groundplan of 30 characters for this apparently monophyletic subfamily. With various rearrangements and reassignments from earlier classifica- tions, he splits the subfamily into eight tribes. An aggregation of “‘lower’’ tribes (e.g., Terelliini) retaining many primitive characters may be paraphyletic, while the *‘higher tephritines”’ (e.g., Tephritini) seem clearly monophyletic. Korneyev provides several independent cladistic analyses of various groups within the subfamily. Some groups are clearly monophyletic, but sister group relationships are uncertain. Homopla- sy is very high, thus he concludes that even a complete data set would not guarantee clearly resolved relationships. All in all, there is a great deal to digest in this chapter, one of the most esoteric in the book. Freid- 462 berg and Norrbom provide a generic re- classification and phylogeny of the tribe Myopitini, one of Korneyev’s lower tephri- tine groups. Here are included 130 recog- nized plus 60 undescribed species. The ge- nus Urophora, important to biological con- trol of weeds, is the tribe’s best known member. The authors removed several gen- era and species previously classified as my- opitines to make a monophyletic group of 11 genera (3 newly described, and 3 ele- vated from subgenus). Relationships among genera are tentatively proposed, as various manipulations of Henning86 suggested nu- merous possibilities. The authors make an interesting comment on their modus oper- andi in attempting this higher level classi- fication using a small number of select taxa, especially the type species of included gen- era: they claim that doing so before the gen- era themselves were revised, forced them to add more and more species, thus “ending up with partial revisions of most genera, which was not [their] original intention.” Merz presents a phylogeny of Palearctic and Afrotropical genera of the Tephritis group (Tephritini), a group of high diversity in the highlands of East Africa. He attempts to determine whether the genera included by Munro and other authors in several poorly defined “series”? or “‘“complexes”’ form a monophyletic group. He had diffi- culty even in deciding an outgroup. His cla- distic analysis of 20 characters and 29 spe- cies leads to a new classification, in which he names five new genera for a total of 20. Given the regional nature of the analysis, this classification must be considered pro- visional. Headrick and Goeden discuss the behav- ior of flies in the subfamily Tephritinae. Their work has been seminal in generating a glossary of terms to standardize behav- ioral descriptions of courtship and a data- base to develop evolutionary and ecological hypotheses on mating behavior in tephritid flies. They observed the mating behavior of 48 species from California under lab con- ditions. They treat 15 genera, providing for PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON each a description of behaviors and com- parisons among its species. This chapter significantly expands the phylogenetic con- text of their earlier papers. Important con- clusions are that life history strategies are major factors in predicting the types of re- productive behaviors a species will exhibit, and even population densities may be im- portant. Life history strategies in turn are critically dependent on host plants and their phenologies. The authors suggest caution in generalizing globally from their behavior studies in southern California, as even these have continuously provided new surprises and exceptions to most rules they have at- tempted to make. VII. Evolution of Behavior (7 chapters) In this section, authors synthesize infor- mation and ideas from previous chapters into a broad view of the evolution of te- phritid behavior. It opens with a chapter by Berlocher on genetic population structure in Tephritidae. Demarcating the interface be- tween populations and species has long been a problem in the Tephritidae, which is rife with “‘sibling species,” “‘cryptic spe- cies,” ““species complexes,”’ “host races,”’ etc. Here is presented a clear, basic review of general concepts, case studies from the tephritid literature, and pitfalls in interpret- ing population genetic data. Tephritid behaviors are subsumed into three major categories—feeding, oviposi- tion, and sexual—for discussion of evolu- tionary patterns. The editors thus manage to keep the three chapters devoted to these behaviors largely non-overlapping; but they create a problem conceptually, because, from an evolutionary perspective, this is an arbitrary division of life’s activities. For ex- ample, Drew and Yuval’s basic assumption in discussing feeding behavior “‘is that the key to the biological success of the Tephri- tidae was the evolution of phytophagy, and that this was made possible primarily due to morphological and behavioral adapta- tions in the female, namely, ovipositors and specific oviposition preferences’’—clearly a VOLUME 103, NUMBER 2 topic for the oviposition chapter as well. The feeding chapter does provide a good summary of what is known of larval and adult nutritional requirements, their feeding mechanisms, resource partitioning, bacterial associations, and effects on reproductive success. The authors speculate on the evo- lutionary ecology of feeding behavior and make some very broad statements relating these to subfamilies, morphology, and to habitats (e.g., rain forest vs. savannah). In discussing the evolution of oviposition be- havior, Diaz-Fleischer, Papaj, Prokopy, No- rrbom, and Aluja take the view that host specialization is the driving force behind speciation and ultimately the rich pattern of diversification observed in the family, but key innovations in ovipositor morphology, Oviposition behavior, and patterns of bac- terial transfer set the stage for specializa- tion. Thus, ecology (opportunism) and mor- phology (key innovation) set the stage, then behavioral changes (host finding and pref- erence) implement the transition. Interest- ingly, they find ‘“‘little correlation between hosts preferred for oviposition and hosts that best support larval growth and devel- opment.’ Also, “‘it is commonly believed that specialists arise from generalist ances- tors and not the other way around. . ..’’, but this may not be true for tephritids, where host ranges tend to expand, not retract, at least in historical times (e.g., R. pomonella, A. ludens); however, dacine generalists may be better preadapted to shift to novel hosts than are dacine specialists. Oviposition be- havior also commonly involves marking the host with pheromones (HMP) or plant prod- ucts, another example of opportunism, in which gut products or oviposition exudates have been commandeered as signals. Hy- pothetical relationships among HMPs, host plant characteristics, adult dispersal and life history traits, and levels of larval competi- tion are explored here. One might expect that levels of cross-recognition of HMPs among species would reflect phylogeny; this seems to be true among Rhagoletis, but not true among Anastrepha. Is this because 463 Rhagoletis, being steno/monophagous, are not much subject to interspecific competi- tion, while the polyphagous Anastrepha are? Who knows?! Numerous topics in the evolution of sex- ual behavior are covered in a multi-au- thored chapter. Sivinski summarizes some theoretical speculations relating resource distribution (also host range and temperate/ tropical habitats) to mating systems. Expla- nations for some differences among tephri- tid groups that made sense at one time, have become less tenable as new observations have filled in the gaps. However, a ‘‘core principle ... that the abundance of hosts relative to the number of female fruit flies influences male distribution, remains an at- tractive vehicle for the exploration of te- phritid mating systems.’ Aluja suggests that the timing of many sexual activities is likely determined by the presence of related species, and that these times diverge to pre- vent interference and hybridization. Dodson applies some predictions from theory to ag- onistic behavior among tephritid flies as displayed by males vs. females, for defen- sible- vs. non-defensible resources (*‘scram- ble competition’’), and to the degree of physical interaction between contestants. Predation risks, as discussed by Landolt, may constrain or funnel the evolution of numerous aspects of behavior and mor- phology, as anything that attracts attention, whether color, movement, or odors, may in- vite not only mates but also predators. Freidberg describes trophallactic behavior, in which males provide females with nup- tial gifts to consume. At least 20 species of tephritids engage in this behavior; wonder- ful details are provided for Schistopterum moebiusi. Headrick presents a behavioral character matrix for 82 species of tephritids vs. their life history strategies, wing dis- plays, trophallaxis, leg and mouthpart movements, male stalking and guarding, multiple mating, etc. Then he discusses mating behavior and the reconstruction of phylogeny, and neatly lays out the promise (rich data set) and the problems (homopla- 464 sy, inconsistency among observers, ecolog- ical context) of using behavior as characters for phylogenetic analysis. Lastly, Kaneshiro discusses theoretical and genetic aspects of sexual selection, which he sees as playing an extremely important role in the initial stages of species formation. Other intriguing and potentially infor- mative characters to analyze relationships are biochemical pathways to and chemical structures of pheromones. A chapter by Heath, Landolt, Robacker, Dueben and Ep- sky explores this possibility. Some compar- isons are provided for Anastrepha species, but these are not concordant with morpho- logical relationships. Unfortunately, the au- thors do not compare Toxotrypana to An- astrepha even though the pheromones of the former are known. Bactrocera oleae is curious in that females produce an attractant for males, while the roles of the sexes are reversed among its congeners. “‘Although there are very minor overlaps in pheromone chemistries of fruit flies in different genera, they are quite different overall with no in- dications yet of any similarities among taxa above the species level.’’ Chemical phylog- enies may be complicated by possible re- lationships between fruit fly pheromones and plant volatiles and even with stinging Hymenoptera. A chapter by Cayol on changes in sexual behavior and life history traits of fruit flies under mass-rearing shows just how geneti- cally labile these traits are, and how quickly profound differences can arise. This cer- tainly suggests that divergence may swiftly result from new ecological opportunities, e.g., as afforded by introduction of flies into new areas, or of novel hosts into existing fly ranges. Kaneshiro discusses sexual selection and speciation in Hawaiian Drosophila, as a model system for research in tephritids. Surely, the genetics, behavior, and evolu- tion of no other family of flies is better known than that of the Drosophilidae, es- pecially the large subset that exists in Ha- waii. Like the tephritids, breeding of the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Hawaiian Drosophila is closely tied to spe- cific plants, and they show a great diversity of morphology, behavior and life histories. Kaneshiro argues for the greater importance of sexual selection than natural selection in driving speciation among Hawaiian Dro- sophila. Much of the chapter presents the- oretical arguments to explain mating asym- metries, mechanisms for maintaining genet- ic and behavioral diversity, and the possi- bility of natural hybridization (and havoc in phylogeny estimation) among Drosophila species. He applies principles of sexual se- lection to the mating system of medfly. His observations suggest that most natural mat- ings are between highly fit males and less choosy females, which perhaps generates a balanced polymorphism of mating types in the population; this in turn, may account for medfly’s great ability to survive population bottlenecks and colonize new areas. Other observations on the importance of oviposi- tional behavior over larval nutritional re- quirements in explaining host specificity, the derivation of generalist breeding species from specialist species, and lekking behav- ior among Drosophila species, will also in- terest tephritid biologists. VIII. Glossary A 45-page glossary by White, Headrick, Norrbom, and Carroll has greatly expanded upon White and Elson-Harris (1992) by ad- dition of many behavioral, ecological, and nomenclatural terms. It includes some de- partures from the terminology used in the Manual of Nearctic Diptera (McAlpine 19S 15): Whew—dquite a book, unlike anything else in the Diptera literature! The one other family for which a similar treatment of phy- logeny and evolution of behavior would be possible is the Drosophilidae, but a com- parable book does not exist. Our knowledge of tephritid physical and behavioral evolution is a work in active progress, and this book will be shaping its future. Students of the family will be mull- ing over and digesting these chapters for VOLUME 103, NUMBER 2 decades to come. The framework laid here by the phylogenetic analyses will likely lead to rapid progress in clarifying evolu- tionary relationships. Despite data gaps and homoplasy problems, these authors have al- ready shown great success in identifying phylogenetically informative characters and in newly defining many monophyletic groups. However, at varying levels, each of the cladistic analyses presented here was unable to identify sister groups, and further improvements in classification based on morphology may be incremental at best. Molecular data will be needed on many lev- els to suggest viable and novel relationships that might not otherwise be discerned. Despite the very esoteric and mind- numbing nature of cladistic character anal- yses, weightings, tree-building methods, etc., I found the taxonomy chapters more satisfying than the behavior chapters (not that the latter weren’t extremely fascinat- ing). That is, the former were able to come to some sort of conclusion about histories and relationships among taxa, along with some measure of confidence in them. That’s because taxonomists can exercise a reduc- tionist approach that delimits characters and states, whereas behaviorists ..., well, it’s not so easy. I think we will see the quickest progress in understanding the origin, func- tion, and limitations of those “‘behaviors”’ that are perhaps more aptly considered physiological traits, such as pheromone and lure responses and host plant associations. These can be considered well-defined char- acters with relatively invariant states within species. Indeed, these are considered so fundamental to our understanding of phy- logeny of fruit flies, that several of the tax- onomy chapters explicitly include these data in their cladistic analyses. Many yet undone studies would be very easy to ac- complish; for example, only a few hours of observation of HMP behavior of common flies such as Zonosemata and Carpomya could add valuable information to relate to so much that is known about their near and well-studied relative, Rhagoletis. Other 465 studies at the interface of physiology and behavior will be more technical, but they are especially amenable to experimentation and likely to be productive of answers in understanding evolution of behavior. Key among all the factors in tephritid evolution are host plant associations and their shifts in space and time. A whole cascade of events, both morphological and behavioral, accompany these shifts. What sensory physiology is involved in host plant finding and acceptance for oviposition? Similarly, there are important synergies between pher- omone chemistry and behavior. These must be involved in the evolution of whole suites of behavior, such as mating, oviposition site selection, and dispersal behavior. What is the sensory basis for pheromone reception and the genetic basis for pheromone pro- duction? What, if any, are the relationships between pheromones and the paraphero- mones that are so important to fruit fly pest detection systems? For many other behaviors we will do well simply to discover proximal causes. If we had perfect knowledge of tephritid phy- logeny, would we then also have an expla- nation for the evolution of these behaviors? Their myriad, fluid, and intricate variations are difficult enough to describe, much less explain. Sure, evolutionary ecologists and behaviorists have many clever ideas to ex- plain why things are as they are and why they should be just so. But it is not produc- tive to assign an adaptive function to ev- erything that we see. Even though we pre- sume that all behaviors work in concert to some end, that is, to achieving reproductive success, this ignores the fact that many if not most individuals in most populations are failures in this regard. So, which out of all the behaviors we observe (not to men- tion the ones we don’t observe) are actually the ‘‘chosen”’ ones? Possibly we will never even correctly guess at the origins of many features, both the ordinary and the bizarre. Some of the tracks to follow are clear: more “horizontal” studies, standardization of terminology and techniques, hypothesis 466 design and testing, development of better classifications of life history and mating strategies, and an experimental mind-set. Anyone who has ever collected and mar- veled at these beautiful insects, or watched some display of their behavior and won- dered what motivated it, will find cause for reflection by reading this book, and hope- fully be inspired to return to the scene to more fully explore and attempt to relate it to what he has read here. This is contribution number 898, Bureau of Entomology, Nematology and Plant Pa- thology-Entomology Section. LITERATURE CITED McAlpine, J. E 1981. Morphology and terminology— adults, pp. 9-63. In McAlpine, J. FE, B. V. Peter- son, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, and D. M. Wood, coords. Manual of Nearctic Dip- tera Vol. 1. Monograph of the Biosystematics Re- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON search Institute, No. 27. Agriculture Canada, Ot- tawa. 674 pp. McPheron, B. A. and G. J. Steck. 1996. Fruit fly pests: A world assessment of their biology and manage- ment. St. Lucie Press, Delray Beach. 586 pp. Robinson, A. S. and G. Hooper. 1989. Fruit flies. Their biology, natural enemies, and control. W. Helle, ed., World crop pests, Vol. 3(A), xii + 372 pp., and Vol. 3(B), xv + 447 pp. Elsevier, Amsterdam. Thompson, E C., ed. 1998. Fruit fly expert identifica- tion system and systematic information database, Myia Vol. 9, Backhuys Publishers, Leiden, viii + 524 pp. White, I. M. and D. L. Hancock. 1997. Indo-Austra- lasian Dacini fruit flies. (Computer Aided Biolog- ical Identification Key). International Institute of Entomology, London. Compact Disk. Gary J. Steck, Florida Department of Agriculture and Consumer Services, Divi- sion of Plant Industry, Gainesville FL 32614-7100, U.S.A. (e-mail: steckg@ doacs.state.fl.us). PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 467-468 REPORTS OF OFFICERS EDITOR Volume 102 of the Proceedings included four issues with a total of 1,100 pages. Eighty-five regular articles, seven notes, five book reviews, two obituaries, Society Meetings, Reports of Officers, Instructions for Authors, and the Table of Contents for Volume 101 were published. About 110 manuscripts were submitted for possible publication during the past year. Memoir 23, ‘“‘Revision of the Net- Winged Midges of the Genus Blepharicera Macquart (Diptera: Blephariceridae) of Eastern North America’? by Gregory W. Courtney will be published in December, 2000. The Proceedings is listed with the Sci- ence Citation Index. I extend thanks to members of the Pub- lications Committee, Wayne Mathis, Tom Henry, and Ray Gagné, Book Review Ed- itor, for their continued advice and support. Marie Blair and Cathy Anderson provided much needed assistance in handling corre- spondence, routing manuscripts, and prep- aration of manuscripts and plates for the printer. Without their help, my job would be much more difficult. I am also grateful to the many reviewers for their time-consuming efforts and con- structive reviews. Their contributions are essential to help increase the quality of pa- pers published in the Proceedings. Respectfully submitted, David R. Smith, Editor TREASURER SUMMARY FINANCIAL STATEMENT FOR 2000 Special General Publication Total Fund Fund Assets Assets: November 1, 1999 $ 2,505.93 $113,340.73 $115,846.66 Total Receipts for 2000 97,974.04 29,694.13 127,568.14 Total Disbursements for 2000 63,617.96 63,617.96 Assets: October 31, 2000 36,862.01 143,034.86 179,896.87 Net Changes in Funds $34,356.08 $ 29,694.13 $ 64,050.21 Audited by the Auditing Committee, December 5, 2000 consisting of Norman E. Woodley, Wayne N. Mathis, and Stephen D. Gaimari, Chairman. Presented to the membership at the meeting of December 14, 2000. Respectfully submitted, Michael G. Pogue, Treasurer 468 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MEMBERSHIP In 2000 the Society received applications for new membership for 23 people as follows: Michael Adams James B. Munro W. Evan Braswell Debra Murray Andrew Brower Tam Nguyen David Campbell Benjamin Normark Gabriela Chavarria Kevin O’Neill Netta Dorchin Daniel E. Perez-Gelabert Arthur V. Evans Gordon E Pratt Qiao Gexia Gale E. Ridge-O’ Connor David Houghton Maria Marta Schang Ken Karns Wayne Wehling James Kruse John D. Zyla Juan Carlos Mariluis Each applicant was sent a letter acknowledging receipt of his/her application, and his/ her name was read at a regular meeting of the Society and repeated in the minutes of the following meeting. The number of applications for new membership equalled those from 1999: Respectfully submitted, Steven W. Lingafelter Membership Chair VISIT THE ENTOMOLOGICAL SOCIETY OF WASHINGTON WEBSITE http://entomology.si.edu:591/ES W/ES W Menus.html1 PROC. ENTOMOL. SOC. WASH. 103(2), 2001, pp. 469-472 SOCIETY MEETINGS Note regarding meeting numbers: An ESW Annual Banquet was thought to not have been counted as a meeting, resulting in erroneus meeting numbers. This was cor- rected in October 1998 by skipping a meet- ing number. After checking ESW archives in preparation for our upcoming website, President David Furth reported at the No- vember 2000 meeting that the October ad- justment was not needed because the Jan- uary 1996 meeting had been cancelled due to inclement weather and associated gov- ernment closure. Consequently, the meet- ings from October 1998 through May 2000 were not, as reported and published, the 1032nd—1048th meetings, but rather the 103 1st—1047th. 1,048th Regular Meeting—June 8, 2000 The 1048th regular meeting, erroneously thought to be the 1049th meeting, of the Entomological Society of Washington (ESW) consisted of the Annual Banquet, held once again at the Uniformed Services University of Health Sciences in Bethesda, Maryland. Approximately 100 members and guests attended. The disorderly social half-hour was called to order by President David Furth at 7:30 p.m to start the meet- ing. After a delicious catered meal, John Brown introduced the invited speaker, Da- vid A. Grimaldi of the American Museum of Natural History, New York. He told us about his exotic travels in his talk entitled ‘“‘New Jersey Mud Pits and the Glories of Fossil Insect Work.” These mud pits hold among the oldest amber insect inclusions in the world, dating from the Upper Cretaceous, about 95 mil- lion years ago. The amber was apparently washed down ancient streams and deposited along the riverbanks, forming layers in the mud that are now easily accessible. The in- clusions are especially significant because of their great age, despite the small size and clouded nature of most amber pieces. Among its treasures are the oldest ants, the oldest mushrooms, and the oldest feather of a North American bird. The meeting was adjourned at 9:30 p.m. Respectfully submitted, Stuart H. McKamey, Recording Secretary 1,049th Regular Meeting—October 5, 2000 The 1049th regular meeting of the En- tomological Society of Washington (ESW) was called to order in the Cathy Kerby con- ference room (CE-340) of the National Mu- seum of Natural History, Washington, D.C., by President Dave Furth at 7:05 p.m. The meeting was attended by 15 members and 5 guests. Stu McKamey read the minutes of the previous two meetings, which were ap- proved with minor modification. Dave Smith, standing in for membership chair Steve Lingafelter, read the names of new applicants for membership: W. Evan Brazwell, Debra Murray, Lic. Maria Marta Schang, Dr. Juan Carlos Mariluis, James B. Munro, Michael Adams, and Andrew Brower. No new members were present. Dave Smith then gave the editor’s report: a new Memoir is coming out soon, by Greg Courtney on the net-winged midges of North America. The Society’s Proceedings are running on schedule. In miscellaneous business, Dave Furth discussed the planned ESW website. We will contract George Venable (Smithsonian Entomology Illustrator) and the site will be posted on the Smithsonian Entomology server. Documents on bylaws, ESW histo- ry, instructions for authors, etc. have been gathered. The site will include a member- ship list (name and city) and fillable fields for additional information if members so choose. For exhibits, Edd Barrows displayed 470 photos from the past few BugFests held on the Mall. Dave Furth had on hand both ab- stract volumes of the XXI International En- tomology Congress and displayed new books: Hawkmoths of the World; Handbook of Palearctic Macrolepidoptera, 1999, Vol. I: Sessiidae; Entomology Contributions in Memory of Byron A. Alexander (Byers et al., eds.); and World Catalogue of Insects, Vol. 2, Hydrophiloidea by Michael Hanson. Our new program chair Ted Schultz in- troduced the night’s speaker, Aurthur V. Evans, a Smithsonian Research Associate and co-author of the popular book “An In- ordinate Fondness for Beetles.”’ His pre- sentation was entitled “Spineless in Rich- mond: An Entomologist Explores Virgin- ia.” A recent immigrant to Virginia, Dr. Evans has discovered that the Virginian fauna is not well known. In the last ten months he has discovered 20 new state rec- ords of scarabs alone, most within walking distance of his home. He wowed the audi- ence with many photos, all of local arthro- pods, including crustaceans, spiders, and representatives of almost every insect order. Appropriately, considering their relative richness and his personal interest, about half the subjects were beetles, but the pho- tos were so nice that there were no com- plaints from the audience. The meeting was adjourned at 8:20 p.m. Refreshments were provided by Ted Schultz. Respectfully submitted, Stuart H. McKamey, Recording Secretary 1050th Regular Meeting—November 2, 2000 The 1050th regular meeting of the En- tomological Society of Washington (ESW) was called to order in the Waldo Schmidt room of the National Museum of Natural History, Washington, D.C., by President Dave Furth at 7:04 p.m. The meeting was attended by 24 members and 12 guests. The PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON minutes of the previous meeting were ap- proved as read. President Furth clarified the meeting numbers. This meeting is the 1050th rather than the 1051st, because the January 1997 meeting had been cancelled by inclement weather and government closure. Steve Lingafelter read the names of two new applicants for membership: Bejamin Normark and Gabriela Chavarria. One new member was present. Terry Erwin moved that the Executive Committee look into spe- cial family membership rates for ESW. The nominating committee, comprised of M. Alma Solis, Mike Pogue, and Marc Epstein, proposed a slate of officers to be voted on at the December meeting, during which ad- ditional candidates can be taken from the floor. Officers from the present year ac- cepted nominations to continue next year, with the exception of new President Elect candidate Gabriela Chavarria and new Pro- gram Chair candidate Ronald Ochoa. Four visitors, from England, Germany, and Scot- land, and one member from Israel were in- troduced. For exhibits and news, Art Evans an- nounced the death of BioQuip founder Rich Fall. Gabriella Chavarria had on hand the new book Bees of the World by Charles Michener. Terry Erwin called attention to the website www.all-species.org, a recent must-see site for all taxonomists and other biodiversitists. David Furth displayed a copy of the book Buzzwords, by May Ber- enbaum, a compilation of her humorous contributions to the ESA magazine Ameri- can Entomologist, and a Washington Post article on a USDA elaterid specialist who had been deaf until a recent operation, which was the subject of the article. Fellow coleopterist Terry Erwin remarked that now he understands how click beetles got their name. Ron Ochoa displayed a copy of the USDA Agricultural Research magazine spread on Cryo-SEM, a new technique he and colleagues are using to get closer and sharper images of mites than ever before possible, frozen in action. VOLUME 103, NUMBER 2 Program chair Ted Schultz reminded us that due to a conflict with the annual na- tional meeting of the Entomological Society of America, December’s meeting will be held on the 14th. He then introduced the evening’s speaker, Jonathan Mawdsley, Smithsonian Postdoctoral Research Fellow, who presented his research on “‘Biogeog- raphy of Checkered Beetles (Coleoptera: Cleridae) in the ‘Sky Islands’ of Southeast- ern Arizona.’’ There are about 4,000 de- scribed species of clerids in the world. Al- though most are predators of scolytids, a surprising variety of prey are becoming known as the life histories of more species are discovered. A hotspot for clerid species richness is Southeastern Arizona, USA, the northern extension of the Sierra Madre de Occidental of Mexico and where the moun- tains are separated by deserts. The check- ered beetle faunas are distinct across the habitats along the altitudinal gradients, from the deserts up to the grasslands, to the oak-juniper woodlands, and finally to the montane forests dominated by ponderosa pines. The woodland foothills hold the greatest number of clerid species and are home to most of the seven endemic clerids known from the region. Dr. Mawdsley has also been involved in an Arizona clerid da- tabase project that interfaces with Geo- graphical Information System software. By simultaneously mapping clerid species lo- calities, vegetation, and protected areas, he found that the oak-pine forest of the Santa Rita Mountains is the most important place to protect, from the perspective of Cleridae. Fortunately, that area is already protected. The meeting was adjourned at 7:53 p.m. Refreshments were provided by Ted Schultz. Respectfully submitted, Stuart H. McKamey, Recording Secretary 1051st Regular Meeting—December 14, 2000 The 1051st regular meeting of the Ento- mological Society of Washington (ESW) 471 was called to order in the Cathy Kerby room (CE-340) of the National Museum of Natural History, Washington, D.C., by President Dave Furth at 7:03 pm. The meet- ing was attended by 19 members and 10 guests. The minutes of the 1050th meeting were approved with minor modification. Membership Chair Steve Lingafelter read the names of two new applicants for mem- bership: Jonathan Mawdsley and Paul Opler, and reported that ESW gained 23 new members this year. President David Furth reported that ESW acquired its own slide projector this year and also said that the website was almost ready. Furth noted the main results of the November 15 ESW Executive Committee meeting: a $2,000 honorarium is now avail- able for the Proceedings editor, compensat- ing for the huge effort required, as is the case for other academic societies; family membership is now available ($10 for ad- ditional family members, without an addi- tional copy of Proceedings); ESW is now officially archived in the Smithsonian Insti- tution, and archiving was added to the ESW Custodian responsibilities; storage of back issues of Proceedings will be an upcoming problem because we will be losing the space due to USDA renovations. Editor Dave Smith reported that ESW published four issues of Proceedings over the year, totalling 1,100 pages, and one Memoir, which he put on display. The Treasurer’s report was read by John Brown, who announced our assets as of Oc- tober 31st, audited December 5th (see Trea- surer’s report, p. 467). The nominations for officers were read, no new nominations were offered from the floor, and the slate was passed unanimously. In miscellaneous business, four visitors were introduced. For exhibits, Jil Swearin- gin showed us the ESA T-shirt and also glow-in-the-dark insect T-shirts, and rec- ommended the Insectarium in Montreal. Ralph Eckerlin displayed the book *“‘Natu- ral History of the Great Dismal Swamp” (R.K. Rose, ed.), which includes many 472 chapters on arthropods. Edd Barrows dis- played the May 2000 issue of BioScience, featuring an article on beetle horn evolu- tion, and passed around some stalk-eyed flies. Dave Furth described the entomo- phagic delights at a booth at the ESA meet- ing in Montreal, including scorpion-on-a- stick, and announced that a two-volume book on American beetles, by R.H. Arnett and M.C. Thomas, is coming out soon. We’ll no doubt have a chance to see it at a later meeting. Furth also displayed the book ‘A Fly for the Prosecution,” by M. Lee Goff, about forensic entomology, and an example of the Smithsonian NHB’s greet- ing card, which features a chrysomelid. Program Chair Ted Schultz introduced the speaker, David W. Roubik, of the Smith- sonian Tropical Research Institute in Pana- ma, who spoke to us about “‘Truth or Con- sequences: What We Know About Bee Population Dynamics and Diversity in Eco- logical Time.”’ Drawn mostly from his own research, simply for paucity of other long- term studies, Dr. Roubik described his find- ings on the population fluctuations of eu- glossines (so-called orchid bees, though only a small percentage visit orchids), tri- gonines (stingless bees), and Africanized PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON honey bees over 20 years in natural, tropi- cal forest habitats. Sampling was usually by scent traps (for euglossines) or light-traps fitted with baffles, which also collected di- urnal species. The only emergent pattern was that population peaks track overall flowering phenology and an El Nimo effect. El Nifio years were abnormally dry, result- ing in more flowers and consequently a huge increase in bees, followed by popu- lation crashes about three months later. Sur- prisingly, in one study of euglossines, a de- cline was detected in one species, but it was the most abundant species. In closing, Dr. Roubik cast doubt on conclusions drawn from any short-term studies and recom- mended a minimum length of at least four years to claim a pattern. At 8:50 p.m. the meeting was handed over to incoming ESW President John Brown, who immediately appeased the masses by adjourning the meeting and start- ing his inaugural bash. Refreshments were provided by Ted Schultz. Respectfully submitted, Stuart H. McKamey, Recording Secretary VISIT THE ENTOMOLOGICAL SOCIETY OF WASHINGTON WEBSITE http://entomology.si.edu:59 1/ES W/ES W Menus. html PUBLICATIONS FOR SALE BY THE ENTOMOLOGICAL SOCIETY OF WASHINGTON MISCELLANEOUS PUBLICATIONS A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera), by E. Eric Grissell and Michael E. 2 EAC SG 0 ob Ee OS Wie Vest Ue RO JA 0 SY ae RT ke AR lee Me LSE Tad CAE GN DAN aed ? A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera): Second Edition, Revised, by E. Eric Grissell and Michael E. Schauff. 87 pp. 1997 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Memoirs 2, 3, 7, 9, 10, 11, and 13 are no longer available. No. 1. The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939 No. 4. _A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 1952 No. 5. A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp. 1957 __.. No. 6 The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi Die tasi e U OO rk ewe tie eri kite FU ela i us Ns eet Meee es oS ee No. 8. The North American Predaceous Midges of the Genus Palpomyia Meigen (Diptera: Cerato- pogonidae), by W. L. Grogan, Jr. and W. W. Wirth. 125 pp. 1979 __. No. 12. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff. CTA By Ny ANS TP A SIRT Rea Ak Sera Set cat tes ty MEM RA Cg LO OU NCR 9 MRL A ef LL No. 14. Biology and Phylogeny of Curculionoidea, edited by R. S. Anderson and C. H. C. Lyal. 174 [af 291A ES) MSE aR ieee 1 OP AN Ee i Bodo) Et AP coe es AAD AP ETD RO RRS Sy yD No. 15. A Revision of the Genus Ceratopogon Meigen (Diptera: Ceratopogonidae), by A. Borkent ANGI Wil Grogan Iie lOS\ppy OO Sire eas eee Oe Sake ae Ns SOE a eee eS No. 16. The Genera of Beridinae (Diptera: Stratiomyidae), by Norman E. Woodley. 231 pp. 1995 ___. No. 17. Contributions on Hymenoptera and Associated Insects, Dedicated to Karl V. Krombein, edited by Bos: Norden and A’ S: Menke! 216 pp: 1996" 22s ie Aa ea A et Net No. 18. Contributions on Diptera, Dedicated to Willis W. Wirth, edited by Wayne N. Mathis and Walia ce Grogan 207 spp LOO ia race i rie Serene MAN oh Oe Pp Le a ae a No. 19. Monograph of the Stilt Bugs, or Berytidae (Heteroptera), of the Western Hemisphere, by hnomas. |, Lichty: LAD pp Oey fao2 7 ajith eee ee RAUL, Se a 8 See a een No. 20. The Genera of Elaphidiini Thomson 1864 (Coleoptera: Cerambycidae), by Steven W. Lin- Palette ULSI. LIOR tte e sae a eels BL B82 5 ad SUN Toe SY pe A i eR ee No. 21. New World Blepharida Chevrolat 1836 (Coleoptera: Chrysomelidae: Alticinae), by David G. eeeraas OOM pp POOR etn aes ee Set te Pe eee ee ee es a es ee ae No. 22. Systematics of the North American Species of Trichogramma Westwood (Hymenoptera: dimchopranunatidae):by, John: Pinto: 28 7,pp..1 999) Was seks eve es eves Ieee eee ey No. 23. Revision of the Net-Winged Midges of the Genus Blepharicera Macquart (Diptera: Blepha- riceridae) of Eastern North America, by Gregory W. Courtney. 99 pp. 2000 __... $10.00 15.00 $15.00 15.00 15.00 15.00 12.00 5.00 25.00 25.00 25.00 25.00 25.00 18.00 12.00 12.00 28.00 14.00 Back issues of the Proceedings of the Entomological Society of Washington are available at $60.00 per volume to non-members and $25.00 per volume to members of the Society. Prices quoted are U.S. currency. Postage extra except on prepaid orders. Dealers are allowed a discount of 10 percent on all items, including annual subscriptions, that are paid in advance. All orders should be placed with the Treasurer, Entomological Society of Washington, % Department of Entomology, Smithsonian Institution, Washington, D.C. 20560-0168. CONTENTS (Continued from front cover) MARSH, PAUL M. and SCOTT R. SHAW—Revision of North American Aleiodes Wesmael (Part 6): The gasterator (Jurine) and unipunctator (Thunberg) species-groups (Hymenoptera: Braconidae: Rogadinae) MARSHALL, S. A.—A review of the southern South American genus Gyretria Enderlein (Diptera: Sphaeroceridae: Limosininae) McCAFFERTY, W. P. and J. R. DAVIS—Texas Caeninae (Ephemeroptera: Caenidae), with description of a new species NGUYEN, TAM C.—A new species of Cladochaeta Coquillett (Diptera: Drosophilidae) and a new record for Cladochaeta sturtevanti Wheeler and Takada in Arizona, with notes on natu- ral history PAPP, LASZLO and WAYNE N. MATHIS—A review of the family Nannodastiidae (Diptera) .. PINTO, JOHN D.—Two new species of Zonitis F. (Coleoptera: Meloidae) from southwestern North America, with comments on generic definitions in the Nemognathinae PODENAS, SIGITAS, GEORGE O. POINAR, JR., and RAIF MILKI—New crane flies (Diptera: Limoniidae) from Lebanese amber PRATT, GORDON F. and CECILIA L. PIERCE—ZJncisalia henrici (Grote and Robinson) (Lepi- doptera: Lycaenidae) reared on reproductive and non-reproductive tissues of three different plant species RIDGE-O’CONNOR, GALE E.—Distribution of the western conifer seed bug, Leptoglossus occidentalis Heidemann (Heteroptera: Coreidae) in Connecticut and parasitism by a tachinid fly, Trichopoda pennipes (F.) (Diptera: Tachinidae) ROBBINS, RICHARD G., LOUIS N. SORKIN, and FRANCOIS VUILLEUMIER—First report of Ixodes auritulus Neumann (Acari: Ixodida: Ixodidae) from the blackish cinclodes, Cinclodes antarcticus (Garnot) (Aves: Passeriformes: Furnariidae), with additional records of parasitism of Cinclodes spp. by this tick species TOGASHI, ICHIJI—Description of a new species of Urocerus Geoffroy (Hymenoptera: Siricidae) from Japan VANWIEREN, BRIAN J., BORIS C. KONDRATIEFF, and BILL P. STARK—A review of the North American species of Megarcys Klapalek (Plecoptera: Perlodidae) YASUNAGA, TOMOHIDE—New records of two plant bug genera (Heteroptera: Miridae: Phylinae: Pilophorini) from Japan, with descriptions of two new species ZHANG, LIKUN, GEXIA QIAO, and GUANGXUE ZHANG—Study on Chinese Melanaphis van der Goot (Homoptera: Aphididae) with descriptions of three new species BOOK REVIEW STECK, GARY J.—Fruit Flies (Tephritidae): Phylogeny and Evolution of Behavior, edited by Martin Aluja and Allen L. 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