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S > * 2 2 SMITHSONIAN INSTITUTION NOlIfUllSNI NVINOSHII WS^S 3 IBVaan^LIBRARIEi m Ti if) ~~ < I m NVINOSHillNS S3iavaai1 libraries SMITHSONIAN INSTITUTION NOiinillSN >2 r* 2 r” 2 ?! O _ X'oo'vnX O 00 70 “1 > CO ” m x^jvosv^^ ^ m SMITHSONIAN INSTITUTION NOliniliSNi “ NVINOSHlIl/^S S3 I dVd a IT LIBRARIES ~ CO 2 CO 2 , 2 o 3: CO ^ ■w I y i NVIN0SH1IPMS^S3 I HVB a 11 LI B RAR I E S^SMITHSONIAN INSTITUTION NOliniliSN ^ to “ CO SMITHSONIAN INSTITUTION NOlinillSNI NVINOSHimS S3!aV^an LIBRARIES ^ ~ — (O _ NVINOSHllINS S3!BVaan LIBRARIES SMITHSONIAN INSTITUTION NOlinillSN ^ 2: . CO 2; V- CO (O ^ 2 ^ ^ SMITHSONIAN_INSTITUTION NOlinillSNI NVINOSHillNS S3iaVdai1 LlBRARIEi 2 ^ _ r; „ CO Journal of the New York Entomological Society published by The New York Entomological Society Contents Volume 97, 1989, Numbers 1-4 Number 1 The biology of Sthenopis auratus (Grote) (Lepidoptera: Hepialidae) Tim L. McCabe and David L. Wagner 1-10 A revisionary study of the neotropical hairstreak butterfly genus Noreena and its new sister genus Contrafacia (Lepidoptera: Lycaenidae) Kurt Johnson 1 1-46 A new Aphaenogaster (Hymenoptera: Formicidae) from southern New Mexico William P. MacKay 47-49 Notes on ant larvae: Ponerinae George C. Wheeler and Jeanette Wheeler 50-55 Biology of Andrena crataegi Robertson (Hymenoptera: Andrenidae), a commun- ally nesting bee Eben A. Osgood 56-64 The genus Metopina (Diptera: Phoridae) from Cretaceous and Tertiary ambers David Grimaldi 65-72 Naucoridae (Heteroptera) of New Guinea. IV. A revision of the genus Cavocoris with descriptions of four new species Dan A. Polhemus and John T. Polhemus 73-86 Cariniocoris, A new phyline plant bug genus from the eastern United States, with a discussion of generic relationships (Heteroptera: Miridae) Thomas J. Henry 87-99 Froeschnerana mexicana, a new genus and species of Deraeocorinae from Mexico (Heteroptera: Miridae) J. C. Schaffner and Paulo Sergio Fiuza Ferreira 100-104 Two new species of Mormidea from Mexico and Guatemala (Heteroptera: Pentatomidae) D. A. Rider and L. H. Rolson 105-110 Notes and Comments New records of Palearctic Heteroptera in New York State: Microphysidae and Miridae Michael D. Schwartz 111-114 Names and authorship of two family-groups in the Ephemeroptera William L. Peters and Michael D. Hubbard 1 1 5 A necessary new name in the family Hebridae (Heteroptera: Gerromorpha) John T. Polhemus 1 1 6 Book Reviews Insect-plant Interactions Hans Damman 1 1 7-1 1 8 Insects and Flowers. The Biology of a Partnership P. J. DeVries 118-120 Pheromone Biochemistry Dietrich Schneider 120-122 Portraits of South Australian Geometrid Moths Frederick H. Rindge Proceedings of the Fifth International Symposium on Trichoptera, Lyon, France 21-26 July 1986 Wolfram Mey Announcement Number 2 A fossil solpugid, Haplodontus proterus, new genus, new species (Arachnida: Sol- pugida) from Dominican Amber George O. Poinar and Jorge A. Santiago-Blay On the abundance and ecology of Ricinulei (Arachnida) from Central Amazonia, Brazil Joachim U. Adis, Norman L Platnick, Jose W. de Morais, and Jose M. Gomes Rodriguez Two new species of Eosentomon from Chickasaw State Park, Tennessee (Protura, Eosentomidae) William A. Outten and Robert T. Allen New species rediscriptions, and cladistics of the genus Pseudocentroptiloides (Ephemeroptera: Baetidae) R. D. Waltz and W. P. McCajferty Review of Daleapidea Knight (Heteroptera; Miridae: Orthotylinae: Orhtotylini) Randall T. Schuh Texocoris nigrellus: Distribution and hosts of an enigmatic plant bug (Heteroptera: Miridae: Orthotylinae) A. G. Wheeler, Jr. Tantysoma diabolica new species (Coleoptera: Carabidae: Platynini) from Baja California and its Biogeographic Significance James K. Liebherr A new microcadddisfly genus (Trichoptera: Hydroptilidae) from the interior high- lands of Arkansas, U.S.A. Michael L. Mathis and David E. Bowles Relationships among Holarctic genera in the Cyrtogaster-gvowp with a review of the species of North American north of Mexico (Hymenoptera: Pteromalidae) Steven L. Heydon Stinging behavior and residual value of worker honey bees {Apis mellifera) Steven A. Kolmes and Linda A. Fergusson-Kolmes Notes and Comments Evaluation of the Spider Steatoda triangulosa (Araneae: Theridiidae) as a Predator of the Red Imported Fire Ant (Hymenoptera: Formicidae) William P. MacKay Book Reviews CLADISTICS IN THE FAST LANE Hennig86. Version 1.5 Kirk Fit zhugh TWO NEW TRUE BUG CATALOGS Catalog and Bibliography of the Leptodomorpha (Heteroptera) Nils M oiler Andersen Catalog of the Heteroptera, or True Bugs, of Canada and the Continental United States Randall T. Schuh LIFE CYCLES AND DIAPAUSE Insect Development: Photoperiod and Temperature Control Jesse A. Logan The Evolution of Insect Life Cycles Lori Stevens 122- 123 123- 124 124 125-132 133-140 141-150 151-158 159-166 167-172 173-186 187-191 192-217 218-231 232-233 234-241 241-242 243-245 245-248 248-249 DARWIN’S INSECTS Darwin’s Insects. Charles Darwin’s Entomological Notes George E. Ball 249-250 Number 3 Annotated checklist of the Thysanoptera of Bermuda Sueo Nakahara and Daniel J. Hilburn Annotated checklist of the whiteflies (Homoptera: Aleyrodidae) of Bermuda Sueo Nakahara and Daniel J. Hilburn Blissus breviusculus: New distribution records of a little-known chinch bug (Het- eroptera: Lygaeidae) A. G. Wheeler, Jr. and Jonathan E. Fetter Three new species of Lincus (Hemiptera: Pentatomidae) from palms L. H. Rolston Reconstitution of Coleometopini, Tenebrionini and related tribes in America north of Colombia (Coleoptera: Tenebrionidae) John T. Doyen Pityogenes bidentatus (Herbst), a European bark beetle new to North America (Coleoptera: Scolytidae) E. Richard Hoebeke New Trichoptera from Alabama S. C. Harris An unusual black fly (Diptera: Simuliidae), representing a new genus and new species B. V. Peterson Butterfly exploitation of an ant-plant Mutualism: Adding insult to herbivory P. J. DeVries and L Baker Sensory structures on the ovipositor of the ball gall fly Eurosta solidaginis (Fitch) (Diptera: Tephritidae) Edward Ritter and Carey E. Vasey Review of Nearctic Rhicnocoelia and Callimerismus with a discussion of their phylogenetic relationships (Hymenoptera: Pteromalidae) Steven L. Heydon Seed predation by a braconid wasp, Allorhogas sp. (Hymenoptera) Margarete V. de Macedo and Ricardo F. Monteiro Book Reviews Asa Fitch and the Emergence of American Entomology: With an Entomological Bibliography and a Catalog of Taxonomic Names and Type Specimens Conner Sorensen Insect Flight: Dispersal and Migration Biogeography and Taxonomy of Honeybees Peter Turchin Lynn S. Kimsey Tree and Shrub Insects of the Prairie Provinces A. G. Wheeler, Jr. and Gregory A. Hoover 251-260 261-264 265-270 271-276 277-304 305-308 309-316 317-331 332-340 341-346 347-357 358-362 363- 364 364- 365 365- 366 367-370 Number 4 Atlas of antennal trichobothria in the Pachynomidae and Reduviidae (Heterop- tera) Pedro W. Wygodzinsky and Sarfraz Lodhi 371-393 Review of the New World species of the genus Neottiglossa Kirby (Heteroptera: Pentatomidae) D. A. Rider 394—408 Orius minutus (Linnaeus) in North America (Hemiptera: Heteroptera: Antho- coridae) John D. Lattin, Adam Asquith and Steve Booth 409-416 Biology of Lopidea nigridea Uhler, a possible aposematic plant bug (Heteroptera: Miridae: Orthotylinae) James D. Mclver and Adam Asquith 4 1 7-429 Redescription of Platynus prognat hus Van Dyke (Coleoptera: Carabidae: Platyn- ini) and circumscription of Lindroth’s Decentis and Hypolithos groups James K. Liebherr 430-437 Aggregation and predator avoidance in whirligig beetles (Coleoptera: Gyrinidae) K. Vulinec and M. C. Miller 438-447 First record of the Palearctic species Oxypoda opaca (Gravenhorst) from North America (Coleoptera: Staphylinidae: Aleocharinae) E. Richard Hoebeke 448-454 Two mouthpart modifications in larval notodontidae (Lepidoptera): their taxo- nomic distributions and putative functions G. L. Godfrey, J. S. Miller and D. J. Carter 455-470 On Venezuelan Leprolochus (Araneae, Zodariidae) Rudy Jocque and Norman I. Platnick 471^74 Karyotypes of three spider species (Araneae: Pholcidae: Physocyclus) James C. Cokendolpher 475^78 Notes and Comments A new structure on the hind legs of male Monalocoris carioca Carvalho and Gomes (Heteroptera: Miridae) Jorge A. Santiago- Blay and Jenaro Maldonado Capriles 479—482 Records of Chimarra soda (Trichoptera: Philopotamidae) from interior highland streams in Arkansas and Missouri Paul K. Lago, Michael L. Mathis and David E. Bowles 482-483 Poecilochirus monospinosus (Acarina: Mesostigmata: Parasitidae), a predator of house fly immatures: new locality records Christopher J. Geden, Donald C. Steinkraus and Donald A. Rutz 483-485 Book Reviews ARTHROPOD OVERVIEWS Spiders. Webs, Behavior, and Evolution Robert J. Raven 486-487 Evolution and Adaptation of Terrestrial Arthropods George C. Eickwort 487-489 Thomas W. Culliney 489-491 ENTOMO-ECOLOGICAL ASSOCIATIONS Coevolution and Systematics Brian D. Earrell 492 Biology of Mutualisms Brian D. Earrell 493-494 Honorary, Life, and Sustaining Members 495 Reviewers for 1989 495 Statement of Ownership, Management, and Circulation 496 Vol. 97 JANUARY 1989 No. 1 Journal of the New York Entomological Society (ISSN 0028-7199) Devoted to Entomology in General JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Editor: Randall T. Schuh, Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024 Book Review Editor: David A. Grimaldi, Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024 Publications Committee: Louis Trombetta, St. Johns University, Queens, New York, Chairman; Alfred G. Wheeler, Jr., Pennsylvania State Department of Agriculture, Harrisburg; Joseph M. Cerreta, St. Johns University, Queens, New York. The New York Entomological Society Incorporating The Brooklyn Entomological Society President: Dennis J. Joslyn, Department of Biology, Rutgers University, Camden, New Jersey 08102 Vice President: Durland Fish, Medical Entomology Laboratory, New York Medical College, Armonk, New York 10504 Secretary: Richard Falco, Westchester County Health Department, White Plains, New York Treasurer: Louis Sorkin, Department of Entomology, American Museum of Natural History, New York, New York 10024 Trustees: Class of 7955— Henry M. Knizeski, Jr., Mercy College, Dobbs Ferry, New York; Michael D. Schwartz, American Museum of Natural History, New York, New York; Class of 7959— Christine Falco, West- chester County Health Department, White Plains, New York; James S. Miller, Department of Entomology, American Museum of Natural History, New York, New York. Annual dues are $23.00 for established professionals with journal, $10.00 without journal, $15.00 for students with journal, $5.00 without journal. Sustaining memberships are $53.00 per year, institutional memberships are $125.00 per year, and life memberships are $300.00. Subscriptions are $40.00 per year domestic and $45.00 foreign. All payments should be made to the Treasurer. Back issues of the Journal of the New York Entomological Society, the Bulletin of the Brooklyn Entomological Society, Entomologica Americana, the Torre-Bueno Glossary of Entomology and other Society publications can be purchased from Lubrecht and Cramer, RD 1, Box 244, Forestburgh, New York 12777. Meetings of the Society are held on the third Tuesday of each month (except June through September) at 7 p.m. in the American Museum of Natural History, Central Park West at 79th Street, New York, New York. Mailed March 29, 1989 The Journal of the New York Entomological Society (ISSN 0028-7199) is published 4 times per year (January, April, July, October) for the Society by Allen Press, Inc., 1041 New Hampshire, Lawrence, Kansas 66044. Second class postage paid at New York, New York and at additional mailing office. Postmaster: Send address changes to the New York Entomological Society, % American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024-5192. Known office of publication: American Museum of Natural History, New York, New York 10024. Journal of the New York Entomological Society, total copies printed 700, paid circulation 602, mail subscription 602, free distribution by mail 19, total distribution 621, 79 copies left over each quarter. THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER. J. New YorkEntomol. Soc. 97(1):1-10, 1989 THE BIOLOGY OF STHENOPIS AURATUS (GROTE) (LEPIDOPTERA: HEPIALIDAE) Tim L. McCabe and David L. Wagner Biological Survey, New York State Museum, State Education Department, Albany, New York 12230, and Department of Ecology and Evolutionary Biology, 75 North Eagleville Rd., Room 312, U-43, Storrs, Connecticut 06268 Abstract. — Larvae of Sthenopis auratus (Grote) were found tunneling in the artichoke-like leaf bases and stems of several ferns: Ostrich Fern (Matteuccia struthiopteris (Linnaeus) Todaro), Marginal Shield Fern {Dryopteris marginalis (Linnaeus) Gray), Mountain Wood Fern {Dryopteris campyloptera (Kunze) Clarkson), and Lady Fern (Athyrium filix-femina (Linnaeus) Roth), all Polypodiaceae. An ichneumonid parasite, Pterocormus devinctor Say, was associated with a cocoon of S. auratus. The early evening calling behavior by males is described. The male has prominent androconia on the hind tibia presumably for dissemination of a pheromone. Males did not seek out females, but rather adopted sessile calling displays on emergent understory vegetation. Additional locality records are given for this rare northeastern moth. The larva and pupa are described and illustrated in detail. Sthenopis auratus (Grote) (1878) is among the rarest of hepialids in North America (Winn, 1909; Beutenmiiller, 1913; Forbes, 1923). Its biology was heretofore com- pletely unknown. Few moths are represented in collections. S. auratus has been reported from St. Johns and Brome County, Quebec; Black Mountains, North Car- olina; Franconia Mountains, New Hampshire; and in New York in Lewis County and at Fentons, Lancaster, Ithaca, McLean, and in the Catskills (Beutenmiiller, 1913; Forbes, 1923). New locality records for New York include Rensselaer (Mill Creek, Rensselaer County) (TLM, Quinter, Dievendorf), Cambridge and Murray Hollow (both Wash- ington County) (Romack, TLM, DLW), Whiteface Mountain (Essex County) (TLM), Indian Lake (Beaver Meadow, Hamilton County) (TLM), Rensselaerville (the Huyck Preserve) (TLM, Quinter, Franclemont) and Albany (Pine Bush) (both Albany Coun- ty) (TLM). In Vermont, it has been recorded from Camels Hump (Chittenden County) (Don Tobi). Larvae of S', auratus were recovered most frequently from Ostrich Fern, Matteuccia struthiopteris (Linnaeus) Todaro. Aderkas and Peterson (1987) provided a partial list of insects associated with Ostrich Fern, but did not report the hepialid, perhaps because their research site was in Nova Scotia, a province for which the insect has yet to be recorded (Douglas Ferguson and Berry Wright, pers. comm.). Nonetheless, searching for larvae may be the most practical way to survey for the presence or absence of many hepialids, including S. auratus. BIOLOGY OF STHENOPIS AURATUS Larval hosts were identified as Ostrich Fern {Matteuccia struthiopteris), Marginal Shield Fern {Dryopteris marginalis L.), Mountain Wood Fern {Dryopteris campy lop- 2 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) tern (Kunze) Clarkson), and Lady Fern (Athyrium filix-femina (L.) Roth), all Poly- podiaceae. Ostrich Fern patches were observed to support the largest populations of the moth, perhaps because single rootstocks could support several larvae. As many as six larvae were recovered from a single rootstock of Ostrich Fern. Ostrich Fern did not appear to suffer from presence of larvae. No more than one larva was associated with a given individual of the other fern species. All four fern species shared a similar artichoke-like rootstock, with imbricated subterranean leaf bases. The rootstocks of many Royal Fern (Osmunda regalis L.), Interrupted Fern {Os- munda claytoniana L.), and Bracken Fern {Pteridium aguilinum (L.) Kuhn) were searched for larvae without success. The size, form, and consistency of the root mass may be a limiting factor. Early instar larvae were not observed. The majority of the larval collections were made in June 1987 at Murray Hollow. At this time, immatures representing two distinct cohorts were noted. The first consisted of middle instar to nearly fully de- veloped larvae, which would not mature until summer 1988. The second cohort consisted of prepupal larvae and pupae, which would produce adults during the later part of June and early July 1987. This observation and our records for adult captures suggest that S. auratus has a two-year life cycle in New York. In Matteuccia, larvae bored in the outer portions of the rootstock, tunneling into the leaf bases of previous years’ growth. A single larva tunneled into several bases. In Dryopteris spp., the larval channel ran down the axis of the subterranean stem. In all ferns, the larval tunnel was relatively free of silk until the last larval instar when an elongate cocoon was spun in the tunnel. The elongate cocoon afforded the pupa considerable mobility. Prior to eclosion the pupa was thrust to the outside. Individual plants that are most likely to host larvae are the older, more mature plants with rootstocks that are most protuberant from the forest floor. On June 12, 1987, both prepupal larvae and pupae were present in the Murray Hollow colony. Adults began flying at Murray Hollow in the later half of June, and no adults were taken after 15 July, which would suggest that the pupal stage lasts less than a month (Howard Romack, pers. comm.). DESCRIPTION OF LAST INSTAR LARVA AND PUPA OF STHENOPIS AURATUS (GROTE) Larval characters for the Hepialidae have been given by Hinton (1 946), Gerasimov (1937), Aitkenhead and Baker (1964), and Wagner (1987). We adopt Hinton’s (1946) nomenclature for the larval chaetotaxy except for the D and SD setae on the pro- thoracic shield where Wagner (1987) is followed. We emphasize characters known or likely to vary among species or genera of Hepialidae. The pupal nomenclature follows Mosher (1916). Late Instar Larva Figs. 3-1 1 Length to 47 mm (preserved), N = 15. Head (Fig. 5) orange; width to 3.7 mm. Labrum shallowly emarginate to cleft, anterior half furrowed with 5 pairs of setae. Antennae as in Figure 1 1 . Membranous frontoproximal portion of hypopharynx (Fig. 7) with sparse vestiture of spinules. Spinneret perpendicular to head, scarcely ex- ceeding posterior margin of labium. 1989 BIOLOGY OF STHENOPIS A URA TVS 3 Figs. 1-3. Sthenopis auratus: 1. Male calling— metatibial scent brush visible just above middle of body; 2. Male at rest; 3. Mature larva shortly before pupation in rootstock of fern (notice silk-lined tunnel). 4 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) Figs. 4-7. Sthenopis auratus larva: 4. Habitus of mature larva; 5. Head capsule, frontal view; 6. Left mandible; 7. Hypopharyngeal complex. 1989 BIOLOGY OF STHENOPIS A URA TVS 5 Figs. 8-11. Sthenopis auratus larva: 8. Integument of thoracic segments which appears white (x 1,000); 9. Microspinules of abdominal segments, which cause segments to appear rusty (x 1,000); 10. Ventral surface of head and prothorax of larva (x25); 11. Antenna of larva (x250). Thorax. Ground color white, setae on large, brown to yellow-brown, plate-like pinacula. Spinules reduced to platelets (Fig. 8). Prothorax with SD setae fine, less than A width and Vi length of D2, subtended by patches of darkened spinules. Distance from D2 to XD2 equal to that between D2 and SD setae. L2 and L3 about Vi length of LI; L3 on small elliptical pinaculum free of dorsal plate. Meso- and metathorax 6 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) with D1 pinacula fused over midline; MDl twice as large as MSD setae on both segments. Mesothorax with D2 and SD setae on single pinaculum; metathorax with both D2 and L3 free of SD pinaculum. Claw with basal tooth ending about halfway to apex. Abdomen. Integument with prominent vestiture of spinules over A1-A9 (Fig. 9). D1 on A1-A7(A8) arising from large, hemispherical, lightly melanized pinaculum; D2 on smaller, elliptical, slightly raised plate. Other setae subtended by small, darkly pigmented pinacula. SD and SV setae on separate pinacula. MV3 elongate on Al- A7 and grouped with SV setae on A3-A6; minute on A8 and A9. Distance from SDl to LI less than that between D1 and D2 on A9. Crochets in multiserial ellipse with hve complete rows and one or two incomplete series. Male Pupa Figs. 12, 13 Cylindrical, 6-7 mm wide by 26-27 mm long, N = 2. Spination reduced; setae short, approximately equal to antennal width. Head. Mandible with one seta. Cuticle over eye and mandible roughened. Labium short, ending before maxillae, broadest at base of palpus. Clypeolabral suture absent; labrum with one medial pair of setae and clypeus with two pairs laterad. Antenna extending to dorsolateral margin of mesotibia. Frons and vertex deeply furrowed. Vertex with small, paired cocoon cutters above base of antenna, each with two setae; and one seta to either side of ecdysial line. Thorax. Ecdysial line extending to Al. Prothorax with prominent deep furrows and six pairs of setae; prothoracic spiracle elongate, nearly twice antennal width. Meso- and metathorax roughened, each with two pairs of setae. Forewings produced at apex. Abdomen. Al with one pair of setae. A2 and A3 with seven setal pairs. Setae arranged as on larva on A3-A8, except with SV seta only on A8, although setal insertion may remain (1 of 2); A9 and AlO without setae; A1-A7 with transverse furrows over dorsum, these bounded cephalad and caudad by raised ridges of darkly pigmented teeth on A3-A7; anterior row extends below spiracle on A5 and A6, and completely encircles A7, where teeth form broad plate ventrad. Venter of A4-A6 with wavy, pigmented, fluted ridges over position of larval prolegs, these connected by thin pigmented ridge. Spiracles nonfunctional on A7 and A8. A9 with dark, Y-shaped genital slit with raised, darkened hemispherical area to either side; and low, broad, three-toothed horn dorsolaterad. Female Pupa. 7x31 mm, N = 1 . SV 1 setal insertion absent on A8. A8 with short, dark genital slit that runs into furrows posteriorad which delimit A8 from A9 and A9 from AlO. DESCRIPTION OF MALE CALLING BEHAVIOR Male calling behavior was studied at Mill Creek, Rennselaer County, where Ostrich Fern was the principal host. Males called for a very short period at evening. They did this by hanging from the tip of a fern frond (rarely using a taller plant growing amongst the ferns), obtaining a foothold with their first two pairs of legs (Figs. 1, 2). Suspended in this position, they next “fanned” their wings, thereby forcing air over 1989 BIOLOGY OF STHENOPIS A ERA TVS 1 Figs. 12, 13. Sthenopis auratus pupa: 12. Ventral view; 13. Lateral view. the metatibial scent brush (Fig. 1). Homologous metatibial scent brushes are found in all Nearctic Sthenopis species and in a number of related hepialid genera (Mallet, 1984; Wagner, 1985). Morphological details of the tibia and androconia have been studied in other ghost moths (Deegener, 1902; Deegener and Schaposchnikow, 1905; Wagner, 1985). Birch and Hefetz (1987) gave an overview of analogous androconial organs in male moths. 8 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) Attraction and courtship were not observed, although a single copulating pair was encountered by Howard Romack in Murray Hollow during the evening period of flight activity. The moths paired end to end, with one moth (male?) suspended only by its genitalia, a mating posture typical for other Hepialidae (Wagner, 1985). The duration of pairing was not noted. Male calling behavior was observed 27 different times over the course of four years. The best viewing was at Mill Creek during a period from 25-30 June 1986. Sunset was at 2 1 :04 EDST during this period of observation. The onset of flight and calling behavior was affected by ambient light intensities. For example, calling began earlier on overcast evenings. Similarly, a moth resting in a shaded recess began calling earlier than other males. Hence, our earliest record for calling, at 20.54 EDST, was of a moth sitting in a shaded recess on a day with an overcast skyline. Males typically began calling at the point at which human vision became insufficient and a flashlight was needed. Many moths were observed to “fan” the wings for a few seconds or a minute or two, then became inactive although still suspended by the first two pairs of legs. Repeated observations suggested that this was due to the disturbance caused by the presence of the observer. Males occasionally flushed when probed, but quickly dis- continued “fanning” if disturbed. Observations from twenty feet (versus 3 feet) in- dicated that males continuously fanned during the entire period of evening crepus- cular activity (N = 9). Similar calling behaviors occurred in Hepialus californicus Boisduval and H. hectoides Boisduval. In both these species, the males called until a female was attracted or, in the absence of a female, for the 15-25 minute duration of the evening courtship and mating flight (Wagner, 1985). Mated females of S. auratus were active later into the evening. Such gravid females constituted the ma- jority of light captures for this species. The longest period of calling observed in a population was on 29 June 1987, when the first male was observed calling at 20:55 and one of several other males called until 21:21. Out of 27 males observed calling, the longest documented from initiation to completion was for 12 minutes and 35 seconds. Six or seven minutes was typical, although some of these moths may have been calling for a few minutes prior to when they were first discovered. On two occasions, a male was observed to fly to a new location and initiate a second display. Three times the male reached a new location but did not take up calling again. Several flushed males were unable to be tracked and were lost. All males observed calling for six minutes or more were finished calling for the night once they quit. They continued to hang from the fern tip (two were located still hanging by 01:00 hr), but changed locations before daybreak (N = 8). Frequently females were observed hovering over the fern patch, but none were observed approaching a calling male. On two occasions males were observed calling within four feet of another calling male. No aggression between males was observed. Both males and females were slow-flying and maintained a low altitude, barely above the tips of the fronds of fern. Moths located in the daylight rested suspended from a twig or leaf, not at the very tip of a leaf as when calling (N = 2). DISCUSSION The larva of Sthenopis auratus possesses several unique characters which serve to distinguish this species from other Nearctic hepialids: (1) L3 on a small, dark pinacu- 1989 BIOLOGY OF STHENOPIS A URA TVS 9 lum separate from the dorsal plate; (2) MD3 is elongate, nearly twice the size of the MSD setae on the mesothorax; (3) similarly, MV3 is elongate on Al, comparable in size to MV3 on A2-A7; (4) D1 on A1-A7 arises from a large, knob-like pinaculum; and (5) on all abdominal segments, both the SD and SV setae are located on separate pinacula. These characters outweigh any comparable list that could be compiled to separate other members of Sthenopis from Holarctic hepialids in the genera Kor- scheltellus Bomer, Phymatopus Wallengren, and Triodia Hiibner (Gerasimov, 1937; Aitkenhead and Baker, 1964). Several species of hepialids are known to feed on ferns (Buckler, 1887; Barrett, 1895; Wagner, 1985; Wagner et al., in press). However, in no other species does the association with pteridophytes seem as pronounced as in Sthenopis auratus, which presently is known to feed only on three fern genera. Larvae of another hepialid, Korscheltellus gracilis (Grote) have been collected from three of the four fern hosts of S. auratus, included are the two Dryopteris species and Athyrium. Larvae of the two moths are readily separable, however. The larvae of S. auratus appear bicolored, the nearly white ground color of the thoracic segments contrasting with the rusty-appearing abdominal segments. In Korscheltellus the ground color is uniformly whitish. In S. auratus the DI setae arise from enlarged knobby pinacula on A1-A7; in K. gracilis the Dl pinacula are unmodified. A female of Pterocormus devinctor Say was found emerging from a recently spun cocoon of S. auratus on 1 2 June 1 987, at Murray Hollow. Elsewhere this ichneumonid has been reared from the pupae of Sthenopis thule (Winn 1912). The entomophagous fungus Beauvaria bassiana (Vuill.) was identified from a late instar larva found mummified within its tunnel during August. ACKNOWLEDGMENTS We thank Howard Romack for his assistance with the Murray Hollow population. We thank Douglas Wolfe of the Atmospheric Science Research Center whose cooperation facilitated field work on Whiteface Mountain in the Adirondacks. Brian Farrell gave permission to collect on his property near Indian Lake. Herb Holzer and Gerald Dievendorf assisted with field collections in the Albany area. The larval habitus, head capsule, hypopharynx, and mandible was illustrated by Patricia Eckel. Mary Ann Tenerio assisted with the larval preparation, scanning electron microscopy, and darkroom work. Henry Townes determined the ichneumonid. Charles Sheviak graciously introduced one of us (TLM) to the Mill Creek site. Eric Quinter shared (with TLM) in the discovery of the larva at Mill Creek. We thank Eric Quinter and Fred Rindge for their review. This work was supported, in part, by a Tilton Fellowship to DLW. Contribution number 581 of the New York State Science Service. LITERATURE CITED Aderkas, P. von and B. V. Peterson. 1987. Chirosia betuleti (Ringdahl) (Diptera: Anthomyi- idae) a gall-former on the ostrich fern, Matteuccia struthiopteris, with notes on other insect-fern associates. Proc. Entomol. Soc. Washington 89:532-547. Aitkenhead, P. and C. R. B. Baker. 1964. The larvae of the British Hepialidae. Entomologist 97:25-38. Barrett, C. G. 1895. Lepidoptera of the British Islands. Vol. II. Reeve and Co., London, pp. 151-168. Beutenmiiller, W. 1913. Notes on Hepialus auratus Grote. Insecutor Inscitiae Menstruus 1: 129-130. 10 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) Birch, M. C. and A. Hefetz. 1 987. Extrusible organs in male moths and their role in courtship behavior. Bull Entomol. Soc. Am. 33:222-229. Buckler, W. 1887. The larvae of the British butterflies and moths. Vol. II. Sphinges or Hawk Moths and part of the Bombyces. Ray Soc., London, 175 pp. Deegener, P. 1902. Das duftorgan von Hepialus hectus L. Zeitz. Wiss. Zool. 71:276-295. Deegener, P. and C. Schaposchnikow. 1 905. Das duftorgan von Phassus schamyl Chr. Zeitschr. Wiss. Zool. 78:245-260. Forbes, W. T. M. 1 923. Lepidoptera of New York and neighboring states. Pt. 1 . Cornell Univ. Agr. Expt. Sta. Mem. 68, 729 pp. Gerasimov, A. S. 1937. Hepialidenraupen (Lepidoptera). Entomol. Zeit. Stuttgart 51:81-84. Grote, A. R. 1878. A new Hepialus from New York. Can. Entomol. 10:18. Hinton, H. E. 1 946. On the homology and nomenclature of the setae of Lepidopterous larvae, with some notes on the phylogeny of the Lepidoptera. Trans. Roy. Entomol. Soc., London 97:1-37. Mallet, J. 1984. Sex roles in the ghost most Hepialus humuli (L.) and a review of mating in the Hepialidae (Lepidoptera). Zool. J. Linn. Soc. 79:67-82. Mosher, E. 1916. A classification of the Lepidoptera based on characters of the pupa. Bull. Illinois State Lab. of Nat. Hist. 12:13-159. Wagner, D. L. 1985. The biosystematics of Hepialus F. 5. lato, with special emphasis on the californicus-hectoides species group. Ph.D. dissertation. University of California, Berke- ley. (Diss. Abs. Intern. Order No. DA86 10260.) Wagner, D. L. 1987. Hepialidae. Pages 347-349 in: F. Stehr (ed.). Immature Insects. Kendall- Hunt, Dubuque, Iowa. Wagner, D. L., D. R. Tobi, W. E. Wallner, B. L. Parker and J. Leonard. In review. The immature stages and natural enemies of Korscheltellus gracilis. Annals of the Entomol. Soc. Amer. Winn, A. F. 1909. The Hepialidae, or Ghost-moths. Can. Entomol. 41:189-193. Winn, A. F. 1912. A hymenopterous parasite of Hepialus thule. Ann. Rep. Entomol. Soc. Ontario (1911) 42:70-71. Received July 5, 1988; accepted August 17, 1988. J. New York Entomol. Soc. 97(l):ll-46, 1989 A REVISIONARY STUDY OF THE NEOTROPICAL HAIRSTREAK BUTTERFLY GENUS NOREENA AND ITS NEW SISTER GENUS CONTRAFACIA (LEPIDOPTERA: LYCAENIDAE) Kurt Johnson Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10025 Abstract.— Yovmtrly monotypic Noreena Johnson, MacPherson & Ingraham (Theclinae, Eu- maeini) is revised to comprise a monophyletic group of nine species distributed from east- central Mexico to northwestern Argentina. Four new species are described, N. guianivaga (Guyana Shield), N. luxuriosa (Maranon area of endemism, Ecuador), N. pritzkeri and N. galactica (Rio de Janeiro area of endemism, Brazil), and four species are transferred from Theda, T. cambes Godman and Salvin, T. comana Hewitson, T. molena Jones and T. lemona Hewitson. Lectotypes are designated for T. lemona and T. molena. From ten eumaeine out- groups studied by numerical cladistic analysis (PAUP, Swofford), the sister genus Contrafacia is described, including four new species, C. rindgei (Sonora, Mexico), C. mexicana (montane central Mexico), C. australis (Paraguay/eastem Bolivia), and C. minutaea (Rio de Janeiro area of endemism, Brazil). The ^"orcynia'" and ‘‘^orios" species complexes of thecline grade genus Theda are indicated as the sister group of Noreena/ Contrafacia and their species composition further defined. Recently, two colleagues and I described a monotypic genus Noreena (Theclinae; Eumaeini) from upland xeric woodland habitats in northwestern Argentina (Johnson, MacPherson, and Ingraham, 1986). This taxon exhibited unusual characters for the tribe. Particularly, the female genitalia were laterally arched and the ductus seminalis disjunct from its usual point of attachment at the distal end of the ductus bursae. Such divergence was clearly suggestive of cladistic significance. However, at the time neither we nor reviewers knowledgeable of the Eumaeini could propose any congeners for the type species of Noreena, N. maria. Subsequently, I initiated a search for Noreena congeners amongst unidentified eumaeine samples and the nearly 750 eumaeine species presently placed in broadly polyphyletic “genus” Theda (Bridges, 1988). The study included specimens from the Allyn Museum of Entomology (AME), American Museum of Natural History (AMNH), British Museum (Natural History) (BMNH), Carnegie Museum of Natural History (CMNH), Field Museum of Natural History (FMNH), Institute Miguel Lillo, Tucuman, Argentina (IML), Milwaukee Public Museum (MPM), and Museum Na- tional d’Histoire Naturelle, Paris, France (MNHN). Four Theda taxa were found to share the generic characters of Noreena. These species were poorly represented in collections and seldom referenced in the systematic literature: T. cambes Godman & Salvin, T. comana Hewitson, T. lemona Hewitson and T. molena Jones. They had been included by Draudt (1919) with T. cupentus Cramer and T. lausus Cramer in his Theda ""cupentus Group.” Also, four undescribed species of Noreena (Guyana Shield, SW Ecuador, and SE Brazil) were discovered. 12 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) Species of Noreena are apparently seldom collected. The early faunal lists (Weeks, 1905; Kohler, 1923, 1928; Schweizer and Webster-Kay, 1941; Zikan and Zikan, 1968) do not include them and recent detailed faunal lists from Vera Cruz State, Mexico (Ross, 1975-1977; Llorente-Bousquets, Garces Medina and Martinez, 1986) do not record T. cambes, though the species is prevalent in early collections. Recent western Argentine collectors R. Eisele and B. MacPherson have collected only six specimens of N. maria in thirty years of concerted sampling. Two southeastern Brazilian species are known only from small samples from the turn of the century and a third from specimens taken exclusively by one collector at one locality. The taxa are apparently extremely habitat restricted in disparate desert, chaparral/chaco, and primary tropical forest biomes. Xeric environs are often poorly sampled by lepidopterists; primary tropical forest appears largely destroyed in the areas where Noreena specimens were originally collected (K. S. Brown, Jr., pers. comm.). To understand the Noreena assemblage among the Eumaeini, and explore the cladistic signihcance of its characters, numerical cladistic analysis (PAUP, Swofford, 1985) was used to study species of Noreena and representatives of ten eumaeine outgroups resembling Noreena. PHYLOGENETIC ANALYSIS OF NOREENA AND EUMAEINE OUTGROUPS Ten eumaeine groups were studied in relation to Noreena (Table [hereafter, “Tb.”] 1). Three included taxa grouped by Draudt (1919) with T. cambes (Tb. IF, G, H). Four included taxa listed by Johnson, MacPherson & Ingraham (1986) as morpho- logically similar to N. maria (Tb. IB, C, D, J). Others were included which had wing patterns or bipartite male forewing androconia (“brands” sensu Eliot, 1973) similar to N. maria (Tb. IE, I). Wing brand occurrence and similarity have been used by certain authors to group eumaeine taxa (Draudt, 1919; Clench, 1961; Field, 1967a, b). In addition, any apparently undescribed taxa resembling Noreena were also stud- ied. Taxa were dehned by examination of type specimens and other material (if dif- ferences between common usage and types indicated additional diversity). Species criteria were derived from standard taxonomic procedures involving consistent dif- ferences in characters of the wing, genitalia and tergal morphology. Taxon/character matrices were prepared and parsimonious distributions of characters constructed using PAUP (Swofford, 1 985). Various hypotheses of relationship were tested, rooting trees by different outgroups and by the Lundberg method (Swofford, 1985). From the ten eumaeine groups studied (Tb. 1 A-J) a monophyletic study set (Tb. 1 , Noreena + A-D) was delimited and its apomorphies (Tb. 2) delineated from the final rooted tree (Figs. 8, 9). This ingroup includes five terminal assemblages: Noreena, a new genus Contrafacia (described herein, including four previously undescribed species), a group of undescribed eumaeine taxa hereafter called “sister group X,” and certain taxa of the ""orcynia" and ""orios'" groups of Draudt (1919) (Tb. 1C, D). Noreena is indicated as sharing nine synapomorphies with Contrafacia (Chs. 1 , 2, 5, 8-12, 15). Noreena is distinguished by five autapomorphies (Chs. 19, 21, 22, 23, 25) and Contrafacia by three autapomorphies (Chs. 18, 20, 24). These include the outstanding modified eighth tergite in males (Chs. 1, 18, 19; Figs. 2, 6) and the laterally arched genitalia in females (Chs. 8-13, 20, 21; Figs. 4, 6) not present in other groups. Noreena and Contrafacia share three structural synapomorphies with 1989 NEOTROPICAI LYCAENIDAE 13 Table 1. Taxa of ingroup and outgroups.’ Ingroup: Noreena (type species N. maria plus new species described herein) plus A: Contrafacia (all new species described herein, Figs. 5, 6, lOB). B: sister group X (one undescribed species, Fig. 7B). C: ‘‘"orcynia Group” including Theda orcynia (Figs. 7A, IOC), T. catharina Draudt, T. ahola Hewitson, T. bassania Hewitson, T. marmoris Druce, T. aunia Hewitson, T. Cordelia Hewitson, T. anthracia Hewitson and two unde- scribed species. D: ""orios Group” including Theda orios (Fig. 7F), and a large number of unde- scribed species distributed from Mexico S to Argentina. Outgroups: E: ""spurina Group” including T. spurina Hewitson (Fig. 7C), T. ericusa Hewit- son (Fig. 7D), T. thoana Hewitson, and T. brescia Hewitson (Draudt placed these latter two in his "'brescia Group”); F: "thyesta Group” including T. lausus (Fig. 7E) (which Draudt placed with the taxa placed here in Noreena), T. pharus Druce (Fig. 7G), and T. radiatio Druce; G: "brescia Group” including T. lyde Godman & Salvin (Fig. 7H) (which Draudt considered similar to T. cupentus), and T cupentus (Fig. 71) (which Draudt included with the taxa placed here in Noreena). T cupentus appears to be one of an assemblage of species, the rest of which are undescribed; H: "avoca Group” including T. olbia (Fig. 7L) (which Draudt compared, in dis- cussion, to the taxa placed here in Noreena)-, I: "echion Group” including T. fabulla Hewitson (Fig. 7K), and T. philinna Hewitson. J: "atrana complex” (one group of taxa placed by Draudt, 1919 in his triphy- letic "americensis Group”) including T. atrana (Fig. 70) (type female, NMNH), T. tegaea (Fig. 7N) and T. tarania (Fig. 7M). ' Outgroups studied have, hitherto, all been placed "Theda'' (Draudt, 1919; Bridges, 1988). For historical purposes, this list cites the group names of Draudt (1919) but, since his groups were often nonmonophyletic, limits each group to taxa determined as probably monophyletic in the present study. Type material has been used in all cases except for taxa of Column D, since this group is under study by another worker. For this group representative material identified at AMNH by R. K. Robbins (NMNH) has been used. All types are BMNH unless otherwise indicated. Outgroups are not in any particular order but simply include taxa whose superficial similarity to Noreena required study (for instance, taxa in outgroup E are often placed in the genus Rekoa Kaye along with numerous taxa varying greatly from their general wing pattern). sister group X (Chs. 3, 7, 16; Figs. IG, 2, 6, 7B). Sister group X is presently monotypic, consisting of a morphologically unusual undescribed species with wing pattern similar to Theda orios Godman & Salvin. This wing pattern is indicated as plesiomorphic (Tb. 3, 31). Since it is anticipated that further members of sister group X will be discovered, it is not formally described here. Noreena, Contrafacia and sister group X share two structural synapomorphies (Chs. 4, 6) with taxa of the "orcynia Group” (Tb. 1C). Noreena and Contrafacia share with the "orcynia Group” the distinctive wing pattern which makes them superficially distinctive among the Eumaeini (the “split-stripe” on the hindwing under the surface, Chs. 17, 25; Fig. 10). The primitive wing pattern is maintained in sister group X 14 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) Table 2. Ingroup characters. The following describes apomorphic (A) and plesiomorphic (P) states for characters used for constructing cladogram in Figure 8, as listed in the matrix of Figure 9.' A. Characters of Figure 9 (e.g., apomorphic states shared by two or more terminal taxa in Fig. 8). Tergal morphology 1. Male, condition of eighth tergite: (A) eighth tergite with “incised posterior cavity” (sensuFidd, 1967a, b; Johnson, 1988; Johnson, Eisele and MacPherson, 1988; Johnson and Matusik, 1988) (Figs. 2, 6). (P) eighth tergite normal {sensu Ehrlich, 1958; Johnson and Matusik, 1988). Genitalia 2. Male, bilobed area of valvae (Fig. 3A): (A) bilobed area robust and strongly angled ventrad the caudal extension (Fig. 3); (P) bilobed area thin to mid-rimmed and generally parabolic (Figs. 6, 7A, B, E, F, I, M). 3. Male, caudal extension of valvae (Fig. 3 A); (A) caudal extension greatly thickened caudad bilobed area, tapering caudad with convex ridge defined along ventral inner margin of the lobes (Figs. 3, 6, 7B); (P) caudal extension greatly constricted terminad the bilobed area (Fig. 7A, C, D, E, F, G, H, I, K). 4. Male, saccus: (A) saccus radically elongate, cephalic expanse exceeding that of entire vincular arc (measured from base of saccus to basal juncture of uncus lobes) (Figs. 6, 7 A, B); (P) saccus short, cephalic expanse never greater, and usually much shorter, than that of entire vincular arc (measured as above) (Figs. 7C, D, E, F, G, H, I, M, O). 5. Male, terminus of saccus: (A) with emphatic terminal knob (Figs. 3, 6); (P) gradually tapered (Fig. 7A, C, D, E, F, G, H, I, M, O). 6. Male, vinculum: (A) ventro-caudal area of vincular arc with spurs abutting or over- lapping the juncture of valve’s bilobed area and caudal extension (Figs. 3, 6, 7A, B); (P) ventro-caudal area of vincular arc smooth and entire (Fig. 7C, D, E, F, G, H, I, K, M, O). 7. Male, vinculum: (A) ventrum of vinculum extremely compact, measure of entire edge not exceeding measure of entire edge of bilobed area of valvae (Figs. 3, 6, 7B); (P) ventrum of vinculum expansive, measure of entire edge exceeding (usually greatly) measure of entire edge of bilobed area of valvae (Fig. 7A, C, D, E, F, G, H, I, K, M, O). 8. Female, ductus bursae: (A) cephalic “ductus” (Fig. 3 A) strongly arched laterally (Figs. 4, 6); (P) cephalic ductus straight or displaced only slightly from plane of caudal “antrum” (Fig. 3A) (Fig. 7A, B, C, D). 9. Female, point of entry of ductus bursae into corpus bursae: (A) point of entry on centro- lateral surface of corpus bursae (Figs. 4, 6); (P) point of entry at distal end of corpus bursae (Fig. 7A, B, C, D, E, F, H, I, J, K, L, N). 10. Female, condition of juncture of ductus bursae and corpus bursae: (A) ductus bursae joins corpus bursae with variously expansive sclerotized “arms” (Figs. 4, 6); (P) ductus bursae joins corpus bursae in a flush manner, without additional sclerotization of the corpus bursae (Fig. 7A, B, C, D, E, F, H, I, J, K, L, N). 1 1 . Female, condition of corpus bursae and ductus seminalis: (A) ductus seminalis ema- nating from a sclerotized shield located on the lateral to disto-lateral surface of the corpus bursae (Figs. 4, 6); (P) ductus seminalis emanating from unsclerotized corpus bursae near flush juncture of ductus bursae and corpus bursae (Fig. 7A, B, C, D, E, F, H, I, L, N). 1 2. Female, condition of juncture between cephalic area of ductus bursae and antrum (Fig. 4 A): (A) transparent juncture, viewed from any angle, with marked “hour-glass”-like constriction (Figs. 4, 6); (P) transparent juncture not constricted (simply part of general, 1989 NEOTROPICAL LYCAENIDAE 15 Table 2. Continued. contiguous, cephalic tapering of ductus) (Fig. 7A, B, C, D). [Note: “transparent” is specified because this condition is indicated as apomorphic to a fully sclerotized con- dition in cladistic analysis among outgroups of the present study group (Fig. 7E-L), see Tb. 2, 9]. 13. Female, condition of sclerotization at juncture between cephalic area of ductus bursae and antrum (Fig. 4 A): (A) juncture of cephalic area of ductus and antrum with closely abutting, fully sclerotized, dorsal surfaces; folded transparent region ventrad (Fig. 7 A); (P) juncture fully sclerotized throughout (Fig. 7E-L). [Conditions oftaxa in Fig. 7B-D have been omitted here from characterization as 13A because, although it is certain that condition 13P is primitive, it is uncertain that the slightly different juncture in taxa of Fig. 7B, C, D is homologous with that of 7A, see Tb. 3, 10]. Internal secondary sexual characteristics 14. Male, vincular brush organ (Fig. 2B): (A) present [see entry 2] (all taxa of Figs. 3, 6); (P) absent. 15. Male, saccal brush organ (Fig. 2B): (A) present (all taxa of Figs. 3, 6); (P) brush organ along vinculum only [Note: to date, in the Eumaeini, saccal brush organs have only been found in taxa also having a vincular brush organ (Johnson, 1989)]. External secondary sexual characteristics 16. Male, dorsal forewing androconial structures (“brands”): (A) androconial on each wing with two sectors (“bipartite”), each occurring on the respective distal and basal sides of the crossvein of the discal cell (Fig. 1 H); (P) androconia absent. Wing pattern 1 7. Both sexes, pattern of hindwing under surface stripe: (A) stripe with two to three parallel, discal cell-end streaks, breaking stripe into caudal and cephalic elements (“split-stripe” of text) (Figs. 1, 5, 10); (P) stripe uniramous across entire wing. B. Characters with apomorphic state unique to one terminal taxon in Figure 8. 18. Male, condition of eighth tergite: (A) Dorsal plate of incised posterior cavity ovate, extending terminally from beneath seventh tergite (Fig. 6). (P) A, entry 1 . 19. Male, condition of eighth tergite: (A) Dorsal plate of incised posterior cavity elongate cephalad and incised or pronged, extending terminally from at least beneath sixth tergite (Fig. 2). (P) A, entry 1. 20. Female genitalia, condition of corpus bursae and ductus seminalis: (A) Sclerotized distal shield and point of attachment of ductus seminalis on corpus bursae conjoined proxad cephalic ductal terminus (Fig. 6). (P) A, entry 1 1 . 21. Female genitalia, condition of corpus bursae ductus seminalis: (A) Sclerotized distal shield and point of attachment of ductus seminalis on corpus bursae detached from juncture of ductus bursae and corpus bursae (Fig. 4); (P) A, entry 11. 22. Male, internal secondary sexual characteristics: (A) Terminus of saccal brush organ extending beyond proximal edge of vinculum and saccus; (P) A, entry 15. 23. Male genitalia: (A) Terminus of aedeagus recurvate (Fig. 3); (P) terminus of aedeagus straight. 24. Male genitalia, bilobed area of valvae: (A) Bilobed area of valvae constricted caudad saccus (Fig. 6); (P) P, entry 2. 25. Both sexes, under surface of hindwing: (A) Limbal spots postmedial-submarginal, cell CuA 1 and at base of anal lobe (Fig. 1 , 1 OA); (P) either or both of above spots reduced or absent (Fig. 5). ' Terminology for tergal and genitalic structures follows introductory section, notations in Figures 3-4, and Johnson (1976, 1978, 1988); additional references regarding some structures are provided as appropriate. 16 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) Table 3. Outgroup characters.' Tergal morphology 1 . Male, eighth tergite unspecialized (outgroups D, E, F, G, H, I): plesiomorphic. 2. Female, eighth tergite unspecialized (outgroups A, B, C, D, E, F, G, H, I): plesiomorphic. Genitalia 3. Male, caudal extension of valvae greatly constricted (Fig. 7 A, B, C, D, E, F, G, H, I, K) (outgroups F, G, H, I): homoplasic. 4. Male, caudal extension smoothly tapered to blunt ends (Fig. 7C, D) (outgroup E): plesio- morphic. 5. Male, bilobed configuration smoothly parabolic without notable sculpturing (Fig. 7C, D) (outgroup E): plesiomorphic. 6. Male, aedeagus caudally straight or only slightly curved (e.g., not radically recurvate cau- dad) (ougroups B, C, D, E, F, G, H, I): plesiomorphic. 7. Male, saccus reduced, greatly constricted, highly sculptured, cephalic expanse barely ex- ceeding cephalic arc of vinculum [(i) undescribed ""brescia Group” taxon like T. cupen- tus type in wing pattern but differing greatly in morphology; (ii) one undescribed species of the orios complex (outgroups D, G)]: homoplasic. 8. Male, saccus smoothly parabolic and without notable sculptured features (Fig. 7C, D, E, F, G, H, I, M, O) (outgroups E, F, G, H): plesiomorphic. 9. Female, sclerotized genital parts occurring as one uniformly sclerotized tapered tube (e.g., not divided into distinct cephalic ductal and caudal antrumal elements) (Fig. 7E, F, H, I, J, K, L) (outgroups F, G, H, I): plesiomorphic. 10. Female, sclerotized genital parts disjunct, occurring as cephalic ductal and caudal antru- mal elements separated by a transparent juncture (Figs. 4, 6, 7A, B, C, D) (outgroups B, C, E): apomorphic to character 9 but homoplasic in several outgroups relative to study set (Fig. 7N, O) (outgroup J). 1 1. Female, sclerotized genital parts as in 10, but oriented uniplanar or only slightly dis- placed laterad (Fig. 7 A, B) (outgroup E): homoplasic. 12. Female, sclerotized genital parts as in 10, but cephalic area of ductus displaced laterally such that juncture of ductus bursae and corpus bursae is slightly behind distal end of corpus bursae (Fig. 7C, D) (outgroup E): apomorphic to entry 9 but homoplasic relative to Tb. 2, Chs. 8-11). [Note that such homoplasy also includes similar, slightly displaced, junctures of the ductus bursae and corpus bursae in certain taxa without lateral displace- ment of the ductus bursae itself (Fig. 7H) (outgroup G). 13. Female, cephalic terminus of ductus bursae adjoining distal end of the corpus bursae (Fig. 7A, B, E-L) (outgroups B, C, F, H, I): plesiomorphic. 14. Female, cephalic terminus of ductus bursae adjoining corpus bursae slightly cephalad caudal end of corpus bursae (Fig. 7C, D, H) (outgroups E, G): homoplasic. 15. Female, corpus bursae without extending “arm” from cephalic terminus of ductus bur- sae (Fig. 7A-N) (outgroups B-I): plesiomorphic. 16. Female, corpus bursae distally unsclerotized (e.g., with no corpus bursae shield, either attached or detached) (Fig. 7C, D) (outgroups D, E, G, H) [Though distinctive of some groups, this condition varies greatly in others; thus it must be studied across the entire species diversity of any group to ascertain its value]; homoplasic. 18. Female, corpus bursae as in Tb. 2, Chs. 8-10 but lacking detached sclerotized shield on corpus bursae (Fig. 70) (outgroup J): homoplasic. Internal secondary sexual characteristics 19. Male genitalia without brush organs (outgroups F, G): plesiomorphic. 20. Male genitalia with vincular brush organ component only (Fig. 3B) (outgroups C, D, E, 1989 NEOTROPICAL LYCAENIDAE 17 Table 3. Continued. F, G, H, I): apomorphic to 19, but plesiomorphic to saccal brush organ (Fig. 3B, No- reena and Contrafacia only). 2 1 . Male genitalia with small dorsal vincular brush organ (attached beneath dorsa-caudad arch of vinculum and barely extending to base of falces) (outgroup G): homoplasic. External secondary sexual characteristics 22. Male, forewings, dorsal androconia (“brands”) on each wing with two sectors (“bipar- tite”), each occurring on the respective distal and basal sides of the crossvein of the dis- cal cell (Fig. IH) (outgroups E, I): homoplasic. 23. Androconial brand as in 22, but with distal sector located more vertically over basal sector (e.g., poised more adjacent radial area of discal cell) (outgroup E): homoplasic. 24. Bipartite androconial brand with sectors non-adjacent and divided widely above and be- low the costal vein of the apical end of discal cell (outgroup I): homoplasic. 25. Bipartite androconial brand divided into cephalic and caudal sectors but within the dis- cal cell (outlying or edging scales may lie variously on the veins of the cell) (outgroup F): homoplasic. 26. Androconial brand bipartite but as two concentric circles inside the apex of discal cell (outgroup G): homoplasic. 27. Androconial brand of each wing undivided (outgroup H): apomorphic to no androconia; plesiomorphic to divided androconia. Wing pattern 28. Under surface wing pattern, hindwing medial band with centrally broken or laterally dis- junct pattern resembling the “split-stripe” of Noreena (Figs. 1, lOA) (outgroups F, G): homoplasic. 29. Upper surface wing pattern, suffused with iridescent coloration fading gradually into dis- tal fuscous (various taxa in outgroups B-I): plesiomorphic relative to distinct patches of iridescent blue. 30. Upper surface iridescent coloration occurring in distinct patches (Figs. IH, 4C) (out- groups F, G, H): homoplasic. 31. Under surface wing pattern, medial or postmedial band not centrally broken (ingroup sister group X, outgroups D, E, F, G): plesiomorphic to ingroup; homoplasic to condi- tion in 28. ' Outstanding characters in outgroups superficially resembling conditions in the ingroup but indicated as relatively plesiomorphic or homoplastic. , 18 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) Fig. 1 . Species of Noreena. Previously described species, A-E, wing under surfaces: A. N. maria (type species), 9, Cornejo, Salta, Argentina (AMNH); B. N. cambes, 6, Presidio, Veracruz, Mexico (AMNH); C. N. comana, holotype 3 [B&W reproduction of color photo]; D. N. molena, syntype 6 [B&W reproduction of Jones (1912) color plate]; E. N. lemona, lectotype 9 [B&W reproduction of color photo]. New species, F-I, upper surface (left), under surface (right): F. N. guianivaga, holotype 3; G. N. luxuriosa, holotype S (bipartite androconial brand illustrated diagrammatically on left forewing); H. N. pritzkeri, holotype 3; I. N. galactica, holotype S. 1989 NEOTROPICAL LYCAENIDAE 19 Fig. 2. Specialized eighth tergites of Noreena. A. N. cambes, holotype; B. N. comana, ho- lotype; C. N. guianivaga, holotype; D. N. luxuriosa, holotype; E. N. pritzkeri, holotype (MPM); F. N. molena, syntype; G. N. maria, paratype, Mosconi, Argentina (AME); H. N. lemona, Castro, Parana, Brazil (BMNH); I. N. galactica, holotype (MPM). (Tb. 3, 31). Thus, the sister group of Noreena, Contrafacia and sister group X is the ""orcynia Group,” not the branded taxa which had been associated with T. cambes by Draudt (1919). Taxa of the "‘‘orcynia Group” lack male forewing brands (basal view, Fig. IOC). It is apparent from the present study that, in some groups of Eumaeini, occurrence of such external secondary sexual characters (Figs. 1, 5, 10) is of limited cladistic sig- nificance (Ch. 16; Figs. 8, 9; Tb. 3, 22-27). Accordingly, taxa of Draudt’ s (1919) ""spurina Group” also do not appear to be closely related to Noreena. The laterally arched ductus bursae in Noreena and Contrafacia (Chs 8-13; Figs. 4, 6) appears to have evolved from the centrally constricted, but uniplanar, ductal structure of the ""orcynia"" and ""orios" groups (Tb. 3, 10, 12, 15, 16; Fig. 7A, F). As indicated using ""spurina Group” taxa as an outgroup for rooting in Figures 8-9, the slightly curvate ductus bursae in ""spurina Group” taxa (Tb. 3, 10, 11, 12, 14, 16; Fig. 7C, D) appears to be homoplasic since these taxa lack all other derived characters of the ingroup. As in sister group X, the simple, postmedial under surface hindwing band occurring in the ^^spurina Group” (and many other Eumaeini) is indicated as plesiomorphic (Tb. 3, 31). The other member of the ingroup of this study is the ""orios Group” (Tb. ID). Taxa 20 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) Fig. 3. Male genitalia of Noreena, ventral view with aedeagus removed and placed vertically alongside. A. N. cambes, holotype; B. N. comana, holotype; C. N. guianivaga, holotype; D. luxuriosa, holotype: E. N. lemona, Castro, Parana, Brazil (BMNH); F. N. molena, syntype; N. maria, paratype, Mosconi, Argentina (AME); H. N. pritzkeri, holotype; I. N. galactica, holotype. p 1989 NEOTROPICAL LYCAENIDAE 21 Fig. 4. Female genitalia of Noreena. A. N. maria, holotype; B. N. cambes, Urualoana, Mexico (AMNH); C. N. comana, Canal Zone, Panama (MNHN); D. N. guianivaga, paratype; E. N. luxuriosa, paratype; F. N. molena, lectotype; G. N. lemona, lectotype; H. N. pritzkeri, allotype. of this group have bipartite male forewing brands and a simple under surface hindwing postmedial band like the ""spurina Group.” Structurally, however, taxa of the "'orios Group” (Fig. 7F) belong in the clade including Noreena, Contrafacia, sister group X and the ""orcynia Group” (Figs. 8, 9). For purposes of this study, the "'orios Group” includes T. orios (as defined by a type, BMNH, Fig. 7F) and a large number of undescribed taxa. The type locality for T. orios is “Guatemala” and the species appears insular. Superficially similar butterflies occur from Honduras to southern Brazil, but vary greatly in morphology. Since cladistic study of these undescribed species will probably result in additional basal branching below the ""orcynia Group” of Figure 8, the ""orios Group” as defined here is probably paraphyletic. Resolution of relationships in this largely undescribed group will be an important future project. 22 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) Fig. 5. Species of Contrafacia, upper surface (left), under surface (right) (except D, F op- posite). A. C. rindgei, holotype $; B. C. mexicana, allotype C. C. mexicana, holotype 9; D. C. australis, allotype 5; E. C. australis, holotype 9; F. C. minutaea, holotype 9. Cladistic analysis in this study indicates many characters used by early workers for grouping eumaeine taxa had little cladistic significance. Relative to a particular cladistic ingroup, many such characters are either primitive or, because of homoplasy, useful for clustering only a few taxa. Table III lists, and cross-references to illustrations from type specimens (Fig. 7), many features previously thought significant for group- 1989 NEOTROPICAL LYCAENIDAE 23 Fig. 6. Tergal morphology and genitalia of Contrafacia. A. C. mexicana, allotype 3; B. C. mexicana, holotype 9; C. C. minutaea, holotype 9; D. C. australis, allotype 5; E. C. australis, holotype 9; F. C. rindgei, holotype 9. ing Neotropical hairsteak butterflies (Draudt, 1919; Clench, 1961; Field, 1967a, b; Bridges, 1988). The table indicates plesiomorphy and/or homoplasy relative to char- acters defining the ingroup of the present study. Of particular interest, and reviewed briefly below, are several parallelisms in extremely salient structures. There are other eumaeine taxa with laterally arched female genitalia. Some of these taxa share no other characters with ingroup or outgroup taxa of the present study. Most notable is the Theda atrana complex (Tb. IJ; Tb. 3, 18, Fig. 7N, O). Draudt (1919) used wing pattern similarity to group these taxa in this Theda ""americensis Group.” Johnson, Miller and Herrera (ms.) demonstrate this group is triphyletic; the 24 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) Fig. 7. Morphology of outgroups (males and females both ventral view, unless indicated otherwise). A. Theda orcynia, left: syntype 3; right: syntype 9 (BMNH) (lateral view of ductus, far right). B. undescribed species of sister group X: left: 3, Caripito, Venezuela (AMNH); right: 9, same data. C. T. spurina, left: 3, Igarapi-Assu, Brazil (AMNH); center: 9, same data (AMNH); right: cephalic ductus terminus and fanlike bursal shield in undescribed Andean species of spurina Group (see Discussion), Cucho, Argentina (nr. Bolivia) (AMNH). D. T. ericusa, left: 3, Hololo Mt. Road, Trinidad-Tobago (AMNH); right: 9, Oropoche, Trinidad-Tobago (AMNH). E. T. lausus, left: lectotype 9 (Johnson and Matusik, 1988) (BMNH); right: 3, Managua, Nic- aragua (CMNH) (i.d. and genitalic preparation, H. K. Clench). F. T. orios, left: syntype 3 (BMNH); right: 9, Tenedores, Guatemala (AMNH). G. T. pharus, left: holotype 3 (BMNH). H. T. lyde, left: syntype 3 (BMNH); right: syntype 9 (BMNH) (lateral view, cephalic ductus terminus, far right). I. T. cupentus, left: syntype 3 (BMNH); right: syntype 3 (BMNH). J. T. thyesta, syntype 1989 NEOTROPICAL LYCAENIDAE 25 Fig. 8. Cladogram of Noreena, Contrafacia and relatives. Cladogram derived from parsi- monious distribution of unweighted characters of Figure 9 rooted by concensus between the Lundberg rooting method and outgroup rooting using outgroups E, F and G (Tb. 1 ) (Consistency Index = .850). Uniquely derived apomorphies are specified by the horizontal bars on intemodes, and represent characters described in Tb. 2 and listed in the matrix of Figure 9. Since data for this cladogram are limited to the character matrix of Figure 9 and Tb. 2, autapomorphies are specified only for the two terminal genera revised in this study. lateral arch in female genitalia of the atrana complex is of independent derivation. There is also slight lateral displacement of the ductus bursae in some taxa of the ""spurina Group” (Tb. IE; Fig. 7C, D). Such lateral displacement causes the ductus bursae to join the corpus bursae just behind the caudal end of the bursal sac. This condition also occurs in Theda lyde of Draudt’s Theda '"brescia Group” (Fig. 7H) and in other, undescribed, taxa of the ""spurina Group.” These structures vary from a simple, flush juncture (as in the taxa of Fig. 7C, D) to a fanlike sclerotized structure extending caudad along the bursal sac (as in Fig. 7C, right). Significantly, the latter structure, which occurs in an undescribed "^spurina Group” species from the north- western Argentine and Bolivian Andes, is also typical of many other Eumaeini— taxa among the seven genera of Callophryina (Johnson, 1981) and certain species of Strymon Hubner (Johnson, Eisele & MacPherson, 1989). Neither callophryines, nor Strymon taxa, are closely related to the "^spurina Group” (Johnson, MacPherson and Ingraham, 1986). This assessment of homoplasy in the various, slightly displaced. 9 (BMNH). K. T.fabulla, left: syntype 3 (BMNH); right: syntype 9 (BMNH). L. T. olbia, holotype 9 (BMNH). M. T. tarania Hewitson, holotype $ (BMNH). N. T. tegaea, syntype 9 (BMNH). O. T. atrana, left: 5, Castro, Parana, Brazil (BMNH); right: holotype 9 (NMNH). 26 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) orios orcynia sister Contra- Group Group group X facia Noreena 1. 2 . 3. 4. 5. 6. 7 . 8. 9. 10. 11. 12 . 13 . 14. 15. 16. 17. 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 0 0 0 0 1 0 1 0 0 0 0 0 0 1 1 0 0 1 0 0 1 1 0 1 1 0 0 0 0 0 1 1 0 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Fig. 9. Character state matrix used for cladogram construction (Fig. 8) of Noreena, Con- trafacia, and relatives (plesiomorphic states of Tb. 2 = 0, apomorphic states of Tb. 2=1). Outgroup consists of groups E, F and G (Tb. 1 ) (from study of their types and other specimens) scored all 0 (characters 1 and 1 1 could be scored as 1 if based on exceptional taxa in a minority of E, F or G). Characters for Lundberg rooting of parsimonious network scored all 0. ductal structures in ""spurina Group” is further supported by the normal eighth tergite, and vincular characters, ofspurina Group” males (Fig. 7C, D). The latter structures do not indicate cladistic affinity to the radically modified eighth tergite in Noreena or Contrafacia, on the constricted vinculum and elongate saccus characteristic of Contrafacia, sister group X and the ""orcynia Group.” From this analysis it is apparent that bipartite androconial brands and laterally arched female genitalia have arisen independently in several groups of the Eumaeini. Bipartite androconial brands vary greatly within small eumaeine assemblages and thus may be of little use as taxonomic characters except within genera or species groups. On the other hand, morphological innovation in disparately evolved laterally arched female genitalia is far more diverse and such structures appear taxonomically useful for differentiating larger assemblages of taxa. Species criteria and distributions. Nine Noreena taxa are accorded species status and represent six allopatric areas of endemism. Four species are regionally sympatric in SE Brazil, two on the Guyana Shield, and it is possible that the Central TAXONOMY OF NOREENA AND CONTRAFACIA 1989 NEOTROPICAL LYCAENIDAE 27 Fig. 10. The “split-stripe” (Ch. 17) and “limbal spots” (Ch. 25) characteristic of Noreena, Contrafacia and the orcynia Group. A. Noreena (N. pritzkeri shown here)— costal portion of medial band junctures variously with parallel cell-end streaks: (i) basal in N. pritzkeri, (ii) at central cell-end streak in most other Noreena taxa, (iii) at distal cell-end streak in N. molena. All taxa have two limbal spots, postmedial to submarginal, cell CuAl, and at base of anal lobe. B. Contrafacia (C. australis shown here)— medial band junctures to distal cell-end streak; CuAl spot (submarginal) and spot at base of anal lobe often reduced or latter missing. C. orcynia complex— medial band junctures to distal cell-end streak; CuAl spot (submarginal) apparent but base of anal lobe often not emphatically marked {Theda ahola, shown here [Orizaba, Mexico, AMNH], also exhibits postbasal stripes characterizing some species of this complex). American/northem South American taxa {N. cambes and N. comand) are sympatric at the Isthmus of Panama. In wing pattern, four species of Noreena are disparately marked; N. molena, N. lemona and N. galactica of SE Brazil and N. guianivaga of the Guyana Shield. The other species (A^. maria, N. cambes, N. comana, N. luxuriosa and N. pritzkeri) are part of a more superficially similar complex of allopatric pop- ulations. Structurally, members of this latter complex diverge as much from each other as from the disparately patterned species. Such divergence probably results from the extreme habitat restriction typifying all of these taxa. Structural divergence in the group exceeds that of eumaeines whose pan-Neotropical allopatric segregations are usually considered complexes of subspecies (see, for instance, Chlorostrymon, Johnson, 1989). Consequently, along with apparent sympatric species, I treat widely distributed allopatric populations of Noreena as species, if morphologically distinct. This view could be modified when enough neotropical Eumaeine groups are revised to allow some overall assessment of nomenclatural status and allopatric divergence across many groups. Contrafacia includes four distinctively marked new species representing three al- lopatric areas of endemism. These species are included in a new genus because they are distinguished by three autapomorphies and, like Noreena, encompass a Pan- 28 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97( 1 ) Neotropical distribution. Contrafacia taxa are currently known from few specimens but it is anticipated that future research will increase knowledge both of species diversity and geographic distribution. At present, one species is known from desert habitat in Sonora, Mexico; another occurs in xeric montane central Mexico and, as in Noreena, two species are sympatric in SE Brazil and Paraguay. Format and terminology. Taxa of Noreena and Contrafacia exhibit a number of wing and morphological features not previously characterized in the literature. Ac- cordingly, in the following diagnostic text, I introduce a number of descriptive terms and phrases used subsequently throughout the taxonomic entries, tables and figures. Aside from these, terminology follows Johnson (1976, 1978, 1988) and Johnson and Matusik (1988). In descriptions and discussions, taxonomic characters are cross- referenced with Tb. 2 and Figures 8 and 9 by referring to character number as “Ch.'#,'” etc., and such references are given only once per taxonomic entry. Cross references in the tables are also provided by notations for table number (“Tb. '#'”) and outgroup number (“Outg. referring to taxa listed in Tb. 1). In the revisionary treatments, species are arranged in north to south geographic order. Noreena Johnson, MacPherson and Ingraham Figs. 1-4, 10 Noreena K. Johnson, MacPherson and Ingraham, 1986:2 Johnson 1988:34; Bridges 1988:11.78. Diagnosis. From other Eumaeine, Noreena can be superficially recognized by the wings’ upper surface “bipartite” scent brands (males, Ch. 16, Fig. IG) and iridescent blue to silvery blue coloration (both sexes) combined with an under surface hindwing medial band split into costal and anal elements by two to three disjunct stripes in the discal cell (the “split-stripe,” Ch. 17, Figs. 1, lOA). In addition, on the hindwing undersurface are bright, red to orange, “limbal spots” in cell CuAl (postmedian to submarginal) and at the base of the anal lobe (Fig. lOA). Morphologically, the genus is characterized by autapomorphic characters 19, 21-23, and 25 of Tb. 2. Description. Johnson et al., 1986, p. 2-3, illustrated in the present study in Figures 1 (imago), 2-3 (male morphology), and 4 (female morphology). Types Species. Noreena maria K. Johnson, MacPherson & Ingraham (Figs. lA, 2G, 3G, 4A) by original designation. Distribution. Nine species distributed from east-central Mexico southward to north- western Argentina. Noreena cambes (Godman & Salvin), new combination Figs. IB, 2 A, 3 A, 4B Theda cambes Godman and Salvin 1879-1901 [1887], vol. 2:53, vol. 3: pi. 54, f 16-18; Draudt 1919:769, pi. 154e; Schaus 1920:176; Comstock and Huntington 1958-1964 [1959]:173, [1964]:64, Bridges 1988:1.68, 11.78; Theda syvix Dyar 1918:3; Schaus 1920:176; Comstock and Huntington 1958-1964 [1959]:174, [1964]:64; Bridges 1988:1.68, 1.338, 11.78. Diagnosis. Above, both sexes silvery blue, female hindwing often uniquely bold blue. Beneath, concolorous, split-stripe and submarginal bands only moderately dis- tinctive, fading in anal area with limbal markings obsolescent. Male androconial 1989 NEOTROPICAL LYCAENIDAE 29 brands and tergal and genital morphologies distinctive as detailed below. Known from Mexico to Costa Rica. Description. MALE: Upper surface of wings: forewing dull blue on basal two thirds, distally fuscous. Bipartite androconial brand with distal sector equilaterally triangular, about half the size of parabolic basal sector. Tail of medium length at terminus of vein CuA2, shorter tail at CuAl. Hindwing dull gray-blue except for darker fuscous base and costal area. Under surface of wings (Fig. 1 B): ground color uniformly brown- hued gray. Forewing with white postmedial line, costa to cell CuAl, angled caudo- basad, not gently rounded; submarginal area with vague white intercellular lines, costa to vein CuA2. Hindwing ground concolorous but markedly obsolescent in limbal area. Medial split stripe and cell-end streaks not outstanding, each whitish and equally emphatic with companion whitish submarginal band extending from costa to cell CuA2; black marginal spots parallel submarginal band in cell interspaces from costa to anal margin. All pattern elements increasingly obsolescent toward anal lobe. Limbal spots orange, dull in cell CuA2, vivid at anal lobe. Length of forewing: X of six males (AME, AMNH) 13.3 mm, range 13.0-13.5 mm. FEMALE: Upper surface of wings: similar to male but with blue much duller on both wings; no androconial brand. Under surface of wings: as on males. Length of forewing: x of three females (AME, AMNH, BMNH) 13.7 mm, range 13.5-14.0 mm. MALE TER- GAL MORPHOLOGY AND GENITALIA: Figures 2A, 3 A. Incised posterior cavity extending only through the seventh abdominal segment, varying infraspecifically in length of cephalic prongs {N. cambes thus sharing shortest incised cavity of the genus with N. molena and N. guianivaga, these two species having cephalic lobes, not prongs). Genitalia, though otherwise typical of genus, only in N. cambes and N. comana showing little tripartite sculpturing at margin of saccus and valvae, N. cambes with margin of valval bilobed configuration irregular. FEMALE GENITALIA: Figure 4B. Cephalic component strongly angled distad, not arched or inclined as in con- geners. Lamellae postvaginalis extremely serrate; apophyses of the papillae analyses shortest of genus. Types. Holotype, 5, BMNH, labelled “Type, Sp. figured, Theda cambes, G & S. B.C.A. Lep. Rhop., Godman-Salvin Coll. 191 1-93” “Cordova, Vera Cruz. Rumeli, BM Type Lep. Rh. 693.” Holotype, 5, T. syvix. National Museum of Natural History (NMNH), labeled “Presidio, Mexico, December 1913, type no. 19,253.” Distribution. Spatial: from Vera Cruz State, Mexico southward at least to Costa Rica (see below). Temporal: specimens (see below) are known from nearly every month of the year. Remarks. A label on the holotype indicates it was figured in Draudt (1919). How- ever, this color figure differs significantly from the holotype, particularly in omitting the CuAl limbal spot and not clearly depicting pattern obsolescence in the limbal area. I have been unable to study the type of T. syvix (NMNH) but its description is unambiguous and it has been listed as a synonym by Comstock and Huntington (1958-1964), Bridges (1988) and correspondence by R. K. Robbins (NMNH) to Bridges (1988, p. IV.76). Material examined. COSTA RICA, San Jose, 28 April, H. Schmidt, \6 (BMNH), “Costa Rica,” A.G.M. Gillott, 1929, 19, (BMNH); MEXICO, Presidio, Vera Cruz State, June 1939, C. C. Hoffman, 16 (AMNH), 12 July 1945 (AME); Urualoana, May, C. C. Hoffman, 19 (AMNH); Catemaco, Vera Cruz State, August 1962, T. 30 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) Escalante, 16 (AME); Santeco Mapan, Vera Cruz State, May 1955, T. Escalante, \6, (AME); Rincon, Guerrero State, 2,800 ft, September, H. Druce, 1(3 (BMNH); Colima, S Mexico, one male, 19 (CMNH). Specimens in NMNH assumed to be N. cambes by distribution (R. Robbins, pers. comm.)— MEXICO, Coatepec, male: Jalapa, 3(3, 19 [one labelled February]; Presidio, December 1913 (type of T. syvix)\ Vera Cruz State, Cordoba, 31 July 1955; COSTA RICA, San Jose, 19. Godman & Salvin (1879- 1901 [1887]) list Cordova, Mexico and Polochic Valley, Guatemala. They question the association of their figured female because it is blue on both wings above; I have seen no completely blue female of N. cambes, but since many are fully blue on the hindwing such wider blue coloration m^ay be possible. Noreena comana (Hewitson), new combination Figs. 1C, 2B, 3B, 4C Theda comana Hewitson 1863-1878 [1867], vol. 1:97, vol. 2: pi. 36, f 87, 88 [or 86, 87]; Kirby 1871:388; Weeks 1911: xiv; Draudt 1919:769, pi. 154e; Comstock and Huntington 1958-1964 [1959]:192, [1962]:107; Bridges 1988:1.68, 11.78. Theda peraltd Moschler 1883:308, pi. 17, f 1; Draudt 1919:769; Comstock and Huntington, 1958-1964 [1959]:192, [1962]:107; Bridges 1988:1.272, 1.68, 11.78. Diagnosis. Above, both sexes silvery blue, less distinctive in female. Beneath, split- stripe emphatically marked, ground color distad and basad strongly contrasted gray and brown, respectively; all limbal and anal markings emphatic, ground nowhere strongly suffused. Male androconial brands and tergal and genital morphologies dis- tinctive as detailed below. Known from Panama through northern South America (see below). Description. MALE: Upper surface of wings: forewing dull blue on basal two thirds, distally fuscous; hindwing dull blue at base, silvery blue on distal two-thirds, margins and costal area fuscous. Bipartite male androconial brand with sectors of about equal size, distal sector nearly oval, basal sector as isosceles triangle. Long tail at terminus of vein CuA2; shorter tail at vein CuAl. Under surface of wings: forewing ground color basally deep brown, distally bright light gray; vivid white postmedial line, costa to cell CuAl, patchy white submarginal line, costa to cell CuA2; hindwing basally deep brown contrasting distal bright gray separated by vivid white split-stripe with emphatic cell-end streaks. Limbal area bright gray, emphatic black spots in cells across entire wing, each with white chevron-like markings at the base; limbal spots bright red-orange blotches. Length of forewing: x of three males (AME, AMNH, BMNH) 12.7 mm, range 12.0-13.0 mm. FEMALE: Upper surface of wings: marked similarly to male but with hindwing duller blue. Under surface of wings: as of males. Length of forewing: one female, MNHN, 12.5 mm. MALE TERGAL MORPHOL- OGY AND GENITALIA: Figures 2B, 3B. Incised posterior cavity generally elongate and extending cephalad beneath sixth and seventh segments; lateral edges with central indentation; cephalad prongs widely bifurcate with rounded ends. Genitalia resemble N. cambes most, but bilobed area in N. comana is smoothly parabolic, not angled along the distal edge. Aedeagus remarkably short, length less than 1.2 times that of entire genitalia from labides tip to terminus of saccus and with more than one-third of the terminal aedeagal length recurved {N. cambes aedeagus length exceeding 1.5 times that of entire genitalia and with less than the terminal one-fourth recurved). N. comana lacks the vincular process for abutment of the brush organs prominent 1989 NEOTROPICAL LYCAENIDAE 31 on N. cambes. FEMALE GENITALIA: Figure 5C. Caudal component elongate with rectangular, flaplike lamellae; cephalic component broadly arched with lateral at- tachment to corpus bursae limitedly sclerotized across center of corpus bursae; de- tached sclerotized shield parabolic, curving slightly proxad cephalic component of ductus. Due to format restriction, corpus bursae of Figure 4C shown folded under. Types. Holotype $ (Fig. 1C), BMNH, labelled “Tapajos, Amazons, H. W. Bates, Coll. Godman-Salvin, B.M. type No. 642.” Holotype, 9, T. peralta, type locality, Paramaribo, Surinam, deposition unknown (Comstock and Huntington, 1958-1964 [1962]). Distribution. Spatial: Widely distributed from Panama across northern South America southward to the Amazon River. Temporal: specimens represent nearly every month of the year. Remarks. Week’s (1911) reference to N. comana in a brief list of Venezuelan specimens collected is the only reference to this species in the classic early faunal lists (see Introduction). With the range of N. cambes being principally Mexican but extending at least to Costa Rica and that of N. comana extending north to Panama, their overall similarity may prove to represent ends of a dine. However, considering the long history of trans-Panamanian disjunctions and the compelling differences in the incised posterior cavities and certain genitalic characters of presently known specimens, it seems more likely the two may prove to be sympatric species in some areas of their respective southward and northward ranges. The specimen figured by Draudt (1919) does not show the bold split-stripe and drastic basal/distal ground color contrast (though somewhat variable) typical of most N. comana and the holotype. Deposition of the type of T. peralta is unknown but its description appears un- ambiguous. It has been listed as a synonym by Draudt (1919), Comstock and Hun- tington (1958-1964) and Bridges (1988). Material examined. BRAZIL, [see type data]; “Vinea, Amazon,” 16 (BMNH); COLOMBIA, Puerto Atlantico, 1 1-12 July 1920, dry hills in thick scrub, 15 (CMNH); PANAMA, La Boca, Canal Zone, 24 January 1908 (AMNH), 15, Canal Zone, 25, 19 (MNHN). TRINIDAD, Maupartius, November 1923 R. Dick (AME), 15; VEN- EZUELA, Naiguata, Federal District, 29 August 1937, Lichy, (AME), 15; Caracas, 15 (MNHN); specimens in NMNH presumed to be N. comana by distribution but not available for study— PANAMA, Canal Zone, Paraiso, 2 June 1979, 25, Panama Province, Cerro Campana, 19, 5 August 1977, San Carlos, 15, 21 July 1973; CO- LOMBIA, Meta, Villavicencio, 55, 49, 4-31 July 1972. Noreena guianivaga, new species Figs. IF, 2C, 3C, 4D Diagnosis. Above, male deep steel blue with thin black marginal lines, distal sector of androconial brands apparent only along outer edge of basal sector; female brown. Beneath, both sexes deep, concolorous, chocolate brown with a bright white costal suffusion framing a deep brown costal patch ground; split-stripe thin, vividly incised gray-white (black basad), with cell-end streaks, submarginal band, and limbal spots the most emphatic of the genus. Tergal and genital morphologies distinctive as detailed below. Known only from localities on the Guyana Shield. Description. MALE: Upper surface of wings: ground color dull deep blue iridescent 32 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) throughout except for blackish marginal lines. Bipartite and androconial patch par- abolicly elongate, light distal component comprising only about one-fourth the length of the remaining dark-colored brand (in N. luxuriosa basal sector is large and oval with distal sector only about one quarter its size and both curvate). Long tail at terminus of vein CuA2, shorter tail at vein CuAl. Under surface of wings: ground, both wings, deep chocolate brown. Forewing with white-blotched postmedian line, costa to cell M3, submarginal line obsolescent. Hindwing split-stripe vivid over dark concolorous ground; prominent bright gray-white distal suffusion in cells Ml and M2; three prominent cell-end streaks. Limbal pattern of concise, white-edged sub- marginal black line, anal area to costa. Limbal spots deep red. Length of forewing: 16.0 mm (holotype). FEMALE: Upper surface or wings: ground color dull brown; lacking androconial brand. Under surface of wings: as on male but undersurface ground color slightly lighter, pattern elements more vivid but costal hindwing suf- fusion reduced. Length of forewing: 16.0 mm (paratype, BMNH). MALE TERGAL MORPHOLOGY AND GENITALIA: Figures 2C, 3C. Incised posterior cavity small- er than on any congener except N. cambes and N. molena, extending only through the seventh abdominal segment. Contrasting N. cambes, but like N. molena, cavity dorsal plate with no cephalad pointing prongs, only two gently rounded lobes. Gen- italia distinctive in tear-drop shaped ventral valval configuration and extreme con- striction of valval terminus. Vincular brush organs in thick bundle, abutting wide area of dorso-cephalic region of vincular arc; saccal brush organs in thick strip anchored between caudal and distal margins of saccus. FEMALE GENITALIA: Figure 4D. Genital plate distinctly massive, both components quite wide relative to length (length of caudad component below lamellae, 2.2 x maximum width; length of cephalad component proxad corpus bursae 3.0 x maximum width). Lamellae angled distad; prominent centrad lobe on the lamella antevaginalis. Types. Holotype <5 (Fig. IF, G), labelled “Caripito, Venezuela, 23 August 1942,” “collection New York Zoological Society, Tropical Research Department,” deposited AMNH. Allotype, 9, labelled “Guyana Frangaise, C. Bar,” deposited BMNH. Para- types, 2$, same data as allotype, BMNH; 16, Upper Putumayo, S.E. Colombia, MNHN. Distribution. Spatial: presently known only from the type locality and the gener- alized data of the paratypes. Temporal: known only from 23 August (holotype). Remarks. It appears biogeographically significant that the type locality of N. gui- anivaga is the same as that of the recently described Heraclides matusiki Johnson and Rozycki (1986) (Papilionidae) which is presently known from a single specimen. The immediate sister species of each of these is western Andean in distribution (see H. isidorus (Doubleday), Johnson and Rozycki, 1986 and N. luxuriosa, below). Etymology. The name, using the Latin suffix "^vaga"" means “Guiana roamer.” Noreena luxuriosa, new species Figs. IG, 2D, 3D, 4E Diagnosis. Above, male iridescence brilliant and in distinct baso-medial patches, androconial brands with basal section abnormally large and ovate; female shiny brown; tails very long, costa vivid red from forewing base beyond expanse of discal cell. Beneath, ground concolorous medium brown, split-stripe vivid, with submar- ginal bands of both wings “exotically” scalloped basad and distad each vein. Male androconial brands and tergal and genital morphologies distinctive as detailed below. 1989 NEOTROPICAL LYCAENIDAE 33 Description. MALE: Upper surface of wings: ground color dominated by patches of brilliant blue— forewing from base to distinct juncture with postmedian fuscous, hindwing from postbasal area to margin; outer margin of wings fuscous, fuscous blotches at margin in cells Ml, M2 and M3. Bipartite androconial brand with basal sector large and ovate, distad sector only about one-third this size and oblong in shape. Very long tail at terminus of vein CuA2, somewhat shorter tail at vein CuAl. Under surface of wings: ground, both wings, deep chocolate brown. Forewing post- medium line scalloped basad and distad along each vein, extending from costa to cell CuA 1 . Hindwing split-stripe vivid, extremely angled along the band (particularly toward anal margin) and with bright postmedian line scalloped basad and distad along each vein. Limbal spots deep red. Length of forewing: 16.0 mm (holotype). FEMALE: Upper surface or wings: ground color shiny brown; faint marginal blue- gray hue; no androconial brand. Under surface of wings: as on male. Length of forewing: 16.0 mm (allotype). MALE TERGAL MORPHOLOGY AND GENI- TALIA: Figures 2D, 3D. Incised posterior cavity large, extending well beneath sixth abdominal segment, widely tear-drop shaped with short rounded prongs formed by a cephalic indentation. Male genitalia with extremely thick terminally knobbed saccus and a rectangular vincular configuration. Valvae widely angled at base, tapering thickly to broad terminus. Vincular brush organs abutting a long triangular flap protruding from a large dorsal vincular spur. Vincular brush cluster consequently thin compared to bulbous clusters of congeners (latter abutting either an ovate flap or occurring variously along the dorsal edge of the vincular arc). Aedeagus elongate (length nearly one-third more than entire length of the genitalia); terminal third recurvate. FEMALE GENITALIA: Figure 4E. Genital plate massive, similar to N. guianivaga, but with central lobe of lamella antevaginalis more prominent. Compared to other congeners, cephalic component very thin, elongately arched so as to signif- icantly displace the corpus bursae laterally. Arched cephalic component adjoining corpus bursae with a fingerlike extension about one-half the lateral expanse of the bursae. Sclerotized shield elongate and bent distinctly around caudal curvature of bursae. Types. Holotype 6 (Fig. IG), Guayquichuma, Dept, el Oro, Ecuador, 1,200 m, August 1980, xeric habitat, leg. Henri Descimon. Allotype, $, “Ecuador,” leg. Carlos Vela 1986, both deposited AMNH. Distribution. Spatial: known only from the type locality (see below) and an asso- ciated specimen with ambiguous data. Temporal: known only from August (holo- type). Remarks. The type locality is xeric habitat in the Maranon area of endemism (sensu Brown, 1982) characterized by a number of highly insular butterfly taxa including Papilio streckerianus Honrath (Papilionidae), Diaethria ceryx Hewitson, and Heli- conius erato himera Hewitson (Nymphalidae). Etymology. The name is taken from the Latin luxuriosus referring to the exotic patterning and other distinctive markings of this species. Noreena pritzkeri, new species Figs. IH, 2E, 3H, 4H Diagnosis. Above, male shiny steel blue fading distally, distal sector of androconial brand extremely large, generally ovate; female dull brown. Beneath, ground concol- orous medium brown; split-stripe cephalad discal cell pronounced, displaced basad 34 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) and thickly white, white extending basad and also distad through cells M 1 and M2 to the margin. Anal area of split-stripe dull, thin and widely angled. Tergal and genital morphologies distinctive as detailed below. Known only from a cluster of localities in southeastern Brazil. Description. MALE: Upper surface of wings: forewing baso-medially dull shiny iridescent steel blue, apex to subapex, brown; hindwing lighter iridescent blue dusted silverish toward the margin. Bipartite androconial brands with distal sector ovate and two or more times size of triangular basal sector. Hindwing with tail at terminus of vein CuA2. Under surface of wings (Fig. IH): ground color deep brown; forewing with submarginal and postmedian lines, costa to at least to vein CuA2. Costal portion of hindwing split stripe thickly white, extending to and surrounding the cell-end streaks; adjacent distal and basal areas of cells Ml and M2 highly suffused white; rest of medial stripe thin and sharply angled. Limbal area suffused; limbal spots small, reddish. Length of forewing: 14.0 mm (holotype), x of 28 paratypes 13.8 mm, range 13.5-15.0 mm. FEMALE: Upper surface of wings: completely brown; no an- droconial brand. Under surface of wings: as on males. Length of forewing: 13.0 mm (allotype). MALE ABDOMINAL MORPHOLOGY AND GENITALIA: Figures 2H, 3H. Incised posterior cavity elongate, extending to beneath sixth segment, steeply tapered cephalically, but with elongate, widely bifurcate prongs. Genitalia with bi- lobed area of valvae basally rounded, caudal extension thin, though recurvate caudad the bilobed extension and then tapering sharply terminad. Saccus elongate, terminally knobbed. Aedeagus longest of genus, with length x 2 remaining genitalic length from tips of labides and saccus. Vincular and saccal brush organs uniquely contiguous. FEMALE GENITALIA: Figure 4H. Antrum elongate, terminal lamellae widely ex- panded dorsad, sculptured ventrad; ductus with mild lateral arch but widely extended sclerotized arms at juncture with bursae. Detached corpus bursal shield small but sclerotized to flap proxad the ductus. Types. Holotype <3, allotype 2, “Morro Dona Martha, Rio de Janeiro” [Brazil] 27 April 1938, 25 April 1938, respectively, MPM. Paratypes: MPM— same locale as primary types, but 10 June 1938, 14 August 1958, 17 October 1936 (2); “Gavea, Rio de Janeiro,” 1 January 1933, 23 June 1936, 1 January 1951; “Rio de Janeiro” 8 May 1961, 1 1 July 1934; “Teatu, Rio de Janeriro” 28 June 1930 (2), 1 June 1932, 12 May 1935, 29 July 1938, 25 March 1939, 23 May 1939; “Castorina, Rio de Janeiro” 1 May 1936, 20 May 1936(3), 6 May 1936; “Collegio Baptista, Rio de Janeiro” 29 May 1938 (2); “[illegible], Rio de Janeiro” 19 May 1939, 8 June 1934, 26 August 1939; Joinville [Brazil], 12 March 1954 (28<3). AMNH: 2<3, same data as primary types but 3 March 1936; “Gavea, Rio de Janeiro” 24 June 1932; AME— 16, same data as primary types but 14 August 1958. Distribution. Spatial: known from several localities in the vicinity of Rio de Janeiro formerly characterized by virgin primary forest (see above and Remarks). Temporal: dates on specimens range from early March to late August. Remarks. The Gagarin Collection (MPM) contains large samples of SE Brazil theclines otherwise known from very few specimens (sometimes only the types). In addition, this collection contains numerous undescribed taxa, ranging from many singletons (see N. galactica, C. minutaea, below) to long series. Such rich collections of high diversity/low density taxa, like theclines, result from long term residence by a collector in a particular area. Indeed, K. S. Brown, Jr., who knew Gagarin, states 1989 NEOTROPICAL LYCAENIDAE 35 that when Gargarin caught a specimen unlike anything previously seen, he would return to its collection site again and again. Brown indicates that the type locality listed above was, at the time of Gagarin’s work, virgin primary forest. Gagarin’s collection sites at Joinville were also virgin primary forest; habitat at Gavea was, at least, transitional from primary to secondary forest. This suggests that the absence of other specimens of N. pritzkeri in any collection probably results from this habitat restriction and subsequent alteration. Etymology. Patronym for Nicholas Pritzker. Noreena galactica, new species Figs. II, 21, 31 Diagnosis. In this small species (forewing base to apex 10.5 mm) the split-stripe converges in a helix-shaped configuration at the discal cell and is complemented distad by a bright white postmedian line extending to the costa. The male eighth tergal plate is cephalically non-bifurcate, valvae of the genitalia caudally constricted and elongate saccus laterally inclined. Description. MALE: Upper surface of wings: forewing dull fuscous, base shiny steel blue; hindwing dull blue. Bipartite scent brands with distal sector ovate, twice size of basal section and more distally detached than in congeners. Under surface of wings: forewing with rather straight white postmedian stripe, costa to cell CuA 1 ; white split- stripe on hindwing converging at discal cell in an ovate shape, making an overall helix-like configuration. White postmedian band proceeding costad from CuAl to margin; limbal spot at cell CuAl large, yellow, with central black spot, limbal spot near anal angle small, orangish. Limbal area unsuffused. Length of forewing: 10.5 mm (holotype). FEMALE. Unknown. MALE TERGAL MORPHOLOGY AND GENITALIA: Figures 21, 31. Plate over eighth tergite extending beneath seventh segment, nonbifurcate cephalad. Valvae with bilobed area steeply parabolic, slightly lobed near saccus; caudal extension extremely thin for genus, adjacent vincular spurs very wide; saccus longest of genus, inclined laterally. Aedeagus length exceeding rest of genitalia by only about one-third, caudal one-third recurved. Brush organs thin and elongate. Type. $, “Collegio Baptista, Rio de Janeiro” 29 May 1938, Gagarin Collection, deposited MPM. Distribution. Known only from type locality from May. Remarks. Remarks concerning N. pritzkeri also apply to this species. Etymology. The name refers to the helix-shaped stripe on the hindwing under surface. Noreena molena (Jones), new combination Figs. ID, 2F, 3F, 4F Theda molena Jones, 1912:899, pi. 97, f. 9; Draudt, 1919:769, pi. 154e; Comstock and Huntington, 1958-1964 [1961]:171; Bridges, 1988:1.231, 11.78. Diagnosis. Both sexes very small {x forewing length of both sexes 1 1.0 mm com- pared to 13.0-16.0 mm for congeners), male above basally dingy iridescent blue, distally fuscous; female, dull brown. Beneath, split-stripe pattern greatly reduced and vague, exceeded in distinction by highly variegated ground color grizzled in bright 36 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) yellow-brown patchwork-like postmedian and submarginal patterns and by light, chevron-like, markings at the forewing subapex and across limbal area of hindwing. Male androconial brands and tergal and genital morphologies distinctive as detailed below. Description. MALE; Upper surface of wings: forewing dull dark blue on basal two- thirds, rest of wing fuscous. Male androconial brand with bipartite sectors oblong, distal sector about one-third size of basal. Hindwing base to submargin dull dark blue, distally fuscous. Short thick tail at terminus of cell CuA2, stubby tail at vein CuAl. Under surface of wings: forewing dull brown, basal two thirds with white postmedian line, costa to cell CuAl. Postmedian ground color yellow-brown; sub- marginal line of gray chevron-like markings, costa to cell CuA2; yellow-gray grizzling along the apex. Hindwing dull mottled brown. Split-stripe pattern reduced, partic- ularly at the cell-end streaks. Limbal areas with bold gray patchlike pattern in each cell, shaped distally into chevrons. Limbal spots reduced, red-orange. Length of forewing: 11.0 mm (syntype). FEMALE: Upper surface of wings: both wings dull brown, only faintly hued blue. Under surface of wings: as on male but limbal gray patchwork pattern more obsolescent. Length of forewing: 1 1 .0 mm (lectotype). MALE ABDOMINAL MORPHOLOGY AND GENITALIA: Figures 2F, 3F. Incised pos- terior cavity with sides of relatively equilateral length compared to congeners, width equalling .75 length, plate extending cephalad beneath the sixth and seventh segments. Cephalic plate margin with two small smoothly rounded lobes, not prongs. Genitalic components bulbously thick and highly sculptured, valvae with bilobed areas tri- angular, caudal extension with raised ventral ridge and terminus tapered to a point. Vincular spurs thick and lobate, nearly bifurcate. Saccus funnel-shaped, distally an- gled. Aedeagus with terminus only slightly recurved. Vincular brush organs abutting entire edge of dorsal vincular wall and along slight, rounded basal anchorage lobe. Saccal brush organs elongate along margin of vinculum and valval base. FEMALE GENITALIA: Figure 4F. More dimunitive than congeners— lamellae nearly absent; ductus bursae small, caudally arched with lateral edge closely paralleling elongate sclerotized shield. Caudal component cephalically tapered, bilobate area adjoining diminutive lamellae. Types. Syntype $ (Fig. ID) and 9, BMNH, female hereby designated lectotype, labelled “Theda molena, type female. Dr. Jones, Castro Parana, 2,900 ft. E. D. Jones, E. D. Jones Collection, Brit. Mus. 1919-295,” syntype 6 labelled “Theda molena, type male. Dr. Jones, Castro Parana, 2,900 ft. 2 May ’10, E. D. Jones, E. D. Jones Coll., Brit. Mus. 1919-295, B.M. Type No. Rh 1086.” Distribution. Spatial: Known only from southeastern Brazil (see Remarks). Tem- poral: known specimens have May, July and November collection dates. Remarks. The figure in Draudt (1919) does not adequately portray the mottled under surface and thin, yet vivid, split-stripe (which is more vivid on the syntype male than on the lectotype). The original Jones (1912) figure (Fig. ID) is more accurate, though the whitish limbal markings on the actual specimen are more chev- ron-like, especially in the lectotype. The species is known only from a few old specimens. These include the types, another female in the BMNH and two additional specimens in the MNHN. Com- ments by K. S. Brown concerning habitats at collection sites of Gagarin may apply to this species as well. N. molena is not included in Gagarin’s collection, but Castro, 1989 NEOTROPICAL LYCAENIDAE 37 Parana was not a major collection area for him. Perhaps, as with unique Noreena taxa in the Gargarin Collection, N. molena originally inhabited primary forest now absent from the type locality. Material examined. BRAZIL [see type data], Caicera, Orinoco, Nov. 1898, 19 (BMNH); Espirito Santo, Brazil, July 1899, 1(5, 19 (MNHN). Noreena lemona (Hewitson), new combination Figs. IE, 2H, 3E, 4G Thecla lemona Hewitson, 1863-1878 [1874], 1:177, 2: pi. 69, f 519, 520; Draudt, 1919:770, pi. 154e; Comstock and Huntington, 1958-1964 [1961]: 108-1 09; Bridges, 1988:1.193, 11.78. Diagnosis. Like N. molena, dingy above and small {N. lemona x forewing length 12.0 mm, others see above) with ground color beneath dominated by variegated coloration; distinguished by large black-brown blotches, subapical on both wings, medial in cell CuA2 and surrounding the limbal spots. Of the tergal and genitalic distinctions noted below, the incised posterior cavity is the most notable of the genus, extending caudad from beneath the juncture of the fifth and sixth abdominal seg- ments. Description. MALE: Upper surface of wings: basal ground color dark iridescent azure blue becoming fuscous distad. Bipartite androconial brand with sectors oblong and of about equal size. Under surface of wings: ground color variegated light brown; outstanding dark blackish brown patches scattered throughout. Hindwing pattern with (a) postmedian band distinctly white, costa to cell CuAl, (b) white subapical line meandering, framing blackish brown blotch from costa to vein CuA 1 ; hindwing with (a) thin split stripe clearly defined, basal area in cell CuA2 with large black- brown blotch, (b) limbal spots large, deep red, surrounded by dark black-brown ground color and (c) limbal area variegated light and dark brown with alternating bright gray-white and dark brown-black submarginal blotches, costa to vein M3. Length of forewing: 12.0 mm (BMNH). FEMALE: Upper surface of wings: ground color brown; lacking androconial scent brand. Under surface of wings: as on males. Length of forewing: 12.0 mm (lectotype). MALE TERGAL MORPHOLOGY AND GENITALIA: Figures 2H, 3E. Incised posterior cavity longest of the genus, extending caudad from juncture of fifth and sixth abdominal segments, uniquely arched dorsad beneath juncture of seventh and eighth segments. Genitalic configuration elongate for genus, saccus of equal length with valvae and thinly parabolic vincular arc. Valvae sharply angled between bilobed configuration and caudal extension; vincular spurs elongate and thin. Longest aedeagus of genus (length exceeding that of vinculum by X 2), terminally recurvate only in caudal one-third; caecum small. Vincular brush organs abutting prominent basal vincular spur (as only in N. cambes) saccal brush organs in short compact bundles. FEMALE GENITALIA: Figure 4G. Cephalic com- ponent straight, corpus bursae consequently removed to position more terminad caudal component. Caudal component elongate, length X3 width (beneath lamellae); lamellal lips wide, cephalically recurved. Sclerotized shield covering only caudal end of bursae, extending laterally only slightly. Types. 9 (Fig. IE), BMNH labelled “Brazil, Hewitson Coll., Thecla lemona (2), B.M. type No. Rh. 694.” hereby designated lectotype (see Remarks). 38 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) Distribution. Spatial: Known only from a few old specimens from southeastern Brazil with data indicating the species is locally sympatric with N. molena (see Remarks thereunder) and regionally sympatric with N. pritzkeri. Temporal: No data available (see Remarks). Remarks. Only one Hewitson specimen appears extant in the BMNH, though the label notation “Theda lemona (2)” on this specimen is standard BMNH procedure indicating there were originally two syntypes. A search of the collection, including the World War II reference collection (which in some cases is still not reincorporated into the general BMNH collection) has revealed only an additional non-Hewitson male. Hence, although some references (e.g., AME type catalogue) treat the extant Hewitson female as a holotype, I designate this specimen as lectotype. The figure in Draudt (1919) does not adequately portray the vividly contrasting ground color patches on this species. Habitat comments under N. molena Remarks may also apply to N. lemona. Material examined. BRAZIL, [see type data], Castro, Parana, Jones 2,900 ft., 16, 19 (BMNH); Espirito Santos [sic], Brazil, 15, 19 (MNHN). See Remarks under N. molena. Noreena maria Johnson, MacPherson and Ingraham Figs. lA, 2G, 3G, 4A Noreena maria K. Johnson, MacPherson and Ingraham, 1986:2; Johnson 1988:34; Bridges 1988:1.214, 11.78. Diagnosis. Resembling the wing pattern group N. cambes, N. comana, N. luxuriosa and N. pritzkeri but small (12-13 mm). Female brown, both sexes with forewing lines beneath convergent caudad, hindwing limbal pattern mottled black and white along the anal angle, and split-stripe rounded at the anal angle, not w-shaped. Male androconial brands and tergal and genital morphologies distinctive as detailed below. Description. MALE: Upper surface of wings: forewing dull brown, apex to subapex, rest of wing iridescent blue; hindwing lighter iridescent blue dusted silverish toward the margin. Bipartite androconial brand with sectors of about equal size, triangular distad, oblong basad. Hindwing with tail at terminus of vein CuA2. Under surface of wings (Fig. 1 A): ground color dull gray; forewing with submarginal and postmedian lines, costa to at least to vein CuA2, often converging caudad. Hindwing with split- stripe emphatic; median area costad vein Ml immaculate. Limbal area with arc of white macules in submargins, each heavily colored black at the vein interspaces, becoming one line costad, engulfing the parallel cell-end streaks and postbasal mark- ings. Limbal spots vividly orange. Length of forewing: 12.0 mm (holotype), 11.5, 12.0 mm (paratypes). FEMALE: Upper surface of wings: as on male, but upper surface completely brown; no androconial brand. Under surface of wings: as on males but with lines of cell-end streaks often parabolic in shape and disjunct from the split-stripe. Length of forewing: 13.0 mm (allotype), 12.0 mm (paratype). MALE ABDOMINAL MORPHOLOGY AND GENITALIA: Figures 2G, 3G. Incised pos- terior cavity wide, oblongly hexagonal in general shape with cephalad prongs widely bifurcate and pointed. Genitalia with lateral surface of vinculum well defined; saccus broadly parabolic. Valval lobes separate; ventrad surface caudally tapered with slope to terminus somewhat jagged-edged on lateral surface. Aedeagus with small caecum. 1989 NEOTROPICAL LYCAENIDAE 39 long slender shaft and recurvate terminus; two comuti. Vincular brush organs densely packed strips abutting the saccal vincular junction. Saccal brush organs as smaller clusters ventral along the proximal sides of the saccus. FEMALE GENITALIA: Figure 4A. Caudal component angled widely toward terminus; lamellal lips prominent. Cephalic component recurved widely distad, joining corpus bursae centrally. Scler- otized shield widely detached; point of attachment of ductus seminalis remote from ductus terminus. Adjoining arms of arched cephalic component meeting corpus bur- sae at midway position along its length. Two thomlike signa in cephalic half of corpus bursae. Types. Holotype $, allotype 9, AME. Type locality. Mosconi, Salta Province, Argentina by original description. Distribution. Spatial: presently known from numerous chapparal and chaco habitats in northwestern Argentina. Temporal: known from mid-May to late July. Remarks. The overall rarity of Noreena taxa seems typified by the experience of Robert C. Eisele and Bruce MacPherson with N. maria. Thirty years of field collecting in northwestern Argentina has yielded only six specimens. The habitat is xeric wood- land on the margin of xeric chaco vegetation. Mosconi is the type locality of several chaco and chaco margin endemic eumaeines (Johnson, 1988; Johnson, Eisele and MacPherson, 1988, 1989). As example of N. maria wing pattern, I have illustrated a recently caught female (Fig. lA). This increases the number of specimens figured in the literature (see Johnson, MacPherson and Ingraham, 1986) and characterizes the marked caudal convergence of the forewing bands apparent on most specimens. Material examined. Paratypes— type locality: 15 May 1976, leg. Bruce Mac- Pherson, 19 (AME, loan by Eisele); June 1975, from Robert Eisele Collection (leg. Bruce MacPherson [correction from OD]), 16, (AMNH); May 1978, from Eisele Collection (leg. MacPherson), 19 (AMNH); other— 2 km NW San Pedro, Jujuy Prov- ince, Argentina, 550 m, 2 May 1979, leg. Eisele, 1(3 (Eisele Collection); 4 km NW San Pedro, at Morro Centinela, SW ridge, 700 m, 24 July 1979, leg. Eisele, 19 (Eisele Collection). Contrafacia, new genus Figs. 5, 6, 10 Diagnosis. Distinctive in resembling taxa of Draudt’s '"spurina Group” above (Tb. IE) (bipartite androconial brands), but his "^orcynia Group” beneath (Tb. 1C, Figs. 8, 9) (light split-striped pattern. Fig. lOB). Differing from all of these in sharing with Noreena nine synapomorphies (Chs. 1, 2, 5, 8-12, 15), most saliently the incised posterior cavity in males (Ch. 1, Figs. 2, 5, 8, 9) and laterally arched genitalic configuration in females (Ch. 8, Figs. 4, 5, 8, 9). Superficially distinguished from Noreena by reduction of the split-stripe pattern and reduced limbal spots (Fig. lOB) more characteristic of the orcynia Group (Tb. 1C, Ch. 17, Figs. 5, 8, 9, IOC). Description. ADULT: Antennae fuscous, finely striped white, length about one third that of forewing base to apex; head with frons uncolored, eyes outlined white; thorax fuscous with gray to fuscous hairs profuse distad; abdomen fuscous, often with scattered blue powdering adjacent the hindwings. MALE: Upper surface of wings (Fig. 5): basal areas of forewings dull dark blue, postmedian and apical areas black. Bipartite androconial brand distad in discal area, basal component widely triangular 40 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) basad cross vein of discal cell, distal component variously oval and variously detached caudad from vein LDC. Hindwing usually of brighter iridescenee than forewing: bright blue-green in one species, azure blue in the other. Long tail at terminus of vein CuA2, shorter tail at vein CuAl. Under surface of wings (Figs. 5, lOB): ground color tawney to brown, pattern with (a) whitish forewing postmedian line, costa to cell CuAl, (b) thin white hindwing medial band, variously meandering or jagged, with two to three cell-end streaks and white colored markings (blotches or dashes) in the submargin. Thecla-spot in eell CuAl an orange orb. FEMALE: Upper surface of wings (Fig. 5): both wings silvery blue to somewhat darker iridescent blue, brown distally from postmedian area; no androconial brands. Under surface of wings (Figs. 4B, F; 5D, lOB): as on males but often with more distinction of cell-end streak component of light split stripe band. MALE TERGAL MORPHOLOGY AND GEN- ITALIA: Figure 6. Eighth tergite specialized into posterior incised subcordate cavity with surrounding generally ovate sclerotized plate extending from beneath caudal one-half of seventh segment, laterally nearly to the spiracles, and usually indented caudo-centrad (Ch. 1, Figs. 6, 8, 9). Male genitalia similar to Noreena in caudal extension and terminus of valvae (Ch. 3, Figs. 3, 6, 8, 9), vincular spurs (Chs. 6, 7, Figs. 3, 6, 8, 9), vincular and saccal brush organs (Ch. 15, Figs. 3, 6, 8, 9) and general form of vinculum and saccus (Chs. 4, 5, 7, Figs. 3, 6, 8, 9) but with saccus extremely elongate as undescribed sister group X and in the ""orcynia Group” (Tb. IB, C). FEMALE GENITALIA: Fig. 6. Female with disjunct genital structure (Ch. 13, Figs. 6, 8, 9) but (Fig. 6) (a) ductus compactly arched proxad corpus bursae {Noreena taxa variously widely arched or angled), (b) ductus cephalic terminus contiguous with corpus bursae’s distal sclerotized shield, (c) ductus seminalis emanating from juneture of ductus terminus and sclerotized shield and (d) cephalic juncture of ductus terminus and corpus bursae with only slight sclerotized arm extending laterally around bursal sac. Types Species. Contrafacia mexicana, new species (Fig. 4C-F). Distribution. Four species, distributed from Sonora, Mexico southward to Para- guay. Etymology. The name, considered feminine, combines the Latin "‘‘contra’'^ and ""facia"" and refers to the disparate wing and genitalic characters compared to outgroup taxa. Contrafacia rindgei, new species Figs. 5A, 6C Diagnosis. Known only from a Sonora, Mexico female with distinctive gray-white under surface ground color, split-stripe divided into a costal line including two, ovate, cell-end streaks and a line extending caudad vein CuAl (w-shaped along the anal margin and two limbal spots on each hindwing reduced to orangish dots). In the genitalia, the caudal component is elongate as in South American C. australis (though lacks the lobate lamellae of this species), not compact as in C. mexicana and C. minutaea. Description. MALE: Unknown. FEMALE: Upper surface of wing: forewing ground color shiney light brown overshadowed with silvery blue basad especially on hind- wing. Long tail at terminus of vein CuA2, short tail at vein CuAl. Under surface of wings: ground color, both wings bright whitish gray with very little variation of hue; 1989 NEOTROPICAL LYCAENIDAE 41 forewing with white postmedian band, bordered gray basad, angled steeply basad from costa to vein CuA 1 ; hindwing with split-stripe restricted to a thin white line running from the costa to and including the cell-end streaks and a disjunct w-shaped line from anal margin to cell CuA 1 . Limbal spots occurring only as vague organgish dots. Submargin with vague line across wing from anal margin and intersecting the limbal spots. Length of forewing: 12.5 mm (holotype). FEMALE GENITALIA: Figure 6C. Caudal component elongate and with bilobate lamellae, similar to C. austmlia. Cephalic component compactly arched close to caudal end of corpus bursae. Juncture of ductus and corpus bursae with only slight lateral sclerotization on bursal sac. Sclerotized plate not detached; rather, conjoined with ductus proxad its cephalic terminus; ductus seminalis emanating from juncture of sclerotized shield and ductus bursae. Type. Holotype $ (Fig. 5 A), Port San Pedro, Sonora, Mexico, 6 January 1939, Frederick H. Rindge, taken at wildflowers in desert gully (F. H. Rindge, pers. comm.), deposited AMNH. Etymology. The species is named for Dr. Frederick H. Rindge who collected the holotype. Contrafacia mexicana, new species Figs. 5B, C, 6A, B Diagnosis. Superficially stands out among Mexican theclines by male’s bipartite androconial brands and under surface (both sexes) exhibiting lightened split-stripe pattern of the brandless ocrynia complex. Compared to congeners, most similar to South American C. australis (Fig. 5) since C. rindgei (Fig. 4A, B) has the medial band broken into costal and anal elements. C. australis (known only from Paraguay/east Bolivia) is much larger {x forewing length 14.3 mm, C. mexicana 1 1.8 mm), has a less jagged split-stripe and a larger, more detached distal sector in the bipartite androconia. C. mexicana also has distinctive morphological characters (see below). Description. MALE: Upper surface of wings: ground fuscous at apex of forewing and distad the submargins of hindwing. Base of forewing dull iridescent blue; hind- wing with central patch of brilliant blue-green. Bipartite androconial scent brand with basal sector large and oblong, distad sector slightly detached costad from vein LDC. Long tail at terminus of vein CuA2; shorter tail at CuAl. Under surface of wings: ground color beige; forewing with vague to obsolescent submarginal band and distinct white postmedian band, costa to cell CuA 1 . Hindwing with moderately bright white split-stripe and prominent, variously detached, single cell-end streaks. Anal band area extremely incised in w-shape from vein CuAl to the anal margin. Submargin, anal lobe to costa, with whitish parallel streaks in each cell, more broadly whitish toward costa. Orange spot at anal lobes and submarginal in cell CuA 1 . Length of forewing: 12.5 mm (allotype). FEMALE: Upper surface of wings: similar to male but duller silvery-blue; junctures of iridescent patches and distal fuscous less distinct; no androconial brand. Under surface of wings: as on males but white markings on hindwing more emphatic and cell-end streaks nearly connected to the split-stripe. Length of forewing: 1 1.0 mm (holotype). MALE TERGAL MORPHOLOGY AND GENITALIA: Figure 6 A. Specialized eighth tergite caudally incised and of oval shape. Genitalia with extremely elongate, terminally knobbed, saccus (length over 1.25 x caudal expanse of vincular arc). Vincular spurs prominent and elongate at caudo- 42 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) ventral margin of vincular arc, overlapping area with marked constriction of valve between its bilobed area and caudal extension. Aedeagus length exceeding entire length of genitalia from tip of saccus to tip of labides by about one-third; caudal terminus very slightly recurved. Vincular brush organs thickly clustered, saccal com- ponent reduced. FEMALE GENITALIA: Figure 6B. Relative to Noreena taxa, dual component conhguration with both components compact; of Contrafacia taxa, caudal component relatively shorter than on congeners. Cephalic terminus of ductus bursae connected to sclerotized shield across area of corpus bursae proximal to ductus bursae, ductus seminalis emanating from this juncture. Corpus bursae with two signum with large sclerotized bases and thin central spines. Types. Holotype 9, allotype $ (Fig. 4C-F), Mexico City, September 1935, leg. Hoffman deposited AMNH. Paratypes: MPM-29, Mexico City, Mexico, Niedhofer Collection; AMNH— 13, Matamoros, Puebla, 14 September 1910, Hoffman Collec- tion. Distribution. Montane central Mexico. Remarks. I have made the female the holotype of the type species, consistent with the gender of C. rindgei and because the female genitalic characters most clearly show the sister group relationship to Noreena. It is unfortunate that Hoffman attached only brief data to the C. mexicana specimens. Though Hoffman usually placed individ- ualized data in specimen papers, he sometimes enclosed large numbers of papers (each individually dated) in a larger envelope indicating the collecting locality. This practice appears more common for collecting areas more familiar to him (“Mexico City,” “Presidio”) or, perhaps, considered by him as less remote. Regarding speci- mens of C mexicana, Hoffman may have assumed them to be T. orcynia by the under surface patterns. Since T. orcynia is considered a rather common butterfly (Draudt, 1919), it is possible that some museums may have specimens of C. mexi- cana, particularly female, included with T. orcynia. The MPM, though having small numbers of Mexican material, has a number of unusual samples, particularly in collections assembled by Moeck. Etymology. The name denotes the Mexican region. Contrafacia australis, new species Figs. 5D, E, 6D, E Diagnosis. Of congeners, C. australis is the largest (x forewing length 14.3 mm, compared to 12.2 mm, C. mexicana', 12.5 mm, C. rindgei', and 10.0 mm, C. minu- taea), has a pronounced, jagged, split-stripe, diminutive limbal spots and (in fresh specimens) brilliant white limbal crescents (Fig. lOB). Of known males of Contrafacia (C mexicana and C. australis), C. australis has a dark azure blue upper surface iridescence and an extremely triangular basal section of the bipartite androconial brand. In addition to generic morphological distinctions, the split-stripe band of C. australis distinguishes it from regionally sympatric outgroup taxa T. ericusa and T. catharina, which each have a simple postmedial band of broken slashes. Known only from southeastern Boliva and Paraguay. Description. MALE: Upper surface of wings: apex and submarginal areas black, remaining basal areas of both wings iridescent deep azure blue. Bipartite scent brand with basal sector an extreme isosceles triangle boldly pointed distad; dorsal sector about half as large and ovate, detached slightly caudo-distally from vein LDC. Long tail at terminus of vein CuA2, short tail at vein CuAl . Under surface of wings: ground 1989 NEOTROPICAL LYCAENIDAE 43 color tawney. Forewing with postmedian band, costa to cell CuAl. Hindwing with thin, jagged mesial band and two to three thin cell-end streaks. Submargin with white parallel slashes in each cell. Length of forewing: 14.0 mm (allotype). FEMALE: Upper surface of wings: basal to medial areas dull silvery blue, distal areas brown. Tailed as male; no androconial brand. Under surface of wings: as on males but with more closure of cell-end streaks with the medial band. Length of forewing: 14.5 (holotype). MALE TERGAL MORPHOLOGY AND GENITALIA: Figure 6D. Incised posterior cavity distinctive, slightly concave at caudal edge. Genitalia similar to C. mexicana but with saccus less knobbed, vincular spurs more elongate, and valvae in ventral view with marked cephalic constriction followed terminally by recurvation to widely bulbous termini. FEMALE GENITALIA: Figure 6E. Disjunct configuration with caudal component elongate, lamellae oblongly ovate and bilobed (similar to C rind- gei). Cephalic component compactly arched, joining corpus bursae with only slight lateral sclerotization, sclerotized shield conjoined proximally to cephalic terminus of ductus bursae. Ductus seminalis emanating from juncture of sclerotized shield and ductus bursae. Two signa with broad sclerotized based constricting surface of corpus bursae. Types. Holotype 9, allotype $ (Fig. 5 D), Cordillera, Santisima-Trinidad, 25°15'S, 57°38'W, Paraguay, August, B. Podtiaguin, deposited AMNH. Paratypes: CMNH— (5, Rio Surutu, E. Bolivia, 350 m, December 1913, leg. Steinbach, deposited CMNH (H. K. Clench genitalic preparation No. 1045); MPM— 19, Nueva Italia, Paraguay, 6 July 1 940; BMNH — 1 9, Paraguay, 1901, Crowley Bequest; MNHN — 1 9, Villa Rica, Paraguay, 10 January 1925 (Fig. lOB). Remarks. Podtiaguin’s collections from the central Cordillera of Paraguay have already been shown to include some very rare and endemic taxa (Johnson, Rozycki and Matusik, 1988). Clench, at CMNH, had separated from the collection a group of male specimens with bipartite scent brands which he tentatively identified as “sp. near Thecla orios" (TL Guatemala) but which evidenced distributions from SE Brazil and Bolivia northward to Honduras. All of these specimens appear morphologically as undescribed species near T. orios or of undescribed sister group X (Tb. 1 B, Figs. 8, 9) except the male designated paratypical of C. australis above. Etymology. The name, using “austral” or “south” denotes the area of geographic occurrence of this Contrafacia species. Contrafacia minutaea, new species Figs. 5F, 6F Diagnosis. The known specimen is extremely small (female, FW, base to apex, 10.0 mm); the medial band from cell M2 is angled distinctly baso-costad and the parallel cell-end streaks detached. Antrum of genitalia with unique dorsal arch, ductus laterally arched; proximad shield on corpus bursae smallest of genus, corpus bursae extremely large, diameter of corpus bursae exceeding length of rest of genital parts. Description. MALE: Unknown. FEMALE: Upper surface of wings: dull brown hued bluish; no androconial brand. Under surface of wings: ground color beige; forewing with vague white postmedian band, costa to cell CuAl. Hindwing with moderately bright, but thin, white medial split-stripe steeply angled baso-costad from cell M2. Parallel cell-end streaks small, detached basad from split-stripe. Limbal spot small, orangish. Limbal area unsulfused. Length of forewing: 10.0 mm (holotype). FEMALE GENITALIA: Figure 6F. Antrum uniquely inclined dorsally at juncture 44 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) with ductus bursae; ductus bursae laterally arched joining corpus bursae disto-centrad. Proximad sclerotized shield on corpus bursae small, extending to distal end of corpus bursae; corpus bursae extremely large, diameter exceeding length of all other genital parts. Type. Holotype 9 (Fig. 5F) labelled “Morro Dona Martha, Rio de Janeiro” [Brazil], 19 October 1938, deposited MPM. Remarks. Remarks under N. pritzkeri pertain. Etymology. The name indicates the extremely small size of the holotype. ACKNOWLEDGMENTS For specimens examined, I am grateful to curators at the following museums: AME, Lee D. and Jacqueline Y. Miller; AMNH, Frederick H. Rindge; BMNH, Philip Ackery and Richard Vane-Wright; CMNH, John E. Rawlins; FMNH, Stephen Ashe; IML, Z. D. Ajmat de Toledo and R. Golbach; MPM, Allen M. Young and Susan Borkin; MNHN, G. Bemardi and Jacques Pierre. Henri Descimon (Universitie de Provence, France) and Nicholas Pritzker (Chicago, Illinois) provided valuable specimens from private collections; Keith S. Brown, Jr. (Univer- sidade Estadual de Campinas, Brazil) answered queries about certain type localities. Susan Borkin and David Matusik (FMNH) kindly prepared and shipped certain specimens. Roberto C. Eisele, Bruce MacPherson (Tucuman and Jujuy, Argentina, respectively) and David Matusik facilitated receipt of the IML material. For review comments I am grateful to two anonymous reviewers and also for initial comments concerning Noreena by J. N. Eliot (Taunton, United Kindgom) and subsequent comments on this paper by John E. Rawlins, Randall T. Schuh (AMNH), and L. D. and J. Y. Miller. For use of technical facilities I am greatly indebted to administration and staff of the CMNH and for financial support from anonymous contributors to the Theclid Research Fund, AMNH. LITERATURE CITED Bridges, C. A. 1988. Catalogue of Lycaenidae & Riodinidae (Lepidoptera: Rhopalocera). Urbana, Illinois, published by the author, vii + 81 1 pp. Brown, K. S. 1982. 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The “hairstreak butterflies” (Theclinae) of northwestern Argentina. I. Introduction, Calycopis, Calystryma, Tergissima and Fem- niterga (Lycaenidae). Bull. Allyn Mus. 123:1-49. Johnson, K., R. Eisele and B. MacPherson. 1989. The “hairstreak butterflies” (Theclinae) of northwestern Argentina. II. Strymon Hiiebner sensu stricto (Lycaenidae). Bull. Allyn Mus. (in press). Johnson, K., B. MacPherson and J. Ingraham. 1986. A new genus and species of Eumaeini from northwestern Argentina (Lepidoptera, Lycaenidae). Bull. Allyn Mus. 102:1-7. Johnson, K. and D. Matusik. 1988. Five new species and one new subspecies of butterflies from the Sierro De Baoruco of Hispaniola. Ann. Carnegie Mus. 57:221-254. Johnson K., L. D. Miller and J. Herrera, [ms.]. Eiseliana and Heoda, high Andean assemblages of the Strymon Hiibner sens. lat. grade of Theclinae, Eumaeini (Lepidoptera). ms. (in review). Johnson, K. and R. Rozycki. 1986. A new species of the anchisiades Group of Heraclides from Venezuela (Lepidoptera; Papilionidae). J. N.Y. Entomol. Soc. 94:383-393. Johnson, K., R. Rozycki and D. Matusik. 1987[1989]. A study of Protesilaus microdamas (Burmeister) and the little-known P. dospassosi (Rutimeyer) and P. huanucana (Varea de Luque) (Papilionidae). J. Res. Lepid. 26(1) (in press). Jones, E. D. 1912. Descriptions of new butterflies of the genus Thecla from S. E. Brazil. Proc. Zool. Soc. London, Unnumbered, pp. 896-903. Kirby, W. F. 1871. A synonymic catalogue of diurnal Lepidoptera. John van Voorst, London, vii -I- 192 pp. Kohler, P. 1923. Fauna Argentina Lepidoptera e collectione Alberto Breyer. Sonderbeilage der Zeitschrift fur wissenschaftliche Insekten-biologie. Bd. XVIII (12), I. Teil, 34 pp. Kohler, P. 1928. Catalogo de Lepidopteros Argentinos. Buenos Aires, Puclicaciones Breyer, 12 pp. Llorente-Bousquets, J., A. Garces Medina and A. L. Martinez. 1 986. Las mariposas de Jalapa- Teocelo, Veracruz. Teocelo 14(4): 14-37. Moschler, H. B. 1883. Beitrage zur Schmetterlings-Fauna von Surinam. V. (Supplement), Verb. Zool.-Bot. Ges. Wien 32:303-362, pis. 17, 18 (continued from 1876). Nicolay, S. S. 1971. A new genus of hairstreak from Central and South America. Supp. J. Lepid. Soc., 25, suppl. 1, 39 pp. Nicolay, S. S. 1976. A revision of the Huebnerian genus Parrhassius with description of the new genus Michaelus (Lycaenidae, Eumaeini). Bull. Allyn Mus. 35:1-30. 46 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) Ross, G. N. 1975-1977. An ecological study of the butterflies of the Sierra de Tuxla in Veracruz, Mexico. J. Res. Lepid. 14(2): 103-124; 14(3): 169-1 88; 15(l):41-60; 15(2):109- 128; 15(3): 185-200; 15(4):225-240; 16(2):87-130. Schaus, W. 1920. Synonymy of some species of Theda (Lepid). Entomol. News 31:176. Schweizer, F. and R. G. Webster-Kay. 1941. Lepidopteres del Uruguay. II. Catalogo siste- matico, parte 1. Rhopalocera and Grypocera. Anales del Museo de Historica Nat. de Montevideo, 2. a Serie-Tomo V, No. 3, pp. 4-24. Swoffbrd, D. 1985. PAUP (Phylogenetic Analysis Using Parsimony) and Users Manual, a computer software package made available by the Illinois Natural History Survey and the author. Weeks, A. G. 1905. Illustrations of diurnal Lepidoptera with descriptions. Boston, Boston Univ. Press, xii + 117 pp. Weeks, A. G. 1911. Illustrations of diurnal Lepidoptera with descriptions. Vol. 2. Boston Univ. Press, Boston, xvi + 37 pp. Zikan, J. F. and W. Zikan. 1968. Insecto-Fauna de Itatiaia e da Mantiquieva. Sc9ao Ento- mologia e Fitopatologia; Pesq. Agropec. Bras. 3:45-109. Received March 24, 1987; accepted September 12, 1988. J. New York Entomol. Soc. 97(l):47-49, 1989 A NEW APHAENOGASTER (HYMENOPTERA: EORMICIDAE) FROM SOUTHERN NEW MEXICO William P. MacKay* Department of Biology, New Mexico State University, Las Cruces, New Mexico 88003 Abstract. —Aphaenogaster punctaticeps, n. sp. is described from the northern Chihuahuan Desert of south-central New Mexico, USA. It is closely related Xo Aphaenogaster texana (Emery). The workers of the two species can be easily distinguished in that the posterior border of the head of A. punctaticeps is moderately pointed (rounded in A. texana) and the dorsum of the head is primarily punctate (rugose with punctures in the interrugal spaces in A. texana.). During an intensive investigation of the ant fauna of a site in the northern Chi- huahuan Desert of south-central New Mexico (Jornada Long Term Ecological Re- search Area), an undescribed species of the genus Aphaenogaster was collected. The following is a description of this species. Aphaenogaster punctaticeps, new species Description. Worker: (Abbreviations as defined by Snelling, 1981.) HL 1.32-1.40 mm, HW 0.94-1.00 mm (at anterior border of eye), SL 1.58-1.70 mm, EL 0.21- 0.23 mm, EW 0.17-0.18 mm. Cl 71, SI 168-170, OI 24-27, PNW 0.23-0.26 mm, PPW 0.32-0.35 mm, WL 1.11 mm. Mandible with large apical tooth, subapical tooth about one half as large, first basal tooth subequal in size to subapical tooth, other teeth poorly defined, mandible striate; clypeus with well defined depression on anterior border, two rugae present posterior to notch; frontal area with median carina, smooth and somewhat shining on both sides of carina; frontal carinae large and protruding almost perpendicular to surface of head; rugae present on malar region, a few poorly defined rugae present posterior to frontal area, but absent on most of rest of dorsum of head, which is strongly punctate; base of scape simple (Fig. 1); eyes exceed margins of head in full face view, posterior margin of head moderately pointed (Fig. 4). Dorsum and sides of mesosoma moderately punctate, including both faces of propodeum; propodeal spines poorly developed (Fig. 6). Petiole and postpetiole (Fig. 7) finely punctate with a few costulae on posterior face of postpetiole. Gaster smooth and shining. Scapes with numerous appressed white setae; dorsum of head with numerous truncated long hairs (max. length 0.25 mm); submentum with a few decumbent hairs; dorsum of pronotum with 8 truncated hairs, the anteriormost weakly spatulate (max. length 0.13 mm); mesonotum with 6 truncated hairs (max. length 0.13 mm) pro- * Present address: Department of Entomology, Texas A&M University, College Station, Texas 88743. 48 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) Figs. 1-7. 1. Base of the scape of the holotype of A. punctaticeps. 2. Base of the scape of a specimen of A. texana (collected in Culberson Co., Texas). 3. Base of the scape of a cotype of A. huachucana. (Figs. 1-3 drawn to same scale.) 4. Outline of the dorsal view of the head of A. punctaticeps. 5. Outline of the dorsal view of the head of A. texana (specimen collected in Culberson Co., Texas). (Figs. 4, 5 drawn to same scale.) 6. Lateral view of the mesosoma of the holotype of A. punctaticeps. 1. Lateral view of the petiole and postpetiole of the holotype of A. punctaticeps. (Figs. 6, 7 drawn to same scale.) podeum with one small hair anterior to base of spine; dorsum of petiole with 6 truncate hairs, postpetiole with 8 similar hairs; gaster with numerous weakly truncated hairs (max. length 0.14 mm). Head and mesosoma concolorous reddish-brown, petiole, postpetiole and gaster slightly darker. Female and male: unknown. Type material. Holotype and paratype worker, collected together at a bait on the Jornada Experimental Range, Dona Ana Co., NM on 7 Aug. 1 979 by Wendy Wisdom. The holotype is deposited in the United States National Museum, the paratype in the Los Angeles County Museum of Natural History. Etymology. From Latin, referring to the punctate head. Remarks. This species is closely related to A. texana (Emery) to which it would go in the most recent key to the North American species (Creighton, 1950). It differs from A. texana in that the posterior border of the head is moderately pointed whereas it is rounded in A. texana (Figs. 4, 5). In addition, the dorsum of the head is almost completely punctate whereas it is rugose with the interrugal spaces coarsely punctate in A. texana. Occasionally the dorsum of the head of specimens of A. texana is not completely rugose, but at least the sides of the head and the region posterior to the frontal area have coarse rugae in such specimens. It is also related to A. huachucana 1989 NEW SPECIES OF APHAENOGASTER 49 Creighton (and A. huachucana crinimera Cole), but can be easily distinguished by the shape of the base of the scape. Aphaenogaster huachucana has an antennal scape with an enlarged blunt angular lobe which projects forward (Fig. 3). The lobe in A. punctaticeps and A. texana is much smaller and sharply pointed (Figs. 1, 2). The propodeal spines are slightly larger than those on the cotypes of A. huachucana, but the spines of specimens of A. huachucana from the Chiricahua Mountains of south- eastern Arizona tend to be larger than those of the cotypes, reducing the importance of this character in defining ^4. huachucana. The shape of the head of^. punctaticeps in dorsal view is more similar to that of A. huachucana than it is to A. texana. The key in Creighton (1950) can be modified as follows to accommodate this new species: 20. Large workers 5.5 mm in length; female 7 mm in length [female unknown in A. punctaticeps] 20a Large workers 4.5 mm in length; female 5.5 mm in length . .texana subsp. carolinensis 20a. Dorsum of head with prominent coarse rugae; common in southern United States texana Dorsum of head without prominent coarse rugae, a few poorly defined rugae present posterior to frontal area, rest of head punctate; rare, collected only in southern New Mexico punctaticeps This is apparently a very rare ant. Despite several years of extensive collecting using a variety of methods and at differing times, we have not seen this species again. ACKNOWLEDGMENTS Dr. David Smith, Systematic Entomology Laboratory, Agriculture Research Service, Wash- ington, D.C. arranged the loan of cotypes of Aphaenogaster huachucana from the United States National Museum. The research was supported by the National Science Foundation Long Term Ecological Research Program, grant #BSR 8 1 14466, Walter G. Whitford principal investigator. LITERATURE CITED Creighton, W. S. 1950. The ants of North America. Bull. Mus. Comp. Zool. 104:1-585 + 57 plates. Snelling, R. R. 1981. The taxonomy and distribution of some North American Pogonomyrmex and descriptions of two new species (Hymenoptera: Formicidae). Bull. Southern Calif Acad. Sci. 80:97-112. Received July 27, 1987; accepted August 4, 1988. J. New YorkEntomol. Soc. 97(l):50-55, 1989 NOTES ON ANT LARVAE: PONERINAE George C. Wheeler and Jeanette Wheeler 3358 NE 58th Avenue, Silver Springs, Florida 32688 Abstract.— larvae of four species of ants in the genera Platythyrea, Plectroctena and Streblognathus are described. The larvae of Streblognathus and Simopelta are characterized for the first time. Included also are a few additional references to ponerine larvae found in the literature. Most of this article is the result of a gift of the larvae of four species of African ants from Martin Villet of the University of Witwatersrand in Johannesburg. All these larvae seem weird even to seasoned students of 800 species in 200 genera. Streblognathus, which has never been previously studied has unique maxillae and tubercles, which are queer even in a tribe noted for peculiar tubercles. Platythyrea lamellosa has hairs which are unique among all known ant larvae; in fact, if they did not have alveolus and articular membrane we would call them tubercles. Even among tubercles they would be unique. Plectroctena conjugata has about 1,600 tubercles, which exceeds by far the number in any other species of ant larvae. Because of Brown’s 1975 revision of Platythyrea the nomenclature of the species we have studied has become quite confused. The following changes should therefore be vndidQ'.— australis in 1971 becomes parallela\ incerta in 1971 becomes pilosula\ Eubothroponera tasmaniensis in 1971 becomes Platythyrea turneri. Under MATE- RIAL STUDIED in our 1976b Memoir (p. 97) change australis Forel to parallela (F. Smith) and incerta Emery to pilosula (F. Smith). In our Ten-Year Supplement (1986b) under MATERIAL STUDIED (p. 699) delete tasmaniensis (Forel) and “de- lete australis, incerta."' To summarize, the six species of Platythyrea that we have studied previously are cribrinodis (Gerstacker), inermis Forel, modesta Forel, par- allela (F. Smith), pilosula (F. Smith) and turneri Forel. In 1976a (p. 59) we used the name Plectroctena sp. It should be changed to Plec- troctena crypt ica Bolton. We described a mature larva of one species of Simopone in 1986a, but we did not characterize the genus, because we hoped that someone would send us the mature larva of another species. Thus far we have hoped in vain. The terms used below for describing profiles and mandible shapes are defined in our 1976 Memoir. Whenever we refer to our own publications we give only the year. TRIBE AMBLYOPONINI Genus PRIONOPELTA Mayr Prionopelta amabilis Borgmeier Holldobler and Wilson ( 1 986:45). “The prey are given directly to the larger larvae.” 1989 LARVAL PONERINAE 51 Fig. 1. Platythyrea lamellosa. a, Prothoracic hair, x254; b, typical body hair in side view, X 254; c, typical body hair in surface view, x 254. TRIBE CYLINDROMYRMECINI Genus Cylindromyrmex Mayr Cylindromyrmex williamsi Wheeler W. M. Wheeler (1924:104) described the larvae as very long and slender, with narrow, curved neck and small head; the body lacked tubercles and was covered with numerous short even hairs. TRIBE PLATYTHYREINI Genus PLATYTHYREA Roger Platythyrea lamellosa Roger Fig. 1 Mature (?) larva. Length (through spiracles) 8.8-12.1 mm. Profile platy thyreoid. Similar to P. inermis (1952:118) except as follows. Ends of body not so strongly curved ventrally and tail blunter, ventral tubercles feeble. Leg and wing vestiges present. Integument spinulose, the spinules numerous and on short transverse ridges. Body hairs sparse, short (about 0.025 mm long), narrower at the base and wider at the top; narrower on T1 (0.005 mm at base and 0.012 mm at the top), elsewhere wider (0.012 mm at base and 0.03 mm at top); top surface covered with rounded, closely packed projections. Head hairs longer (about 0.025 mm long), moderately numerous, unbranched, slender. Labrum longer than broad, with apex narrowly paraboloidal; posterior surface densely spinulose, the spinules so long and the rows so close together that the spinules overlap. Mandible with middle half of all surfaces spinulose, the spinules isolated and rather small. Maxillary apex with spinules large and isolated; palp with 5 sensilla. Opening of sericteries a narrow transverse slit. Hypopharynx densely spinulose, the spinules coarse and isolated. Material studied. 4 larvae from E. Sigodink, Zimbabwe, courtesy of Martin Villet. Platythyrea schultzi Forel Length (through spiracles) 5.7-7. 1 mm. Profile platythyreoid, i.e., similar to P. cribrinodis (1971:1198). Similar to P. inermis except in the following details. Swellings on venter of AIV-AVII only. Integument with 2 sizes of spinules: (1) numerous, minute and frequently in rows; (2) fewer, larger and isolated. Body hairs 0.013-0.075 52 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) mm long. Head hairs few (about 36) and about 0.013 mm long. Labrum with breadth equal to length; anterior and ventral surfaces with about 40 hairs (0.013 mm long) and spinulose sensilla; the spinules on the posterior surface isolated and rather large. Maxillary palp with 5 (3 terminal and 2 lateral) sensilla. Labial palp with 5 (2 apical, 2 subapical and 1 basal) sensilla. Material studied. 5 larvae from Republic of South Africa, courtesy of Martin Villet. TRIBE PONERINI Genus HYPOPONERA Santschi Characterization. 1971:1210. Change “Antennae large” to “Antennae minute.” Genus PLECTROCTENA F. Smith In our 1986b Ten- Year Supplement the specialization index should be changed to 14. Plectroctena conjugata Santschi Mature larva. Length (through spiracles) 9.6-14.2 mm. Similar to P. cryptica (1976a: 59) except as follows. Neck more slender and curved. Body beset with about 1,600 tubercles, most numerous on AIV-AVI, 0.03-0.39 mm tall, hairs 0.06 mm long. Spiracles on T2 0.054 mm in diameter, decreasing posteriorly. Spinules on integu- ment isolated and coarser posteriorly and dorsally. Body hairs about 0.025 mm long, unbranched, smooth, confined to each ventrolateral surface (8 on Tl, 4 on T2, 4-6 on T3). Head hairs moderately numerous (about 44) and longer (0.025 mm long). Apical tooth of mandible curved abruptly medially. Maxilla with large isolated spi- nules apically, smaller near base of palp and galea; palp a tall cylinder; galea taller and with more slender apex. Labial palp taller. Submature larva. Length (through spiracles) 6. 9-9.0 mm. Similar to mature larva except as follows. Tubercles fewer (about 1,400). Head hairs fewer (about 30). Medial teeth of mandible reduced. Ventral border of labium trilobed; opening of sericteries not salient. Material studied. 5 larvae from Mkuzi Game Reserve, Natal, courtesy of Martin Villet. Genus SIMOPELTA Mayr Profile myrmecioid (i.e., elongate and rather slender; curved ventrally; without a differentiated neck; diameter greatest at sixth abdominal somite, decreasing gradually toward the anterior end and more rapidly toward the posterior end). Spiracles minute. Body hairs lacking. Cranium longitudinally subelliptical and sclerotized. Head hairs lacking. Mandible dinoponeroid (i.e., narrowly subtriangular, distal portion strongly curved posteriorly and slightly curved laterally, with one blunt medial tooth). In our 1986b Ten-Year Supplement this genus keys to “Profile 5. Myrmecioid” to which must be added a new rubric “1 g. Mandibles dinoponeroid.” The specialization index is 18. 1989 LARVAL PONERINAE 53 Genus STREBLOGNATHUS Mayr Profile pogonomyrmecoid but more slender and with a long slender neck. Body beset with numerous (about 300) tubercles; typical tubercle a slender spirelike sub- cone; integument with isolated spinules. Body hairs lacking. Cranium subhexagonal. Head hairs moderately numerous, minute, unbranched. Mandible ectatommoid; blade small; apical tooth curved slightly posteriorly; anterior surface sparsely spinulose. Maxilla divided into (1) a raised area bearing palp and galea and (2) a lobose apex; palp a dorsally curved subcylinder with 1 3 apical sensilla. Labial palp with 5 sensilla. In our 1976 Memoir this genus keys to “Profile 1 . Pogonomyrmecoid” under which it fits rubric 23 c. The specialization index is 14. Streblognathus aethiopicus (F. Smith) Figs. 2-4 Mature larvae. Length (through spiracles) 13.5-16.8 mm. Profile pogonomyrme- coid (neck formed from thorax, AI and All); ventral profile of remainder of abdomen nearly straight, anus ventral. Leg and wing vestiges present. Body beset with numerous (about 300) tubercles; typical tubercle a slender spirelike subcone; integument with isolated spinules; tubercles shortest on thorax (up to 0.4 mm) and longest (0.78 mm) near anus. Ten pairs of spiracles of equal size. Integument spinulose, the spinules more numerous and in transverse rows ventrally, elsewhere isolated, and less nu- merous posteriodorsally, absent around bases of tubercles. Body hairs lacking. Cra- nium subhexagonal. Antennae at midlength of cranium. Head hairs moderately nu- merous (about 60), minute (0.003-0.008 mm long), unbranched, smooth and slender. Labrum subrectangular but with base feebly constricted, with about 25 sensilla con- centrated near lateral surfaces; with each ventrolateral surface slightly raised and bearing 3 scattered sensilla; ventral surface feebly convex and densely spinulose with 4 isolated and 2 clusters of 3 sensilla each; posterior surface spinulose, the spinules large and isolated basomedially and ventrolaterally, elsewhere slender and in short arcute rows. Mandible ectatommoid, slender; heavily sclerotized; blade slender; api- cal tooth curved medially and slightly posteriorly; subapical tooth partially anterior to apical tooth, small basal tooth blunt; anterior surface with short isolated spinules directed laterally. Maxilla divided into (1) a raised area (bounded basally by a scler- otized band) bearing palp and galea and (2) a lobose apex; entire integument spinulose, the spinules rather coarse and isolated; palp a dorsally curved subcylinder with about 13 (12 with a spinule each and 1 encapsulated) sensilla; palp digitiform with 2 apical sensilla. Labium densely spinulose, the spinules large and isolated; transverse basal welt densely spinulose, the spinules slender and in short transverse rows; palp pax- illiform with 5 (4 with a spinule and 1 encapsulated) sensilla; opening of sericteries wide and salient. Hypopharynx densely spinulose, the spinules short and in short to long transverse rows. Young larvae. Length (through spiracles) 5. 1-5.9 mm. Similar to mature larva except as follows. Body with about 250 tubercles which are conoidal or subcylindrical with rounded top, lacking integumentary spinules. Spiracles set on small tubercles. Dorsal profile of head concave. Mandible lacking spinules on anterior surface. Max- illary spinules smaller; palp a short frustum directed slightly ventrally. Labium tri- 54 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) Figs. 2-4. Streblognathus aethiopicus. 2. MATURE LARVA, a, Head in anterior view; x 2 1 ; b, left mandible in anterior view, x 50; c, AX tubercle, x41; d, thoracic tubercle, x41; e, larva in side view, x 6; f, left maxilla in anterior view, x 50. 3. YOUNG LARVA, a. Head in anterior view, x21; b, left mandible in anterior view, x50; c, AX tubercle, x41; d, thoracic tubercle, x41; e, larva in side view, x6. 4. VERY YOUNG LARVA, a, Head in anterior view, x21; b, left mandible in anterior view, x 50; c, AX tubercle, x41; d, thoracic tubercle, x41; e, larva in side view, x 6. lobed and with smaller spinules; palp paxilliform. Hypopharynx with smaller spi- nules. Very young larvae. Length (through spiracles) 4.0-4. 3 mm. Similar to mature larva except as follows. About 160 tubercles, reduced to rounded knobs. Spiracles set on rounded tubercles similar to other tubercles. Mandible lacking medial tooth and surface spinules. Maxillary palp a rounded knob; galea a short subcylinder. Labium bilobed; palp a slight elevation; opening of sericteries a narrow slit. Hypopharynx with numerous minute spinules. Material studied. 12 larvae from Vernon Crookes N. R. nr. Scotburgh, Natal, Republic of South Africa, courtesy of Martin Villet. LITERATURE CITED Brown, W. L. 1975. Contribution toward a reclassification of the Formicidae. V. Ponerinae, tribes Platythyreini, Cylindromyrmecini, Acanthostichini, and Aenictogetini. Search. [Cornell Univ. Agric. Exp. Sta.] 5(1):1-115. 1989 LARVAL PONERINAE 55 Holldobler, B. and E. O. Wilson. 1986. Ecology and behavior of the primitive cryptobiotic ant Prionopelta amabilis. Insectes Sociaux 33:45-58. Wheeler, G. C. and Jeanette Wheeler. 1952. The ant larvae of the subfamily Ponerinae. Amer. Midland Nat. 48:1 1 1-144, 604-672. Wheeler, G. C. and Jeanette Wheeler. 1971. Ant larvae of the subfamily Ponerinae: second supplement. Ann. Entomol. Soc. Amer. 64:1197-1217. Wheeler, G. C. and Jeanette Wheeler. 1976a. Supplementary studies on ant larvae: Ponerinae. Trans. Amer. Entomol. Soc. 102:41-64. Wheeler, G. C. and Jeanette Wheeler. 1976b. Ant Larvae: Review and Synthesis. Mem. Entomol. Soc. Washington No. 7, 108 pp. Wheeler, G. C. and Jeanette Wheeler. 1 986a. Supplementary studies on ant larvae: Ponerinae. Trans. Amer. Entomol. Soc. 112:85-94. Wheeler, G. C. and Jeanette Wheeler. 1986b. Ten-year supplement to “Ant Larvae: Review and Synthesis.” Proc. Entomol. Soc. Washington 88:684-702. Wheeler, W. M. 1924. The Formicidae of the Harrison Williams Galapagos Expedition. Zoologica 5:101-122. Received May 18, 1988; accepted July 21, 1988. J. New York Entomol. Soc. 97(l):56-64, 1989 BIOLOGY OF ANDRENA CRATAEGI ROBERTSON (HYMENOPTERA: ANDRENIDAE), A COMMUNALLY NESTING BEE Eben a. Osgood Department of Entomology, University of Maine, Orono, Maine 04469 Abstract. — The seasonal history, nesting activity and other aspects of the biology of Andrena (Plastandrena) crataegi Robertson are described. This species is the second North American species of Andrena known to nest communally. Nest structure, orientation by females to the nest and nest site, and parasitism are discussed. The genus Andrena contains 51 1 species in America north of Mexico (Krombein et al., 1979). The biology of most species is unknown. This paper describes the nest, life history, and other aspects of the biology of Andrena {Plastandrena) crataegi Robertson. A. crataegi is transcontinental in southern Canada and the northern United States, and its range extends to Georgia and New Mexico. LaBerge (1969) gave details of its range and floral records and redescribes the species. Rau (1922) removed a bee of this species from a shallow burrow which was evidently under construction. No further biological information is available. Known cases of communal nesting behavior in the genus Andrena are rare. A British species A. bucephala Stephens was found by Perkins (1917) to have a large number of females per nest. Yarrow and Guichard (1941) found the same for A. ferox Smith, another British species. These are probably communal but detailed studies and dissections of ovaries were not made. In North America, A. erythronii Robertson usually has one female per nest but occasionally a nest is used simulta- neously by two females (Michener and Rettenmeyer, 1956). A. accepta Viereck was determined to be communal by Rosen (1973). A. crataegi is the second North Amer- ican species known to have communal nests. It was one of 20 species of Andrena collected on the low-bush blueberry complex by Boulanger et al. (1967), and was the second most abundant species of Andrena collected on this complex in Maine. Specimens associated with this study are in the collection of the Department of Entomology at the University of Maine. STUDY SITE AND METHODS The communal nesting behavior of A. crataegi was first observed on 16 June 1966, when 1 1 females emerged from a single nest. A second nest was located nearby in 1967. This latter area was the principal study site and was located in a commercial low-bush blueberry field (Fig. 1) in Deblois, Washington County, Maine. Vegetation consisted principally of Vaccinium angustifolium Ait. and grasses, and cover was rather sparse. There was little surface organic matter, and the leached E layer was visible on the surface. Similar nesting areas are fully described by Osgood (1972). 1989 BIOLOGY OF ANDRENA CRATAEGI 57 Figs. 1,2. 1 . Nesting site of A. crataegi in a commercial low-bush blueberry field. 2, Female pausing at the nest entrance. 58 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) The A. crataegi population increased in subsequent years and field data were collected at this site in 1968, 1973 and 1987. Commercial blueberry fields in this area are burned every other year; therefore, nests were marked with a steel wire placed 5 cm due north of the entrance. Small screen cones (Linsley et al., 1952) were used to collect adults of A. crataegi and its parasites as they emerged in the spring. Females of A. crataegi which were to be dissected were placed in alcoholic Bouin’s solution for 24 hours and then transferred to 70% ethyl alcohol. Plaster of Paris was poured into nests to aid in nest excavation for life history studies. Temperature readings were taken with a mercury bulb ther- mometer in the shade 5 cm above ground level. BIOLOGY Seasonal history and nesting activity. Spring emergence data of A. crataegi from a large communal nest observed in 1973 shows conclusively that the species is proter- androus. The site was visited at 2-4 day intervals and on the morning of 18 May, 28 males were released from the screen cone covering the nest entrance. Most of these probably emerged on 1 7 May as the soil under the cone contained many burrows approximately 1.25 cm in depth. Most burrows contained one or occasionally two males which had spent the night in these shallow excavations. Six males of Nomada cressonii Robertson also emerged from the nest on this date. On 20 May 12 males of A. crataegi emerged and were released and one emerged on 23 May. The next visit was on 27 May. Ten males and 19 females of A. crataegi had emerged at 10:45 a.m. and were released. The screen cone was then removed from the nest entrance which was observed closely. By 1 :00 p.m. twenty one additional females had emerged for a total of 40. By 1:10 p.m. emerged females had returned to the nesting area and began to enter the nest. Females continued to emerge but since newly emerging bees could not be distinguished from those entering, counts ceased. Two specimens of Nomada sp. also emerged the morning of 27 May. One was released. The other was collected and determined to be a female of N. cressonii suggesting that this species is also proterandrous. The first blooms of pin cherry, Prunus pennsylvanica L., in the immediate area were observed on 18 May. Male bees were collected resting on pin cherry blooms on this date, and a mating pair collected on pin cherry blooms on 27 May were identified as A. crataegi. Other matings of A. crataegi were observed on pin cherry and at the nesting site. Females of A. crataegi often use the same nest entrance year after year. Nest construction was first observed on 30 May 1973 when a small tumulus was observed at one of the entrances to the previous year’s nest. On 1 June the tumulus, which was on a level area, was 3 cm in diameter, round, with the entrance in the center. The first females laden with pollen were also observed entering the nests on this date. Six pollen laden females were collected on 20 June. These were dissected and all contained 2-6 well-developed oocytes, indicating that each was capable of ovipos- iting. Observations on daily and seasonal activity of a rather large communal nest were made on several days in 1 987. Females were provisioning nests on 28 May and daily activity was observed periodically thereafter. On a clear morning the first female emerged from the nest on a foraging trip as early as 7:34 a.m. and at a temperature 1989 BIOLOGY 0¥ ANDREN A CRATAEGI 59 as low as 15.2°C. On a foggy overcast morning the first foraging trip did not begin until clearing occurred at 8:55 a.m. at a temperature of 20°C. The latest the last female was observed entering the nest was 7:05 p.m. at a temperature of 24°C. Light intensity and humidity were not measured. During active foraging periods females pause in the nest entrance (Fig. 2) for 5- 30 seconds before taking flight. Females orient to the nest and nesting site before and during flight (Figs. 3-6). Upon emerging the female turns and faces the nest entrance (Fig. 3). She then hovers 1-2 cm above the ground near the nest entrance (Fig. 4) and slowly increases the height and distance from the nest (Figs. 5, 6). Figure 5 also shows another female in the nest entrance. They then fly at a height of 1-2 meters and 3-4 meters on all sides of the nest before leaving on a foraging trip. Early in their season of foraging activity (last of May) nearly all females orient to the nest on the first trip of the day but none orient on subsequent trips that day. Observations on 1 1 June showed that 13 of 29 females oriented on the first trip and for three of these the orientation time spent was relatively brief. On 1 6 June a second nest entrance had been constructed 5 cm from the original, and was being used. From 16 June to the end of their activity period in July only an occasional female was observed to exhibit nest orientation behavior and for some of these it was quite brief. On their return to the nest with pollen and nectar females flew directly to the nest entrance and quickly entered. The daily tripping rates on 3 1 May for a single nest containing 44 females were recorded (Fig. 7). The first female appeared at the nest entrance at 8:50 a.m. and emerged at 8:55 a.m. Individual females continued to leave the nest, usually at 1-2 minute intervals, until all 44 had emerged by 11:23 a.m. The first female entered the nest with pollen at 1 1:45 a.m. Assuming the first out was the first to return, trip time was 2 hours 50 minutes. The first female left the nest for the second trip at 12:36 p.m. Assuming the first female completing the first foraging trip was the first female emerging for a second trip, time in the nest was 5 1 minutes. Eighteen females did not make a third trip as it began to darken and rain at 5:40 p.m. Under these conditions, foraging bees entered the nest at short intervals, ten entering in the final 9 minutes of activity (Fig. 7). The recorded time elapsed on different days for a single foraging trip varied con- siderably from 2 hours 50 minutes to a maximum of 4 hours 43 minutes. On one occasion, when over 4 hour trips were recorded, the foliage was wet in the morning from rain the previous night, and there was a strong breeze all day. But later in their active foraging period on 2 July it was warm, foliage was dry, winds were nearly calm and trips of over 4 hours also occurred. This may have been due to the lack of pollen and nectar resources. As expected, the number of foraging females decreased as the season progressed. From 44 on 31 May they decreased to 29 on 1 1 June, 20 on 16 June, 19 on 17 June, 11 on 24 June and 2 on 2 July. In other years, females have been collected in the area on Spiraea latifolia (Ait.) Borkh. as late as 20 July. Nest construction and contents. Three nests were excavated on various dates to gain information on seasonal development of A. crataegi and its parasites and on nest architecture of^. crataegi. Nests varied in size but the architecture was similar. The largest nest (Fig. 8) contained four entrances and is used to illustrate nest ar- chitecture. It was excavated following completion of the seasonal development of A. 60 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) Figs. 3-6. Orientation of A. crataegi female to the nest and nesting site. 3. Female on the ground facing the nest entrance. 4-6. Females hovering at an increasing height and distance from the nest entrance. Figure 6 with arrows showing nest entrance (A), bee (B), and shadow of bee (C). 1989 BIOLOGY 0¥ ANDREN A CRATAEGI 61 Fig. 7. Daily tripping rates for a communal nest of A. crataegi containing 44 females. crataegi and is, therefore, discussed last. The two nests excavated earlier in the season were smaller, contained a single entrance, and the architecture was similar to the left portion of Figure 8. Individual cells removed from a nest excavated on 3 July 1973 during the adult female activity period contained pollen and several life history stages. Eleven females of A. crataegi were still active in provisioning this nest on 3 July, and one adult of Nomada sp. was also observed. Twenty nine cells were recovered; 1 9 contained larvae of A. crataegi, 2 contained larvae of Nomada sp., and 8 cells were in various stages of being provisioned. Cells were most abundant at a depth of approximately 38 cm. In order to avoid disturbing nearby nests to be studied later, some cells were probably missed. Cells were essentially horizontal (Fig. 8) and varied in length from 13-14 mm (7 measurements). Cells were wider near the rear, symmetrical around their long axis, and tapered gradually to the cell entrance. Maximum diameter was 7. 3-7. 5 mm (5 measurements), 5 mm from the rear of the cell. They were 5 mm in diameter (4 measurements) at the cell entrance. Cell closure was concave on the inside but the spiral pattern described by Rozen (1973) for A. accepta was not evident. Cells were lined with a shiny, waterproof substance, and the soil within 0.5 cm of the cells was more compact than the surrounding soil. The complete pollen mass (one observed) was yellow, spherical, firm, moist and placed near the rear of the cell. The egg taken from this cell was white, slightly curved, and 1 .8 mm in length and 0.4 mm in diameter. Other eggs taken from ovarioles after they had been preserved were of similar shape and color and were 2.2 mm in length and 0. 5-0.6 mm in diameter. Larvae from the first to probably the last instar were 62 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) Fig. 8. Two dimensional diagram of a large communal nest of A. crataegi showing 4 nest entrances. Distances shown are between entrances at soil surface. present on this date, and feces were observed as early as 1 3 July from larvae brought back to the laboratory. A second nest with a single entrance was excavated in early September. The number of females originally active in provisioning this nest is not known, but the combined length of the burrow and branches was 1.08 meters and varied in diameter from 0.6- 0.8 cm with some areas considerably enlarged. Of 54 cells recovered, 15 contained adult males of A. crataegi (3 callow), 14 contained adult females of A. crataegi (1 callow), 8 contained adults of N. cressonii (4 male and 4 female) and 1 7 contained larvae of an undetermined species of bombyliid. Adults of^4. crataegi and N. cressonii remain in the cells and emerge the following spring. The largest nest was excavated on 20 September 1973, and the number of pro- visioning females could not be determined. It contained four entrances (Fig. 8) which were relatively close together. The main burrow and its branches were open. This nest had been used for more than one year, and the nonuniformity of the burrow diameters was striking. Burrows varied from 0.7- 1.0 cm in diameter in the upper and lower extremities of the burrows with several enlarged areas ranging from 1989 BIOLOGY OF ANDRENA CRATAEGI 63 1.8-2. 4 cm in diameter at the confluence of burrows. The purpose of these enlarged areas is not known but may allow for easier passage of entering and exiting bees in these nests containing a large number of active individuals. The interconnected burrows descended and began to ramify from one of these enlarged areas at a depth of about 30 cm. The burrow extended to a depth of 60 cm and contained 2.44 meters of open burrow. From the extensiveness and pattern of the burrow, it seems most likely that the Plaster of Paris did not reach all of the open burrows. No attempt was made to recover all cells belonging to this nest. Lateral connectives leading from the main burrow were 5-6.4 cm in length and filled with loose soil. It was impossible in most instances to connect cells with the proper branch of a burrow. Burrows of other nests of^4. crataegi were nearby, further complicating the situation. Cells were found at depths from 33-53 cm with the largest number being found at approximately 38 cm. Of the cells recovered, 16 contained adults of A. crataegi (1 1 males and 5 females), two contained adults of N. cressonii ( 1 male and 1 female) and one contained a bombyliid larva. Adults may emerge in the spring from the same nest entrance used during the previous season, and as previously mentioned, females often use the same nest entrances year after year. Often they emerge 8-10 cm from the original entrance. Some establish new nests in the immediate vicinity from whence they emerge, and others may nest elsewhere. The population of A. crataegi was observed to increase in the study area. When the nesting area was located in 1967 only 1 nest entrance was present. There were 7 in 1972 and 11 in 1973 all within a radius of approximately 2 meters. Three were determined to be occupied by lone females in 1973. Parasitism. Since all cells were probably not recovered from excavated nests, quantitative data on parasitism rates could not be determined, and, in any event, data on 1-2 nests in any one year probably are not significant. The incidence of N. cressonii and an undetermined species of bombyliid have been noted. No antagonism was shown by females of A. crataegi toward females of N. cressonii as they passed in the nest entrance. The number of parasites varies greatly in different years (Perkins 1917) and judged by the adult activity of N. cressonii that was certainly true in this study. In 1973 several adults of N. cressonii were over the nest site and periodically entered the nests during most of the activity period of A. crataegi, but during 54 hours of observation on nine different days in 1987 not one was seen. Adults of Sphecodes sp. and an unidentified anthomyiid also occasionally entered the nests and remained for long periods of time, but none were found in the cells. A crab spider, Misumena vatia (Clerck), was collected from Rosa virginiana Mill, feeding on a female of A. crataegi. ACKNOWLEDGMENTS 1 thank Wallace E. LaBerge, Illinois Natural History Survey, for verification of the identifi- cation ofT. crataegi, G. I. Stage, U.S.D.A., Systematics Entomology Laboratory, for verification of the identification of its parasite N. cressonii, Daniel T. Jennings, F.S. U.S.D.A., for identi- fication of the spider, Robert V. Rourke, University of Maine, for soil information, John B. Dimond, University of Maine, and Wallace E. LaBerge for their helpful comments on the manuscript. 64 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97( 1) LITERATURE CITED Boulanger, L. W., G. W. Wood, E. A. Osgood and C. O. Dirks. 1967. Native bees associated with the low-bush blueberry in Maine and eastern Canada. Maine Agri. Exp. Sta. Tech. Bull. 26:22 pp. Krombein, Karl V., Paul D. Hurd, Jr., David R. Smith and B. D. Burks et al. 1979. Catalog of Hymenoptera in America north of Mexico. Smithson. Inst. Press, Washington, D.C. LaBerge, Wallace E. 1969. A revision of the bees of the genus Andrena of the western hemi- sphere. Part II. Plastandrena, Aporandrena, Charitandrena. Amer. Entomol. Soc. Trans. 95:1-47. Linsley, E. G., J. W. MacSwain and R. F. Smith. 1952. Outline for ecological life histories of solitary and semi-social bees. Ecology 33:558-567. Michener, Charles D. and Carl W. Rettenmeyer. 1956. The ethology of Andrena erythronii with comparative data on other species (Hymenoptera, Andrenidae). Univ. Kansas Sci. Bull. 37, Pt. 2, No. 16:645-684. Osgood, E. A., Jr. 1972. Soil characteristics of nesting sites of solitary bees associated with the low-bush blueberry in Maine. Maine Agri. Exp. Sta. Tech. Bull. 59:8 pp. Perkins, R. C. L. 1917. Andrena bucephala Steph. and Nomada bucephalae Perk, in Dev- onshire, and notes on their habits. Entomol. Monthly Mag. 53 (Ser. 3, Vol. 3), pp. 198- 199. Rau, Phil. 1922. Ecological and behavior notes on Missouri insects. Trans. Acad. Sci., St. Louis 24:1-71. Rozen, Jerome G., Jr. 1973. Biology notes on the bee Andrena accepta Viereck (Hymenoptera, Andrenidae). Jour. N.Y. Entomol. Soc. 81:54-61. Yarrow, I. H. H. and K. M. Guichard. 1941. Some rare Hymenoptera Aculeata, with two species new to Britain. Ent. Monthly Mag. 77 (Ser. 4, Vol. 3), pp. 2-13. Received June 17, 1988; accepted July 26, 1988. J. New YorkEntomol. Soc. 97(l):65-72, 1989 THE GENUS METOPINA (DIPTERA: PHORIDAE) FROM CRETACEOUS AND TERTIARY AMBERS David Grimaldi Department of Entomology, The American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024 Abstract. — Six specimens of the genus Metopina are reported from the late Oligocene amber of Chiapas, Mexico, the lower Miocene amber of the Dominican Republic, and with one of the specimens, Metopina goeleti, n. sp., the oldest known phorid, from the late Cretaceous amber of New Jersey. Details of the morphology and distribution are discussed with respect to living members of the genus. Species distinctions between the specimens in the Oligomiocene ambers are not apparent. Taxonomic placement of the Sciadoceridae previously described from Cretaceous amber of Canada is discussed, and it is hypothesized that the Phoridae s.s. appeared much earlier than the late Cretaceous. While sorting and identifying Diptera in ambers recently acquired by the American Museum, the distinctive genus Metopina Macquart was discovered in pieces from Chiapas, Mexico and the Dominican Republic. The finding was made more significant by a specimen in a piece of amber from the late Cretaceous of New Jersey, in which there are about a dozen arthropods, including the oldest known bee, Trigona prisca (Michener and Grimaldi, 1988). The geology of the New Jersey fossil resins is dis- cussed elsewhere (Grimaldi et al., 1989). Not only does this extend the geological record of the Phoridae, but it provides a record ranging from the present and through the Tertiary to the Mesozoic, which is more complete than for any other group of phorids. Strangely enough, Metopina is unknown from the well-studied, huge deposits of Baltic amber. Due to the small size of these phorids, it is quite likely that any specimens of them in this amber have been overlooked by earlier workers. Metopina also might be rare in most ambers because of their flower- visiting habits. Methods used in studying the amber are given in Grimaldi (1987). References to geological ages of the various amber deposits are given in the sections dealing with the various specimens. THE TERTIARY AMBER SPECIMENS Figs. 1-7 Five specimens of Metopina were found in Tertiary ambers, four from Chiapas, Mexico, and one from the Dominican Republic. The Mexican amber is considered to be late Oligocene by Hurd et al. (1962) and studies on the lower Miocene ages of at least some of the Dominican amber is briefly discussed in Grimaldi ( 1 987). Detailed locality information is given under the account of each specimen. The five Tertiary specimens collectively have more features preserved than are seen in the Cretaceous specimen, but any species distinctions between the Chiapas and Dominican amber specimens have not been found. Chiapas amber specimens: University of California (Berkeley) Museum of Pa- 66 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97( 1 ) Figs. 1-5. Metopina in amber from Chiapas, Mexico. 1, 2. UCMP B-7046-32, lateral view of habitus and oblique dorsal view of head and notum. 3. UCMP B-7048, lateral view. 4. Dorsal view of portion of head of UCMP B-7045-58, showing the supra-antennal setae and the pair of setae that flank them (the dorsal portion of the head and thorax of this specimen were missing). 5. AMNH Ch47, lateral view. Figures 1 through 8 are to the same scale. leontology (UCMP) nos. B-7046-32, B-7045-58, B-7048, and American Museum of Natural History no. AMNH Ch47. All are from the mines near Simojoval, Chiapas State, the deposits of which are discussed in Hurd et al. (1962). Well-preserved, taxonomically important features are the following. 1989 AMBER METOPINA 67 Figs. 6-8. Metopina in amber from the Dominican Republic and the late Cretaceous of New Jersey. 6, 7. AMNH 1 1848B, from the Dominican Republic, frontal and oblique lateral views. 8. Metopina goeleti, n. sp., holotype (AMNH C88720), from New Jersey. Lateral view, with most of the venation labeled. Dorsal portion of the head, and the notum, halter, and abdominal tergites are darkest portions of the body. Head: eyes higher than long; front broad and with numerous fine interfrontal setulae (cf. Figs. 13, 14); apparent on at least UCMP B-7048 is a pair of parallel supra- antennal bristles, lying on the dorsal margin of the ptilinal suture (cf. Figs. 13, 14), and in UCMP B-7045-58 a pair of slightly divergent bristles flank this pair (this 68 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) feature is more obvious in this specimen, because of the orientation); apparent in UCMP B-7048 and B-7046-32 is a pair of inclinate (convergent) orbitals (the anterior fronto-orbital bristles in Figs. 1 3 and 1 4), at about the middle of the front and slightly lateral to the supra-antennal bristles; postocellar setae parallel or subparallel, flanked by a pair of inclinate inner verticals and a pair of lateroclinate (divergent) outer verticals; flagellomere 1 spherical, and the arista (flagellomeres 2-4) is Vh to 3 times the length of fl- 1 ; deep cavities of smooth contour house the pedicels (cf. Fig. 1 3) and are separated by a very thin carina (seen in B-7048); palp is longer than the pedicel, with 3 strong, straight setae at or near the apex; proboscis, seen in UCMP B-7048, thin, projected anteriad. Specimen 7046-32 possesses a median furrow along the length of frons (cf Figs. 13, 14) (this feature is obscure in the other specimens, if present). Thorax: notum evenly covered with fine, short setulae; fore coxa of B-7048 as wide as fore femur and slightly concave; hind tibia with an apical seta, length about equal to width of tibia; lateral surface of hind tarsomere 1 with 2 hair seams (cf , Fig. 12); in B-7048 and AMNH Ch47 hind tarsomere 1 slightly wider than the tibia, but not so in the other Chiapas specimens; mesal surface of hind tarsomere with 5 rows of ctenidial combs (cf Fig. 1 0); lateral surfaces of hind tarsomeres 2-4 with a single hair seam each extended along most of their lengths; wing with arcuate M2 and CuAi veins (veins M3+4 and A,, respectively, in Disney [1985]; and veins 5 and 6, re- spectively, of other papers by Disney), but not as strongly curved as in the Cretaceous species; no evidence of a small fork at the apex of vein R4+5; M2 and R4+5 separated by a very small gap; scutellum with a pair of apical, convergent setae; pair of slightly convergent prescutellar setae on posterior margin of notum and slightly lateral to scutellars (not all scutellars are presented in the illustrations, as they were not apparent on all of the specimens). Abdomen: details of the terminalia are not apparent, but UCMP B-7046-32 can be identified as a female and B-7048 as a male. The female lacks the distinctive flap on tergite 5 (Fig. 9). Dominican amber specimen: AMNH 1 1 848, from mines in the vicinity of Santiago, Santiago Province. The distal portion of the abdomen of this specimen is missing, and some other portions of the body are obscured by concoidal fractures. Although a fully frontal view of the head is available, the orbital setae are so fine in Metopina that they are virtually invisible in this specimen. Only the long, stout palpal setae and the pair of nearly parallel postocellars are apparent on the head. Veins M2 and CuAi are arcuate, the former slightly more so than in the Chiapas specimens (but not as strongly as in the Cretaceous specimen), and the latter vein with about as much curvature as in the Chiapas specimens. Metopina goeleti, new species Fig. 8 Diagnosis. The most distinctive features are in the wing venation: apical portion of R4+5, distal to M2, is straight and meets the costal vein at about a 45° angle; M2 strongly arcuate, its main axis almost parallel to the main axis of R4+5; CuAj very strongly arcuate, the distal portion at nearly a right angle to middle portion of this vein. Description. The right side of the specimen shows the best view with most of the 1989 AMBER METOPINA 69 wings and legs intact and their detailed features visible. Some clearing of the specimen has occurred, but the darkest (most melanized) portions are the notum, scutellum, tergites, and halter knob. Apparently the legs, pleura, and palpi were light yellowish. Although there is only an oblique lateral view of the head, visible are 2 straight, parallel postocellars flanked by divergent vertical setae, and 3 straight, stout setae on the apex of the palp. No long setae were seen on any segments of the forelegs. Midtarsomere 1 with a terminal seta on ventral surface. Hind leg with coxa about one-half the length of the femur, femur nearly twice as wide as tibia and about as long; combined lengths of tarsomeres slightly longer than length of the tibia. Relative lengths of hind tarsomeres: tarsomere 1 > 2=3 > 4=5. Mesal surface of hind femur with small, heavily sclerotized tubercle or spine near distal margin, which is appar- ently the same structure as that shown in Fig. 1 1 for M. oligoneum. Apex of hind tibia with 5 setae of lengths one-half to full width of tibia, the seta in the most ventral/ anterior position being longest. Lateral surface of hind tarsomere 1 with 2 hair seams along the middle for most of length of segment, 5 rows of ctenidia on the mesal surface, and 4 setae at apex, of lengths approximately equal to those on tibia. Mesal surface of hind tarsomere 2 with 4 short, stout setae that project anteriad. Lateral surfaces of hind tarsomeres 2, 3, and 4 each with a single hair seam along most of their lengths. Wing venation distinctive, as described in the diagnosis. Holotype. Female, in amber piece AMNH C88720, from upper Cretaceous of New Jersey, found at Kinkora, New Jersey by Alfred C. Hawkins. In the amber collection of the American Museum of Natural History. The date of collection and exact strati- graphic position is unknown, but as discussed in Michener and Grimaldi (1988) and in Grimaldi et al. (1989), this deposit is probably from the upper Magothy Formation, or about 80 million years old. Etymology. Named for Mr. Robert Goelet, former president of the American Museum, whose generosity made this and other studies of amber possible. Discussion. This is the oldest known Phoridae, sensu stricto. The previously oldest known specimens, disregarding the Sciadoceridae, are from Baltic amber. Because the source deposits for most of the Baltic material have not yet been discovered, it is presumed to range in age between the upper Eocene and lower Oligocene. One sciadocerid and a phorid are known from the Canadian Cretaceous amber, of an age approximately 72 million years old (McAlpine and Martin, 1966), which accords with dating based on the austral distribution of the living representatives. While I agree with Hennig (1973) that Prioriphora canadambra McAlpine and Martin should be placed in the Phoridae, Sciadophora bostoni McAlpine and Martin certainly should not. Prioriphora canadambra has the thickened, dark, radial veins which are greatly shortened like the costal vein, with unbranched M,, M2, and M3 veins that are nearly parallel, as is diagnostic for most of the Phoridae. Sciadophora bostoni venation is plesiomorphic with respect to phorid venation at the family level in virtually every respect: veins C and R4+5 reach nearly to the apex of the wing, the stem and fork of Ml and M2 is retained, and vein M3 +4 originates at a distinctive basal-medial cell. CONCLUSIONS The genus Metopina presently consists of 33 species, 31 of them being described. The other two, undescribed species, are from Malaysia and Sulawesi (R. H. L. Disney, pers. comm.). The most recently described species is Metopina ciceri, the only known 70 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) Figs. 9-14. Scanning electron micrographs of various features of living Metopina. 9. Flap on tergite 5 of female M. oligoneura. 10. Ctenidial comb, on mesal surface of hind tibia, M. recurvata. 11. Spur on mesal surface of hind femur, M. oligoneura. 12. Hair seams, lateral surface of hind tibia, M. oligoneura. 13. Head of critical point-dried male of M. oligoneura. 14. Frontal surface of head of air-dried female of M. recurvata (eyes are collapsed), afo, anterior fronto-orbital seta; mf, median furrow; pfo, posterior fronto-orbital seta; p, palp; po, postocellar seta; sa, supra-antennal seta; ui, upper interfrontal seta. Scales are 40 gm, except Figure 1 1 , which is 10 gm. member of the genus from Asia (Disney, 1 988). The New World tropics is apparently the most diverse area for this genus. There are 7 species from South America, 3 from Central America, and two from the Caribbean. The two Caribbean species are Me- topina reflexa (from Puerto Rico) and M. recurvata (from Dominica). No species are 1989 AMBER METOPINA 71 as yet known from the Dominican Republic, but this may just as well reflect little collecting effort there as it may a lack of the genus on Hispaniola. The only North American species is M. subarcuata, ranging from Idaho to Canada. In the only other paper treating Oligomiocene amber phorids (Disney, 1987), the taxa treated were a species of Dohrniphora and 3 species of Megaselia in Dominican amber. Given the minute size of Metopina specimens, the amber specimens could not be examined for genitalic features, number of palpal segments, the presence of bifid tibial/tarsal comb hairs (Disney, 1983), and various aspects of chaetotaxy. Access to such characters would be necessary in order to accurately place the fossil species among the extant ones. The fossils possess a combination of features indicating their placement in the genus Metopina, as well as having some features plesiomorphic with respect to at least some Metopina. One of these plesiomorphic features is the absence of a flap-like abdominal tergite 5 in the females (Fig. 9). This flap was interpreted by Borgmeier (1963) as the basal portion of tergite 5, with tergites 5 and 6 being fused, although this latter interpretation is not what Disney (1986) concludes. Given this plesiomorphic feature it might be reasonable to name a new subgenus or genus for these amber specimens. In a transformation series towards reduction in the length and ultimately the number of radial veins, it must be assumed that absence of the small vein R2+3 is a loss, and therefore apomorphic with respect to those taxa (most other metopinine and non-metopinine phorids) that possess the full complement of venation. The presence of at least two Phoridae in the late Cretaceous, one of which is apparently of a recently-derived group {Metopina), strongly suggests the family to be older than the fossils. Speculations on such ages should await detailed phylogenetic and bio- geographic studies, in conjunction with the discovery of new fossils from various Cretaceous ambers. Other than Prioriphora canadambra, no other phorids are known from the Cretaceous ambers (McAlpine and Martin, 1 969; Schlee and Dietrich, 1 970). ACKNOWLEDGMENTS I thank David Lindberg of the Museum of Paleontology at Berkeley for loaning to me the Chiapas specimens, to F. Christian Thompson for loaning specimens of Metopina recurvata, to Mr. Robert Goelet for his generous support of this research, and especially to R. H. L. Disney, for providing many informative comments on the manuscript and phorids in general, and for a series of Metopina oligoneura. B. V. Peterson and an anonymous reviewer also provided useful comments on the manuscript. LITERATURE CITED Borgmeier, T. 1963. Revision of the North American phorid flies. Part 1. The Phorinae, Aenigmatiinae and Metopininae, except Megaselia (Diptera, Phoridae). Studia Entomol. 6:1-256. Disney, R. H. L. 1983. A useful new character in the giant genus Megaselia (Diptera: Phoridae), with two new species from Britain. Zeit. Angew. Zool. 70:225-234. Disney, R. H. L. 1985. Re-interpretation of the wing veins in the Phoridae (Diptera). Entomol. Mon. Mag. 121:55-58. Disney, R. H. L. 1986. Morphological and other observations on Chonocephalus (Phoridae) and phylogenetic implications for the Cyclorrhapha (Diptera). J. Zool. 210:77-87. Disney, R. H. L. 1987. Four species of scuttle fly (Diptera: Phoridae) from Dominican amber. Pan-Pac. Ent. 63:377-380. 72 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97( 1 ) Disney, R. H. L. 1988. A remarkable new species of scuttle fly (Diptera: Phoridae) whose larvae infest chickpea root nodules in India. J. Nat. Hist. 22:61 1-616. Grimaldi, D. A. 1987. Amber fossil Drosophilidae (Diptera), with particular reference to the Hispaniolan taxa. Amer. Mus. Novit. 2880:1-23. Grimaldi, D., C. Beck and J. Boon. 1988. Occurrence, chemical characteristics, and paleon- tology of fossil resins from New Jersey. Amer. Mus. Novit. (manuscript). Hennig, W. 1973. Diptera. Kukenthal’s Handbuch der Zoologie 31:1-337. Hurd, P. D., Jr., R. F. Smith and J. Wyatt Durham. 1 962. The fossiliferous amber of Chiapas, Mexico. Ciencia (Mex.) 21:107-118. McAlpine, J. F. and J. E. H. Martin. 1966. Systematics of Sciadoceridae and relatives with descriptions of two new genera and species from Canadian amber and erection of family Ironomyiidae (Diptera: Phoroidea). Can. Entomol. 98:527-544. McAlpine, J. F. and J. E. H. Martin. 1 969. Canadian amber— a paleontological treasure-chest. Can. Entomol. 101:189-838. Michener, C. D. and D. A. Grimaldi. 1988. A Trigona from late Cretaceous amber of New Jersey (Hymenoptera: Apidae: Meliponinae). Amer. Mus. Novit. 2917:1-10. Schlee, D. and H-G. Dietrich. 1970. Insektenfiihrender Bernstein aus der Unterkreide des Libanon. N. Jb. Geol. Palaont. Mh. 1970:40-50. Received June 30, 1988; accepted August 19, 1988. J. New York Entomol. Soc. 97(l):73-86, 1989 NAUCORIDAE (HETEROPTERA) OF NEW GUINEA. IV. A REVISION OF THE GENUS CAVOCORIS WITH DESCRIPTIONS OF FOUR NEW SPECIES Dan a. Polhemus and John T. Polhemus University of Colorado Museum, 3115 S. York St., Englewood, Colorado 801 10 Abstract.— The genus Cavocoris is revised based on recent collections from Papua New Guinea, and its generic characters are compared to other endemic Papuan genera. Four new species are described: C rotundatus, C. ibatiri, C. minor, and C ismayi. A key to species, a species distribution map, and illustrations of key characters are provided. The monotypic genus Cavocoris was described by La Rivers (1971) to hold C. bisulcus, based on a single female type from what is now the Indonesian province of Irian Jaya. This species has a deep pit on the posterior margin of abdominal paratergite III, a character that La Rivers considered to be diagnostic for the genus as a whole. Our recent collections in Papua New Guinea, however, have revealed four additional species of Cavocoris, two of which lack pits on paratergite III. Furthermore the pits are a sexually dimorphic character, being present only in the females of those species which possess them. These pits are not associated with the spiracle on paratergite III and must instead have a separate function, possibly sensory or secretory. In particular, the presence of a tapering glabrous channel leading outward from each pit suggests a flow path for secretory products. This “flow channel” leads to one of two paired ovate glabrous patches set into the hydrofuge pile of paratergite III (Fig. 3); these ovate patches, whose function is not known but is presumed to be sensory on the basis of their location and appearance, occur on other abdominal paratergites which lack pits, and in many other naucorid taxa. The structure and function of similarly located “sense organs” in several plastron breathing naucorid genera was discussed by Polhemus (1986), but in these swift water genera the abdominal and thoracic paratergites bear ovate depressions that are not paired or glabrous as in Cavocoris and other slow water species. It is hypothesized that the structure of these “sense organs” is closely linked to habitat, and that modified types may have evolved independently in several naucorid lineages whose members have shifted from an- cestral slow water habitats to swift streams (Polhemus and Polhemus, 1986a). La Rivers (1971) compared C. bisulcus to Sagocoris biroi Montandon, noting that it differed from the latter in the presence of abdominal pits, by certain head modi- fications discussed below, and by the shape of the posterolateral angles of the prono- tum. The only specimen at his disposal, however, was the female type of C. bisulcus, which is macropterous and thus has a pronotal shape different from typical bra- chypterous morphs (Figs. 1, 4). The brachypterous forms, which we have found to be far more common, generally have a broadly arcuate pronotal margin that forms an acute angle posterolaterally. Cavocoris does share with Sagocoris and Aptinocoris a reduced and truncate male left paramere and a distinctly asymmetrical vesica with a membranous tip, but we have found that far better key characters for distinguishing 74 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) Cavocoris from these genera lie in the structure of the head, particularly in the anteclypeal margin which forms a smooth projecting lip (Fig. 2) without a distinct notch above the base of the labrum as is encountered in Sagocoris and Aptinocoris. The labrum itself is also characteristic, coming to a point apically in a condition reminiscent of the New World Limnocoris. These head modifications and the presence of short pale club-like setae on the hemelytra of certain species represent synapo- morphies allying Cavocoris to the Papuan Nesocricos, these two genera apparently representing sister groups that have diverged from the more plesiomorphic Sagocoris lineage. Nesocricos is further defined by several autapomorphies, including an elon- gate body shape reminiscent of Belostomatidae, and the presence of cup-like sense organs at the anterolateral angles of the proepimeron. La Rivers separated Cavocoris from Warisia, another related genus in the Sagocoris complex, by the number of transverse rows of spines on the mesotibia. We feel this character is rather subjective, however, since these spine rows are variable in size and often difficult to count, and would note that the head structures of the two genera once again provide a much more reliable method of separation. In Warisia the anteclypeal structure is intermediate between Sagocoris-Aptinocoris and Cavocoris- Nesocricos, with a shallow indentation in the anteclypeal margin above the extremely recessed labrum, flanked by large ovate depressions of a presumably sensory nature. The abdominal venter of Warisia is also different, bearing a longer and less densely appressed hydrofuge pile than in Cavocoris, causing it to appear dark brown instead of shining gold. In addition, the “sense organs” on the abdominal paratergites in Warisia consist of one to four irregular elongate glabrous patches rather than the paired ovate patches as discussed above for Cavocoris. Finally, the male parameres in Warisia are of approximately equal size, while in Cavocoris the left paramere is reduced. All of the above genera are classified as members of the subfamily Cheirochelinae by La Rivers (1971), but as we have indicated previously (Polhemus and Polhemus, 1986b; D. Polhemus, 1986) this subfamily is almost undoubtedly polyphyletic and the assignation of these taxa to it is dubious. The vertex of Cavocoris is not produced posteriorly behind the eyes, thus it will not even key to the Cheirochelinae in Usinger’s (1941) key. All measurements and proportions are given in millimeters. CL numbers refer to codes used by the authors to reference ecological data. Specimen depository abbre- viations are indicated in the acknowledgments. This research was sponsored in part by a grant from the National Geographic Society, Washington, D.C. KEY TO THE SPECIES OF CAVOCORIS LA RIVERS Note: the keys to males and females contain different sets of species, since certain species are known only from a single sex. Males 1. Length less than 9.5 C. minor - Length exceeding 10.0 2 2. Pronotum set with short stout pale setae; overall length exceeding 1 1.5 C ibatiri - Pronotum lacking short stout pale setae; overall length less than 11.0 C. rotundatus 1989 REVISION OF CA VOCORIS 75 Females 1 . Abdominal paratergite III bearing a distinct pit (Fig. 3) 2 - Abdominal paratergite III not bearing a pit C. ismayi 2. Stemite VI with projections on posterior margin at base of subgenital plate (Fig. 14) C. bisulcus - Sternite VI lacking projections on posterior margin (Figs. 11,12) 3 3. Subgenital plate with indentation at tip (Fig. 1 1); pit on abdominal paratergite III deep C. rotundatus - Subgenital plate lacking an indentation at tip (Fig. 1 2); pit on abdominal paratergite III shallow C. ibatiri Cavocoris bisulcus La Rivers Figs. 4, 14 Cavocoris bisulcus La Rivers, 1971, 2:57. Diagnosis. This species may be recognized by the pair of rounded projections on the posterior margin of abdominal stemite VI in females and by the deeply indented apex of the female subgenital plate (Fig. 1 4). The third abdominal paratergite bears a deep pit, and the posterolateral angles of abdominal tergites III-V are moderately produced. The male is unknown. Discussion. We have examined the type and only known specimen of this species, a macropterous female taken by Evelyn Cheesman in the Cyclops Mountains behind present day Jayapura in Indonesian New Guinea. The distinctive female subgenital morphology will separate it from all other known Cavocoris species. Material examined. INDONESIA, Irian Jaya: 1 female, Sabron, Cyclops Mtns., 930 ft (283 m), VI-36, B.M. 1936-271, L. E. Cheesman (holotype, BMNH). Cavocoris rotundatus, new species Figs. 1-3, 5, 6, 1 1 Diagnosis. C. rotundatus may be recognized by the deep pit along the posterior margin of abdominal paratergite III in females (Fig. 3), the absence of protuberances on the posterior margin of female abdominal stemite VI, the indentate apex of the female subgenital plate (Fig. 1 1), and the structure of the male parameres (Figs. 5, 6). Description. Brachypterous form: Of moderate size, ovate, basic coloration dull yellowish brown with scattered dark brown or black markings. Male length 10.46; maximum width (across abdomen) 7.49 mm; female length 10.56; maximum width 7.68. Head dark yellowish, with longitudinal dark brown stripe medially and transverse dark markings at posterior margin of vertex, width/length = 3.50/2.21; eyes brown, shining, width/length = 0.77/1.54, dorsal surfaces flat, not rising above plane of vertex, inner margins divergent anteriorly, separated from vertex by shallow furrows set with very short pale setae, anterior/posterior interocular width = 2.06/ 1 .92, lateral flange small, glabrous; posterior margin of vertex weakly and broadly rounded, barely produced behind eyes; anteclypeus with anterior margin broadly rounded, barely projecting ahead of eyes, produced beyond base of labmm for distance less than 76 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) length of labrum, bearing shallow depressions to either side of midline; labrum triangular, coming to acute point distally, light brown; maxillary plates moderately developed, oriented at angle approximately 45° from vertical, anterior margins gla- brous, carinate, forming sides of rostral cavity; rostrum yellowish basally, second segment gold, glabrous, extending beyond labrum; antennae slender, filiform, yel- lowish, segment IV glabrous, barely extending beyond lateral eye margin. Pronotum dark yellowish, mottled centrally with dark brown and black at muscle attachments, weakly depressed medially behind vertex, width/length (midline) = 6.53/1.73, lateral margins narrowly glabrous, broadly rounded, posterolateral angles acutely rounded, posterior margin nearly straight, not sinuate. Scutellum dark brown, width/length (midline) = 2.88/1.44, lateral margins weakly sinuate, transverse sulcus present along anterior margin. Hemelytra dark brown, lighter brown bordering scu- tellum, posteromedially, and along inner margin of embolium, embolium dark yel- low, surface of corium coarsely rugose, entire hemelytral surface bearing fine pale granular microstructure and scattered short stout erect pale setae, tips of hemelytra rounded, extending to base of genital segment, embolium demarcated by deep narrow furrow on inside margin, posterior margin obscure, lateral margin narrowly glabrous, bearing long recumbent gold setae, hemelytral commissure with small triangular tab on left hemelytron fitting into corresponding triangular indentation on right heme- lytron. Abdomen with lateral portions of segments II-VIII exposed, dark yellow, with dark brown markings along anterior margins, lateral margins of all segments bearing recumbent gold setae, these setae becoming longer and forming tufts at posterolateral angles of segments V-VIII, posterolateral angles of segments III and IV moderately produced and spinose, angles of segments V-VII acute, tips of projections on segment VIII rounded. Ventral surface light brown, with head, prostemum, mesostemum centrally, and abdomen covered with thick recumbent gold hydrofuge pile; head with prominent glabrous median longitudinal keel lacking dentation and having weak posterior pro- jection over similar and continuous structure on prostemum; proepimeron densely covered with very short fine recumbent gold setae, inner projections not touching medially; mesostemal plate sharply reflexed along anterior margin, coming to acute subconical point anteromedially, point separated by transverse sulcus from broad tumescence posteromedially; females with deep pit present centrally along posterior margin of abdominal paratergite III and anterior margin of paratergite IV; abdominal paratergites II-V with paired ovate glabrous pits adjacent to spiracle, spiracle rep- resented by small raised protruberance thickly covered with gold hydrofuge setae, paratergites VI and VII each with single glabrous pit, all paratergites with lateral margins narrowly glabrous. Legs dark yellowish, anterior femora with thick pad of gold setae along anterior margin, fringe of long fine gold setae along posterior margin; anterior tibia slender, gently curving, with short gold setae on inner face, anterior tarsi single segmented, claw tiny, obscure, fused to tarsus; middle and hind coxae each bearing single glabrous tubercle distally; middle and hind trochanters with narrow longitudinal fringe of short thick gold setae distally on posterior margins; middle and hind femora bearing scattered short reddish spines along anterior margins, continuous longitudinal rows of short sharp reddish spines along posterior margin on dorsal and ventral faces, single small combs of reddish spines distally on posterior 1989 REVISION OF CA VOCORIS 77 Figs. 1-10. 1-3. C. rotundatus new species 1. Female, dorsal habitus. 2. Head, anterior view, showing lack of notch above labrum and triangular labrum coming to acute point. 3. Ventral view of abdominal paratergites III and IV, showing location of deep pit, spiracles, and small ovate glabrous patches in hydrofuge pile. 4. C. bisulcus La Rivers. Lateral margin of pronotum in macropterous individual. 5-10. Cavocoris species, male parameres. 5. C. rotun- datus new species, left paramere. 6. C. rotundatus new species, right paramere. 7. C. ibatiri new species, left paramere. 8. C. ibatiri new species, right paramere. 9. C. minor new species, left paramere. 10. C. minor new species, right paramere. 78 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) margins; middle femur with thick pad of short gold setae on posterior face; middle and posterior tibiae and posterior tarsi thickly set with longitudinal rows of stout reddish spines, these spines longer and more dense distally; middle tarsi lacking spines dorsally, bearing longitudinal rows of short reddish spines ventrally; middle and posterior femora, tibia and tarsi set with gold swimming hairs on posterior margins; claws gold, sharply bent; parempodia setiform. Female subgenital plate trapezoidal on basal half, apical half narrowed with sides parallel, tip broadly and shallowly indented; abdominal stemite VI lacking projections on posterior margin adjacent to base of subgenital plate (Fig. 1 1). Male parameres asymmetrical; left paramere smaller, leaf shaped; right paramere gently curving, with rounded projection laterally (Figs. 5, 6). Macropterous form: unknown. Discussion. The type locality was a clear rocky tributary to the Wampit River, descending from the Herzog Mountains through lush primary rain forest. The type series was taken in a shallow reach with a gravel substrate and moderate current. Etymology. The name “rotundatus” refers to the nearly circular shape of this species when viewed from above. Holotype. Male, allotype, female: PAPUA NEW GUINEA, Morobe Province, stream 17.8 km N of Mumeng on Wau rd., September 19, 1983, CL 1835, J. T. and D. A. Polhemus (BPBM). Paratypes. 1 male, 2 females, same data as holotype (JTPC). Cavocoris ibatiri, new species Figs. 7, 8, 12 Diagnosis. This species, the largest in the genus, may be recognized by its size (length exceeding 1 1.50), the presence of erect pale peg-like setae on the pronotum and scutellum in addition to the hemelytra, the presence of a pit on abdominal paratergite III in females, the structure of the female subgenital plate (Fig. 12), and the shapes of the male parameres (Figs. 7, 8). Description. Brachypterous form: Large for genus, shape elongate ovate, basic coloration dull yellowish brown broadly and diffusely marked with dark brown. Male length 11.81; maximum width (across abdomen) 8.06; female length 12.48 mm; maximum width 7.97. Head dark yellowish, with three longitudinal dark brown stripes medially and transverse dark markings at posterior margin of vertex, width/length = 4.03/2.26; eyes brown, shining, width/length = 0.96/1.92, dorsal surfaces flat, not rising above plane of vertex, inner margins divergent anteriorly, separated from vertex by shallow furrows set with very short pale setae, anterior/posterior interocular width = 2.30/ 2.06, lateral flange small, glabrous; posterior margin of vertex weakly and broadly rounded, barely produced behind eyes; anteclypeus with anterior margin broadly rounded, barely projecting ahead of eyes, produced beyond base of labrum for dis- tance less than length of labrum, bearing shallow depressions to either side of midline; labrum triangular, acutely rounded apically, light brown; maxillary plates moderately developed, oriented at angle approximately 45° from vertical, anterior margins gla- brous, carinate, forming sides of rostral cavity; rostrum yellowish basally, tip glabrous, distal two segments extending beyond labrum; antennae slender, filiform, yellowish, segment IV barely extending beyond lateral eye margin. 1989 REVISION OF CAVOCORIS 79 Figs. 1 1-14. Cavocoris species, female subgenital plates. 1 1. C. rotundatus new species. 12. C. ibatiri new species. 13. C. ismayi new species. 14. C. bisulcus La Rivers. Pronotum yellowish brown, mottled centrally with dark brown at muscle attach- ments, weak ovate depression present centrally, width/length (midline) = 7.15/2.21, lateral margins narrowly glabrous, broadly rounded, posterolateral angles acutely rounded, posterior with weak sinuations to either side of midline at point of contact with lateral embolar margins; entire pronotal surface set with short erect pale peg- like setae. Scutellum dark brown, width/length (midline) = 3.46/1 .82, surface coarsely rugose, set with short erect pale peg-like setae, lateral margins weakly sinuate, trans- verse sulcus present along anterior margin. Hemelytra dark brown, lighter brown in narrow irregular patches bordering scutellum and posteromedially, anterior portion of embolium dark yellow, membrane black, surface of corium coarsely rugose, entire hemelytral surface bearing fine pale granular microstructure and scattered short erect pale peg-like setae, tips of hemelytra rounded, extending to posterior margin of abdominal tergite V in males and to posterior margin of abdominal tergite VI in females, embolium demarcated by deep narrow furrow on inside margin, posterior margin obscure, lateral margin narrowly glabrous, lacking long recumbent gold setae, hemelytral commissure with small triangular tab on left hemelytron fitting into cor- responding triangular indentation on right hemelytron. Abdomen with lateral portions of segments II-VIII exposed, brown, with darker markings along posterior and lateral margins, lateral margins of all segments bearing fringe of gold setae, with tufts of long gold setae arising inside of lateral margins near middle, posterolateral angles of segments II- VI acute, not produced or spinose, angles of segments VII and VIII in females and VIII in males rounded. Ventral surface light brown, with head, prostemum, basal portions of propleurae, mesostemum centrally, hind coxae and abdomen covered with thick recumbent gold hydrofuge pile; head with prominent glabrous median longitudinal keel lacking den- tation and evenly meeting similar and continuous structure on prostemum; ventral margin of prostemal keel rising to gentle obtuse angle when viewed laterally; proepi- meron densely covered with very short fine recumbent gold setae, inner projections not touching medially; mesostemal plate sharply reflexed along anterior margin. 140 80 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) Fig. 15. Distribution of Cavocoris species in New Guinea. Areas above 2000 m stippled. ■— C. bisulcus La Rivers. •— C ibatiri new species. — C. rotundatus new species. T— C. minor species. ♦ — C. ismayi new species. 1989 REVISION OF CA VOCORIS 81 coming to acute subconical point anteromedially, point separated by transverse sulcus from broad tumescence posteromedially; females with deep pit present centrally along posterior margin of abdominal paratergite III; abdominal paratergites IV-VII with paired ovate glabrous pits adjacent to spiracle, spiracle represented by small raised protruberance thickly covered with gold hydrofuge setae, paratergites II and III each with single glabrous pit, all paratergites with lateral margins narrowly glabrous. Legs dark yellowish, anterior femora with thick pad of gold setae along anterior margin, fringe of long fine gold setae along posterior margin; anterior tibia slender, gently curving, with short gold setae on inner face, anterior tarsi single segmented, claw tiny, obscure, fused to tarsus; middle and hind coxae each bearing single glabrous tubercle distally; middle and hind trochanters with narrow longitudinal fringe of short thick gold setae distally on posterior margins; middle and hind femora bearing scattered short reddish spines along anterior margins, continuous longitudinal rows of short sharp reddish spines along posterior margin on dorsal and ventral faces, single small combs of reddish spines distally on posterior margins; middle femur with narrow longitudinal patch of thick short gold setae along posterior face, middle and posterior tibiae and posterior tarsi thickly set with longitudinal rows of stout reddish spines, these spines longer and more dense distally; middle tarsi lacking spines dorsally, bearing longitudinal rows of very short spines on ventral face; middle and posterior femora, tibia and tarsi set with gold swimming hairs on posterior margins; claws gold, sharply bent; parempodia setiform. Female subgenital plate trapezoidal on basal half with glabrous patches laterally, apical half narrowed with sides parallel, tip broadly rounded; abdominal stemite VI lacking projections on posterior margin adjacent to base of subgenital plate (Fig. 12). Male parameres asymmetrical; left paramere truncate, with blunt projection at tip; right paramere gently curving, with slender apical projection (Figs. 7, 8). Macropterous form: unknown. Discussion. C. ibatiri is by far the most distinctive species in the genus, and differs from other species of Cavocoris in several respects. In all other species in which the female is known the subgenital plate is indented apically, whereas in C. ibatiri it is broadly rounded at the tip. The male right paramere is also much different in shape than in the other species for which this character is known (Figs. 6, 8, 10), and the overall shape of the body is massive and elongate, rather than rotund when viewed from above. The dorsum is set with short pale club-like setae, a character state found in no other Papuan genus except Nesocricos, and this, in concert with the somewhat elongate body shape mentioned above, indicates that C. ibatiri is an annectant taxon linking Nesocricos and Cavocoris. The type locality was a small ankle deep stream approximately 3 meters wide mostly shaded by primary rain forest. The type series was taken in a shallow rocky riffle where a gap in the surrounding forest allowed sunlight to reach the stream. Etymology. The name “ibatiri” is a noun in apposition and refers to a water spirit believed to dwell in the rivers of the Papuan highlands. Holotype. Male, allotype, female: PAPUA NEW GUINEA, Western Highlands Province, small stream at Baiyer River Bird of Paradise sanctuary, September 8, 1983, CL 1792, J. T. and D. A. Polhemus (BPBM). Paratypes. PAPUA NEW GUINEA. Western Highlands Province: 3 males, 1 female, same data as holotype (JTPC); 1 male, 1 female, Trauna River, nr. Baiyer 82 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) River, September 8, 1983, CL 1793, J. T. and D. A.. Polhemus (JTPC). Morobe Province: 1 female, Bulolo River, E of Wau, 900 m (2,950 ft), October 25, 1964, W. L. and J. G. Peters (LACM); 2 females, stream 17.8 km N of Mumeng on Wau rd., September 19, 1983, J. T. and D. A. Polhemus (JTPC). Cavocoris minor, new species Figs. 9, 10 Diagnosis. C. minor, the smallest species so far known in the genus, may be recognized by its small size (length less than 10.00), narrow pronotum (less than 6.00 in width), the tufts of setae on the posterolateral angles of the abdominal segments, and the structure of the male parameres (Figs. 9, 10). The female is presently un- known. Description. Brachypterous male: Of moderate size, ovate, basic coloration dull yellowish brown with scattered dark brown or black markings. Length 9.12; maxi- mum width (across abdomen) 6.62. Head dark yellowish, with longitudinal dark brown stripe medially and transverse dark markings at posterior margin of vertex, width/length = 3.22/2.02; eyes black, shining, width/length = 0.72/1.39, dorsal surfaces flat, not rising above plane of vertex, inner margins divergent anteriorly, separated from vertex by shallow furrows set with very short pale setae, anterior/posterior interocular width = 1.92/1.73, lateral flange small, glabrous; posterior margin of vertex weakly and broadly rounded, barely produced behind eyes; anteclypeus with anterior margin broadly rounded, barely projecting ahead of eyes, produced beyond base of labrum for distance less than length of labrum, bearing shallow depressions to either side of midline; labrum triangular, coming to acute point distally, light brown; maxillary plates moderately developed, oriented at angle approximately 45° from vertical, anterior margins gla- brous, carinate, forming sides of rostral cavity; rostrum yellowish basally, second segment gold, glabrous, extending beyond labrum; antennae slender, filiform, yel- lowish, segment IV glabrous, barely extending beyond lateral eye margin. Pronotum dark yellowish, mottled centrally with dark brown at muscle attach- ments, weakly depressed medially behind vertex, width/length (midline) = 5.57/1 .54, lateral margins narrowly glabrous, broadly rounded, posterolateral angles acutely rounded, posterior margin nearly straight, not sinuate. Scutellum dark brown, width/ length (midline) = 2.88/1.25, lateral margins weakly sinuate, transverse sulcus present along anterior margin. Hemelytra dark brown to black, lighter brown bordering scutellum, embolium dark yellow, surface of corium coarsely rugose, entire hemelytral surface bearing fine pale granular micro structure and scattered short stout erect pale setae, tips of hemelytra rounded, extending to base of genital segment, embolium demarcated by deep narrow furrow on inside margin, posterior margin obscure, lateral margin narrowly glabrous, bearing long recumbent gold setae, hemelytral commissure with small triangular tab on left hemelytron fitting into corresponding triangular indentation on right hemelytron. Abdomen with lateral portions of segments II-VIII exposed, dark yellow, with dark brown markings along anterior margins and centrally, lateral margins of all segments bearing recumbent gold setae, these setae becoming longer and forming tufts at posterolateral angles, posterolateral angles of segments III and IV moderately 1989 REVISION OF CA VOCORIS 83 produced and spinose, angles of segments V-VII acute, tips of projections on segment VIII rounded. Ventral surface light brown, with head, prostemum, mesostemum centrally, and abdomen covered with thick recumbent gold hydrofuge pile; head with prominent glabrous median longitudinal keel lacking projections and evenly meeting similar and continuous structure on prostemum; proepimeron densely covered with very short fine recumbent gold setae, inner projections not touching medially; mesostemal plate sharply reflexed along anterior margin, coming to acute subconical point an- teromedially, point separated by transverse sulcus from broad tumescence postero- medially; abdominal paratergites II-V with paired ovate glabrous pits adjacent to spiracle, spiracle represented by small raised protruberance thickly covered with gold hydrofuge setae, paratergites VI and VII each with single glabrous pit, all paratergites with lateral margins narrowly glabrous. Legs dark yellowish, anterior femora with thick pad of gold setae along anterior margin, fringe of long fine gold setae along posterior margin; anterior tibia slender, gently curving, with short gold setae on inner face, anterior tarsi single segmented, claw tiny, obscure, fused to tarsus; middle and hind coxae each bearing single glabrous tubercle distally; middle and hind trochanters with narrow longitudinal fringe of short thick gold setae distally on posterior margins; middle and hind femora bearing scattered short reddish spines along anterior margins, continuous longitudinal rows of short sharp reddish spines along posterior margin on dorsal and ventral faces, single small combs of reddish spines distally on posterior margins; middle femur with thick pad of short gold setae on posterior face; middle and posterior tibiae and posterior tarsi thickly set with longitudinal rows of stout reddish spines, these spines longer and more dense distally; middle tarsi lacking spines dorsally, bearing longitudinal rows of short reddish spines ventrally; middle and posterior femora, tibia and tarsi set with gold swimming hairs on posterior margins; claws gold, sharply bent; parempodia setiform. Male parameres asymmetrical; left paramere truncate, with blunt projection at tip; right paramere gently curving, with rounded projection laterally (Figs. 9, 10). Macropterous form: unknown. Discussion. The type locality was a deep, swift stream with steep banks flowing through disturbed rain forest. The type series was taken from under a partially submerged tree trunk swept by the current. Etymology. The name “minor” refers to this species’ small size. Holotype. Male: PAPUA NEW GUINEA, Morobe Province, stream 39 km SW of Lae on Wau rd., September 15, 1983, CL 1 8 1 2, J. T. and D. A. Polhemus (BPBM). Paratypes. 3 males, same data as holotype (JTPC). Cavocoris ismayi, new species Fig. 13 Diagnosis. Females of C. ismayi may be recognized by the small projections on the posterior margin of abdominal stemite VI, the shape of the subgenital plate (Fig. 13), the small pointed projection on the posterior end of the gular keel on the bottom of the head, and the absence of pits on abdominal paratergite III. The male of this species is presently unknown. Description. Brachypterous female: Of moderate size, ovate, basic coloration dull 84 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) yellowish brown with scattered dark brown or black markings. Length 10.56; max- imum width (across abdomen) 7.68. Head dark yellowish, with longitudinal dark brown stripe medially and transverse dark markings at posterior margin of vertex, width/length = 3.60/2.06; eyes black, shining, width/length = 0.86/1.44, dorsal surfaces flat, not rising above plane of vertex, inner margins divergent anteriorly, separated from vertex by shallow furrows set with very short pale setae, anterior/posterior interocular width = 2.02/1.78, lateral flange small, glabrous; posterior margin of vertex weakly and broadly rounded, barely produced behind eyes; anteclypeus with anterior margin broadly rounded, barely projecting ahead of eyes, produced beyond base of labrum for distance less than length of labrum, bearing shallow depressions to either side of midline; labrum triangular, coming to acute point distally, light brown; maxillary plates moderately developed, oriented at angle approximately 45° from vertical, anterior margins gla- brous, carinate, forming sides of rostral cavity; rostrum yellowish basally, second segment gold, glabrous, extending beyond labrum; antennae slender, filiform, yel- lowish, segment IV glabrous, barely extending beyond lateral eye margin. Pronotum dark yellowish, mottled centrally with dark brown at muscle attach- ments, weakly depressed medially behind vertex, width/length (midline) = 6.72/1.63, lateral margins narrowly glabrous, broadly rounded, posterolateral angles acutely rounded, posterior margin nearly straight, not sinuate. Scutellum dark brown, dark yellowish at basal angles, width/length (midline) = 3.36/1.44, lateral margins weakly sinuate, transverse sulcus present along anterior margin. Hemelytra dark brown to black, lighter brown bordering scutellum and in irregular patches posteromedially, embolium dark yellow, surface of corium coarsely rugose, entire hemelytral surface bearing fine pale granular microstructure and scattered short stout erect pale setae, tips of hemelytra rounded, extending to base of abdominal segment VIII, embolium demarcated by deep narrow sinuate furrow on inside margin, posterior margin ob- scure, lateral margin narrowly glabrous, bearing long fine recumbent gold setae, hemelytral commissure with small triangular tab on left hemelytron fitting into cor- responding triangular indentation on right hemelytron. Abdomen with lateral portions of segments II-VIII exposed, dark yellow, with dark brown markings along anterior margins and adjoining lateral margins, lateral margins of all segments bearing recumbent gold setae, these setae becoming longer and forming tufts at posterolateral angles of segment VII, posterolateral angles of segments III-V weakly produced, angles of segments VI and VII acute, tips of pro- jections on segment VIII acutely rounded. Ventral surface light brown, with head, prostemum, mesostemum centrally, and abdomen covered with thick recumbent gold hydrofuge pile; head bearing prominent glabrous median longitudinal keel on gula with small pointed projection on posterior end, evenly meeting similar and continuous keel on prostemum; proepimeron densely covered with very short fine recumbent gold setae, inner projections not touching medially; mesosternal plate sharply reflexed along anterior margin, coming to acute subconical point anteromedially, point separated by transverse sulcus from broad tumescence posteromedially; abdominal paratergite III lacking pit along posterior margin; abdominal paratergites II-V with paired ovate glabrous pits adjacent to spiracle, spiracle represented by small raised protruberance thickly covered with gold hydrofuge setae, paratergites VI and VII each with single glabrous pit, all paratergites 1989 REVISION OF CA VOCORIS 85 with lateral margins narrowly glabrous, stemite VI with two (1 + 1) small rounded projections to either side of midline on posterior margin. Legs dark yellowish, anterior femora with thick pad of gold setae along anterior margin, fringe of long fine gold setae along posterior margin; anterior tibia slender, gently curving, with short gold setae on inner face, anterior tarsi single segmented, claw tiny, obscure, fused to tarsus; middle and hind coxae each bearing single glabrous tubercle distally; middle and hind trochanters with narrow longitudinal fringe of short thick gold setae distally on posterior margins; middle and hind femora bearing scattered short reddish spines along anterior margins, continuous longitudinal rows of short sharp reddish spines along posterior margin on dorsal and ventral faces, single small combs of reddish spines distally on posterior margins; middle femur with thick pad of short gold setae on posterior face; middle and posterior tibiae and posterior tarsi thickly set with longitudinal rows of stout reddish spines, these spines longer and more dense distally; middle tarsi lacking spines dorsally, bearing longitudinal rows of short reddish spines ventrally; middle and posterior femora, tibia and tarsi set with gold swimming hairs on posterior margins; claws gold, sharply bent; parempodia setiform. Subgenital plate roughly trapezoidal basally, narrowing on apical half with lateral margins parallel, tip with weak indentation (Fig. 1 3). Macropterous form: unknown. Discussion. Eio Creek at the type locality was a clear rocky stream descending from the Sogeri Plateau through primary rain forest. The type specimens were taken amid rocks and gravel in shallow water along the edge of a deep, unshaded, flowing pool. Etymology. This species is named in honor of John Ismay, who showed us many interesting collecting localities in the vicinity of Port Moresby. Holotype. Female: PAPUA NEW GUINEA, Central Province, Eio Creek, nr. Ba- ruanumu, September 22, 1983, CL 1840, J. T. and D. A. Polhemus (BPBM). Paratypes. 1 female, same data as holotype (JTPC). ACKNOWLEDGMENTS We thank the following individuals for the opportunity to examine specimens held under their care (abbreviations following institutional names are those used in the text): Dr. C. L. Hogue, Los Angeles County Museum (LACM); W. R. Dolling, British Museum (Natural His- tory), London (BMNH). All additional material is held in the J. T. Polhemus collection, En- glewood, Colorado (JTPC); types of new species are deposited in the Bernice P. Bishop Museum, Honolulu (BPBM). Special thanks are also due to Dr. J. W. Ismay, formerly associated with the Dept, of Primary Industry, Konedobu, Papua New Guinea, who helped us in many ways during the course of our research in that country. LITERATURE CITED La Rivers, I. 1971. Studies of Naucoridae (Hemiptera). Biol. Soc. Nevada Mem. 2:1-120. Polhemus, D. A. 1986. A review of the genus Coptocatus Montandon (Hemiptera: Naucor- idae). Pan-Pac. Entomol. 62:248-256. Polhemus, D. A. and J. T. Polhemus. 1986a. Naucoridae (Hemiptera) of New Guinea. 2. A review of the genus Idiocarus Montandon, with descriptions of three new species. J. New York Entomol. Soc. 94:39-50. Polhemus, D. A. and J. T. Polhemus. 1986b. Naucoridae (Heteroptera) of New Guinea. III. 86 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) A review of the genus Tanycricos La Rivers with the description of a new species. J. New York Entomol. Soc. 94:163-173. Usinger, R. L. 1941. Key to the subfamilies of Naucoridae with a generic synopsis of the new subfamily Ambrysinae. Ann. Entomol. Soc. Amer. 34:5-16. Received March 10, 1988; accepted May 18, 1988. J. New YorkEntomol. Soc. 97(l):87-99, 1989 CARINIOCORIS, A NEW PHYLINE PLANT BUG GENUS FROM THE EASTERN UNITED STATES, WITH A DISCUSSION OF GENERIC RELATIONSHIPS (HETEROPTERA: MIRIDAE) Thomas J. Henry Systematic Entomology Laboratory, Plant Sciences Institute, Agricultural Research Service, USDA, % National Museum of Natural History, Washington, D.C. 20560 Abstract. — The new genus Cariniocoris is described to accommodate Plagiognathus geminatus Knight, P. ilicis Knight, and the new species C. nyssae from Florida and Maryland. Photographs of adults, illustrations of male genitalia, and micrographs of certain other structures are given and distributions and host plants are outlined. A neotype is designated for P. ilicis. Evidence supporting the group’s monophyly and its relationship to several other phyline genera are discussed. Keys are provided to facilitate recognition of the genus and species. Recent collecting and discovery of a plant bug feeding on ornamental black gum, Nyssa sylvatica Marsh., and subsequent efforts to place it in an existing genus, have revealed the need for a new genus. Further research shows that two other species, described in the genus Plagiognathus, require transfer to this new genus. Herein, Cariniocoris is described to accommodate Plagiognathus geminatus Knight, P. ilicis Knight, and the new species C. nyssae. Evidence for its monophyly and relationship to several other phyline genera are discussed. A neotype is designated for C. ilicis. Photographs of adults, illustrations of male genitalia, and micrographs of certain other structures are given. Keys to males and females of the genus and modifications for Knight’s (1941) key to the genera of Phylinae are provided to facilitate recognition of the genus and species. The following abbreviations are for institutions cited in this paper: AMNH (Amer- ican Museum of Natural History, New York); CU (Cornell University, Ithaca, New York); JTP (John T. Polhemus and University of Colorado Museum, Englewood, Colorado); PDA (Bureau of Plant Industry, Pennsylvania Department of Agriculture, Harrisburg); and USNM (U. S. National Museum of Natural History, Washington, D.C.). Cariniocoris, new genus Type species. Cariniocoris nyssae, new species. Diagnosis. Sexes dimorphic (males slender, color variable; females broader, color mostly green), head shorter in lateral aspect than distance from eye to apex of tylus, male genital capsule with a distinct mesal carina on the ventral surface, bases of tibial spines immaculate or nearly so, and male vesica C-shaped with a spiculate base on the primary spiculum and a middorsal flange. Description. Small to medium sized species, length 3. 0-4.0 mm, with recumbent simple setae. Head impunctate, much broader than long, width nearly 2 x length; 88 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97( 1 ) compound eye prominent; in lateral aspect, width of eye distinctly broader than distance from anterior margin of eye to tylus (Figs. 7-9); antennal fossa contiguous with and set into distinct emargination on inner lower Vs of eye. Rostrum relatively stout, segment I not extending beyond posterior margin of head, segment IV not extending beyond apices of mesocoxae or bases of metacoxae. Antenna slender; segment I shortest and thickest, but apex of segment II in some species approaching equal thickness; segment II longest, length greater than combined lengths of segments III and IV. Pronotum trapeziform, impunctate; calli weakly raised, slightly depressed behind; mesoscutum prominent; scutellum equilateral. Hemelytron macropterous and dimorphic; slender and subparallel in males, weakly arcuate or broadly oval in females. Ostiolar evaporative area (Fig. 10). Ventral surface smooth, semishiny, abdomen shiniest. Legs slender; tibial spines without or with only vague spots at bases; claws with short arolia attached for entire length on basal halves, parempodia simple, setiform, apices somewhat widened (Figs. 11-12). Ventral surface of male genital capsule with a distinct median carina or keel, arising on anterior Vs and extending to or near posterior margin (Figs. 13-16). Left paramere (Figs. 17, 21, 25) somewhat mitt-shaped with two parallel arms: right arm slender, elongate; left arm short, pointed, dorsal edge finely serrate, basal edge raised from connecting edge of two arms to form a blunt tubercle. Right paramere (Figs. 18, 22, 26) simple, elongate oval. Vesica of aedeagus (Figs. 19, 23, 27) twisted, C-shaped, having two apical spiculi and a distinct subapical secondary gonopore. Phallotheca (Figs. 20, 24, 28). Etymology. The generic name of this taxon is taken from the Latin word carina meaing carina, ridge, or keel (and coris meaning bug) and refers to the unusual carina found on the ventral surface of the male genital capsule. The gender is masculine. Discussion of generic relationships. The presence or absence of spots at the bases of the tibial spines is considered useful in distinguishing many genera of Phylini. I have examined the majority of New World and many Old World phylines in search of genera which contain species lacking these basal spots, as well as those having similar sexual dimorphism, a carina or keel on the male genital capsule, and other features possessed by species of Cariniocoris. Based on this comparison, evidence supporting the monophyly of the species being placed in Cariniocoris includes the following synapomorphies: 1) strong sexual dimorphism (males slender, subparallel, color variable; females broadly oval, color mostly pale); 2) head short in lateral aspect, area from anterior margin of eye to apex of tylus shorter than lateral width of eye; 3) rostrum relatively stout and short, not extending beyond apices of mesocoxae or bases of metacoxae; 4) ventral aspect of male genital segment with a long, narrow median carina or keel (homologous in general form with several other taxa but differing in exact structure); 5) ventral half of genital opening with a thickened rim, the right side with a broad, blunt protuberance; 6) vesica C-shaped with 2 apical spiculi, the primary one with variable patterns of spicules at base, a middorsal flange, and a subapical secondary gonopore; and 7) left paramere with right arm elongate and left arm serrate on dorsal edge. In the Nearctic, Icodema nigrolineatum (Knight), the only North America species of Icodema Reuter, has a large keel on the male genital capsule and lacks spots at the bases of the tibial spines, suggesting a relationship somewhere near Cariniocoris. However, the pale and more elongate body in both sexes, diagnostic linear black markings on the antennae and legs, the elongate structure of the head, and the different 1989 CARINIOCORIS, A NEW GENUS OF PHYLINI 89 Figs. 1-6. Adults of Cariniocoris spp.: 1) geminatus, male. 2) geminatus, female. 3) ilicis, male. 4) ilicis, female. 5) nyssae, male. 6) nyssae, female. 90 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) Figs. 7-10. 7-9. Head micrographs of adult of Cariniocoris spp., lateral aspect: 7) geminatus (113x). 8) ilicis (114x). 9) nyssae (113x). 10. Ostiolar opening of C. geminatus (221 x) in horizontal position, dorsum directed to the right. structure of the male genitalia, including the S-shaped (rather than C-shaped) vesica and a broader, more flattened genital carina serve to distinguish it from species of Cariniocoris. In the Palearctic, several genera in Wagner’s (1975) Phylus group, including at least species of Phylus Hahn and Icodema infuscatum (Fieber), have a distinct keel on the male genital capsule and lack spots at the bases of the tibial spines. Beyond these characters these taxa do not appear to share any of the other characters that define Cariniocoris. Species of Phylus are relatively large, slender bugs, having elongate, acutely produced heads, subparallel hemelytra in both sexes, and significantly dif- ferent types of male genitalia, including an S-shaped vesica. Phylus coryli (Linnaeus) and P. melanocephalus (Linnaeus), unlike species of Cariniocoris, have the genital keel reduced and very slender and lack tibial spots. Although I have not examined specimens of I. infuscatum, Wagner (1975) noted it possesses a genital keel and lacks tibial spots. This species, however, has an S-shaped vesica (figured by Wagner, 1975: 292, fig. 830h) and lacks other features shared by species of Cariniocoris. 1989 CARINIOCORIS, A NEW GENUS OF PHYLINI 91 From this preliminary review of phyline genera, the lack of distinct spots at the bases of the tibial spines and the presence of a median carina on the male genital capsule indicate that Cariniocoris, Icodema, and Phylus may form a natural group and be part of a larger clade yet to be documented. Further phylogenetic analysis of the phyline taxa sharing these and some other corroborating characters is needed, but is beyond the limits of the present study. Remarks on identification. Species of Cariniocoris will key to the genera Micro- phylellus Reuter and Plagiognathus Fieber in Knight (1941) or to Icodema Reuter in Slater and Baranowski (1978) primarily because of the dorsal coloration (dark vs. pale) and the pale yellowish tibial spines without or with only vague pale-brown spots at the bases. Knight (1925) said in describing the species Plagiognathus ilicis: “In some respects . . . intermediate between Plagiognathus and Microphylellus, while the left genital clasper is of a form quite different from either genus. The general form, color and pubescence is that of Plagiognathus while the tibial characters approach Microphy- lellus', the yellowish brown spines with brownish at base of each which scarcely forms distinct spots, may cause some difficulty in tracing ilicis through the generic key.” To facilitate generic placement of Cariniocoris, Knight’s (1941:22-25) key to the genera of Phylinae, beginning with couplet 1 7, should be modified as follows [couplets 18-19 not repeated]: 17. Hind tibia with black spines lacking dark spots at bases 18 Hind tibia with pale yellowish-brown to almost colorless spines, sometimes with pale brown spots at bases, or dark spines with dark spots at bases 20 20. Mesopleuron with flattened scalelike pubescence Psallus Mesopleuron always without flattened, scalelike pubescence 21 21. Tibial spines dark with distinct dark spots at bases; ventral surface of male genital capsule smooth, without a median carina Plagiognathus - Tibial spines pale yellowish brown, sometimes with brownish spots at bases of meso- and metatibiae; ventral surface of male genital capsule with a distinct median carina Cariniocoris The following keys to males and females will allow recognition of the species of Cariniocoris', females are best identified by their association with males, especially when determining generic placement. KEY TO MALES OF CARINIOCORIS 1 . Dorsal coloration uniformly pale green to greenish yellow (Fig. 5); host Nyssa sylvatica nyssae - Dorsal coloration predominantly brown to fuscous; hosts Ilex spp 2 2. Hemelytron uniformly brown to fuscous (Fig. 1); length of second antennal segment nearly equal to basal width of pronotum; hosts Ilex spp geminatus - Hemelytron not uniformly brown, clavus and basal half of corium distinctly paler (Fig. 3); length of second antennal segment much shorter than basal width of pronotum; host Ilex verticillata ilicis KEY TO FEMALES OF CARINIOCORIS 1 . Inside of areoles or membranal cells fumate or black (Fig. 6), dark color sometimes fading to brown but always darker than remainder of membrane; body width 1.52- 1.60 mm nyssae 92 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) - Hemelytral membrane uniformly pallid or, at most, uniformly translucent or smoky brown; body width greater than 1.60 mm 2 2. Length of antennal segment II subequal to length of rostrum; hemelytra uniformly pale green (Fig. 2); body width 1.84-1.92 mm geminatus - Length of antennal segment II distinctly shorter than length of rostrum; apical half of corium infuscated (Fig. 4); body width 1.68-1.72 mm ilicis Cariniocoris geminatus (Knight), New Combination Figs. 1-2, 7, 10, 13-14, 17-20 Plagiognathus geminatus Knight, 1929:265; Carvalho, 1958:102; Henry and Wheel- er, 1988:486. Plagiognathus illicis [5zc]: Khalaf, 1971:340. Diagnosis. Color dimorphic; males with hemelytra uniformly dark brown, base of corium sometimes paler, females uniformly pale greenish; primary spiculum with spicules along ventral surface of basal Vs and middorsal flange elongate with anterior edge pointed. Males can be distinguished from those of C. ilicis and C. nyssae by the uniformly dark-brown hemelytra and the longer 2nd antennal segment. Females differ from those of C. ilicis by the uniformly pale-yellow to yellowish-green body and the proportionately longer 2nd antennal segment. From females of C. nyssae, this species is separated by the uniformly pale smoky-brown hemelytral membrane and larger body size; females of C. nyssae are more slender and have the insides of the areoles fuscous. Description. Male (N = 10): Length 3.20-3.52 mm, width 1.44-1.52 mm. Head: Width 0.74-0.80 mm, vertex 0.30-0.34 mm. Rostrum: Length 0.94-0.96 mm, ex- tending to mesocoxae. Antenna: Segment I, length 0.22-0.24 mm; II, 1.14-1.20 mm; III, 0.52-0.54 mm; IV, 0.28-0.34 mm. Pronotum: Length 0.62-0.64 mm, basal width 1.14-1.24 mm. Female (N = 10): Length 3.66-3.88 mm, width 1.84-1.92 mm. Head: Width 0.78- 0.80 mm, vertex 0.38-0.40 mm. Rostrum: Length 1.02-1.04 mm, extending to mesocoxae. Antenna: Segment I, length 0.22-0.24 mm; II, 1.02-1.12 mm; III, 0.54- 0.58 mm; IV, 0.30-0.32 mm. Pronotum: Length 0.70-0.72 mm, basal width 1.38- 1.42 mm. General coloration brown to dark brown, clothed with recumbent brown simple setae. Head brown. Antenna pale yellowish brown, segment III and IV becoming infuscated. Pronotum brown, areas anterior to calli sometimes paler brown; scutellum dark brown. Hemelytron uniformly brown to dark brown, sometimes having the base of corium paler; cuneus pale at base; membrane dark smoky brown. Under- surface yellowish brown, genital segment frequently darker. Legs yellowish brown; metafemur dark brown to fuscous, paler at base and apex, anterior surface with a few small fuscous spots; mesofemur yellowish brown but sometimes infuscated, anterior surfaces with a few small brown spots; tibial spines pale brown without basal spots or with only very indistinct pale-brown spots. Genital segment with narrow mesal keel ending before touching transverse swollen ridge around basal margin of genital opening (Figs. 1 3-14). Left paramere (Fig. 1 7); right paramere (Fig. 1 8); vesica with dorsal surface of apical V2 having a small, but distinct, backward-curving flange 1989 CARINIOCORIS, A NEW GENUS OF PHYLINI 93 Figs. 1 1-12. Claws of Cariniocoris spp.: 1 1) ilicis (694x). 12) nyssae (694x). and the primary spiculum with numerous spicules along bottom of basal V-i (Fig. 19); phallotheca (Fig. 20). Females differ from males in the uniformly pale-yellowish to yellowish-green body and legs, paler smoky-brown hemelytral membrane, and the distinctly more broadly oval body form. Remarks. Knight (1929) called this species the “southern twin” of C. ilicis and, indeed, it is quite similar in size and sexual dimorphism. However, he commented when describing C. geminatus that “It seems rather remarkable that not a single male can be found in the large series studied.” The pronounced sexual dimorphism found in this species apparently accounted for his not associating the dark males with the uniformly yellowish-green, paratype females taken on the same hosts at the type locality [he later associated these males with females in his collection]. Although I have not seen Khalafs (1971) specimens reported as Plagiognathus ilicis from Mississippi, distribution and extensive fieldwork indicate that he actually had C. geminatus. Specimens examined. DELAWARE: 1$, New Castle Co., Newark, University of Delaware, 28 May 1984, A. G. Wheeler, Jr. coll., taken on Ilex opaca (PDA). FLOR- IDA: 9(5(3, 999, Alachua Co., Austin Cary Mem. Forest, 10 mi NE Gainesville, 4 May 1982, T. J. Henry coll., taken on Ilex glabra (USNM); \6, Gulf Co., Rt. 30, 8 mi S Port St. Joe on St. Joe Peninsula, 1 May 1984, T. J. Henry and A. G. Wheeler, Jr., taken on Ilex glabra (USNM); 166(5, 1099, Walton Co., 3 mi W Freeport, Rt. 20, 9 May 198 1, T. J. Henry coll., taken on flowers of Ilex glabra (USNM). MARYLAND: 2566, 2299, Prince Georges Co., Beltsville, 17-22 May 1982, T. J. Henry coll., taken on Ilex opaca (USNM). MISSISSIPPI: 16, Wiggins, 5 May 1931, H. G. Johnston coll. (USNM); 266, 399, Washington Co., 3 mi SE Leland, 9 May 1983, T. J. Henry and G. L. Snodgrass colls., taken on Ilex decidua (USNM). PENNSYLVANIA: 1066, 599, Dauphin Co., Harrisburg, 25 May 1983, A. G. Wheeler, Jr. coll., taken on male inflorescences of Ilex opaca (PDA). SOUTH CAROLINA: 966, 499, Richland Co., Columbia, Univ. South Carolina campus, 9 Apr. 1988, A. G. Wheeler, Jr. coll., taken on Ilex vomitoria (PDA). TEXAS: Holotype 9 and 31 paratype 99 (and 1766 not mentioned in original description). College Station, 7-12 Apr. 1928, H. G. Johnston 94 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) colL, taken on Ilex decidua and Ilex vomitoria. VIRGINIA: 26<$, 299, Lancaster Co., White Stone, 26 May 1984, A. G. Wheeler, Jr. colL, taken on Ilex opaca (PDA); 7(36, 399, Westmoreland Co., Montross, 25 May 1984, A. G. Wheeler, Jr. coll., taken on Ilex opaca (PDA). Distribution. Known in the literature only from Mississippi (as ilicis by Khalaf, 1971) and Texas (Knight, 1929). New state records are Delaware, Florida, Maryland, South Carolina, and Virginia. Hosts. Recorded from Ilex decidua Walt, and I. vomitoria Ait. (Knight, 1929). New host records are Ilex glabra (L.) A. Gray and /. opaca Ait. This species prefers the male flowers of its host. At Beltsville, Maryland, many thousands of adults and nymphs could be collected on male flowers of a large American holly, I. opaca, but only a few adults could be found on a profusely flowering female tree no more than 25 feet away. Cariniocoris ilicis (Knight), New Combination Figs. 3-4, 8, 11, 15, 21-24 Plagiognathus ilicis Knight, 1925:305; Blatchley, 1926:928; Henry and Wheeler, 1988:486. Plagiognathus illicis [sic]: Carvalho, 1958:103. Diagnosis. Recognized by the brown coloration becoming dark brown on the apical half of the hemelytron in both sexes, but smaller and paler in females, and the structure of the vesica with numerous spicules around the basal V2 of the primary spiculum and a large, triangular, middorsal flange. Cariniocoris ilicis is similar to C. geminatus in general form and coloration. Males are separated from those of C. geminatus by the paler brown clavus, basal Vi of the corium, and cuneus, and by the much shorter second antennal segment; from C. nyssae they differ by the extensive brown coloration of the dorsum. Females can be separated from C. geminatus females by the paler dorsum, having the apical Vi of the corium infuscated, and the shorter second antennal segment; from females of C. nyssae they differ by the apically infuscated corium and uniformly pale smoky-brown membrane. Description. Male (N = 10): Length 3.32-3.60 mm, width 1.48-1.52 mm. Head: Width 0.74-0.76 mm, vertex 0.34-0.36 mm. Rostrum: 1.06-1.10 mm, extending to apices of mesocoxae or bases of metacoxae. Antenna: Segment I, length 0.22-0.24 mm; II, 0.96-0.98 mm; III, 0.50-0.54 mm; IV, 0.28-0.34 mm. Pronotum: Length 0.60-0.64 mm, basal width 1.22-1.24 mm. Female (N = 10): Length 3.24-3.76 mm, width 1.68-1.72 mm. Head: Width 0.74- 0.76 mm, vertex 0.38-0.40 mm. Rostrum: 1.12-1.18 mm, extending to bases of metacoxae. Antenna: Segment I, length 0.22-0.24 mm; II, 0.88-0.92 mm; III, 0.50- 0.52 mm; IV, 0.30-0.32 mm. Pronotum: Length 0.66-0.68 mm, basal width 1.36- 1.40 mm. General coloration brown to dark brown, clothed with recumbent pale-brown setae. Head brown. Antenna pale yellowish brown, segment III and IV becoming infuscated. Pronotum brown, paler brown on posterior V3; scutellum brown. Hemelytron brown to dark brown with the clavus, basal V3 of corium, and inner angle of cuneus pale yellowish brown; membrane pale smoky brown, veins yellowish. Ventral surface 1989 CARINIOCORIS, A NEW GENUS OF PHYLINI 95 Figs. 13-16. Male genital capsules of Cariniocoris spp.: 1 3) geminatus, ventral aspect ( 1 05 x ). 1 4) geminatus, caudal aspect ( 1 1 0 x ). 15) ilicis, ventral aspect ( 1 1 5 x ). 16) nyssae, ventral aspect (120x). yellowish brown. Legs yellowish brown; meso- and metafemora sometimes with a few tiny brown spots on anterior surfaces; tibial spines pale brown without basal spots or, at most, with some basal spines on the pro- and mesotibiae having small indistinct pale brown spots; claws pale brown. Genital capsule with a relatively thickened mesal keel extending to a wide transverse ridge around basal Vi of genital opening (Fig. 15). Left paramere (Fig. 21); right paramere (Fig. 22); vesica (Fig. 23) with dorsal surface of apical V3 having a large triangular flange, and the primary spiculum with numerous spicules around swollen basal V?, to V2; phallotheca (Fig. 24). Females differ from males by the overall paler coloration and the smaller, less distinct dark apex of the hemelytra, the often contrastly dark scutellum, and the much more broadly oval form. Coloration of the membrane, undersurface, and legs similar to that of males. Remarks. Based on distribution, as noted under C. geminatus, I am referring Khalaf ’s (1971) records of Plagiognathus ilicis to geminatus. Type designation. Although the pin and labels for the holotype of this species are present in the USNM type collection, a note placed by R. C. Froeschner states: “Holotype missing from point— 1976.” For nomenclatural stability I designate one 96 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) Figs. 17-28. Male genitalia of Cariniocoris spp.: C. geminatus: 17) left paramere. 18) right paramere. 19) vesica. 20) phallotheca. C. ilicis: 21) left paramere. 22) right paramere. 23) vesica. 24) phallotheca. C. nyssae: 25) left paramere. 26) right paramere. 27) vesica. 28) phallotheca. 1989 CARINIOCORIS, A NEW GENUS OF PHYLINI 97 of Knight’s male paratypes as a neotype to be placed in the USNM type collection. Label data as follows: label 1, “Ringwood, Ithaca, N. Y., 13-VII 1920”; 2, “H. H. Knight Collector”; 3 (red label), PARATYPE Plagiognathus ilicis by H. H. Knight”; 4, “H. H. Knight Collection 1976”; 5 (here added; white label with red border), “Neotype: Plagiognathus ilicis Knight by T. J. Henry.” Other specimens examined. NEW YORK: 27 paratype 66, 21 paratype 99, same locality data as for neotype, 26 June-13 Jul. 1920, H. H. Knight coll., taken on Ilex verticillata (USNM); 2>66, 1899, Tompkins Co., Ithaca [Cornell Univ. Campus], 26 June 1983 & 7 Jul. 1984, E. R. Hoebeke coll., taken on Ilex verticillata (CU); 1355, 899, Tompkins Co., Ithaca, Cornell Univ. Campus, 26 June 1987, A. G. Wheeler, Jr., taken on male flowers of Ilex verticillata (PDA, USNM). Distribution. This species is known only from New York and Wisconsin (Henry and Wheeler, 1988). Hosts. Recorded only from Ilex verticillata A. Gray. Cariniocoris nyssae, new species Figs. 5-6, 9, 12, 16, 25-28 Diagnosis. Recognized by the uniformly green coloration in both sexes, and fuscous membrane in males and fuscous areoles in females, small size, and the vesica having the spicules of the primary spiculum limited to the ventral surface of the base and the shallow, middorsal flange having a V-shape notch anteriorly. Cariniocoris nyssae is noticeably the smallest and most slender species of the genus; size differences are most readily apparent when series of each are compared side by side. Males can be separated from males of C. geminatus and C. ilicis by the uniformly yellow to greenish-yellow body having only the membrane smoky black or fumate. Females of all three species have the same general pale coloration, but those of C. nyssae have only the insides of the areoles or membranal cells smoky black or fumate, rather than the entire membrane uniformly pale or smoky brown. Additionally, females of C. ilicis have the apical Vi of each corium infuscated. Description. Male (N = 10): Length 3.04-3.52 mm, width 1.36-1.48 mm. Head: Width 0.74-0.76 mm, vertex 0.28-0.30 mm. Rostrum: Length 0.86-0.90 mm, ex- tending to mesocoxae. Antenna: Segment I, length 0.22-0.24 mm; II, 0.96-1.02 mm; III, 0.44-0.46 mm; IV, 0.30-0.32 mm. Pronotum: Length 0.54-0.58 mm, basal width 1.10-1.14 mm. ■ Female (N = 10): Length 3.40-3.64 mm, width 1.52-1.60 mm. Head: Width 0.72- 0.76 mm, vertex 0.32-0.34 mm. Rostrum: Length 0.98-1.00 mm, extending to mesocoxae. Antenna: Segment I, length 0.20-0.22 mm; II, 0.96-1.02 mm; III, 0.42- 0.50 mm; IV, 0.28-0.30 mm. Pronotum: Length 0.62-0.66 mm, basal width 1.28- 1.34. Males uniformly pale yellow to pale greenish yellow, clothed with recumbent yellowish or pale-brown setae. Head yellowish green. Antenna pale yellow, segment III and IV infuscated; segment II gradually thickened toward apex, diameter of apical half equal to diameter of segment I. Pronotum and scutellum yellowish, tinged with green laterally and on and posterior to calli. Hemelytron pale, somewhat translucent, yellow; membrane smoky or fumate, veins yellowish. Ventral surface pale yellow, tinged with green, especially on abdomen. Legs uniformly pale yellow; tibial spines 98 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) pale brown, without spots at bases or rarely with vague pale brown spots on basal spines; claws pale brown. Genital keel or carina extending to near rim of genital opening (Fig. 16). Left paramere (Fig. 25); right paramere (Fig. 26); vesica (Fig. 27) with a shallow, subdorsal flange, which is notched anteriorly, and the primary spi- culum with a few, distinct spicules restricted to ventral edge of swollen base; phal- lotheca (Fig. 28). Female coloration much as in males, but with hemelytral membrane pale trans- lucent, having only the inside of the areoles smoky black or fumate (dark area appearing to the naked eye as a black bar at base of the membrane). Also the general body form is more broadly oval and the apical half of antennal segment II is distinctly more slender than segment I. Type specimens. Holotype USA, Maryland, Prince Georges Co., College Park, University of Maryland Campus, 23 May 1987, T. J. Henry and A. G. Wheeler, Jr. colls., taken on blackgum, Nyssa sylvatica (USNM). Paratypes: 23$$, 3329, same data as for holotype (AMNH, PDA, and USNM); 2$$, 1599, Florida, Orange Co., N of Sand Lake Rd., Orlando, 30 Apr. 1984, J. T. Polhemus coll. (JTP, USNM). Etymology. The specific epithet of this species is taken from the generic name of its host, blackgum or tupelo, Nyssa sylvatica Marsh. Distribution. Florida and Maryland. Hosts. This species was collected in Maryland on the male flowers of a large, open- growing, ornamental blackgum, along with many adults of Lepidopsallus nyssae Johnston, previously known only from Illinois and Texas. ACKNOWLEDGMENTS I thank R. C. Froeschner (USNM), R. W. Poole (Systematic Entomology Lab., PSI, ARS, USDA, % USNM), M. E. Schaulf (SEL, PSI, ARS, % USNM), M. D. Schwartz (Amer. Mus. Nat. Hist., New York), A. G. Wheeler, Jr. (PDA), and Brian Wiegmann (Dept. Ent., Univ. Maryland, College Park) for reviewing the manuscript and offering useful suggestions; Wheeler, E. R. Hoebeke (CU), J. T. Polhemus (JTP), R. T. Schuh (AMNH), and G. M. Stonedahl (AMNH) for lending specimens cited in this study; and Rebecca Stanger (SEL, PSI, ARS, USDA, % USNM) for help with the electron micrographs. LITERATURE CITED Blatchley, W. S. 1926. Heteroptera or True Bugs of Eastern North America. Nature Publ. Co., Indianapolis, Indiana. 1,116 pp. Carvalho, J. C. M. 1958. Catalogue of the Miridae of the World. Part II. Subfamily Phylinae. Arq. Mus. Nac., Rio de Janeiro 45:1-216. Henry, T. J. and A. G. Wheeler, Jr. 1988. Family Miridae. Pages 251-507 in: T. J. Henry and R. C. Froeschner (eds.). Catalog of the Heteroptera, or True Bugs, of Canada and the Continental United States. E. J. Brill, Leiden and New York. 958 pp. Khalaf, K. T. 1971. Miridae from Louisiana and Mississippi (Hemiptera). Fla. Entomol. 54: 339-342. Knight, H. H. 1925. Description of a new species of Plagiognathus from the Eastern United States (Hem., Miridae). Entomol. News 36:305-306. Knight, H. H. 1929. The fourth paper on new species of Plagiognathus (Hemiptera: Miridae). Entomol. News 40:263-268. Knight, H. H. 1941. The plant bugs, or Miridae, of Illinois. 111. Nat. Hist. Surv. Bull. 22:1- 234. 1989 CARINIOCORIS, A NEW GENUS OF PHYLINI 99 Slater, J. A. and R. M. Baranowski. 1978. How to Know the True Bugs (Hemiptera-Heter- optera). Wm. C. Brown, Publ., Dubuque, Iowa, 256 pp. Wagner, E. 1975. Die Miridae Hahn, 1831, des Mittelmeerraumes und der Makaronesischen Inseln (Hemiptera, Heteroptera). Entomol. Abh. 40(Suppl.): 1-483. Received June 3, 1988; accepted August 8, 1988. J. New YorkEntomol. Soc. 97(1): 100-104, 1989 FROESCHNERANA MEXICANA, A NEW GENUS AND SPECIES OF DERAEOCORINAE FROM MEXICO (HETEROPTERA: MIRIDAE) J. C. SCHAFFNER* AND PaULO SeRGIO FiUZA FeRREIRA^ ‘Department of Entomology, Texas A&M University, College Station, Texas 77843, and ^Departamento de Biologia Animal, Universidade Federal Vicosa, Minas Gerais, Brazil Abstract.— Fweschnerana mexicana, a new genus and new species, is described from the Mexican states of Aguascalientes, Guerrero, Jalisco, Oaxaca and Puebla. Its relationship to the genus Bothynotus (Clivinemini) is discussed. The review of the genus Bothynotus by Henry (1979) prompted our attention to this previously undescribed genus found in Mexico. Although distinctive, the genus shares with Bothynotus the character of having scattered setae on the hemelytral membrane. Froeschnerana, new genus Description. Deraeocorinae, Clivinemini. Characterized by its relatively large size (8. 0-9.0 mm), lack of clearly defined collar, narrow embolium, and setose hemelytral membrane. Head smooth, shining, declivous, vestiture consisting of scattered short setae above and more dense longer setae ventrally; vertex slightly rounded not margined poste- riorly; frons vertical; clypeus prominent and projecting more anteriorly than frons; gena prominent; buccula clearly delimited; eyes located anteriorly on head, distance between posterior margin of eye and anterior margin of prothorax slightly less than length of eye in dorsal view, height of eye in lateral view about equal to gena. Margin of antennal socket not touching eye; antennal segment I shorter than vertex width; segments linear; relative lengths of segments from shortest to longest 4- 1-3-2; ves- titure of segment I somewhat sparse, that of II more dense with length of longer setae not longer than diameter of segment, those of III and IV with occasional setae longer than diameter of segment. Rostrum reaching or almost reaching mid coxae. Pronotum appearing punctate due to color pigmentation around setal bases, however only very indistinctly punctate under reflected light; disc with shallow middorsal sulcus; lateral margins rounded, posterior margin slightly concave; humeral comers rounded; calli flattened, smooth and shining, impressed line from anterolateral comer of pronotum to posterior margin of calli weakly delimited; collar not delimited dorsally; anterior margin of pronotum not cystiform; vestiture consisting of short erect or semi-erect setae. Scutellum very lightly transversely rugulose, parallel-sided, slightly convex, raised above hemelytra. Hemelytra rugose, parallel-sided; sloping downward later- ally; vestiture consisting of fairly dense semi-erect setae; embolium narrow; cuneus longer than wide; membrane with two cells, minutely and densely pilose with scattered 1989 A NEW DERAEOCORINE 101 longer setae. Femora linear; tibiae without spines, setae about as long as diameter of tibia. Setae on underside of abdomen fairly dense. Type species. Froeschnerana mexicana, n. sp. Discussion. The presence of elongate setae on the hemelytral membrane suggests that Froeschnerana is closely related to Bothy notus since no other genera of the tribe exhibit this specific character. The shape of the head is the same for members of both genera. Among the several differences is the large size (8. 0-9.0 mm) and elongate 102 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) Figs. 2-4. Male genitalia of Froeschnerana mexicanus (nr. Autlan, Jalisco). 2. Vesica. 3. Left paramere. 4. Right paramere. body form of Froeschnerana which contrasts with the somewhat smaller size (2.4- 4.7 mm) and usually more robust body form of Bothynotus. Bothynotus is also characterized by having a pronotum which is strongly punctate whereas the pronotum of Froeschnerana is very weakly and sparsely punctate. Although variable, members of the genus Bothynotus tend to have many long erect or semierect setae on the dorsum of the head whereas Froeschnerana specimens have scattered, short semi- decumbent setae. We take pleasure in naming this genus in honor of Richard C. Froeschner, a personal friend and former fellow graduate student of the senior author while at Iowa State University. Dr. Froeschner is the Curator of Hemiptera at the Smithsonian Institution and was honored in the April, 1986 issue of the Journal. Froeschnerana mexicana, new species Figs. 1-6 Description. MALE (Measurements (in millimeters) taken from 7 specimens; ho- lotype first followed by average and range in parentheses): Length, 8.60 (8.43, 8.00- 8.60); width, 2.80 (2.85, 2.68-3.00). Head length, 0.56 (0.54, 0.52-0.60); width through eyes, 1.40 (1.40, 1.36-1.48); vertex width, 0.76 (0.77, 0.72-0.80). Length of antennal segment I, 0.72 (0.71, 0.64-0.76); II, 2.28 (2.30, 2.20-2.44); III, 1.28 (1.28, 1.20-1.36); IV, 0.64 (0.57, 0.48-0.64). Pronotal length, 1.52 (1.52, 1.44-1.68); width across base, 2.48 (2.50, 2.28-2.68). Cuneal length, 1 .44 ( 1 .43, 1 .32-1 .48); width across base, 0.80 (0.76, 0.68-0.84). General coloration dark fuscous to black with red or orange areas; shining. Head yellow to red, usually orange; clypeus at least basally and apically, underside of head, area behind eyes and band running posteriorly from each antennal socket over frons and coalescing posteriorly on vertex, fuscous to black; rostrum and antennae black. Collar dark fuscous to orange or reddish fuscous, area of calli fuscous to black with 1989 A NEW DERAEOCORINE 103 Figs. 5, 6. Female genitalia of Froeschnerana mexicanus (nr. Acatepec, Puebla). 5. Sclero- tized rings, dorsal view. 6. Posterior wall, dorsal view. remainder of pronotum orange or red, mesoscutum and scutellum dark fuscous to orange or reddish fuscous; hemelytra dark fuscous to black. Ventral aspect ranging from almost entirely dark fuscous or reddish fuscous to orange. Coxae testaceous to orange at least apically, underside of femora usually testaceous ranging to testaceous or orange on entire basal half, tibiae and tarsi dark fuscous. Morphological characters as given for genus. Genitalia as in Figures 2-4. FEMALE (Measurements taken from 8 specimens; average hrst followed by range in parentheses): Length, 8.57 (8.16-8.96); width, 2.85 (2.68-3.08). Head length, 0.60 (0.52-0.68); width through eyes, 1.44 (1.40-1.48); vertex width, 0.85 (0.80-0.92). Length of antennal segment I, 0.73 (0.68-0.80); II, 2.39 (2.32-2.60); III, 1.34 (1.28- 1.40); IV, 0.58 (0.56-0.64). Pronotal length, 1.52 (1.40-1.64); width across base, 2.57 (2.44-2.72). Cuneal length, 1.38 (1.28-1.48); width across base, 0.77 (0.68-0.88). Similar to male in color, lacking pale areas on underside of femora; two females with dark red pronotal discs; membrane of hemelytra with fewer long setae. Genitalia as in Figures 5-6. Holotype. Male, Mexico: Puebla, 4 mi W. Acatepec, 26 July 1973, Mastro & Schaffner. Deposited in the National Museum of Natural History, Washington, D.C. Paratypes. Mexico. Aguascalientes: 16, 8 mi E Calvillo, 11 July 1983, Kovarik, Harrison, Schaffner; 1$, 6 mi E Calvillo, 1 1 July 1983, Kovarik, Harrison, Schaffner. 104 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) Guerrero: 1$, 4 mi W of Chilpancingo, 15 July 1984, Carroll, Schaffner, Friedlander. Jalisco: 36S, 19, 16 km N Autlan, 12-14 July 1983, Kovarik, Harrison, Schaffner. Oaxaca: 19, 4.2 mi N Tonaltepec, 21 July 1987, Kovarik, Schaffner. Puebla: \6, 19, 4.4 mi SW Acatepec, 9 July 1977, J. C. Schaffner; 299, same locality, 26 July 1974, Clark, Murray, Ashe, Schaffner; 16, same locality, 9 July 1981, Bogar, Schaffner, Friedlander; 19, same locality, 21 July 1984, Carroll, Schaffner, Friedlander; 16, 19, 6.3 mi N Tehuacan, elev. 5,900 ft, 22 July 1987, Kovarik, Schaffner. Deposited in the collections of the Institute de Biologia, Universidad Nacional Autonoma de Mexico, Mexico City, D.F. and Department of Entomology, Texas A&M University, College Station, Texas. Discussion. The non-fuscous areas of the individual specimens are usually either orange or red. Both color variations were collected together near Acatepec, Puebla. All specimens taken from near Autlan, Jalisco were primarily orange whereas the individuals from Cavillo, Aguascalientes, were red. The distribution pattern of this species is unusual. We know of no other mirids (with the possible exception of a species of Proba Distant which occurs more or less throughout Mexico) that exist in this fairly well collected area of Puebla (Tehuacan- Acatepec) that also occur in western Mexico (Jalisco, Aguascalientes). All material was collected from sites located on the edges of the plateaus in the region of central Mexico. ACKNOWLEDGMENTS Appreciation is extended to H.R. Burke for critically reading this manuscript. The illustrations were prepared by Mrs. Nancy A. Browning and Mrs. Nannette Goyer Davis. LITERATURE CITED Henry, T. J. 1 979. Review of the new world species ofBothynotus Fieber (Hemiptera: Miridae). Florida Entomol. 62:232-244. Received June 3, 1988; accepted November 2, 1988. J. New YorkEntomol. Soc. 97(1):105-1 10, 1989 TWO NEW SPECIES OF MORMIDEA FROM MEXICO AND GUATEMALA (HETEROPTERA: PENTATOMIDAE)i D. A. Rider and L. H. Rolston Department of Entomology, Louisiana Agricultural Experiment Station, Louisiana State University Agricultural Center, Baton Rouge, Louisiana 70803 Abstract. —Mormidea polita n. sp. and M. guatemalensis n. sp. are described from Mexico and Guatemala, respectively. A key is provided for the identification of the 14 species of Mormidea Amyot and Serville known to occur in Mexico, Belize and Guatemala. The genus Mormidea Amyot and Serville, which is restricted to the western hemi- sphere, belongs in the Pentatomini among those genera lacking a medial spine or tubercle at the base of the abdominal venter. The genera in this group that occur in the western hemisphere north of South America were keyed by Rolston and Mc- Donald (1984), and the genus Mormidea was revised, with a subsequent emendation, by Rolston (1978, 1984). Here, a new species from Mexico and a new species from Guatemala are added to the genus. A key is provided to facilitate identification of the species of Mormidea that are known from Mexico, Belize and Guatemala. KEY TO MEXICAN, BELIZIAN AND GUATEMALAN SPECIES OF MORMIDEA 1 . Middle half of pronotum at posterior margins of cicatrices traversed by narrow, ivory callus (subgenus Melanochila St&l) lugens (F.) Pronotum with calloused spot (usually ivory) at posterior margin of each cicatrice; or with large, irregularly shaped, calloused, ivory macule covering much of anterior pronotal disk; or without callus (subgenus Mormidea) 2 2( 1 ). Dark spot on each mesopleuron at distal end of supracoxal cleft larger than least diameter of tibiae; endocoria nearly transparent excepting punctation; last antennal segment without pale, basal band angustata St^l - Dark spot smaller than least diameter of tibiae, or absent, or obscured by dark punctation and infuscation of mesopleuron; endocoria translucent or obscure; at least last antennal segment bicolored with pale, basal band (except laevigata) ... 3 3(2). Connexiva entirely pale, or with broad, entirely pale lateral borders on at least last four connexival segments (Fig. 2) 4 Connexiva dark with pale areas confined to lateral edges or to marginal scallops, each between transverse sutures (Fig. 1) 11 4(3). Abdominal venter pale with dark medial vitta (sometimes in addition a vague lateral vitta on each side) or a medial row of spots 5 - Abdominal venter pale and without vitta (occasionally a thin, dark medial line on last stemite and sometimes at base of one or more of preceding stemites) ... 10 5(4). Submarginal, ivory callus along frena continuing to scutellar apex as marginal ivory band with very few or no punctures laevigata Distant ‘ Approved for publication by the Director of the Louisiana Agricultural Experiment Station as manuscript number 88-17-2387. 106 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) - Sides of scutellum not or incompletely ivory bordered, or if completely bordered, numerous dark punctures present at distal end of each frenum 6 6(5). Ground color of dorsum light to dark brown; body length without hemelytral membranes usually less than 7 mm; spiracles concolorous with surrounding area 7 Ground color of dorsum black or fuscous; body length without hemelytral mem- branes usually more than 7 mm; spiracles usually black 8 7(6). Small, sharp tooth on posterior margin of pygophore at beginning of each arm of chevron-shaped carina on posterior surface of pygophore (Fig. 3); basal plates mostly fuscous pictiventris St^l Posterior margin of pygophore unarmed (Fig. 4); basal plates entirely or almost entirely light brown faisana Rolston 8(6). Pale markings on pronotum, scutellum and hemelytra limited to spot on scutellar apex and small medial spot or dot at base of scutellum 9 - A pale spot present on apex of scutellum and medial spot or dot at base (latter rarely obscure) plus some or all of following: spot at posterior margin of each cicatrice, in each basal angle of scutellum (sometimes extending along frenum), and on disk of each hemelytron notulata (Herrich-Schaffer) 9(8). Middle half of posterior pygophoral margin evenly concave from ventral view (Fig. 15) polita, n. sp. - Concavity in middle half of posterior pygophoral margin truncate ventrally (Fig. 22) guatemalensis, n. sp. 10(4). Dark spot present at distal end of each supracoxal cleft; from caudal view, dorsal border of pygophore conspicuously impressed medially (Fig. 5); basal plates finely and densely punctate collaris Dallas Distal ends of supracoxal clefts immaculate, or each with minute dark dot; from caudal view, dorsal border of pygophore smoothly contoured or slightly flattened medially (Fig. 6); basal plate finely striate, a few punctures at apical angles .... ypsilon (L.) 1 1(3). Abdominal venter dark, or pale with mostly rufous to black punctation; a broad, irregularly shaped, dark medial vitta usually accompanied by lateral vitta on each side at base of abdominal venter 12 Venter pale with concolorous punctation; abdominal vitta absent or at most a dark, medial discontinuous line on last three stemites (Fig. 7) lunara Rolston 12(11). Anterolateral margins of pronotum strongly concave (Fig. 8); spiracles black; basal angles of scutellum usually immaculate, occasionally each with pale dot discoidea (Dallas) - Anterolateral margins of pronotum weakly concave (Fig. 9); spiracles and sur- rounding areas of stemites concolorous; basal angles of scutellum each with pale spot which often continues as callus along frenum 13 1 3( 1 2). Posterior margin of pygophore from ventral view conspicuously notched medially (Fig. 10); basal plates deeply impressed at posterolateral angles, their mesial mar- gins straight, contiguous pama Rolston Posterior margin of pygophore from ventral view shallowly notched medially (Fig. 1 1); basal plates not impressed at posterolateral angles, their margins sinuous and contiguous only at base (Fig. 12) cubrosa (Dallas) Mormidea polita, new species Figs. 14-20 Description. Dorsum fuscous to black, rather polished, occasionally with interstitial areas of pronotum and hemelytra pale brown; cordiform ivory spot covering scutellar 1989 NEW SPECIES OF MORMIDEA 107 Figs. 1-13. 1. Connexivum with pale scallops. 2. Connexivum with entirely pale border. 3. M. pictiventris. Pygophore, caudal view. 4. M. faisana. Pygophore, caudal view. 5. M. collaris. Pygophore, caudal view. 6. M. ypsilon. Pygophore, caudal view. 7. M. lunara. Abdominal venter. 8. M. discoidea. Anterolateral margin of pronotum. 9. M. pama. Anterolateral margin of pronotum. 10. M. pama. Pygophore, ventral view. 1 1. M. cubrosa. Pygophore, ventral view. 1 2. M. cubrosa. Basal plates. 1 3. M integella. Inferior ridge (ir) and posterior pygophoral margin, dorsal view. apex and usually a medial ivory dot present at scutellar base; connexiva ivory; hemelytral membranes fumose. Venter stramineous to pale brown, sometimes with reddish suffusion, and with broad, fuscous to black medial vitta on abdomen; punc- tures fuscous, usually becoming crowded on abdomen into vague, lateral vitta on each side between medial vitta and spiracles; spiracles dark, usually black; thoracic sterna black. Basal segment of each antenna pale brown, usually with dorsolateral dark streak; segments 2-3 varying from entirely light brown to fuscous with basal ivory band on each segment; segments 4-5 fuscous except ivory band on basal one- 108 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) fifth of segment 4 and basal one-third of segment 5. Rostrum pale brown with segment 4 largely fuscous. Femora and tibiae pale brown with numerous fuscous spots. Each humeral angle narrowly rounded to angulate, produced beyond base of corium (Fig. 1 8); anterolateral margins of pronotum concave. Scutellum slightly longer than wide at base; apex narrowly rounded. Middle one-half of posterior pygophoral margin concave in ventral and caudo- ventral views (Figs. 14, 1 5); inferior ridge produced caudad, sides arcuate with small, troughed, apical projection (Fig. 19). Paramere stout, posterior edge with convex prominence (Fig. 20). Basal plates impunctate except in apical angles; mesial margins straight, slightly separated anteriorly and posteriorly; posterior margins convex, apical angles rounded, fuscous (Fig. 1 7). Measurements (mm) (Holotype in parentheses). Total length excluding hemelytral membranes 6. 9-8. 3 (8.2); width across humeri 4. 6-5. 2 (5.1). Mesial length of prono- tum 1.5-1. 7 (1.7). Length of scutellum 2. 8-3. 3 (3.2); basal width 2. 7-3. 2 (3.0). Length of head from apex to posterior margin of ocelli 1.5-1. 7 (1.6); width across eyes 2.0- 2.2 (2.1). Length of antennal segments 1-5 about 0.4 (0.4), 0.6-0. 7 (0.7), 0.7-0. 8 (0.7), 1.0-1. 1 (1.1), and 1.1-1. 3 (1.2). Length of labial segments 1-4 about 0.8-0. 9 (0.85), 1.1-1.25 (1.25), 0.5-0.6 (0.55), 0.45-0.6 (0.55). Distribution. Mexican states of Hidalgo and Oaxaca. Holotype. Male, labeled “MEX: Hgo., 2.7 mi N. Tlanchinol, 14-V-83, C.W. & L.B. O’Brien & Marshall.” Deposited in the U.S. National Museum of Natural History, Washington, D.C. (USNM). Paratypes. 1 1 specimens. 2 males, labeled as holotype. Female, 3 males, labeled “MEX: Hgo., 2.4 mi N Tlanchinol, 5000' Hwy 105. Aug. 1, 82 O’Briens & G. Wibmer.” 2 males, labeled “MEXICO, Hgo. Hwy 105 2.7 mi. N Tlanchinol 5000' 2 Aug. 1982 C.W. & L. O’Brien & G. Wibmer.” Female, male, labeled “MEXICO, Oax., Hwy 175 18 mi. S. Valle Nacional, 4600' 25 Aug. 1982 C. & L. O’Brien & G. Wibmer.” Male labeled “MEXICO: Oaxaca Vista Hermosa 3 July 1982 William Abies.” Paratypes deposited in the American Museum of Natural History, USNM, and the author’s collections. Comments. Mormidea polita will key to M. notulata in the revision by Rolston ( 1 978). It can be separated from that species by the reduced amount of ivory markings on the dorsum and by differences in the male and female genitalia. In the structure of the male genitalia it more closely resembles M. integella (Distant), particularly in the posteriorly projecting inferior ridge (Figs. 13, 19). Mormidea polita has a heart- shaped spot on the scutellar apex, and dark, usually black spiracles. Mormidea in- tegella, which is known only from Costa Rica and Panama, lacks ivory marks on the apex of the scutellum, and the spiracles and surrounding surface are usually concolorous. Mormidea guatemalensis, new species Figs. 21-25 Description. Dorsum fuscous to black with some interstitial areas on corium, scutellar tongue, and posterior half of pronotum dark brown; scutellar apex and small medial spot along base ivory; connexiva ivory; hemelytral membranes fumose. Venter stramineous with some reddish infusion and black medial vitta; punctures black. 1989 NEW SPECIES OF MORMIDEA 109 Figs. 14-25. 14-20. M. polita. 14. Pygophore, caudoventral view. 15. Pygophore, ventral view. 16. Pygophore, lateral view. 17. Genital plates. 18. Anterolateral pronotal margin. 19. Genital cup; inferior ridge (ir). 20. Right paramere, lateral view. 21-25. M. guatemalensis. 21. Pygophore, caudoventral view. 22. Pygophore, ventral view. 23. Pygophore, lateral view. 24. Anterolateral pronotal margin. 25. Right paramere, lateral view. slightly crowded laterally. Antennal segment 1 pale with small fuscous apical band and fuscous dorsolateral streak; segment 2 pale, becoming infuscated apically; seg- ments 4 and 5 fuscous except basal one-fourth of segment 4 and basal one-half of segment 5 pale. Rostrum pale brown, segment 4 fuscous. Femora and tibiae pale brown with numerous fuscous spots. Each humeral angle acute, nearly spinose, produced beyond base of adjacent cori- um; anterolateral margins of pronotum concave (Fig. 24). Scutellar length and basal width subequal. Posterior margin of pygophore truncately concave in ventral and caudoventral views (Figs. 2 1 , 22); inferior ridge as in polita. Paramere relatively slender, posterior edge undulating, without marked prominence (Fig. 25). Female unknown. 110 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) Measurements (mm). Total length excluding hemelytral membranes 6.9; width across humeri 4.8. Mesial length of pronotum 1.4. Length of scutellum 2.7; basal width 2.7. Length of head from apex to posterior margin of ocelli 1.6; width across eyes 2.0. Length of antennal segments 1-5 about 0.4, 0.6, 0.7, 1.0, and 1.3. Length of labial segments 1-4 about 0.85, 1.25, 0.55, 0.55. Distribution. Guatemala. Holotype. Male, labeled “GUATEMALA, 32mi. SE. Coban, 6000' VII26-1974 C.W. & L. O’Brien & Marshall.” Deposited in the U.S. National Museum of Natural History, Washington, D.C. No paratypes. Comments. This species will key to M. notulata in the revision by Rolston (1978). The absence of most of the dorsal pale markings and the appearance of the male genitalia will easily separate these two species. Mormidea guatemalensis is most closely related to M. integella and M. polita. It can be separated from both species by differences in the male genitalia. It also differs from M. integella by the presence of an ivory spot on the scutellar apex. LITERATURE CITED Rolston, L. H. 1978. A revision of the genus Mormidea (Hemiptera: Pentatomidae). J. New York Entomol. Soc. 86(3): 16 1-2 19. Rolston, L. H. 1984. New synonymy and a new species in the genus Mormidea (Hemiptera: Pentatomidae). J. New York Entomol. Soc. 92(4):342-343. Rolston, L. H. and F. J. D. McDonald. 1984. A conspectus of Pentatomini of the western hemisphere. Part 3 (Hemiptera: Pentatomidae). J. New York Entomol. Soc. 92(1): 69-86. Received July 1, 1988; accepted August 23, 1988. NOTES AND COMMENTS J. New YorkEntomol Soc. 97(1):1 1 1-1 14, 1989 NEW RECORDS OF PALEARCTIC HETEROPTERA IN NEW YORK STATE: MICROPHYSIDAE AND MIRIDAE In the course of studying Miridae in New York I have collected two Palearctic Heteroptera species previously unrecorded from this state, Myrmedobia coleoptrata (Fallen) and Stethoconus japonicus Schumacher, as well as Camptozygum aequale (Villers) not known from the lower Hudson Valley and Long Island. A diagnosis, description, and photographs of the adult male of M. coleoptrata are given to facilitate recognition of this species. Myrmedobia coleoptrata (Fallen) (Microphysidae) Figs. 1, 2 Loricula pselaphiformis Curtis (Kelton, 1980), Mallochiola gagates (McAtee and Malloch) (McAtee and Malloch, 1924; Slater and Baranowski, 1978), and Myrme- dobia exilis Fallen (Kelton, 1981) are the only Microphysidae known to occur north of Mexico. Chiniola quericola Blatchley (Blatchley, 1928), described from a single female collected in Dunedin, Florida and deposited in the collection of the Depart- ment of Entomology, Purdue University, is presumably a microphysid; however, it was destroyed in a flood in 1981 (A. Provonsha, pers. comm). Herein, I report M. coleoptrata (Fallen) new to our fauna. On 10 June 1986, I collected a single male of M. coleoptrata by sweeping grasses beneath low hanging branches of Scotch pine. Firms sylvestris L., at the Nassau County Museum of Fine Art (formerly the Bryce/Frick Estate) near Rt. 25 A in Roslyn on the north shore of Long Island, New York. No other specimens could be found by beating the same tree later that season or in 1987. However, subsequent sweeping under the tree on 20 June 1988 yielded two males. To verify my determination of the specimen, I examined material of M. coleoptrata, as well as several other related species of Myrmedobia identified by J. Pericart. In the British Isles M. coleoptrata has been found beneath the bark of various trees, especially spruce, and occasionally in tufts of moss and grass around the bases of the trunks (South wood and Leston, 1959). The distribution of this species extends from southern Scandinavia south through western Europe (including southern Great Brit- ain) and to the north coast of Africa at Morocco and Tunisia (Pericart, 1972). Diagnosis. M. coleoptrata is distinguished from the other microphysids known from North America by these features: More elongate habitus (Fig. 1); anterior angle and lateral margin of pronotum not strongly marginate, with collar visible in lateral 112 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) Figs. 1, 2. Myrmedobia coleoptrata (Fallen). 1. Dorsal habitus. 2. Lateral view. view (Fig. 2); labium not reaching beyond procoxae; and antennae with long, erect setae, and segment IV longer than segment II. Description of adult male. General aspect: Length 1.90 mm; length from apex of tylus to cuneal fracture 1.39 mm; elongate and slightly flattened; coloration black, with lateral portion of hemelytra translucently fuscous and membrane slightly dark- ened; vestiture with moderately distributed suberect to erect, golden-fuscous setae; surface texture with head shining and smooth, pronotum, mesoscutellum, and scu- tellum shagreened, and hemelytra slightly shining and slightly shagreened. Head: Width across eyes 0.38 mm; interocular width 0.23 mm; length from apex of tylus to between ocelli 0.23 mm; ovoid, short in dorsal view; eyes large and subcontiguous to pronotum; labium robust, just attaining procoxae; antennae with long, suberect, shining setae, length of segment I 0.13 mm; II 0.30 mm; III 0.25 mm; IV 0.33 mm. Pronotum: Posterior width 0.58 mm; trapeziform; collar slightly wider than width of antennal segment II, reaching lateral margin of pronotum, visible in lateral view; lateral margin not carinate, slightly arched dorsad of coxal cleft; ostiolar peritreme obsolete. Hemelytra: Length to apex of cuneus 1.06 mm; length of cuneus 0.25 mm; embolium thickened or beadlike, forming slightly arcuate lateral margin; membrane with one ovoid cell. Genitalia: Not examined. Stethoconus japonicus Schumacher (Miridae: Deraeocorinae: Hyaliodini) Henry et al. (1986) first reported the predatory Japanese plant bug, Stethoconus japonicus, in the Western Hemisphere, based on specimens collected at four localities 1989 NOTES AND COMMENTS 113 in Maryland on ornamental azaleas infested with the azalea lace bug, Stephanitis pyrioides (Scott). Their observations indicated that both adults and nymphs prey on the lace bugs, and suggested that S. japonicus could prove useful in the biological control of this pest. Herein, I report the first occurrence of S. japonicus outside of Maryland. On 3 1 July 1987, I noticed an unusual bug attracted by house lights to the kitchen screen of my residence in South Nyack (Rockland Co.), New York. I identified it as an adult male of S. japonicus. Later I surveyed the ornamental plantings of the immediate vicinity for additional specimens. This search (6 August 1987) turned up one female beaten from the base of an azalea in poor condition, apparently severely damaged by the feeding of Stephanitis pyrioides. This record indicates that S. japonicus is much more widespread than previously reported. Camptozygum aequale (Villers) (Miridae: Mirinae: Mirini) Wheeler and Henry (1973) first reported Camptozygum aequale in North America from central and western Pennsylvania. They collected this bug from several species of introduced pine in nurseries, ornamental plantings, and Christmas tree plantations. Subsequent to their note, C. aequale was collected in central New York, New Hamp- shire, and Nova Scotia and Ontario, Canada (Henry and Wheeler, 1979; Wheeler, 1979; Kelton, 1983). At the same time, locality, and on the same host that I collected the microphysid reported above, I also found 5 male and 5 female C. aequale (on 10 June 1987, I collected 1 1 teneral males and 6 teneral females on the same tree). A study of the mirine collection at the American Museum of Natural History (AMNH) also revealed a series of 44 male and 83 female C. aequale collected by R. T. Schuh at the Huyck Preserve in Rensselaerville (Albany Co.), New York, 29 June-2 July 1977, from Pinus banksiana. DISCUSSION Previous reports mentioned that the possible method of introduction of these European bugs was via the importation of horticultural materials (Wheeler and Henry, 1973; Kelton, 1980, 1981; Henry et al., 1986). The particular section of the Nassau County Museum of Fine Art from which I collected the specimens of C. aequale and M. coleoptrata may be a rich source of introduced conifer inhabiting Heteroptera. It is part of a “pinetum” initiated in 1919, by Childs and Frances Frick, and contains more than 450 varieties of conifers (Metcalf and Libby, 1986). Likewise, the Huyck Preserve, the former estate of the Huyck family, in addition to containing several hundred acres of mixed second growth forests and old fields, incorporates many introduced ornamental comi^r^.— Michael D. Schwartz, Department of Entomology, American Museum of Natural History, New York, New York 10024. ACKNOWLEDGMENTS I thank J. Pericart, Montereau, France, for not only providing Myrmedobia specimens for my comparison, but for donating that material to the AMNH; T. J. Henry, USDA, Washington, D.C., also loaned microphysids from the National Collection, and commented on the manu- 114 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) script; and A. Provonsha, Purdue University, West Lafayette, Indiana, for clarifying the status of C quericola. All specimens mentioned herein are retained as vouchers in the collections of the Department of Entomology, AMNH. LITERATURE CITED Blatchley, W. S. 1928. Two new anthocorids and a new microphysid from Florida (Heter- optera). Entomol. News 39:85-88. Henry, T. J., J. W. Neal, Jr. and K. M. Gott. 1986. Stethoconus japonicus (Heteroptera: Miridae): a predator of Stephanitis lace bugs newly discovered in the United States, promising in the biocontrol of azalea lace bug (Heteroptera: Tingidae). Proc. Entomol. Soc. Wash. 88:722-730. Henry, T. J. and A. G. Wheeler, Jr. 1979. Palearctic Miridae in North America: records of newly discovered amd little-known species (Hemiptera: Heteroptera). Proc. Entomol. Soc. Wash. 81:257-268. Kelton, L. A. 1980. First record of a European bug, Loricula pselaphiformis, in the Nearctic Region (Heteroptera: Microphysidae). Can. Entomol. 112:1085-1087. Kelton, L. A. 1981. First record of a European bug, Myrmedobia exilis (Heteroptera: Micro- physidae), in the Nearctic Region. Can. Entomol. 1 13:1 125-1 127. Kelton, L. A. 1 983. European Pseudoloxops coccineus found in Canada, and additional records of Camptozygum aequale in the Nearctic region (Heteroptera: Miridae). Can. Entomol. 115:107-108. McAtee, W. L. and J. R. Malloch. 1 924. Some annectant bugs of the superfamily Cimicoideae (Heteroptera). Bull. Brooklyn Entomol. Soc. 21:69-83. Metcalf, P. C. and V. Libby. 1986. The House and Garden. Nassau County Museum of Fine Art, Roslyn, New York, 40 pp. Pericart, M. J. 1972. Hemipteres Anthocoridae, Cimicidae et Microphysidae de I’Ouest- Palearctique. Fauna de I’Europe et du bassin mediterraneen, vol. 7. Massin et Cie, Paris, 402 pp. Slater, J. A. and R. M. Baranowski. 1978. How to Know the True Bugs (Hemiptera-Heter- optera). Wm. C. Brown Co. Publ., Dubuque, Iowa, 256 pp. Southwood, T. R. E. and D. Leston. 1959. Land and Water Bugs of the British Isles. F. Wame and Co. Ltd., London, 436 pp. Wheeler, A. G., Jr. 1979. A comparison of the plant-bug fauna of the Ithaca, New York area in 1910-1919 with that of 1978. Iowa St. J. Res. 54:29-35. Wheeler, A. G., Jr. and T. J. Henry. 1973. Camptozygum aequale (Villers), a pine-feeding mirid new to North America (Heteroptera: Miridae). Proc. Entomol. Soc. Wash. 75:240- 246. Received April 12, 1988; accepted May 6, 1988. 1989 NOTES AND COMMENTS 115 J. New YorkEntomol Soc. 97(1):115, 1989 NAMES AND AUTHORSHIP OF TWO FAMILY-GROUPS IN THE EPHEMEROPTERA The Ephemeroptera superfamily encompassing the genera Prosopistoma Latreille and Baetisca Walsh has traditionally been referred to as Prosopistomatoidea Lameere, 1917. Edmunds and Traver (1955) first applied the superfamily name Prosopisto- matoidea to this taxon. Article 23a of ICZN (International Code of Zoological Nomenclature) requires that the valid name for a family-group taxon be based on the oldest available family- group name. The family-group name Prosopistomatidae Lameere was established in 1917 (as Prosopistomidae). Baetiscidae was established as a family-group name by Banks in 1900 (as tribe Baetiscini). Therefore, based on the name Baetiscidae Banks, 1900, the valid name of this superfamily is Baetiscoidea Banks, 1900. Authorship of the family Ephemeridae and related family-group names (Ephem- eroidea, Ephemerinae) have often been attributed to Leach (1815) (cf. Hubbard, 1 982). However, Latreille (1810) established the family “Ephemerinae” in the section Subulicornes of Neuroptera. Based on the principle of priority required by the ICZN (Art. 36a and 50d), the author of the family Ephemeridae is Latreille (1810). The other coordinated family-group taxa also take the same date and authorship. — Wil- liam L. Peters and Michael D. Hubbard, Department of Entomology, Florida A&M University, Tallahassee, Florida 32307. LITERATURE CITED Banks, N. 1910. New genera and species of Nearctic neuropteroid insects. Trans. Am. Entomol. Soc. 26:239-259. Edmunds, G. F., Jr. and J. R. Traver. 1954. An outline of a reclassification of the Ephem- eroptera. Proc. Entomol. Soc. Wash. 56:236-240. Hubbard, M. D. 1982. Catalog of the Ephemeroptera: family-group taxa. Aquat. Insects 4: 49-53. International Trust for Zoological Nomenclature. 1985. International Code of Zoological Nomenclature, 3rd Edition. London. Lameere, A. 1917. Etude sur 1’evolution des Ephemeres. Bull. Soc. Zool. Fr. 42:4 1-59, 61-81. Latreille, P. A. 1810. Considerations generates sur I’Ordre naturel des Animaux composant les Classes des Crustaces, des Arachnides, et des Insects; etc. Chez F. Schoell, Paris. Leach, W. E. 1815. Entomology. Brewster’s Edinburgh Encyclopaedia 9:57-172. 116 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) J. New YorkEntomol. Soc. 97(1):116, 1989 A NECESSARY NEW NAME IN THE FAMILY HEBRIDAE (HETEROPTERA: GERROMORPHA) Hebrus obscurus Cobben and Linnavuori [Cobben, R. H. and R. Linnavuori. 1983. Some new Hebridae from the Ivory Coast (Heteroptera, Gerromorpha). Ann. Ento- mol. Fenn. 49:83-86; obscurus on p. 85] is preoccupied by Hebrus obscurus Polhemus and Chapman [Polhemus, J. T. and H. C. Chapman. 1966. Notes on some Hebridae from the United States with the description of a new species (Hemiptera). Proc. Entomol. Soc. Wash. 68:209-212; obscurus (as obscura) on p. 210]. I here propose the replacement name Hebrus linnavuorii J. Polhemus for Hebrus obscurus Cobben and Linnavuori.— 7(9/2 « T. Polhemus, University of Colorado Museum, 3115 S. York, Englewood, Colorado 801 10. Correction: The following figure from Wheeler, A. G., Jr. and E. R. Hoebeke, “Biology and seasonal history of Rhopalus (Brachycarenus) tigrinus, with description of im- mature stages (Heteroptera: Rhopalidae).” J. New York Entomol. Soc. 96(4):385, is printed as a correction. BOOK REVIEWS J. New York Entomol. Soc. 97(1):1 17-1 18, 1989 Insect-Plant Interactions.— J. R. Miller and T. A. Miller, (eds.) 1986. Springer- Verlag, New York, xii + 342 pp. $67.00 U.S. The study of insect-plant interactions draws on a diversity of biological subdis- ciplines—from plant chemistry to plant and animal physiology to behavior— each with its own technological and methodological quirks. Investigations of insect-plant interactions, as a consequence, regularly stray into territory requiring an understand- ing of experimental techniques with which the researcher is unfamiliar. While the necessary information is undoubtably to be found somewhere, it is often difficult to track down a clear description and evaluation of basic techniques. A broad overview of the current approaches employed in the many areas of insect-plant interactions would therefore be of great help to anyone involved in this sort of work. Although the title of this book suggests a collection of papers documenting the latest insights into the interactions between insects and plants, the book in fact represents an attempt to provide an overview of research methodologies such as I have described above. In assembling this volume. Miller and Miller have brought together a collection of papers that are intended to provide an understanding of the diverse techniques currently used to explore the interactions between insects and plants. A comprehensive list of references accompanies each chapter, providing an easy “in” to the literature for those who require a more comprehensive introduction. The opening chapters consider the host-finding behavior of insects. While each of these three chapters has a different feel to it, they do a good job of surveying the approaches used to study host-finding behavior. Opp and Prokopy describe a variety of techniques for monitoring insect behavior in the field. Finch broadly surveys techniques ranging from the extraction of plant volatiles to the design of wind tunnels and insect traps involved in the investigation of the olfactory and visual cues used by searching insects. Singer wraps up this segment of the book by using his own approach to the study of insect oviposition behavior as an example of the method- ology involved. The middle four chapters of the book consider the effect of plant quality on insect development and feeding behavior. Lewis and van Emden (bioassays), Berenbaum (quantifying toxicity and repellency), and Kogan (nutrition) all thoroughly evaluate the design, interpretation, and weaknesses of various experimental approaches to analyzing the responses of herbivores to plant quality. The chapter by Ishaaya on the interaction of nutritional quality and secondary chemistry strays from the path taken in the rest of the book by presenting a literature review of the topic rather than an introduction to the methodology. The book closes with three chapters that cover a mixed bag of topics. Kubo and Hanke attempt an ambitious overview of phytochemical techniques that is not likely to be of much use to beginners because it skips the basics of extraction and identi- fication, and is not likely to satisfy those more proficient in chemical techniques because it provides little more than a taste of what some of the more advanced techniques can do. Tingey, in the only chapter that takes a plant-centered view of 118 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) the methodology, summarizes techniques used to evaluate the resistance of plants to herbivory in the field and laboratory. Frazier and Hanson conclude by surveying the methods used to investigate the function of insect sensillae at a level accessible to novice neurophysiologists and by providing an overview of the use of microcom- puters to interpret the complex signals commonly generated by sensillae in response to stimulation by plant chemicals. Miller and Miller, in their brief introduction to this volume, state that their ob- jective is to fill a void in the literature by producing a resource of interest to both the expert and non-expert. I suspect that most of the chapters will reveal little new to an expert in any of the fields reviewed. Those same experts, however, will find the book more useful when forced to wear the hat of a non-expert trying to decide on a methodology appropriate to answering a question that lurks on the fringes of their own area of expertise.— Damman, Department of Biology, Carleton Uni- versity, Ottawa KIS 5B6, Canada. J. New York Entomol. Soc. 97(1):1 18-120, 1989 Insects and Flowers. The Biology of a Partnership.— FreidrichG. Barth. 1985. (Trans- lated by M. A. Biederman-Thorson.) Princeton University Press, Princeton, v + 297 pp. $35.00 U.S. The pollination of plants by insects is a process that occurs quietly day and night. Although it goes unnoticed by most people, pollination of plants by insects is none- theless one of those things that makes the earth habitable for the majority of organ- isms, including humans. The subject of this book is about the process of pollination of plants by insects, but as a whole it is decidedly about the dynamics and evolution of insect-plant interactions. Barth uses our oldest insect friend the honey bee {Apis mellifera) as a guide through the hows and whys of pollination biology and also gives the reader an insight into how experimental biology is conducted. The book is divided into 30 short chapters which, in my opinion, should have been collapsed into less than half that number of longer ones. Chapter 1 uses the pollination of Ficus to illustrate both a historical setting for the study of pollination biology, and just how intricate the interaction between pollinator and plant can be. Chapters 2 and 3 are brief outlines of the nuts and bolts of floral function, morphology and ontogeny. Elegantly encapsulated by the topic sentence, “It is not irrelevant where pollen comes from” (p. 21), chapter 4 gives examples of how the sexual mechanics of plant species favor cross pollination and inhibit selfing. After setting up the idea that cross pollination is the norm, chapters 5 and 6 show that pollen transfer in the majority of plants depends upon the intimate relationship between plants and insects. Here we are introduced to those insect groups that have evolved adaptations specifically for working with pollen and transferring it: Hymenoptera, Lepidoptera, Diptera and Coleoptera, with odds favoring bees as the supremely important pollinators of the world. For reasons unclear to me, chapter 7 is devoted to telling us that the evolution of bees may be explained, in part, by the theories of sociobiology and kin selection. Not a word about pollination biology to be found here. 1989 BOOK REVIEWS 119 Chapters 8 through 26 decidedly focus on pollination biology by showing how different insects respond to flower rewards and displays and form the real substance of this book. Chapters 8 and 9 are devoted to the size and food value of pollen and some morphological adaptations that insects have for collecting it. In a similar vein, chapters 1 0 and 1 1 briefly cover nectar as a food reward, followed by biochemical descriptions of the different types of mechanisms insects have for contending with secondary chemicals and assimilating nutrients. The visual spectra and recognition of form in insect pollinators is treated in chapters 12 through 17. Here Barth gives us an account of those elegant experiments that lead to our understanding of insect vision, and delves into the anatomy and physiology of the insect eye. Barth then shows that the colors, uv-patterns, and morphology of flowers can attract, trigger the appropriate pollinator responses, and reinforce these responses. By describing the organs insects have for detecting chemical compounds and the experiments for dem- onstrating their acute sensitivity to minute quantities of chemical substances, chapters 18 through 22 not only make one appreciate how intimately involved and evolved flowers and their insect pollinators are, but also drives home just how pathetically developed taste and smell are in Homo sapiens. Chapters 23 and 24 go down the well-trodden paths of specialized sex in the garden of orchids: scent gathering male euglossine bees and pseudocopulation of Ophrys by male sphecid wasps. Barth gives succinct accounts of how pollination is affected by euglossine bees, the evidence for why bees visit the flowers, and steers clear of the speculations common to much of the literature regarding what the male bees do with the scent. The accounts of Ophrys and its pollinators are beautiful examples of experiments in evolutionary biology. Chapters 25 through 28 integrate pollination biology with a focus on experiments conducted mostly on honey bees that cover learning, time keeping, language, and evolution of the “bee dance,” while chapter 29 gives us an account on the thermal biology of the bumble bee. Chapter 30 shows that pollination biology is a non-static science devoted to the study of natural history, genetics, and evolution— the stuff of life. Richly filled with black and white photographs, color photographs, scanning elec- tron micrographs, drawings, this well bound book is clearly worth the publisher’s retail price. Barth has produced a well written book that will be useful as a general text for undergraduate and graduate students of pollination biology. I have two complaints about this book. It was disappointing to see so few examples of tropical insect-plant interactions mentioned. Some of the more spectacular ex- amples that spring to mind for use in a book such as this are: the stamen and stigmatic morphology of bee flowers of the Lecithycidaceae; anthophorid bee leks generated by mass-flowering canopy trees; the consequences (both to insects and plant) of Heliconius butterfly specialization on flowers of Psiguria and Gurania (Cucurbita- ceae); floral mimicry among different plant families and their shared pollinators; and the classic story of yucca moths. Such examples not only excite the imagination and stimulate evolutionary thought, but bolster the principles of pollination biology anas- tomosing throughout this book. However, such omissions are forgivable since the author was concerned mainly with highlighting European workers. Nonetheless, one should be made aware that the tropics are where the highest diversity of insect-plant interactions on the planet occur. My most serious criticism regards the omission of the pioneering and classic work 120 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) done by Herbert and Irene Baker on pollination syndromes and flower constituents. To my mind the Bakers’ work is a sterling example of the high quality science that the study of pollination biology is capable of generating. Not mentioning their work is a disservice and deprives potential students of pollination biology of a great re- source. The following papers (and the references therein) by the Bakers’ should help fill the void for those potential students, and the present and future owners of this book: Baker and Baker, 1973, 1975, 1977, 1978; Baker et al., 1978. Oh yes, and remember, one can find the last ten years of the Bakers’ work in their local library. — P. J. DeVries, Smithsonian Tropical Research Institute, Box 2032, Balboa, Panama. LITERATURE CITED Baker, H. G. and Irene Baker. 1973. Amino acids in nectar and their evolutionary significance. Nature 24:543-545. Baker, H. G. and Irene Baker. 1975. Studies of nectar constitution and pollinator-plant coevolution. Pages 100-140 In: L. E. Gilbert and P. H. Raven (eds.), Coevolution of Animals and Plants. Univ. of Texas Press, Austin. Baker, H. G. and Irene Baker. 1977. Intraspecific constancy of floral nectar amino acid complements. Bot. Gaz. 138:183-191. Baker, H. G. and Irene Baker. 1978. Sugar analyses of floral nectar and their significance. In: T. S. Elias and B. L. Bentley (eds.), The Biology of Nectaries. Columbia Univ. Press, New York. Baker, H. G., P. A. Oplerand Irene Baker. 1978. A comparison of the amino acid complements of floral and extrafloral nectars. Bot. Gaz. 139:322-332. J. New YorkEntomol. Soc. 97(1):120-122, 1989 Pheromone Biochemistry. — Glenn D. Prestwich and Gary J. Blomquist (eds.). 1987. Academic Press. 565 pp. Cloth, $85.00 U.S. Pheromones are the chemical signals of the most elementary communication sys- tem. After the first identification of such a substance, namely bombykol (1 959), earlier hopes for a highly specific pest control with attractant pheromones were nourished. In the meantime, with modern chemical methods available, hundreds of such sub- stances became known, mostly as rather precise mixtures of related or analogous compounds. Unfortunately, pheromones turned out not to be a panacea against pests, but at least have become useful to predict the threat of pest outbreaks. World- wide, research on pheromones has mainly been done by the pragmatic entomologists, who are rightly worried about protection of plant, animal, and man, and less so by people in the basic disciplines. Pheromones occur all over the animal kingdom, yet, except for the insects and vertebrates, rather few have been studied in detail. What could “pheromone biochemistry” mean? We are, on the sender side, dealing with glandular systems which produce these “exocrines or ektohormones,” as they were called earlier. Is there anything functionally specific to be expected from such a wide variety of barely analogous glands? The book overcomes this dilemma by only treating selected pheromone-producing systems in some insects and ticks, with emphasis on female Lepidoptera. Just here— and also in the bark beetles— the in- teresting question arises about the biochemical basis of the specificity of the pher- 1989 BOOK REVIEWS 121 omone odor cocktails of related and/or sympatric species. Several chapters of the book deal with the fascinating biosynthetic machinery which is responsible for this. The expected diversity of these processes hnds its parallel in the fact that different endocrine systems control the pheromone glands: neurohormones in the Lepidoptera, juvenile hormone in the bark-beetles and ecdy steroids in the flies and ticks. On the receiving side, pheromone-specific biochemical function cannot only be expected in relation to the specificity of the receptor binding but also with other transducing steps. Here, the state of our insight is even less advanced than on the sender side. The first part of the book is introduced by two elegant chapters on “Structure and Function” of insect pheromones (J. H. Tumlinson and P. E. A. Teal) and the “Biology and Ultrastructure” of the respective insect glands (J. E. Percy-Cunningham and J. A. MacDonald). These are useful overviews which set the stage. The following chap- ters on pheromone production and hormonal control are important progress reports. “Desaturation and Chain Shortening” occurs during the biosynthesis of Lepidoptera pheromones from longer fatty acids (L. B. Bjostad, W. A. Wolf, and W. R. Roelofs). The “Enzymes” which serve this function are described by D. Morse and E. Meighen, and the respective “Endocrine Regulation” by A. K. Raina and J. J. Menn. “Bio- synthesis of Pheromones and Endocrine Regulation” in the beetles and flies is treated by D. Vanderwel and A. C. Dehlschlager, and by D. J. Blomquist, J. W. Dillwith, and T. S. Adams, respectively. Then follows a report on “Alkaloid Derived Phero- mones and Sexual Selection” in some male Lepidoptera, which do not rely on the normal metabolites for the production of their pheromones as do their females, but ingest these plant products for their own protection, as nuptial gifts, and eventually as precursors for the biosynthesis of their signals (T. Eisner and J. Meinwald). “Neu- roendocrine Regulation” of the pheromone related behavior of ticks is adequately treated by D. E. Sonenshine. The final chapter of this first part of the book describes “Cantharidine Biosynthesis and Function in Meloid Beetles” (J. P. Mccormick and J. E. Carrel). This is a fascinating defensive substance, yet doubtful as a pheromone. This long, laboratory manual style chapter is not well placed here. How could “biochemistry” of the pheromone receptor organs differ from such a function in other olfactory receptors? Obviously by their specificity but not by their elementary properties. No doubt, electrophysiological studies are an essential first step to understand the specificity on the level of the receptor cells and this was the reason why I started 35 years ago to study olfaction in the silkmoth. The final third of the book begins with a fine treatment of the “Functional Morphology” of those antennal insect sensilla which respond to pheromones (R. A. Steinbrecht). These are the structures and compartments where the transducing processes occur. The cor- responding “Neurobiology of Pheromone Reception” by J. J. DeKramer and J. Hemberger presents not only the established electrophysiological facts but also dis- cusses controversial points, most of which are only distantly related to biochemistry and not too well placed in this book. Two chapters deal with olfactory biochemistry. R. G. Vogt (“Molecular Basis of Pheromone Reception”) describes in a lengthy chapter his and other peoples’ efforts to understand transduction phenomena in the sensillum lymph space. Degrading enzymes have been known for many years and the existence of pheromone binding proteins (with uncertain function) is rather well established. When these chapters were written, receptor protein identification was not yet successful. Yet recently these proteins have been “visualized” by using ra- diolabelled photoaffinity analogs (R. G. Vogt, G. D. Prestwich and L. M. Riddiford. 122 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) 1988. J. Biol. Chem. 263:3952-3959). This publication provides remedies for some of the uncertainties of another rather long, often programmatic and technical chapter by G. D. Prestwich (“Reception and Catabolism”). Hopefully, some of his other claims on transducing functions in a number of species will also be verified soon. The final chapter is a good, compact treatment of the “Molecular Mechanism of Vertebrate Olfaction” with no specific relation to pheromones (nobody ever found specific pheromone receptor cells in noses, in contrast to insects). Interestingly, this chapter is meant to be an “introduction” for the insect people, yet is wrongly placed in this book. In my personal view, the most important future result of pheromone biochemistry will be the understanding of how, and under what evolutionary pressure, speciation took place. The “royal” question is a behavioral one: any mutation which changes the composition of the signal or its receptor risks a misunderstanding or even a total blockage of the communication. There are indications that we are beginning to understand the respective mechanism on the sender side, yet we are still far from this goal on the receiving side. Since many biochemical functions in pheromone systems are still largely unknown, it must have been difficult for the editors to plan this book. The result is a not too satisfying compromise. The book contains good general chapters with little biochem- istry, a series of interesting biosynthesis chapters (some too packed with methods), confusing or even premature olfactorial chemistry, and two chapters which do not relate to the theme. One also wonders, whether the editors wanted essay-style chap- ters, reviews or methodological introductions? For the insider, most chapters will be useful, for they tell in what direction the field is moving. Newcomers and students might like the introductory and overview parts. Unfortunately, some of the chapters even lack a summarizing paragraph and everybody will miss a final author index, which would help to use this book as a reference souycq.— Dietrich Schneider, Max- Planck- 1 nstitut fur Verhaltensphysiolo- gie, D-8131 Seewiesen, Fed. Rep. Germany. J. New YorkEntomol. Soc. 97(1):122-123, 1989 Portraits of South Australian Geometrid Moths.— Noel McFarland. 1988. Published by the author, iv + 400 pp., over 1,400 figs. $80.00 soft cover (includes packing and postage). Copies may be obtained from the author: P.O. Box 1404, Sierra Vista, Arizona 85636. The author has had a long time interest in rearing Lepidoptera, especially members of the Geometridae. This goes back to when he was a young boy living in southern California; he became adept not only at life history work but also became well acquainted with the local fauna. During 1965-1969 he lived in South Australia, and continued his rearings of geometrids. But he was almost completely unprepared for “the mind-boggling array of incredible forms” that he found there, both as larvae and adults. Further, the great majority of these geometrids had never been studied or documented. It was at this time that he learned the necessary photographic tech- niques to be used to document the rearings. And, as they say, the rest is history, as seen in this most impressive volume. 1989 BOOK REVIEWS 123 Seventy-two different species are fully illustrated and described, usually with several views of the adults, eggs (when available), mature larvae and pupae; each species is covered in a separate chapter in the book. Also included are references to the original descriptions and pertinent literature, locations of the type specimens and of preserved material, localities, food plants, habits and general remarks. The author’s original descriptive notes were made while he was rearing each individual species; these have been rewritten for this book. The organization of the volume may be thought of as rather unorthodox; for example, the separate indices for zoological and botanical names are at the front. The heart of the book consists of two sections with the photographs and descriptions of the species; the first 46 chapters cover the members of the Ennominae and Oe- nochrominae, while the remaining 26 are devoted to the remaining subfamilies. The first portion of the book includes, in addition to the indices, the preface, introduction, locality maps and habitat photos. Between the two main sections will be found general comments on the subfamilies, food plants, and three appendices, one each on the resting positions of adults, of geometrid larvae, and of the forms of crypsis. Following the second section there is another appendix, very helpful discussions of rearing and preserving techniques, photography, acknowledgments, glossary and bibliography. This ordering of subjects takes a bit of getting used to, but it works out rather well after you get the hang of it. Do not expect setal maps, drawings of cremasters, genitalia, and the like; this volume was never meant to be a revisionary work. McFarland set out to rear as many geometrids as possible— many were from his back yard or from a short distance from his home— and to obtain a photographic and descriptive record of the living organisms, showing them as they appeared, not as the usual preserved specimens. In this he has done a superb job. Life history material was preserved, much of it being deposited in the South Australian Museum. Thus it is available for future workers, who will need to study the details of the structures of the early stages. From this, considerable light may be shed upon the higher classification of the Geometridae, something that is badly needed, as the author points out. It is extremely valuable having these 72 life histories in a single volume rather than scattered throughout the literature. McFarland is to be congratulated on handling his material in this manner, and in such a professional way. This book sets high standards; it is to be hoped that it will encourage other workers to do life history work in such a thorough and complete manner. The author has earned a “well done” for all the work he has done, both on the life histories themselves and for publishing such a valuable addition to the extremely small amount of our knowledge on geometrid life hisioYiQs.— Frederick H. Rindge, Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024. J. New York Entomol. Soc. 97(1):123-124, 1989 Proceedings of the Fifth International Symposium on Trichoptera, Lyon, France 21- 26 July 1986. M. Boumard, and H. Tachet (eds.) Dr. W. Junk Publishers, Dordrecht 1987, xxiii + 397 pp. 250.00 Dfl. 124 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(1) The International Symposia on Trichoptera are held every three years with Lyon as the site of the fifth meeting. Eighty-nine caddisfly specialists from around the world attended, representing 23 countries. Like the previous symposia it was once again a week of camaraderie, ideas, and with large amounts of original and new information exchanged. The scientific program contained poster displays, two field trips and 57 lectures, the results of which are published in the proceedings. The contributions are arranged under the following topics (numbers of papers in parentheses): Morphology and Anatomy (3), Taxonomy (6), Fossil Forms (3), Faunistics (denoted as Biogeog- raphy) ( 1 0), Phylogeny (4), Biology and Physiology (5), Light Traps and Flight Patterns (7), and Ecology (24). The subjects are generally treated with a thoroughness and competence such that the reader may feel safe to obtain a representative view on the diverse subjects. Among the numerous contributions several would be of particular interest to a broad audience, e.g., “Caddisflies and quaternary palaeoecology— what have we learned so far?” (N. E. Williams), “Techniques for demonstrating sex pher- omones in Trichoptera” (V. Resh, H. Jackson, J. K. and J. R. Wood), “Caddisfly adaptations to the variable habitats at the land-water interface” (N. A. Erman), “Seston quality as a factor influencing Trichoptera populations” (R. C. Peterson, Jr.), “A new ordination technique for ecological purposes applied to caddis larvae in ditches” (L. W. G. Higler, R. Torenbeck, and P. F. M. Verdonschot), “Effects of an artificially silted stream bottom on species composition and biomass of Trichoptera in Breitenbach” (R. H. Wagner), “Studies of Plectrocnemia conspersa (Curtis) in copper contaminated streams in south West England” (S. T. Darlington, A. M. Gower, and L. Ebdon), “Recovery of the Trichoptera fauna near Mt. St. Helens five years after the 1980 eruption” (N. H. Anderson and R. W. Wisseman), and the paper on “Trichoptera of regulated Rocky Mountain streams” (J. V. Ward). The book con- cludes with four indices, dealing with authors, geographic names, subjects, and species, respectively. The book should serve as a valuable and up-to-date reference, not only for caddisfly workers, but also for other students of freshwater benthos, and partic- ularly for freshwater ecologists. — IF6>//r(2m Mey, Entomologie, Museum fur Natur- kunde der Hombolt-Universitdt zu Berlin, Zoologisches Museum, Invalidenstr. 43, DDR-Berlin 1040. Announcement:— The Eagle Hill Wildlife Research Station offers advanced and professional seminars in natural history, ecology, and related fields, May-September. Of particular interest may be: “The Lepidoptera: Moths, Advanced Techniques,” July 2-8, 1989. For more information, contact the Station in: Steuben, Maine 04680; phone 207-546-2821. INSTRUCTIONS TO AUTHORS The Journal of the New York Entomological Society is devoted to the advancement and dissemination of knowledge of insects and related taxa. The costs of publishing the Journal are paid by subscriptions, membership dues, page charges, and the proceeds from an endowment established with bequests from the late C. P. Alexander and Patricia Vaurie. The Journal will consider for publication manuscripts of any length dealing with original research in entomology. Longer papers will be printed as articles, shorter ones as “scientific notes.” Book reviews will be solicited by the Book Review Editor. Manuscripts should be submitted in duplicate to: Dr. Randall T. Schuh, Editor, Journal of the New York Entomological Society, c/o Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024. All material must be double-spaced (including references, tables, captions, etc.) and prepared in the format of a recent issue of the Journal. Longer manuscripts intended for submission as articles should be accompanied by a brief abstract. Footnotes should be avoided. Tables should be prepared as separate pages; they should be kept to a minimum because of the high cost of typesetting, but may be submitted as photographically reproducible material (see below). The list of references is headed “Literature Cited” and should follow the format indicated in the CBE Style Manual or as found in a recent issue of the Journal. Illustrations (originals whenever possible) should be submitted flat, never rolled or folded, with the manuscript. 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Society members will be charged a fee of $20.00 per printed page and $5.00 per plate of figures. Non-members will be charged $45.00 per printed page and $5.00 per plate of figures. Member authors who do not have institutional funds may petition to the Society for waiver of page charges for no more than eight pages on a once a year basis. Because of limited funds, all such requests will be handled on a first-come first-serve basis. Authors will receive a reprint order blank with the proofs. Reprints are ordered directly from the printer with no benefit accruing to the Society. Journal of the New York Entomological Society VOLUME 97 JANUARY 1989 NO. 1 CONTENTS The biology of Sthenopis auratus (Grote) (Lepidoptera: Hepialidae) Tim L. McCabe and David L. Wagner A revisionary study of the neotropical hairstreak butterfly genus Noreena and its new sister genus Contrafacia (Lepidoptera; Lycaenidae) Kurt Johnson A new Aphaenogaster (Hymenoptera: Formicidae) from southern New Mexico William P. MacKay Notes on ant larvae: Ponerinae George C. Wheeler and Jeanette Wheeler Biology of Andrena crataegr Ro};^Qrtson (Hymenoptera: Andrenidae), a commun- ally nesting-bee Eben A. Osgood N The genus Metopina (Diptera: Phoridae) from Cretaceous and Tertiary ambers N David Grimaldi ♦- .V- ' • Naucoridae (Heteroptera) of New Guinea. IV. A revision of the genus Cavocoris w^ descriptions of four new specif * - Dan A. Polhemus and John T. Polhemus A. Cariniocoris, A' new«phy line plant bug genus from the eastern United States, with . a‘"discussion of generic relationships (Heteroptera: Miridae) Thomas J. Henry Froeschnerana mexicana, a new genus and species of Deraeocorinae from Mexico (Heteroptera: Miridae) J. C. Schajfner and Paulo Sergio Fiuza Ferreira Two new species of Mormidea from Mexico and Guatemala (Heteroptera: Pentatomidae) D. A. Rider and L. H. Rolson Notes and Comments New records of Palearctic Heteroptera in New York State: Microphysidae and Miridae Michael D. Schwartz Names and authorship of two family-groups in the Eph^meroptera William L. Peters and Michael D. Hubbard A necessary new name in the family Hebridae (Heteroptera: Gerromorpha) John T Polhemus Book Reviews Insect-plant Interactions Hans Damman Insects and Flowers. The Biology of a Partnership P. J. DeVries Pheromone Biochemistry' Dietrich Schneider Portraits of South Australian Geometrid Moths Frederick H. Rindge Proceedings of the Fifth International Symposium on Trichoptera, Lyon, France 21-26 July 1986 Wolfram Mey 1-10 11-46 47-49 50-55 56-64 65-72 73-86 87-99 100-104 105-110 111-114 115 116 117- 118 118- 120 120-122 122- 123 123- 124 124 Announcement S'‘ii S' -70(^7% !£^ Vol. 97 APRIL 1989 No. 2 Journal of the New York Entomological Society (ISSN 0028-7199) Devoted to Entomology in General JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Editor: Randall T. Schuh, Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024 Book Review Editor: David A. Grimaldi, Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024 Publications Committee: Louis Trombetta, St. Johns University, Queens, New York, Chairman; Alfred G. Wheeler, Jr., Pennsylvania State Department of Agriculture, Harrisburg; Joseph M. Cerreta, St. Johns University, Queens, New York. The New York Entomological Society Incorporating The Brooklyn Entomological Society President: Dennis J. Joslyn, Department of Biology, Rutgers University, Camden, New Jersey 08102 Vice President: Durland Fish, Medical Entomology Laboratory, New York Medical College, Armonk, New York 10504 Secretary: Richard Falco, Westchester County Health Department, White Plains, New York 10601 Treasurer: Louis Sorkin, Department of Entomology, American Museum of Natural History, New York, New York 10024 Trustees: Class of 7955— Henry M. Knizeski, Jr., Mercy College, Dobbs Ferry, New York; Michael D. Schwartz, American Museum of Natural History, New York, New York; Class of 7959— Christine Falco, West- chester County Health Department, White Plains, New York; James S. Miller, Department of Entomology, American Museum of Natural History, New York, New York. Annual dues are $23.00 for established professionals with journal, $10.00 without journal, $15.00 for students with journal, $5.00 without journal. Sustaining memberships are $53.00 per year, institutional memberships are $125.00 per year, and life memberships are $300.00. Subscriptions are $40.00 per year domestic and $45.00 foreign. All payments should be made to the Treasurer. Back issues of the Journal of the New York Entomological Society, the Bulletin of the Brooklyn Entomological Society, Entomologica Americana, the Torre-Bueno Glossary of Entomology and other Society publications can be purchased from Lubrecht and Cramer, RD 1, Box 244, Forestburgh, New York 12777. Meetings of the Society are held on the third Tuesday of each month (except June through September) at 7 p.m. in the American Museum of Natural History, Central Park West at 79th Street, New York, New York. Mailed June 20, 1989 The Journal of the New York Entomological Society (ISSN 0028-7199) is published 4 times per year (January, April, July, October) for the Society by Allen Press, Inc., 1041 New Hampshire, Lawrence, Kansas 66044. Second class postage paid at .New York, New York and at additional mailing office. Postmaster; Send address changes to the New York Entomological Society, % American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024-5192. Known office of publication: American Museum of Natural History, New York, New York 10024. Journal of the New York Entomological Society, total copies printed 700, paid circulation 602, mail subscription 602, free distribution by mail 19, total distribution 621, 79 copies left over each quarter. THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER. THE NEW YORK ENTOMOLOGICAL SOCIETY in cooperation with the AMERICAN MUSEUM OF NATURAL HISTORY announces A Revised and Expanded Edition of THE TORRE-BUENO GLOSSARY OF ENTOMOLOGY Compiled by Stephen A. Nichols Managing Editor Randall T. Schuh First published in 1937, the Glossary of Entomology by J. R. de la Torre-Bueno, with some 8500 entries, has long been the standard for usage of entomological terms in the English language. Now available, after 50 years, is a revised and expanded edition, containing approximately 16,000 terms in 840 pages. The new Glossary serves as a basic reference in general entomology, listing thousands of new terms and definitions in the fields of insect morphology, behavior, classification, physiology, medical entomology, pathology, and applied entomology. In conjunction with the Glossary the Society has prepared an entomological spell- checker file containing over 9000 terms. This ASCII file can be added to most word processors and serve as part of your personal dictionary of technical terms. To be published on June 1, 1989, the Glossary can be ordered directly from the New York Entomological Society at the prepublication discount prices which follow. After September 15, 1989 the Glossary will be available to members of the New York Entomological Society for $40.00 and to non-members for $45.00. Members of the NYES $35.00 □ Non-members $40.00 □ 1990 Membership + Glossary $58.00 □ Library orders $45.00 □ Entomological Spell-Checker $15.00 □ 3V2 inch disc □ SVi inch disc □ To take advantage of the discount price, please check the appropriate box above and return the form with your check or money order (payable in U.S. dollars on a U.S. bank). Please enter your correct address in the space provided below. Send to; New York Entomological Society c/o Department of Entomology The American Museum of Natural History Central Park West at 79th Street New York, N.Y. 10024-5192 ■r •IV”''': -v v."^^'v - ... * - ■"*\ ** ! 1 (' M y' ; ' 'i- J. New York Entomol. Soc. 97(2): 125-1 32, 1989 A FOSSIL SOLPUGID, HAPPLODONTUS PROTERUS, NEW GENUS, NEW SPECIES (ARACHNIDA: SOLPUGIDA) FROM DOMINICAN AMBER George O. Poinar, Jr. and Jorge A. Santiago-Blay Department of Entomological Sciences, University of California, Berkeley, California 94720 Abstract.— \ new genus and species of fossil solpugid, Happlodontus proterus, is described from Dominican amber. This is the first Tertiary record for the order, the second described fossil solpugid and the first solpugid fossil described from amber. On the basis of this fossil and the present distribution of the family Ammotrechidae, phylogenetic aspects of the group are discussed. The windscorpions or Solpugida (=Solifugae) compose an order of the Class Arach- nida characterized by large forwardly directed chelicerae, stout pedipalpi with a blunt end bearing an adhesive organ, slender, tactile front legs, clear abdominal segmen- tation and presence of malleoli (racket organs) on the fourth legs (Petrunkevitch, 1955; Roewer, 1934). Their distribution is mainly tropical and subtropical with most species found in Africa. They are among the fastest invertebrate runners which enables them to be effective predators of other small animals. The fossil record of the Solpugida has, up to the present, been limited to a single species, Protosolpuga carbonaria Petrunkevitch (1913) from the Pennsylvanian of North America (ca. 285-320 million years ago). This paper describes the second known fossil solpugid which represents the only known specimen from Tertiary deposits (ca. 25-40 mya) and the first known solpugid in amber. MATERIALS AND METHODS The piece of amber containing the solpugid was made available to us by Dr. James Truman, Department of Zoology, University of Washington, Seattle. The amber was roughly obovate in outline, measuring 42 mm in length by 27 mm in width and 6 mm in depth. It was mounted within a silver frame in the form of a pendant. The weight of the amber and mounting was 1 0 grams. The amber was clear and yellow and the solpugid was clearly visible toward the wider end. The piece of amber originated from mines in the Cordillera Septentrional of the Dominican Republic. These mines are located in the Altimira facies of the El Mamey Formation which is a shale-sandstone interspersed with conglomerate of well rounded pebbles dating from the Upper Eocene (Eberle et al., 1980). Additional studies in- dicate that the ember in these mines ranges from the Lower Miocene (25 my) to the Upper Eocene (40 my) in age (Lambert et al., 1985). 126 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Table 1. Measurements of Happlodontus proterus gen. n. sp. n. Character Measurement (mm) Chelicera Length, width 1.60, 0.42 Propeltidium Length, maximum width 1.09, 0.92 Diad width 0.25 Pedipalpus Trochanter length, width 0.34, 0.13 Femur length, width 0.84, 0.25 Tibia length, width 1.34, 0.17 Metatarsus length, width 0.97, 0.17 Tarsus length, width 0.29, 0.17 Abdomen I Length, width 0.38, 0.50 II Length, width 0.28, 0.71 III Length, width 0.50, 0.92 IV Length, width 0.29, 0.88 V Length, width 0.46, 0.92 VI Length, width 0.67, 1.13 VII Length, width 0.67, 0.97 VIII Length, width 0.34, 0.88 IX Length, width 0.34, 0.55 X Length, width 0.21, 0.46 Total length 6.83 RESULTS Happlodontus, new genus Order Solpugida Leach, 1815 as defined by Petrunkevitch (1955) Family Ammotrechidae Roewer, 1934 Subfamily Ammotrechinae Roewer, 1934 Description. Movable finger of chelicerae with a single major tooth; fixed finger with three distinct teeth, fingers relatively slender; claws absent on first pair of legs, present on legs 2-4; tarsi on legs 2-3 composed of a single segment; tarsi on leg 4 with 3 segments; ventral spinelike setae on distal segment of tarsi 4 vary from one to six; anus subterminal. Type species. Happlodontus proterus, new species, by monotypy. Specimen de- posited in the private collection of James W. Truman, Seattle, Washington. Happlodontus proterus, new species Figs. 1-12 Type data. From Cordillera Septentrional, Dominican Republic. Estimated age, 25-40 million years. 1989 FOSSIL SOLPUGID 127 Figs. 1 3. 1- Dorsal view of 2. Ventral. 3. Chelicera, dorsal, slight laterally (arrow shows single ventral tooth). 128 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Figs. 4-7. 4. Chelicera, lateral (arrow shows single ventral tooth). 5. Propeltidium. 6. Pedi- palpus and distal segments of right leg. Arrow points tarsus of pedipalpus. 7. Distal segments, right leg 4. 1989 FOSSIL SOLPUGID 129 Figs. 8, 9. 8. Abdominal stemites setation. 9. Subterminal anus (arrow), ventral view. 130 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Figs. 10-12. 10. Dorsal view of propeltidium and anterior appendages. 11. Retrolateral view of left chelicera. 12. Distal segments, left leg 4. Description. With the characters of the genus. Body form as in Figures 1 and 2. Overall coloration pale brown. Measurements as in Table 1. Chelicera. Fingers attenuated distally, darker; with all teeth distinct, well developed (Figs. 3, 4, 11) and a tubercle located retrolaterally, near base of movable finger (Fig. 1 1). Propeltidium. Anterior margin convex; scarcely hirsute (Figs. 5, 10). Pedipalpus. Hirsute; tarsi and metatarsi with several setae much longer and stronger than others but lacking ventral spine-like setae and claws (Figs. 6, 10). Legs. Count of spine-like setae on terminal tarsomeres as follows: leg 1-0, leg 2-5, leg 3-5(7), leg 4- from 1- 6; as illustrated in Figures 6, 7, 12. Counts not given could not be obtained. (See Discussion for usefulness of this character.) Abdomen. Hirsute; stemite setae arranged in three primary transversal rows located anteriorly, medially and posteriorly (Fig. 8); sternites without comb-like setal groups; genital operculi and other modifications on anteroventral region obscure. Anus subterminal, bioperculated (Fig. 9). Discussion. Discovery of a solpugid in amber is interesting because normally this group is associated with hot, dry habitats. However, this find suggests that the arboreal habit known today in some of the Central American species was already established in the Tertiary. Although H. proterus is considered a juvenile, the adults were probably relatively small, which could be advantageous in an arboreal habitat. The size of solpugids 1989 FOSSIL SOLPUGID 131 varies from 9 to 70 mm (Petrunkevitch, 1955) and the fossil speeimen (6.8 mm) probably represents an advanced juvenile stage. According to Muma (personal com- munication, June 1988), the number of teeth on the chelicerae does not change from one stage to another or immatures to females nor does the number of ventral tarsal spines differ between juveniles and adults. Extant species with a single major tooth on the ventral finger are unknown. The fossil species does illustrate the wide variation that can occur regarding the number of spines on the fourth and other tarsal segments. If such variation can occur on the same individual, then the wide use of tarsal spination for the determination of solpugid genera and species is questionable, unless variability studies are first undertaken. The extant distribution of West Indian solpugids includes representatives of the four genera: Ammotrechella Roewer, 1934; Ammotrechona Roewer, 1934; Ammo- trecha Banks, 1 900; and Ammotrechinus Roewer, 1934 of the Ammotrechidae (Muma, 1970, 1987). This report represents the first solpugid from the Dominican Republic although Roewer (1934) reported Ammotrechinus gryllipes (Gervais) from Haiti. There are some curious distributional patterns of solpugids. They are most speciose in Africa (Muma, 1976) and that continent may have been their center of origin. However, there are no known records in Madagascar. They occur in dry, subdesert regions in the New and Old World but are absent in Australia. Representative of the families Eremobatidae Roewer, 1934, Ammotrechidae Roewer, 1934, Mummiciidae Roewer, 1 934 and Daesiidae Roewer, 1 934 occur on the American continent (Maury, 1984). Muma (1971) described the Amacataidae from Chile but Maury (1980) syn- onymized it with Daesiidae. Representatives of the first two (with some 25 genera) are uniquely Nearctic. Representatives of the last two families also occur in Africa, Arabia, Asia Minor, Persia and Spain (Millot and Vachon, 1949). It would appear that the Ammotrechidae constitute a phylogenetically recent group which evolved no earlier than the late Mesozoic after the African and South American continents had separated. ACKNOWLEDGMENTS We wish to thank Dr. James W. Truman for allowing us to study the amber piece. We are also grateful to Dr. Martin H. Muma (Portal, AZ) for his advice, supplying us with pertinent literature and reading the completed manuscript. Drs. Rosen Rosenblatt (Scripps Institution of Oceanography, La Jolla) and William Eschmeyer (California Academy of Sciences, San Francisco) provided us with nomenclatural recommendations. LITERATURE CITED Banks, N. 1900. Synopses of North American invertebrates. IX. The scorpions, solpugids and pedipalpi. Amer. Nat. 34:421^27. Eberle, W., W. Hirdes, R. Muff and M. Pelaez. 1980. The geology of the Cordillera Septen- trional. Proc. 9th Caribbean Geol. Conf August 1980. Santo Domingo, D.R. pp. 619- 632. Lambert, J. B., J. S. Frye and G. O. Poinar, Jr. 1985. Amber from the Dominican Republic: analysis by nuclear magnetic resonance spectroscopy. Archaeometry 27:43-51. Maury, E. A. 1980. Presencia de la familia Daesiidae en America del Sur con la descripcion de un nuevo genero (Solifugae). J. Arachnol. 8:59-67. 132 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Maury, E. A. 1 984. Las familias de Solifugos americanos y su distribucion geografica (Arach- nida, Solifugae). Physis (Seccion C) 42:73-80. Millot, J. and M. Vachon. 1 949. Ordre des Solifuges. In: P. P. Grasse (ed.), Traite de Zoologie, Vol. 6:482-519. Masson et Cie, Paris. Muma, M. H. 1970. A synoptic review of North American, Central American and West Indian Solpugida (Arthropoda: Arachnida). Arthropods of Rorida and adjacent land areas. 5:1-62. Muma, M. H. 1971. The solpugids (Arachnida, Solpugida) of Chile, with descriptions of a new family, new genera, and new species. Amer. Mus. Novit. no. 2476, 23 pp. Muma, M. H. 1976. A review of Solpugid families (with an annotated list of Western Hemi- sphere solpugids). Publ. Office of Research, Western New Mexico Univ. 2:1-31. Muma, M. H. 1 987. New species and records of Solpugida (Arachnida) from Mexico, Central America and the West Indies. Southwest Offset, Silver City, New Mexico. 24 pp. Petrunkevitch, A. 1 9 1 3. A monograph of the terrestrial Palaeozoic Arachnida of North Amer- ica. Trans. Conn. Acad. Arts. Sci. 18:1-137. Petrunkevitch, A. 1955. Arachnida. Pages 154-155 in: Treatise on Invertebrate Paleontology Part P. Arthropods 2. Geol. Soc. Amer. & Univ. Kansas Press. Roewer, C. Fr. 1934. Solifugae, Palpigradi. In: H. G. Bronn (ed.), Klassen und Ordnungen des Tierreichs. Leipzig. Vol. 5, part 4, book 4:1-723. Received July 29, 1988; accepted September 26, 1988. J. New York Entomol. Soc. 97(2): 133-140, 1989 ON THE ABUNDANCE AND ECOLOGY OF RICINULEI (ARACHNIDA) FROM CENTRAL AMAZONIA, BRAZIL Joachim U. Adis,' Norman I. Platnick,^ Jose W. de Morais,^ and Jose M. Gomes Rodrigues^ 'Tropical Ecology Working Group, Max-Planck-Institute for Limnology, Postfach 165, D-2320 Plon, FRG, in cooperation with the National Institute for Amazonian Researeh (INPA), Manaus, Brazil ^Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024 ^Instituto Nacional de Pesquisas da Amazonia (INPA), Caixa Postal 478, BR-6901 1 Manaus/ AM, Brazil — Ricinuleids from primary and secondary dryland forest soils in the region of Manaus represented two species: (1) Cryptocellus becki with 94 specimens, mostly juveniles, maximum abundance 38 ind./m^, and predominantly obtained during the dry season and (2) Cryptocellus adisi with 17 specimens, mostly juveniles, up to 10 ind./m^, and more frequent during the dry season. These sympatric species appear to be separated in their mutual habitat by spatial differences, with C. becki inhabiting mostly the organic soil layer and the smaller C adisi the mineral subsoil. C becki showed no distinct reproductive period and its first stage of development occurred throughout the year. Neotropical ricinuleids are litter and soil inhabitants of dryland forests in Central Amazonia (Morais, 1985; Rodrigues, 1986). They have not been found in caves (Karmann, 1986; Adis, unpubl.) or in inundation forests along rivers in the Amazon Basin (Adis, 1981). A comparison of the abundance and ecology of Cryptocellus becki Platnick and Shadab (1977) and C. adisi Platnick (1988) from primary and secondary dryland forest sites in the region of Manaus is now possible, as the taxonomic evaluation has been completed. STUDY AREA AND METHODS Ricinuleids were collected between 1982 and 1983 in the course of ecological studies on Central Amazonian arthropods from two previously investigated and fully described forest types, all within 30 km of Manaus: (1) in a primary dryland forest (terra firme forest) at Reserva Florestal A. Ducke (2°55'S, 59°59'W) on the Manaus- Itacoatiara highway (AM-010 at km 26), study area of Adis and Schubart (1984), Adis et al. (1984), Morais (1985), Penny and Arias (1982) and others; (2) in a cut but unbumed secondary dryland forest (capoeira forest) adjacent to an inundation forest at Rio Taruma Mirim (03°02'S, 60®17'W), a tributary of the Rio Negro, study area of Adis and Schubart ( 1 984) and Rodrigues ( 1 986). Both forest types were subject to a rainy season (December-May: average precipitation 1,550 mm) and a dry season (June-November: average precipitation 550 mm; cf Ribeiro and Adis, 1984). 134 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) The yellow latosoil of the primary and secondary forest supported a 2-3 cm thick humus layer (AJ, interspersed with fine roots, and a thin, surface covering leaf litter. Morais (1985) and Rodrigues (1986) provided data on the presence of ricinuleids in the soil between August, 1982 and August, 1983. Once a month they took twelve soil samples (in each forest) along a transect, selected at random, with a split corer (a steel cylinder with lateral hinges, diameter 2 1 cm, length 33 cm), which was driven into the soil by a mallet. Each sample of 7 cm depth was then divided into two subsamples of 3.5 cm each. Animals were extracted from subsamples following a modified method of Kempson (Adis, 1987). 1989 RICINULEI IN AMAZONIA 135 Fig. 2. Cryptocellus adisi Platnick, male, dorsal view. Additional ricinuleids were obtained in a cut but unbumed secondary dryland forest at the campus of INPA, Manaus (03°08'S, 60°0rW), study area of Prance (1975). One sampling each was carried out during the dry season in September, 1985 and during the rainy season in April, 1986 (Adis et al., 1987a, b). Six soil samples were taken, respectively, to a depth of 14 cm, divided into four subsamples of 3.5 cm each, and animals extracted as described above. All ricinuleids collected were identified to species and classified as juveniles (larvae, protonymphs, deutonymphs, tritonymphs) or adults (males and females), according to the number of legs and tarsal segments per stage (Mitchell, 1970; Pittard and Mitchell, 1 972). A correlation between the population density and weather conditions of C. becki in the secondary dryland forest was statistically investigated with the linear correlation-test (Cavalli-Sforza, 1972), using the original field data. RESULTS AND DISCUSSION In total, 1 1 1 ricinuleids have been evaluated. Cryptocellus becki (Fig. 1 ) represented 85% of the total catch. About 79% of all ricinuleids came from the secondary forest 136 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) C ryptocellus becki TOO 60 20 -7 0 Cryptocellus odisi 0-3.5 3.5-70 cm 11111111 = larvae = adults L= larvae, P= protonymphs D = deutonymphs T = tntonymphs, A = adults Fig. 3. Distribution of Cryptocellus becki and Cryptocellus adisi in the soil according to soil depth, and percentage of all developmental stages in the secondary dryland forest at Rio Taruma Mirim near Manaus. (Total catch = 100% for each species.) Samples taken monthly at 0-3.5 and 3.5-7 cm depths between August, 1982 and August, 1983. N = total number of specimens. at Rio Taruma Mirim, where abundance was highest, with 36 ind./m^ for the larger species C. becki and 10 ind./m^ for the smaller C. adisi (Table 1). In the primary forest abundance was lower with 7 ind./m^ for C. becki and 5 ind./m^ for C. adisi. In both forest types ricinuleids represented fewer than 0.1% of all arthropods extracted from the soil during 13 months (Morais, 1985; Rodrigues, 1986). In the secondary forest at INPA, only C. becki was collected with 10 ind./m^ during the dry season and 38 ind./m^ during the rainy season (Adis et al., 1987 a, b). In the primary and secondary forests under study, ricinuleids were only found in the soil and never caught in traps on tree trunks (Adis and Schubart, unpubl.). No specimens were caught in emergence traps on the forest floor (see Penny and Arias, 1989 RICINULEI IN AMAZONIA 137 Table 1. Length of carapace and total length of Cryptocellus becki and Cryptocellus adisi according to developmental stages from primary and secondary dryland forests in the region of Manaus. Measurements (in mm) taken for all specimens caught per species (C. becki = 94; C. adisi = 17). Stage of development C. becki C. adisi Length of carapace Total length* Length of carapace Total length* Larvae 0.60-0.72 1.38-2.10 0.60-0.66 1.44-1.50 Protonymph 0.78-1.02 2.22-2.94 0.72^ 1.56-1.68 Deutonymph 1.08-1.26 3.06-4.32 0.84-0.90 1.98-2.52 Tritonymph (1.02)*’ 1.32-1.68 {2.%2f 4.44-5.22 1.02 2.64-3.00 Male (2.05)*^ 2.34-2.40 5.42-6.00 1.08-1.14 3.15-3.30 Female 1.80-(1.84)" 5.28^ 1.11-1.26 3.16-3.54 * Excluding pygidium. Literature data from the same habitat (Platnick and Shadab, 1977). Exceptional small specimens (cf. Fig. 4). Only one specimen sampled. 1982; Adis, unpubL), indicating that C. becki and C. adisi are not active on the soil surface. This conclusion is supported by another study in the primary forest, in which no ricinuleids were collected from 20 baited pitfall traps (see Penny and Arias, 1982). In all forests under study, most specimens of C. becki inhabited the organic layer (cf. Fig. 3: 0-3.5 cm), and a few the mineral subsoil. Almost 92% of all C. becki specimens obtained in the secondary forest at Rio Taruma Mirim were juveniles, with more than half representing the hrst instar, i.e., larvae (Fig. 3). Almost two thirds of these were collected during the rainy season (December-May). No adults were obtained during the dry season. C. adisi from the same habitat was mostly collected during the dry season (71% of the total catch; N = 50). Adults were only obtained from the organic layer (Fig. 3) and 73% of the juveniles inhabited the mineral subsoil (3.5-7 cm depth). The two sympatric Cryptocellus species appear to be sep- arated in their habitat by spatial and probably temporal differences. The smaller size of C. adisi (Table 1) certainly favors euedaphic life, i.e., inhabitation of lower soil layers, which was also reported for Palpigradi from the same forest (Rodrigues, 1986) and for Pseudoscorpiones and Symphyla from inundation forests (Adis and Mahnert, 1985; Adis and Scheller, 1984). Soil arthropods of dryland forests were shown to have no distinct reproductive period (Adis et al., 1988; Adis and Sturm, 1987). This also holds for ricinuleids, at least for C. becki, as larvae were found throughout the year (Fig. 4). Measurements based on carapace size are justihed, as increase of the somewhat varying body length per developmental stage is correlated with the more constant carapace length (Fig. 5), with exceptions in the size of some female tritonymphs disregarded (cf. Fig. 4). Distinct correlations between the population densities of C. becki and weather con- ditions in the secondary forest at Rio Taruma Mirim (see Rodrigues, 1986) could not be found. Only the larvae were somewhat more abundant during lower maximum temperatures of the air near the forest floor {P < 0.05; r > 0.5703, N = 13). No observations were made on the food spectrum of C. becki and C. adisi (cf. Cooke, 1967; Mitchell, 1970). An unexpected enemy of Central Amazonian ricinu- 138 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Fig. 4. Developmental stages of Cryptocellus becki extracted from soil samples of a secondary dryland forest at Rio Taruma Mirim near Manaus. Data arranged by length of carapace (1 gradation = 0.6 mm) and capture date. Samples taken monthly at 0-7 cm depth between August, 1982 and August, 1983. N = total number of specimens caught per month. Fig. 5. Relation between length of carapace and total length of body during post-embryonic growth of Cryptocellus becki (1 gradation = 0.6 mm). Seventy-one specimens extracted from soil samples taken monthly at 0-7 cm depth between August, 1982 and August, 1983 in a secondary dryland forest at Rio Taruma Mirim near Manaus (cf. Fig. 4). 1989 RICINULEI IN AMAZONIA 139 leids is Peripatus sp. (Onychophora). One was observed which covered a male of C becki with filaments from its mucus glands, killing the ricinuleids within a few hours. ACKNOWLEDGMENTS Dr. Vernon Thatcher kindly corrected the English manuscript, Ricardo H. M. Figueroa made the illustrations of C. becki and C adisi (Figs. 1, 2) and Jorge Soares Darcio (all of INPA, Manaus) made the drawings of Figures 3-5. Irmgard Adis is thanked for typing the manuscript. LITERATURE CITED Adis, J. 1981. Comparative ecological studies of the terrestrial arthropod fauna in Central Amazonian inundation-forests. Amazoniana 7(2):87-173. Adis, J. 1987. Extraction of arthropods from neotropical soils with a modified Kempson apparatus. J. Trop. Ecol. 3(2): 131-138. Adis, J. and V. Mahnert. 1985. On the natural history and ecology of Pseudoscorpiones (Arachnida) from an Amazonian blackwater inundation forest. Amazoniana 9(3): 297- 314. Adis, J. and U. Scheller. 1984. On the natural history and ecology of Hanseniella arborea (Myriapoda, Symphyla, Scutigerellidae), a migrating symphylan from an Amazonian black-water inundation forest. Pedobiologia 27(1):35-41. Adis, J. and H, O. R. Schubart. 1 984. Ecological research on arthropods in Central Amazonian forest ecosystems with recommendations for study procedures. Pages 1 1 1-144 in: J. H. Cooley and F. B. Golley (eds.). Trends in ecological research for the 1980s. NATO Conference Series, Series I: Ecology, Plenum Press, New York, London. Adis, J. and H. Sturm. 1987. On the natural history and ecology of Meinertellidae (Archaeog- natha, Insecta) from dryland and inundation forests of Central Amazonia. Amazoniana 10(2): 197-2 18. Adis, J., Y. D. Lubin and G. G. Montgomery. 1 984. Arthropods from the canopy of inundated and terra firme forests near Manaus, Brasil, with critical considerations on the pyrethrum- fogging technique. Stud. Neotrop. Fauna Environ. 19(4):223-236. Adis, J., J. W. de Morais and E. F. Ribeiro. 1987a. Vertical distribution and abundance of arthropods in the soil of a neotropical secondary forest during the dry season. Trop. Ecol. 28(1):1 74-181. Adis, J., J. W. de Morais and H. G. de Mesquita. 1 987b. Vertical distribution and abundance of arthropods in the soil of a neotropical secondary forest during the rainy season. Stud. Neotrop. Fauna Environ. 22(4): 189-1 97. Adis, J., V. Mahnert, J. W. de Morais and J. M. G. Rodrigues. 1988. Adaptation of an Amazonian pseudoscorpion (Arachnida) from dryland forests to inundation forests. Ecol- ogy 69(0:287-291. Cavally-Sforza, L. 1972. Grundziige biologisch-medizinischer Statistik. G. Fischer, Stuttgart, 209 pp. Cooke, J. A. L. 1967. Observations on the biology of Ricinulei (Arachnida) with descriptions of two new species of Cryptocellus. J. Zool. 151:31-42. Karmann, Ivo. 1986. Caracterizagao geral e aspectos geneticas da gruta arenitica “Refugio do Maroaga,” AM-2. Espeleo-Tema 15:9-18. Mitchell, R. W. 1970. Population size and dispersion and species associations of a Mexican cavernicole ricinuleid (Arachnida). Ciencia 27(2-3):63-74. Morais, J. W. de. 1985. Abundancia e distribui9ao vertical de Arthropoda do solo numa floresta primaria nao inundada. M.Sc. Thesis, INPA/Manaus (Brasil), 92 pp. Penny, N. D. and J. R. Arias. 1982. Insects of an Amazon Forest. Columbia University Press, New York, 269 pp. 140 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Pittard, K. and R. W. Mitchell. 1972. Comparative morphology of the life stages of Cryp- tocellus pelaezi (Arachnida, Ricinulei). Graduate Studies Texas Tech Univ. 1:1-77. Platnick, N. I. 1988. A new Cryptocellus (Arachnida: Ricinulei) from Brazil. J. New York Entomol. Soc. 96(3):363-366. Platnick, N. I. and M. U. Shadab. 1977. On Amazonian Cryptocellus {Avdichnid^., Ricinulei). Amer. Mus. Novit. 2633:1-17. Prance, G. T. 1975. The history of the INPA capoeira based on ecological studies of Lecy- thidaceae. Acta Amazonica 5(3):26 1-263. Ribeiro, M. de N. G. and J. Adis. 1984. Local rainfall variability— a potential bias for bioeco- logical studies in the Central Amazon. Acta Amazonica 14(1/2): 159-1 74. Rodrigues, J. M. G. 1986. Abundancia e distribuigao vertical de Arthropoda do solo, em capoeira de terra firme. M.Sc. Thesis, INP A/Manaus (Brazil), 80 pp. Received June 10, 1988; accepted October 12, 1988. J. New York Entomol. Soc. 97(2):141-150, 1989 TWO NEW SPECIES OF EOSENTOMON FROM CHICKASAW STATE PARK, TENNESSEE (PROTURA, EOSENTOMIDAE) William A. Outten' ^ and Robert T. Allen^ '(formerly) East Tennessee State University, Johnson City, Tennessee ^Department of Entomology, University of Arkansas, Fayetteville, Arkansas 72701 Abstract.— \ study of 140 soil and leaf litter samples from Chickasaw State Park, Hardeman County, Tennessee resulted in the collection of four previously described species of Protura and two new species. The previously described species were: Eosentomon montanum Copeland, E. pallidum Ewing, E. wheeleri Silvestri and Styletoentomon rostratum. The two new species are described and named Eosentomon chickasawensis and E. hunnicutti. During June of 1964 and 1965 Mr. George Hunnicutt collected 140 soil and leaf litter samples from Chickasaw State Park, Hardeman County, Tennessee. The sam- ples were processed through modified Berlese funnels and individuals of the order Protura were sorted from other arthropods and prepared for study. Several species of Protura were found in the samples. Among these were four previously described species, viz., Eosentomon montanum Copeland (1964), E. pallidum Ewing (1921), E. wheeleri Silvestri ( 1 909), and Styletoentomon rostratum Ewing ( 1 92 1 ). In addition, two forms were found which had not been recognized by Copeland (1964) and appear to be undescribed species. These two forms are described and named in this paper: Eosentomon chickasawensis and E. hunnicutti. The terminology used in describing these species agrees with that of Tuxen (1964) with the exception of PR, which was determined by dividing the length of the pseudoculus into the total length of the head including the labrum. Eosentomon chickasawensis, new species Figs. 1-10 Holotype. Female. Slide GSH 132-9: Chickasaw State Park, Tennessee, June 1965, G. Hunnicutt. Type Deposition: United States Natural History Museum, Washing- ton, D.C. Paratypes. 3356 2699. One specimen sent to each of the following collections and/ or individuals: American Museum of Natural History, New York, USA; Gentaro Imadate, Tokyo Medical and Dental University, Konodai College, Chiba, Japan; Yin Wen-Ying, Shanghai Institute of Entomology, Academia Sinica, Shanghai, China; S. L. Tuxen Collection, Zoologisk Museum, Universitetsparken 15DK 2100 Copen- hagen, Denmark; 55 paratypes retained in the University of Arkansas Insect Collec- tion, Entomology Department, University of Arkansas, Fayetteville, Arkansas, USA. Etymology. The species is named for the area in which it was collected, Chickasaw State Park, Tennessee. Description. HEAD: Head capsule slightly longer than wide (109:97 microns). 142 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Labrum (Fig. 1)13 microns long, with a broad, apical, V-shaped notch and a semilu- nar shaped apodeme proximal to the notch. Labral setae present. Rostral setae I winged basally, extending to the tip of the labrum. Rostral setae III shorter than rostral setae I (10:14). Mandibles with three teeth each. Lacinia I and II hooked distally. Clypeal apodemes prominent and connected anteriorly. Tentorium distinct, maxillary ramus broader than the cardo. Head ratios: PR 245:13 = 18.8; LR 218: 27 = 8.0. THORAX: Mesothoracic seta P 1 (Fig. 2) inserted on the sclerotized portion of the notum and shorter than the distance to its corresponding primary (13:23) on the opposite side. Seta P P inserted on the unsclerotized portion of the notum and slightly longer than its corresponding primary (15:13). The terminal bulbs of flamento di sostegno in the prothorax. TARSI: Foretarsus (Figs. 3, 4) short and broad, 78 microns long and 23 microns wide at broadest part. Sensillum t 1 (Fig. 3) possessing a large club, short shank, and inserted slightly proximal to a 3'. Sensillum t 2 lanceolate or very narrowly clavate, with a long shank and inserted on the level of a 4. Sensillum a' broad, lanceolate- clavate, its length less than the distance to r 2 ( 1 0: 1 4). Sensillum b' 1 absent. Sensillum b' 2 lanceolate, inserted on the level of y and extending to the insertion of 1 3. Sensillum 5 with a small club, less than twice the width of its shank. Empodium subequal to the unguis and slightly clavate. Sensillum e (Fig. 4) inserted on the level of (3 6 and with a huge club longer than its shank. Sensillum g inserted proximal to (3 ^ and in line between (3 8 and y 4, possessing a huge club that is longer than the shank. Sensillum/ 1 setiform. Sensillum a distinct and extending two-thirds the distance to y 2. Sensillum b short but length equalling the distance to d 6. Sensillum c with a long slender club only slightly broader than its shank and its tip on the level of pit II. Tarsal ratios: EU 27:31 = 0.87; BS 90:64 = 1.40. Metathoracic tarsus (Fig. 5) possessing a short empodium and the unguis with no teeth. ABDOMEN: Posterior row of setae on tergum I (Fig. 6) with three primary setae, one accessory seta, and one microchaeta on each side. Setae P 1 on tergum IV (Fig. 7) much shorter than P P (31 :49). Setae P P on tergum VII (Fig. 8) filamentous and inserted midway between a line drawn connecting the p 1-P 2 and the posterior edge of the tergum. Sternum VIII (Fig. 9) possessing two anterior and seven posterior setae. Sterna IX and X (Fig. 9) with six setae each. The abdominal chaetotaxy is shown schematically in Table 1 with the figure above the dash representing the anterior row and the figure below the dash representing the posterior row. GENITALIA: The female squama genitalis (Fig. 10) shows no unusual features. Discussion. Twenty-one females and thirty-two males were examined. No variation was found in: (a) the 2-7 arrangement of setae on sternum VIII, (b) the length of the Figs. 1-4. Eosentomon chickasawensis. 1. Labrum and anterior portion of head. 2. Tarsus 111. 3. Foretarsus (dorsal view); setae and sensilla as indicated. 4. Foretarsus (ventral view); setae and sensilla as indicated. 1989 NEW PROTURA 143 144 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) 8 1989 NEW PROTURA 145 Figs. 9, 10. Eosentomon chickasawensis. 9. Sterna VIII-XII; SMG, sternal microgland pores. 10. Female genitalia. primary setae much less than that of the accessory setae on tergum IV, (c) sensillum t 1 inserted a little proximal to a i', (d) length of sensillum a' less than the distance to the insertion of t 2, and (e) sensillum / 1 setiform. Nine individuals were oriented in such a manner that it could not be determined whether rostral setae I were winged or not. This species possesses two anterior and seven posterior setae on sternum VIII. This places it in the wheeleri group of Bonet and Tuxen (1960). It also belongs to the 3:1:1 subgroup proposed by Copeland (1962) because of the presence of three primary setae, one accessory seta and one microchaeta on each side in the posterior setal row on tergum I. Within the 3:1:1 subgroup E. chickasawensis and the other new species described in this paper, E. hunnicutti, seems to be most closely related to E. pussilum Ewing. This new species differs from E. pusillum in that sensillum t 1 is inserted closer to a3' than to a2>. Distribution. Known only from the type locality, Chickasaw State Park, near Bo- livar, Hardeman County, Tennessee. Figs. 5-8. Eosentomon chickasawensis. 5. Mesothorax; A1-A4, setae of the anterior row; P1-P3, setae of the posterior row. 6. Abdominal tergum I; A1-A2, setae of the anterior row; P1-P3, setae of the posterior row; M, microchaeta. 7. Abdominal tergum IV; A1-A5, setae of the anterior row; P1-P5, setae of the posterior row; TMG, tergal microgland pore. 8. Abdominal tergum VII; A2-A5, setae of the posterior row; TMG, tergal microgland pore. 146 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Table 1. Schematic representation of the abdominal chaetotaxy of Eosentomon chickasaw- ensis. Abdomen I II III IV V VI VII VIII IX X XI 4 10 10 10 10 10 6^ 6 10^ 16 16 16 l6 Tb l6 9 ^ ^ ^ 46666662 444ToToToTo7^^^ Three primary setae, one accessory seta, and one microchaeta on each side. Setae A 1 and A 3 missing. Tergum Sternum XII 9 12 Eosentomon hunnicutti, new species Figs. 11-20 Holotype. Female. Slide GSH 1 10-9. Chickasaw State Park, Tennessee, June 1965. Type Deposition: United States Natural History Museum, Washington, D.C. Paratypes. 1 166, 1 899. One specimen sent to each of the following collections and/ or individuals: American Museum of Natural History, New York, USA; Gentaro Imadate, Tokyo Medical and Dental University Kandai College, Chiba, Japan; Yin Wen-Ying, Shanghai Institute of Entomology, Academia Sinica, Shanghai, China; S. L. Tuxen Collection, Zoologisk Museum, Universitetsparken, 15DK, 2100 Copen- hagen, Denmark; 30 paratypes retained in the University of Arkansas Insect Collec- tion, Entomology Department, University of Arkansas, Fayetteville, Arkansas, USA. HEAD: Head capsule 103 microns long and 75 wide. Labrum (Fig. 11)10 microns long with a broad V-shaped notch at its tip and a subterminal semilunar shaped apodeme. Labral setae present. Rostral setae I winged and slightly longer than the labrum. Rostral setae III shorter than rostral setae I (8:1 1 microns). Each mandible possessing three teeth. Clypeal apodemes prominent and connected anteriorly. Parts of the tentorium not distinct, cardo not as broad as the maxillary ramus. Head ratios: PR 225:14 = 16.1; LR 205:20 = 10.3. THORAX: Mesothoracic seta P 1 (Fig. 12) length less than the distance to its homologue on the opposite side (15.25) and shorter than P P. The latter inserted in the unsclerotized posterior portion of the notum. Terminal bulbs of the filamento di sostegno in the prothorax. TARSI: Foretarsus (Figs. 13, 14) 75 microns long and 23 microns wide at the broadest point. Sensillum / 1 (Fig. 13) inserted proximal to seta a 3' but much closer to it than to a i and possessing a large club with short shank. Sensillum t 2 narrowly lanceolate with the length of the shank less than that of the blade and inserted on Figs. 11-15. Eosentomon hunnicutti. 11. Labrum and anterior portion of head. 12. Me- sothorax; A1-A4, setae of the anterior row; P1-P3, setae of the posterior row. 13. Foretarsus (dorsal view); setae and sensilla as indicated. 14. Foretarsus (ventral view); setae and sensilla as indicated. 15. Tarsus III. 1989 NEW PROTURA 147 148 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Figs. 16-18. Eosentomon hunnicutti. 16. Abdominal tergum I; A1-A2, setae of the anterior row; P1-P3, setae of the posterior row; M, microchaeta. 17. Abdominal tergum IV; A1-A5, setae of the anterior row; P1-P5, setae of the posterior row; TMG, tergal microgland pore. 18. Abdominal tergum VII; A2-A5, setae of the anterior row; P1-P5, setae of the posterior row; TMG, tergal microgland pore. the level of seta a 4. Sensillum a' broadly lanceolate, length equalling the distance to sensillum t 3 (12:12 microns). Sensillum b' 1 absent. Sensillum b' 2 lanceolate and inserted on the level of 7 3. Sensillum / 1 very narrowly clavate-lanceolate. Club of sensillum .s large and approximately twice the width of the shank. Sensillum e (Fig. 14) inserted on the level of seta (3 6, possessing a large pointed club that is slightly longer than the shank. Sensillum g clavate with club longer than its shank and inserted proximal to 8. Tarsal ratios: BS 81:65 = 1.24; EU 27.31 = 0.87. 1989 NEW PROTURA 149 Figs. 19, 20. Eosentomon hunnicutti. 19. Sterna VIII-XII; SMG, sternal microgland pores. 20. Female genitalia. Metathoracic tarsus (Fig. 15) possessing a short empodium and the unguis with no teeth. ABDOMEN: The posterior row of setae on tergum I (Fig. 16) with three primary setae, one accessory seta, and one microchaeta on each side. Setae P 1 on tergum IV (Fig. 17) shorter than the P V (18:25). Setae P P on tergum VII (Fig. 18) filamentous and inserted midway between a line drawn connecting setae P 1 and P 2 and the posterior margin of the tergum. Two anterior and seven posterior setae on sternum VIII (Fig. 19). Sterna IX and X (Fig. 19) possessing six setae each. The abdominal chaetotaxy is shown schematically in Table 2. GENITALIA: The female squama genitalis is represented in dorsal view in Figure 20. It shows no unusual features. Table 2. Abdomen Tergum Sternum Schematic representatation of the abdominal chaetotaxy of Eosentomon hunnicutti. i ii m iv v vi vii vin ix x xi 4 10 10 8‘’ 8^* 6^ 6 T^T6l6T6T6T6T6 9 46666662 4 4 4 10 10 10 10 7 8 8 8 9 6 6 8 12 Three primary setae, one accessory seta, and one microchaeta on each side. Setae A 3 missing. Setae A 1 and A 3 missing. 150 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Discussion. Eighteen females and seventeen males were examined and no variations were found in: (a) the 3:1:1 and 2-7 arrangements of setae on tergum I and sternum VIII respectively, (b) the relative lengths of P 7 and P V on tergum IV, (c) the insertion of sensillum t 1 slightly proximal to seta a 3', (d) sensillum a' length never greater than the distance to sensillum 1 2 and, (e) sensillum ^always inserted proximal to (3 8. The terminal bulbs of the filamento di sostegno are in the prothorax of 33 specimens and near the anterior margin of the mesothorax in two specimens. These apparent discrepancies were probably caused by the two individuals being squashed in the mounting process. Rostral setae I are longer than their respective rostral setae III and winged in all but three individuals. The orientation of these three setae makes it impossible to determine if they are winged or not. The club of sensillum g on the holotype was positioned in a manner which made it impossible to show its shape; consequently, it has been drawn as it appeared on other individuals of this group. No setal abnormalities were noted. Eosentomon hunnicutti is a member of the wheeleri group of Bonet and Tuxen (1960), and the 3:1:1 subgroup proposed by Copeland (1962). It is closely related to E. chickasawensis, differing chiefly in setae number in the anterior rows of terga IV- VI, E. chickasawensis having ten and E. hunnicutti eight setae per row per segment. Distribution. Known only from the type locality, Chickasaw State Park, near Bo- livar, Hardeman County, Tennessee. ACKNOWLEDGMENTS The senior author would like to express appreciation to the members of his graduate com- mittee, Dr. Wallace A. Tarply, Dr. J. D. Moore, and Dr. T. P. Copeland. Appreciation is also extended to Mr. George Hunnicutt for use of the specimens. The junior author wishes to express his thanks to Dr. T. P. Copeland for the opportunity to participate in this project. The manuscript was reviewed by J. R. Phillips, A. J. Mueller and D. T. Johnson, Entomology Department, University of Arkansas. Published with the approval of the Director, Arkansas Agricultural Experiment Station, Fayetteville, Arkansas 72701. Manuscript originally submitted in partial fulfillment of the requirements for the Degree of Master of Science, East Tennessee State University, Johnson City, Tennessee, directed by Dr. T. P. Copeland. LITERATURE CITED Bonet, F. and S. L. Tuxen. 1960. Re-examination of species of Protura described by H. E. Ewing. Proc. U. S. Nat. Mus. 112:265-305. Copeland, T. P. 1962. A taxonomic treatment of Eosentomon Berlese (Protura) of East Ten- nessee. Unpublished dissertation. University of Tennessee, 160 pp. Copeland, T. P. 1964. New species of Protura from Tennessee. J. Tenn. Acad. Sci. 39:17- 29. Received July 1, 1988; accepted October 25, 1988. J. New YorkEntomol. Soc. 97(2): 15 1-1 58, 1989 NEW SPECIES, REDESCRIPTIONS, AND CLADISTICS OF THE GENUS PSEUDOCENTROPTILOIDES (EPHEMEROPTERA: BAETIDAE) R. D. Waltz' and W. P. McCafferty^ 'Division of Entomology, Indiana Department of Natural Resources, 613 State Office Building, Indianapolis, Indiana 46204 ^Department of Entomology, Purdue University, West Lafayette, Indiana 47907 Abstract.— Two new species of the baetid genus Pseudocentroptiloides Jacob are described: P. usa from the Nearctic, and P. christineae from the Orient. The genus and its type species P. shadini (Kazlauskas) are redescribed in a comparative format, and larval keys to species are provided. The Oriental species P. christineae and P. ceylonica Glazaczow are sister species with several synapomorphies. The Palearctic species P. shadini is the most closely related to this pair, sharing certain other apomorphies with them. The most basally derived species is P. usa. Neither cladistic nor phenetic relationships of species support subgeneric classification as pre- viously proposed. Larvae of the genus Centroptilum Eaton are poorly known in North America, as only seven of the 25 recognized species are known as larvae. Comparative larval descriptions are nonexistent, rendering specific diagnosis of this stage virtually im- possible. A review of Nearctic larvae that have been placed in Centroptilum, together with larva to adult rearings conducted at Purdue University, revealed a new genus and species distinct as both larva and adult from Centroptilum and other Baetidae. We then found that certain larvae collected by G. F. and C. H. Edmunds in Maylasia represented an additional species belonging to this new genus. Meanwhile, the genus Pseudocentroptiloides i?iCob (Jacob and Glazaczow, 1986) was established, based on a Palearctic species and a new Oriental species. It was apparent when the description became available in 1987 that our distinctive new material was assignable to that genus. We here describe our two new species. Also, as part of our ongoing revision of baetid genera, we comparatively redescribe Pseudocentroptiloides and its type species P. shadini (Kazlauskas), provide larval keys to all species, and discuss cladistic re- lationships within the genus. Jacob and Glazaczow (1986) and Keffermiiller and Sowa (1984) (as Centroptilum) presented useful figures of Pseudocentroptiloides. We include citations of these as JG and KS respectively in our descriptions. Pseudocentroptiloides Jacob Description. Larva. Labrum (Figs. 1,13; KS 29; JG la, 2a) deeply and triangularly notched on anterior margin. Mandibles (Figs. 2, 3, 14, 15; KS 30; JG 2b, c) with incisors widely separated apically; tuft of setae present between prostheca and molar region; thumb of left mandible parallel to slightly elevated above plane of incisor bases. Hypopharynx as in Figures 19; KS 32; JG lb, 2d. Maxillae (Figs. 4, 16, 17; 152 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) KS 31; JG 2e) with broadened galealacinea and prominent crest of setae proximal to outer margin of apical denticles. Labium (Figs. 5, 18; KS 33; JG Ic, 3) with three- segmented palps; terminal segment of palps very broad distally and truncate; palpal segment 2 without inner apical lobe; glossae truncate distally, with numerous setae, and shorter than paraglossae; paraglossae truncated distally, with numerous setae apically, and with subparallel margins. Margins of femora (Figs. 6, 7; JG 4) subparallel, without ventral femoral patch and without long, dorsal bristles; stout ribbed bristles sometimes present ventrally and laterally; vertical row of long, hne setae (Fig. 6) present on posterior face. Tibiae (Figs. 6, 7) with subproximal arc of long, hne setae. Claws (Figs. 6, 7, 20; KS 28; JG 4) edentate and elongate, 0.5 x or more length of respective tarsi. Abdominal terga (Fig. 21) with hne setae and broadly pointed, cresentic based scales with median length subequal to basal width; posterior marginal spines (Figs. 21; JG 7) present on all terga, alternately large and small in series, widely spaced and spike-like. Sternal surfaces with hne setae and scales similar to tergal scales. Pleural spines as in Figure 22. Gills (Figs. 23; KS 27; JG 5) 1-7 asymmetric, ovate and simple in presently included species. Median terminal hlament subequal to cerci. Adult Male. Forewings (Fig. 8) with single marginal intercalaries and relatively few crossveins. Hindwings (Figs. 9; KS 25) elongate, fore and hind margins subparallel or somewhat broadened medially and with distinctly hooked costal process. Genitalia (Figs. 1 2; KS 26; JG Id) with forceps segments 2-3 fused, appearing three segmented; (in presently included species, forceps with distinct inner tubercle on segment 2); penes plate between forceps bases broad and battened apically. Included species. Pseudocentropti/oides usa (Nearctic), P. christineae (Oriental), P. shadini (Kazlauskas 1964) (type species by original designation) (Palearctic), and P. ceylonica Glazaczow (Oriental). Pseudocentroptiloides usa, new species Diagnosis. Larva. Body length 7-8 mm. Terminal hlaments ca. 2 mm. Antennae extend to middle coxae. Labrum (Fig. 1) slightly wider than deep. Mandibles (Figs. 2, 3) with relatively broad bases (compared to P. ceylonica). Maxillae (Fig. 4) with crest of hne setae restricted to apical width of galealacineal crown and shorter in height than galealacineal denticles; broad, digitate setae of galealacineal comb not present. Maxillary palp three segmented. Labium as in Figure 5. Claws (Figs. 6, 7) subequal in length to respective tarsi. Adult male. Body length ca. 5 mm. Forewing (Fig. 8) length 4.5 mm. Hindwings (Fig. 9) with fore and hind margins subparallel. Turbinate eyes (Fig. 10), yellow to yellow-orange, slightly divergent anteriorly, and on low stalks. In dorsal view, tur- binate eyes (Fig. 11) ca. 2.0 x longer than wide, hemispherical with medial margin relatively straight. Genitalia as in Figure 12. Holotype. USA, Indiana, Pulaski Co., Tippecanoe River at Co. Rd. 1.5 mi S Tippecanoe River St. Prk., VI-30- 1978, A. V. Provonsha, D. Bloodgood, H. Hollis, male larval exuviae (slidemounted in euparal; solvent abs. ale.) and its reared male adult (body stored in alcohol; forewing, hindwing, foreleg, and genitalia slidemounted on one slide; foreleg and genitalia in euparal). Paratypes. USA, Indiana, Pulaski Co., Tippecanoe River at Co. Rd. 1.5 mi S Tippecanoe River St. Prk., VI-30- 1978, A. V. Provonsha, D. Bloodgood, and H. 1989 PSE UDOCENTROPTILOIDES 153 Figs. 1-7. Pseudocentroptiloides usa, larva. 1. Labrum (left side, ventral; right side, dorsal). 2. Left mandible. 3. Right mandible. 4. Maxilla (cr— crest of setae; cmb— galealacineal comb; apd— apical denticles). 5. Labium. 6. Foreleg. 7. Flindleg. 154 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Figs. 8-12. Pseudocentroptiloides usa, male imago. 8. Forewing. 9. Hindwing. 10. Turbinate eyes, lateral view. 1 1. Turbinate eyes, dorsal view. 12. Genitalia, ventral view. Hollis. 1 female exuviae (slidemounted as above) and its reared female adult (in alcohol); 3 larvae, same locale as holotype, VHI-4-1976, A. V. Provonsha and M. Minno. Holotype and paratypes deposited at Purdue University Entomological Re- search Collection, West Lafayette, Indiana. Etymology. The species epithet, usa, is an arbitrary combination of letters based on the acronym for the United States of America. Pseudocentroptiloides shadini (Kazlauskas) Pseudocentroptilum shadini Kazlauskas, 1964. Centroptilum shadini, Keffermiiller and Sowa, 1984. Pseudocentroptiloides shadini, Jacob and Glazaczow, 1986. Diagnosis. Larva. Body length 5-6 mm. Antennae elongate, reaching to middle coxae. Labrum (hgs. KS 29; JG ia) slightly wider than deep. Hypopharynx as in ngs. KS 32 and JG lb. Mandibular incisors (hg. KS 30) broad based (compared to P. christineae). Maxillae (hg. KS 31) with crest of hne setae extended beyond ga- 1989 PSE UDOCENTROPTILOIDES 155 Figs. 13-15. Pseudocentroptiloides christineae, larva. 13. Labrum (left side, ventral; right side, dorsal). 14. Left mandible. 15. Right mandible. lealacineal crown, and longer than galealacineal denticles; broad, digitate setae of galealacineal comb present and well developed. Maxillary palp three segmented, segment 3 poorly differentiated. Labium as in hgs. KS 33 and JG Ic. Claws (fig. KS 28) subequal in length to, or longer than, respective tarsi. Adult male. Body length 5. 0-5. 5 mm, cerci ca. 9 mm. Turbinate eyes, yellow- orange, slightly divergent anteriorly; oval, ca. 1 .5 x longer than broad, in dorsal view. Hindwings (fig. KS 25) concave from wing base to costal process, convex from costal process to wing apex. Forceps (figs. KS 26; JG Id) with segment 1 broad, slightly tapered distally; segments 2 and 3 fused, with prominent inner tubercle at base and distally swollen; segment 4 broad apically with narrow base. Distribution. USSR (Oka River) and Poland (Warta River and Bug River) (Kef- fermiiller and Sowa, 1984). Pseudocentroptiloides christineae, new species Diagnosis. Larva. Body length 4-5 mm. Length of terminal filaments 1.0- 1.5 mm. Antennae shortened (compared to P. shadini and P. usa), extend to vertex of head capsule. Labrum (Fig. 1 3) elongated, ca. 2.0 x deeper than wide. Mandibles (Figs. 14, 15) with incisors narrowly based (compared to P. shadini and P. usa). Maxillae (Figs. 16, 17) very broad, subquadrate, with crest of fine setae distinctly longer than galealacineal denticles; broad, digitate setae of galealacineal comb highly developed and apparent. Maxillary palp three segmented, segment 3 poorly differentiated. La- bium as in Figure 18. Hypopharynx as in Figure 19. Claws (Fig. 20) shorter in length than respective tarsi, ca. 0.5 x length of tarsi. Hindwing pads absent (as in fig. JG 6a). Adult. Unknown. Holotype. West Malaysia, Trengganu Kampong Sungai Tong, 9-IX-1978, G. F. and C. H. Edmunds, male larva, in alcohol. Holotype deposited at the Purdue Uni- versity Entomological Research Collection (PERC), West Lafayette, IN. 156 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Figs. 16-23. Pseudocentropti/oides christineae,\sLTYa. 16. Maxilla. 17. Maxilla enlarged (cr— crest setae; ds— digitate setae). 18. Labium (dorsal view). 19. Hypopharynx. 20. Claw. 21. Tergal surface. 22. Ventral abdomen (p— paraprocts; pis— pleural spines). 23. Gill (Abd. 4). Figs. 16, 18-20, 22, 23 scale bar = 100 )u; Figs. 17, 21 scale bar = 10 /i. 1989 PSE UDOCENTROPTILOIDES 157 Paratypes. West Malaysia, Trengganu Kampong Sungai Tong, 9-IX-1978, G. F. and C. H. Edmunds, 3 larvae: 1 (in alcohol) deposited at PERC (mouthparts slide- mounted); 1 (on SEM stub used for SEM studies) deposited at PERC; 1 (in alcohol) deposited United States National Museum. Etymology. This species is named in honor of Christine Edmunds, who along with her husband, the renowned ephemeropterist George F. Edmunds, Jr., collected the material upon which this species is based. LARVAL KEY TO THE SPECIES OF PSEUDOCENTROPTILOIDES 1. Labrum (Fig. 13) elongated, elearly longer than wide and deeply notched distally; maxillae with crest of setae highly developed as in Figures 16, 17 2 1 ' Labrum (Fig. 1) length subequal to width and not notched as deeply as above; maxillae with crest of setae not so prominently developed (Fig. 4) 3 2. Maxillae quadrate (Fig. 16), galealacineal width subequal to length. Crest of setae on maxillae strongly hooked distally (Fig. 17) P. christineae T Maxillae not so quadrately formed as above (fig. JG 2e). Crest of setae on maxillae not hooked distally P. ceylonica 3. Maxillae (Fig. 4) with crest of setae restricted to crown of galealacinea, shorter in length than denticles of galealacinea; broad, digitate setae of galealacineal comb not developed P. usa 3' Maxillae (fig. KS 3 1 ) with crest of setae extending basally beyond crown of galealacinea, subequal to or longer than denticles of galealacinea; broad, digitate setae of galealacineal comb well developed P. shadini DISCUSSION Pseudocentroptiloides is most similar to complexes within the large genus Cen- troptUum, as the latter is currently composed. Centroptilum is used as the outgroup for proposing apomorphies for cladistic analysis. Thus, the apomorphic character states common to all Pseudocentroptiloides and its hypothetical common ancestor are 1) labrum triangularly and deeply notched, 2) maxillae broadened, 3) prominent crest of setae present on the crown of the galealacinea, 4) segment 3 of labial palps greatly broadened, 5) glossae truncate and shorter than paraglossae, 6) paraglossae truncate, subparallel, 7) glossae and paraglossae with numerous apical setae, 8) claws edentate, and 9) claws subequal to or greater than 0.5 x length of respective tarsi. The above synapomorphies clearly show Pseudocentroptiloides to be monophyletic. We propose cladistic interrelationships of Pseudocentroptiloides species based on several additional larval apomorphies and one adult apomorphy. The earliest derived species within the group is P. usa, which remains essentially similar to the hypothetical ancestor of the genus. The remaining three species share further apomorphies of increased development of the maxillary crest, extension of the line of crown setae basally on the galealacinea, and a galealacineal comb consisting of digitate setae. The ancestor with these character states gave rise to one lineage leading to P. shadini and one leading to the common ancestor of P. ceylonica and P. christineae. These sister species share apomorphic elongation of the labrum and a more deeply recessed labral notch, further elongation of the crest of setae of the maxillae, and loss of the hind wings (wingpads in larvae). Jacob and Glazaczow (1986) proposed two subgenera for the two species of Pseu- 158 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) docentwptiloides known to them based only on the presence or absence of hindwings. Ironically, in this same paper these authors rejected this character for generic criteria, citing observations of its taxonomic unreliability by Edmunds et al. (1976). Although we have presented characters, including the loss of hindwings, that show P. ceylonica along with P. christineae to be a derived sister pair, the intermediate phyletic position of P. shadini between them and P. usa, and the gradational differences within the genus, do not support a subgeneric division. ACKNOWLEDGMENTS We sincerely thank Dr. George F. Edmunds, Jr., University of Utah, for his encouragement in our studies of Baetidae and his most generous loan of material he has personally collected from throughout the world and has recognized as systematically significant. We also thank Dave McShaffery and Arwin Provonsha, Purdue University, for assistance with SEM work and illustrations, and E. R. Hoebecke, Cornell University, and Lewis Berner, University of Florida, for loan of Centroptilum type material. Research funding was in part provided by National Science Foundation grant PCM-8400133. This paper is Purdue University Experiment Station Journal No. 1 1745. LITERATURE CITED Edmunds, G. F., Jr., S. L. Jensen and L. Berner. 1976. The Mayflies of North and Central America. Univ. Minn. Press, Minneapolis, 330 pp. Jacob, U. and A. Glazaczow. 1986. Pseudocentroptiloides, a new baetid genus of Palearctic and Oriental distribution (Ephemeroptera). Aq. Insects 8:197-206. Kazlauskas, R. S. 1964. Materialy k poznaniju podenok reki Oki. Trudy Zool. Inst. Akad. Nauk USSR. 32:164-176. Keflermuller, M. and R. Sowa. 1984. Survey of central European species of the genera Cen- troptilurn Eaton and PseudocentroptUum Bogoescu (Ephemeroptera, Baetidae). Polsk. Pismo Entomol. 54:309-340. Received July 21, 1988; accepted September 7, 1988. J. New York Entomol. Soc. 97(2): 159-1 66, 1989 REVIEW OF DALEAPIDEA KNIGHT (HETEROPTERA: MIRIDAE: ORTHOTYLINAE: ORTHOTYLINI) Randall T. Schuh Department of Entomology, American Museum of Natural History, New York, New York 10024 Abstract.— A revised diagnosis is provided for Daleapidea Knight. A key, diagnoses, distri- butional data, and illustrations of the male habitus and genitalia are provided for the three species currently placed in the genus. Hadronema decorata (Uhler) is transferred to Daleapidea and a lectotype is designated. Knight (1968) described the genus Daleapidea to include two species, albescens (Van Duzee) and daleae Knight. Recent collecting and examination of existing col- lections reveal that Hadronema decorata Uhler also belongs to this colorful group of Orthotylini which breeds on the leguminous plant genus Psorothamnus in the American southwest and Baja California. Daleapidea Knight Daleapidea Knight, 1968:101 (n. gen., key). Type species: Daleapidea daleae Knight. Diagnosis. Orthotylinae: Orthotylini: Recognized by the structure of the fore tibia, which is flattened distally and covered with thickly set short setae on the ventral surface of the flattened area (Figs. 4A, B), the first antennal segment which is relatively long and inflated subbasally (Figs. 1-3), the rather strong sexual dimorphism with the males elongate and slender and the females much more robust, and the structure of the male genitalia (Fig. 5), with two vesical spines, the longer with two recurved branches, the shorter simple, and the hatchet-shaped right paramere. All three species are strikingly colored, and the males, particularly of daleae, are wasp-like in their movements. Discussion. Knight ( 1 968) in his description and key related Daleapidea to Lopidea Uhler and Lopidella Knight based on the presence of a “suture extending down from the base of the jugum to a point near middle of gena.” It is not clear whether the structure Knight referred to is actually a suture, and it furthermore appears that it occurs widely outside of Lopidea and its close relatives. My examination of the external morphology and male and female genitalia, suggest that Daleapidea is cer- tainly a member of the Orthotylini; however, its relationship to Lopidea or Hadro- nema (in which two of the species were originally described) is not evident. I have found no other generic grouping in North America that shares any unique characters with Daleapidea that are not also shared with some more inclusive taxon, and suggest further searching be pursued in the relatively poorly studied faunas of Mexico and South America. 160 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Daleapidea albescens (Van Duzee). Fig. 2. Daleapidea daleae Knight. Fig. 3. Daleapidea decorata (Uhler). 1989 REVIEW OF DALEAPIDEA 161 Fig. 4. Distoventral view of foretarsus of female Daleapidea decorata. A. Overall view. B. Detail of tibial comb and modified setae. KEY TO SPECIES OF DALEAPIDEA KNIGHT 1. Dorsum charcoal with white cuneus and white longitudinal markings along claval suture; scutellum bright red; all appendages charcoal; male genitalia as in Figures 5I-L decorata - Dorsum never largely charcoal; scutellum sometimes orange; appendages either char- coal or in part lighter in coloration 2 2. All appendages charcoal; scutellum gray with some charcoal markings on anterior margin; antennal segment 2 distinctly clavate in males; male genitalia as in Figures 5E-H daleae - All femora and scutellum with some orange coloration, dorsum largely pale; antennal segment 2 not distinctly clavate in males; male genitalia as in Figures 5A-D . . . albescens Daleapidea albescens (Van Duzee) Figs. 1, 5A-D, 6 Hadronema albescens Van Duzee, 1918:297 (n. sp., host). Daleapidea albescens Knight, 1968:102 (distr., hosts). Holotype. $, Palm Springs, Calif., 50-21-17, EP Van Duzee collector; deposited in the CAS. Diagnosis. Recognized by the generally pale dorsum with an orange scutellum and the endocorium infuscate posteriorly, the orange femora (Fig. 1), and the structure of the male genitalia (Fig. 5 A-D), in which the left paramere has an elongated sensory lobe bearing bristle-like setae arranged in a cluster and a lateral process with a hammerhead-like apex. Distribution. See Figure 6. Hosts. Psorothamnus emoryi, P. polydenius. Specimens examined: Same data as holotype (CAS), 11(5(5, 1229. MEXICO: Baja California Norte: San Luis, April 1889 (Haines; CAS), 19; 26 mi S of San Felipe, 15 April 1965 (Ross et ah; CAS), 2(5(5, 492. USA: Arizona: Yuma Co.: Ligurta, 8 April 1942 (Stitt; OSU), 1(5, 19. California: Imperial Co.: 5.4 mi NW of Ocotillo on 52, 23 April 1980, ex Psorothamnus emoryi (Russell and Schwartz; AMNH), 10(5(5, 699. Inyo Co.: Antelope Springs, 14 June 1961 (Toschi; CAS, UCB), 1(5, 399; 21.7 mi E 162 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Fig. 5. Male genitalia of Daleapidea. A-D. albescens. A. Left paramere. B. Right paramere. C. Large spiculum. D. Small spiculum. E-H. daleae. E. Left paramere. F. Right paramere. G. Large spiculum. H. Small spiculum. I-L. decorata. I. Left paramere. J. Right paramere. K. Small spiculum. L. Large spiculum. of Rt 395 on Westgard Pass Road, ca 1 560 m, 2 July 1 980, ex Psorothamnus polydenius (Schuh; AMNH), 21(56, 30$$; Deep Spring Lake Flats on Rt 168, 1644 m, 12 July 1980, ex Psorothamnus polydenius (Schuh and Stonedahl; AMNH), 366, 12$$; Deep Spring Lake Flats, on Hwy 168, 5260 ft, 12 July 1980 (Stonedahl; AMNH), 1266, 19$$. Marin Co. ?: Mill Valley, May 9, 1927 (Van Dyke; CAS), 16. Mono Co.: 6 mi N of Bishop, fish slough, 15 June 1973 (Pinto; AMNH), 266, 1$; Benton Hot Springs, 1989 REVIEW OF DALEAPIDEA 163 8 June 1966 (Gagne; UCB), IS. Riverside Co.: Blythe, 24 April 1939 (Bliven; CAS) 2SS, 12; Palm Springs, 18-21 May 1917 (Van Duzee; CAS), 29$S, 1822; Coachella Valley, 9 May 1927 (Van Dyke; CAS), 322. San Diego Co.: Anza-Borrego State Park, Carrizo Creek, 10.2 mi NW of Ocotillo on 52, 23 April 1980, ex Psorothamnus emoryi (Russell and Schwartz; AMNH), 366, 222; Borrego, 28 April 1955 (Schuster; UCB), 16. Nevada: Lyon Co.: Yerington, 9 July 1909, 5100 ft (Baumberger; CAS), 16. Nye Co.: Nevada Atomic Test Site, Rock Valley on Jackass Flats Rd, 3300 ft (A25), 6 June 1 983, ex mercury vapor lamp (Schuh, Schwartz, and Stonedahl; AMNH), 266; Nevada Atomic Test Site, 7.5 mi W of Mercury Hwy on Cane Springs Rd, 3800 ft (A26), 6 June 1983, ex Psorothamnus polydenius (Schuh, Schwartz, and Stonedahl; AMNH), 1266, 4622; Beatty, 23 June 1967, at night (Gagne; UCB), 422. Discussion. The Mill Valley record, the label for which did not explicitly indicate a county, is almost certainly in error, or it represents another “Mill Valley” not indicated on maps available to me. Daleapidea daleae Knight Figs. 2, 5E-H, 6 Daleapidea daleae Knight, 1968:102 (n. sp., hosts). Holotype. Not examined; deposited in the USNM. Diagnosis. Recognized by the generally gray coloration with contrasting charcoal appendages, head, and calli, and the clavate second antennal segment (Fig. 2); the male genitalia (Fig. 5E-F) are very similar to those of decor ata, with the sensory lobe of the left paramere not elevated and the setae scattered and the lateral process not developed into the hammerhead-like form found in albescens. Distribution. See Figure 6. Hosts. Psorothamnus emoryi, P. fremontii, P. polydenius, and P. schottii. Specimens examined. USA: California: Imperial Co.: 5.4 mi NW of Ocotillo on 52, 23 April 1980, ex Psorothamnus emoryi (Russell and Schwartz; AMNH), 366, 522. Inyo Co.: 21.7 mi E of Rt 395 on Westgard Pass Road, ca 1560 m, 2 July 1980, ex Psorothamnus polydenius (Schuh; AMNH), 466, 422; Deep Spring Lake Flats on Rt 168, 1644 m, 12 July 1980, ex Psorothamnus polydenius (Schuh; AMNH), 46, 42; Deep Spring Lake Flats on Rt 168, 1644 m, July 12, 1980, ex Psorothamnus polydenius (Schuh and Stonedahl; AMNH), 322; Deep Spring Lake Flats, on Hwy 168, 5260 ft, 12 July 1980 (Stonedahl; AMNH), 622; Antelope Springs, 14-15 June 1961 (Toschi; UCB), 266. Mono Co.: 6 mi N of Bishop, fish slough, 15 June 1973 (Pinto; AMNH), 16. Riverside Co.: 10 mi E of Mecca Box Canyon, 7 April 1966, ex Dalea schottii (Turner; UCB), 366, 12, Boyd Desert Res. Center, 4 mi S of Palm Desert, 6-12 April 1963 (Hurd; UCB), 16, 12. San Diego Co.: Borrego, 24 April 1955 (Wasbauer; UCB), 16, 12; Anza-Borrego State Park, Carrizo Creek, 10.2 mi NW of Ocotillo on 52, 23 April 1980, ex Psorothamnus emoryi (Russell and Schwartz; AMNH), 266, 222. Nevada: Clark Co.: 5.8 mi NW of Valley of Fire State Park, 845 m, 17 May 1978, ex Psorothamnus fremontii (Schuh; AMNH), 2266, 3622. Nye Co.: 2.6 mi W of Mercury Hwy on Cane Springs Rd, 3400 ft, 6 June 1 983, ex Psorothamnus fremontii (Schuh, Schwartz, and Stonedahl; AMNH), 666, 1 122; Nevada Atomic Test Site, 7.5 mi W of Mercury Hwy on Cane Springs Rd, 3800 ft (A26), 6 June 1983, ex Psorothamnus polydenius (Schuh, Schwartz, and Stonedahl; AMNH), 366, 522; 164 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) 1989 REVIEW OF DALEAPIDEA 165 6.8 mi SE of Mercury Hwy on Orange Blossom Rd (A1 1), 4000 ft, 8 June 1983, ex Psowthamnus polydenius (Schuh, Schwartz, and Stonedahl), 3$$. Daleapidea decomta (Uhler), new combination Figs. 3, 5I-L, 6 Hadronema decorata Uhler 1894:251 (n. sp.). Lectotype. 9, San Luis, Lower Cal., Mex., Chas. D. Haines, April 1889; deposited in the USNM. Diagnosis. Recognized by the generally charcoal coloration with a bright red scu- tellum, and the white cuneus and white areas along and adjacent to the claval suture (Fig. 3); the male genitalia (Figs. 5I-L) are very similar to those of daleae. Distribution. See Figure 6. Hosts. Psorothamnus emoryi. Specimens examined. MEXICO: Baja California Norte: San Luis, April, 1889 (Haines; USNM), 16, 19, Calamalli Mines, April 1889 (Haines; CAS, USNM), 299; San Jose de Gracias, April 1889 (Haines; CAS), 299; 24 mi N of Punta Prieta, 2 April 1973 (Powell; UCB), 16; 53 km W of Punta Prieta toward Bahia de los Angeles, 340 m, 22 April 1985, ex Psorothamnus emoryi (Schuh and Massie; AMNH), 9466, 8599. Baja California Sur: 2.3 mi W of Mexico 1 on Bahia Tortugas rd, 19 March 1981 (Andrews and Faulkner; SDNM), 266, 19. Discussion. Uhler ( 1 894) described decorata on the basis of several male and female specimens from three localities in Baja California, Mexico. I located seven specimens matching Uhler’s locality information in the California Academy of Sciences and the National Museum of Natural History. Of the three California Academy speci- mens, one bears a lectotype label and two bear paratype labels, probably affixed by Van Duzee, although he never published the designation. All of the specimens in both museums are in poor condition, and the abdomen is missing from the only male. I have selected a female from the National Museum as the lectotype, because it bears the Uhler label ''"Hadronema decorata Uhler.” I have been reminded by T. J. Henry that according to Steyskal (1973) the sup- posedly correct name is Hadronema decoratum, not decorata. Because Uhler gave no indication of derivation of the name Hadronema, Steyskal (1973) was left to presume that the name “may be derived from Greek nema . . . ,” a neuter noun. Steyskal opted to treat Hadronema as neuter, even though the type of the genus, militaris Uhler, suggested otherwise. Steyskal offered similar arguments for other genera of Miridae, such as Campylomma Reuter, a taxon proposed by a classical author who prepared literally hundreds of descriptions in Latin, and whom we are now supposed to believe— 100 years later— was not able to determine the correct gender for the type species. It is my belief that nomenclature is ill served through the institution of such purely academic changes, and I therefore choose to disregard them. ACKNOWLEDGMENTS Thanks to the following individuals and institutions for the loan of specimens: P. H. Amaud, Jr., California Academy of Sciences, San Francisco (CAS); J. D. Lattin, Department of Ento- mology, Oregon State University, Corvallis (OSU); D. Faulkner, San Diego Natural History 166 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Museum (SDNM); J. Chemsak, Department of Entomology, University of California, Berkeley (UCB); and T. J. Henry, USDA Systematic Entomology Laboratory, % National Museum of Natural History, Washington, D.C. (USNM). Specimens are also deposited in the American Museum of Natural History (AMNH). I thank Michael D. Schwartz and Gary M. Stonedahl for discussion of Orthotylini morphology and classification and for cooperation in the field and MDS for assistance with manuscript preparation and for helpful comments on the final version. I also thank T. J. Henry for useful comments on the manuscript. Funds for field work and artistic services were provided by NSF grants DEB-81 13431 and BSR-8606621. LITERATURE CITED Knight, H. H. 1968. Taxonomic Review: Miridae of the Nevada Test Site and the Western United States. Brigham Young University Science Bulletin, Biol. Ser., vol. 9(3):282 pp. Steyskal, G. C. 1973. The grammar of names in the Catalogue of the Miridae (Heteroptera) of the World by Carvalho, 1957-1960. Studia Entomol. 16:203-208. Uhler, P. R. 1 894. Observations upon the heteropterous Hemiptera of Lower California, with descriptions of new species. Proc. Calif Acad. Sci., ser. 2 4:223-295. Van Duzee, E. P. 1918. New species of Hemiptera chiefly from California. Proc. Calif Acad. Sci., ser. 4, 7:271-308. Received August 31, 1988; accepted September 19, 1988. J. New YorkEntomol. Soc. 97(2):167-172, 1989 TEXOCORIS NIGRELLUS: DISTRIBUTION AND HOSTS OF AN ENIGMATIC PLANT BUG (HETEROPTERA: MIRIDAE: ORTHOTYLINAE) A. G. Wheeler, Jr. Bureau of Plant Industry, Pennsylvania Department of Agriculture, Harrisburg, Pennsylvania 17110 Abstract.— Nine new state records are given for the seldom-collected plant bug Texocoris nigrellus (Knight). Bloodroot, Sanguinaria canadensis L. (Ranunculaceae), is the only previously recorded host, but it is questioned whether this native North American herb is an important host or whether the rearing from this plant was merely accidental. Nymphs are reported from 4 exotic ornamental shrubs; Abelia grandiflora (Andre) Rehd. (Caprifoliaceae), Ilex crenata Thunb. (Aquifoliaceae), and Pyracantha coccinea M. J. Roem. and Spiraea thunbergii Siebold ex Blume (Rosaceae). Texocoris nigrellus, belonging to a monotypic genus, does not show a close relationship to other North American orthotylines. Although certain of its attributes fit those of an immigrant, it is concluded that without further data T. nigrellus should be considered a North American endemic. Knight (1939) described Parthenicus nigrellus from Illinois, Iowa, and Texas, plac- ing the new species in the orthotyline tribe Halticini. Schalfner (1974) described the new genus Texocoris in the Orthotylini, with T. secludis from Texas as the only included species. Henry (1982), however, determined that secludis was conspecilic with Knight’s nigrellus. Because nigrellus was found not to belong in Parthenicus Reuter, Henry retained the monotypic genus Texocoris Schalfner, recognizing the combination T. nigrellus (Knight). Since Knight’s (1939) original description, only Georgia (Knight, 1941), Missouri (Froeschner, 1949), and Wisconsin (Akingbohungbe et al., 1972) have been added to the known distribution of T. nigrellus. Little information is available on this bug’s habits. In Wisconsin, Akingbohungbe et al. (1972) reported that nymphs were col- lected and reared on “blood root” [apparently Sanguinaria canadensis L. (Ranun- culaceae)]. Schalfner (1974) noted that in Texas he had taken a single nymph but had been unable to determine the host owing to this mirid’s occurrence in “dense vegetative undergrowth under a rather thick stand of secondary growth trees.” In Georgia, Henry and Smith (1979) reported its collection from a Malaise trap, and Blinn and Yonke (1985) collected 10 adults from a fruiting mulberry. Moms sp., and one on Ohio buckeye, Aesculus glabra Willd., in Missouri. Until recently, I had not seen nymphs of this species and had rarely collected adults, and then mainly on ornamental plants that seemed unlikely to have served as breeding hosts. Herein, I give new distribution records of T. nigrellus and report the abundance of nymphs on four exotic ornamental shrubs. These data raise ques- tions concerning the geographic origin and native hosts of this orthotyline. 168 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Fig. 1 . Texocoris nigrellus, dorsal habitus. Texocoris nigrellus (Knight) Fig. 1 Distribution. In addition to the states Henry and Wheeler (1988) listed in the recent catalog of North American Heteroptera— Georgia, Illinois, Iowa, Missouri, Texas, and Wisconsin— nine new state records are available. The Michigan and Clemson, South Carolina, records are based on specimens in the collection of the National Museum of Natural History, Washington, DC (USNM); remaining records, except one for Auburn, Alabama from G. L. Miller and Raleigh, North Carolina, from R. L. Blinn, are based on my recent collecting in the eastern United States. Voucher specimens have been deposited in the Cornell University, Pennsylvania Department of Agriculture, and USNM collections. Hosts are not listed below but are mentioned in the discussion of host plants that follows. The following records extend the known distribution of T. nigrellus. ALABAMA: Lee Co., Auburn University campus and Davis Arboretum, Auburn, 8-9, 11 May 1986, A. G. Wheeler, Jr.; Auburn Univ., 19 May 1988, G. L. Miller. ARKANSAS: Washington Co., Univ. of Arkansas, Fayetteville, 15 June 1987, AGW and T. J. Henry. KENTUCKY: Warren Co., Western Kentucky Univ., Bowling Green, 5 June 1985, AGW and T. J. Henry. MICHIGAN: Midland Co., 15 July 1947, R. R. Dreisbach. NORTH CAROLINA: Guilford Co., Univ. of North Carolina— Greens- boro and Greensboro College, Greensboro, 16 May 1988; Rowan Co., Catawba Coll., Salisbury, 16 May 1988, AGW; Wake Co., Raleigh, 6-8, 15 June 1987 and 28-30 1989 TEXOCORIS: DISTRIBUTION AND HOSTS 169 May, 6 June 1988, R. L. Blinn. PENNSYLVANIA: Centre Co., Pennsylvania State Univ., University Park, 25 June 1988, AGW. SOUTH CAROLINA: Greenwood Co., Lander Coll., Greenwood, 14 May 1988, AGW; Laurens Co., Presbyterian Coll., Clinton, 15 May 1988, AGW; Newberry Co., Newberry Coll., Newberry, 15 May 1988, AGW; Pickens Co., Clemson Univ., Clemson, 27 May 1955, D. Dunavin; Spartanburg Co., Converse Coll, and Wolford Coll., Spartanburg, 15 May 1988, AGW. TENNESSEE: Knox Co., Univ. of Tennessee, Knoxville, 27 May 1985, AGW and T. J. Henry; Rutherford Co., Middle Tennessee State Univ., Murfreesboro, 28 May 1985, AGW and TJH. VIRGINIA: Albermarle Co., Univ. of Virginia, Char- lottesville, 21 May 1988, AGW. Host plants and habits. Before 1988, I had collected adults of T. nigrellus by sweeping mixed herbaceous vegetation (Arkansas), a flowering rubiaceous weed grow- ing in a lawn (Tennessee), Croton alabamense E. A. Sm. ex Chapman (Alabama), Fagus sp. and Ilex sp. (Kentucky), and by beating various shrubs and small trees in an arboretum (Alabama). Particular shrub and tree species generally were not re- corded in a field notebook because occurrence of the bugs on those plants seemed accidental or, at most, to reflect dispersal to inflorescences for adult feeding. In addition, T. nigrellus sometimes was mistaken in the field for the common phyline plant bug Criocoris saliens (Reuter) (cf Knight, 1941: Fig. 91). In May 1988, nymphs were encountered on several ornamental shrubs, sometimes in large numbers, during field work in South Carolina. The striking, saltatorial nymphs are bright red with the tip of the head and abdomen whitish; Akingbohungbe et al. (1973) briefly described the fifth instar. After the discovery of host plants, surveys for T. nigrellus were made in other eastern states by concentrating on the same plant species. The following ecological notes pertain to collections made from four principal hosts; botanical information was taken from Everett (1981) and Dirr (1983). Abelia x grandiflora (Andre) Rehd. (Caprifoliaceae): This hybrid between the Chinese A. chinensis R. Br. and A. unijlora R. Br. ex Wallich. was a common host of T. nigrellus. Adults and fifth instars were collected on glossy abelia on two college campuses in South Carolina, and fourth and fifth instars on three North Carolina campuses during mid-May. Small numbers of second through fourth instars were beaten from this plant in Virginia in late May. Nymphs developed on A. x grandiflora before flowers were present. In Alabama, G. L. Miller took adults on glossy abelia in mid-May. Ilex crenata Thunb. (Aquifoliaceae): Fifth instars were found on Japanese holly, including the cultivars ‘Convexa’ and ‘Microphylla,’ on five South Carolina campuses in mid-May. Nymphs were beaten mainly from plants bearing staminate flowers. Pyracantha coccinea M. J. Roem. (Rosaceae): On the University of Virginia cam- pus, small numbers of nymphs (< 10) were collected on a specimen plant of scarlet firethorn in late May. Much larger numbers (> 100) of instars II-IV were present on a hedge of this ornamental that is native from Italy to western Asia. Plants in the hedge were not flowering; nymphs were observed on the new growth. Spiraea thunbergii Siebold ex Blume (Rosaceae): Nymphs were common in mid- May on this Asian spirea on two South Carolina college campuses. Development on this early-blooming shrub (flowers are present before the leaves) apparently takes place on the foliage. On the Penn State campus, adults were common on a honeysuckle, Lonicera 170 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) morrowii A. Gray, in late June. Because some of the specimens obviously were teneral (one female was mostly reddish), it is likely that nymphal development had occurred on this Japanese shrub of the Caprifoliaceae. Nymphs were not observed feeding in the field. In the laboratory, fourth and fifth in stars were reared on foliage of glossy abelia, and third instars were reared on excised terminals of scarlet firethorn. The bugs also fed on crushed caterpillars and on dead nymphs of their species. In South Carolina (Presbyterian College) and on the University of Virginia campus, nymphs of T. nigrellus were collected on glossy abelia with late instars of the darker red, nonsaltatorial Rhinocapsus vanduzeei Uhler. In Virginia, nymphs of both species were present on the single scarlet firethorn plant, and both were abundant on the firethorn hedge. Nymphs of the phyline R. vanduzeei frequently develop on cultivated azaleas. Rhododendron spp. (Ericaceae) (Wheeler and Herring, 1979). Abelia x gran- diflora and Pyracantha coccinea are new host records for R. vanduzeei. In Pennsylvania, an adult was collected on a flowering privet, Ligustrum sp., and R. L. Blinn took adults on azalea in Raleigh, North Carolina (pers. comm.). These plants, and others on which T. nigrellus has been collected, possibly serve as hosts, but it is apparent that adults disperse to various plants following development on their breeding hosts. My observations support Schaffner’s (1974) suggestion that populations of this plant bug are univoltine. DISCUSSION The distribution and habits of T. nigrellus are now better known. Its recorded distribution has been increased from six to 15 states, and four ornamental shrubs have been identified as common hosts. In many respects, though, this plant bug remains a puzzling species. Despite extensive collecting of Miridae in the eastern states since 1972, I have not encountered this species outside landscape plantings. In fact, I have taken T. nigrellus only on college campuses, and nymphs have been collected only on exotic plants. The four shrubs found to serve as hosts are all Old World, mainly Asian, species; Lonicera morrowii, a probable host, also is Asian. What are the native hosts of T. nigrellusl Akingbohungbe et al. (1972) reported bloodroot, an indigenous herb of rich woods, as a host in Wisconsin. Sanguinaria canadensis may indeed be a host plant, but it might be predicted that a mirid developing on this alkaloid-rich member of the Ranunculaceae would be a specialist herbivore unlikely to adapt to exotic shrubs belonging to three unrelated families. Akingbohungbe et al. listed only Dane County as the collection site; three specimens in the University of Wisconsin insect collection bear label data indicating that Ak- ingbohungbe collected T. nigrellus in the University’s arboretum at Madison. Could nymphs have dropped or been dislodged from some ornamental shrub such that their presence on bloodroot in the understory vegetation was merely incidental? Schalfner (1974) reported a series of specimens from a state park in southcentral Texas and a nymph from dense vegetation. The Texas host, as yet undiscovered, may represent a native plant species. The known distribution of T. nigrellus— V\liscons,m and Michigan south to Georgia and west to Texas— is greater than for many mirids. Its abundance on ornamental shrubs, including the widely planted glossy abelia, Japanese holly, and scarlet fire- 1989 TEXOCORIS: DISTRIBUTION AND HOSTS 171 thorn, suggest that its movement in shipments of nursery stock has broadened the native range. Where did this taxon originate? the southwestern United States? perhaps Mexico? Unfortunately, the nearest relatives of this monotypic genus are unknown. Schaffner (1974) stated that despite the enlarged hind femora, a character generally associated with the Halticini, other characters affirmed placement in the Orthotylini. T. J. Henry (pers. comm.) indicates that the male genitalia are not characteristic of the Halticini and agrees with Schaffner that T. nigrellus is quite distinct among North American Orthotylini. In view of its association with several exotic Asian shrubs and consistent occurrence in landscape plantings, another alternative is possible: that T. nigrellus is not native. Although most immigrant mirids detected in North America are indigenous to central and western Europe (e.g., Henry and Wheeler, 1979), a Japanese plant bug, Stetho- conus japonicus Schumacher, has recently been reported from Maryland (Henry et al., 1986) and New York (Schwartz, 1989), apparently having been introduced on azalea nursery stock. A mirid conspecific with T. nigrellus is not included in Hsiao’s ( 1 942) list of Chinese Miridae or Lee’s (1971) Korean list. Only a fragmentary knowl- edge of the plant bug fauna is available for the Oriental Region and, even if T. nigrellus is native there, it may be unknown. Without additional evidence, however, T. nigrellus should be considered a North American endemic. Only further collecting and study will help elucidate host relationships, nearest relatives, and origin of this interesting plant bug. ACKNOWLEDGMENTS I gratefully acknowledge T. J. Henry (Systematic Entomology Laboratory, USDA, ARS, % U.S. National Museum) for commenting on the relationship of Texocoris to other Orthotylinae and reading an early draft of the manuscript, E. R. Hoebeke (Department of Entomology, Cornell University) for his review of the manuscript, J. E. Fetter (Department of Entomology, University of Wisconsin) for providing label data from Akingbohungbe’s specimens, R. L. Blinn (Department of Entomology, North Carolina State University) for allowing me to refer to his collection of T. nigrellus from NC, and W. L. Mountain (PDA, BPI) for identifying some of the host plants. This study was supported in part by a grant from the South Carolina Heritage Trust Program. LITERATURE CITED Akingbohungbe, A. E., J. L. Libby and R. D. Shenefelt. 1972. Miridae of Wisconsin (He- miptera: Heteroptera). Univ. Wisconsin-Madison, Coll. Agric. Life Sci. R 2396, 24 pp. Akingbohungbe, A. E., J. L. Libby and R. D. Shenefelt. 1973. Nymphs of Wisconsin Miridae (Hemiptera: Heteroptera). Univ. Wisconsin-Madison, Coll. Agric. Life Sci, R 2561. 25 pp. Blinn, R. L. and T. R. Yonke. 1985. An annotated list of the Miridae of Missouri (Hemiptera: Heteroptera). Trans. Mo. Acad. Sci. 19:73-98. Dirr, M. A. 1983. Manual of Woody Landscape Plants: their Identification, Ornamental Characteristics, Culture, Propagation and Uses, 3rd Edition. Stipes Publ. Co., Cham- paign, Illinois, 826 pp. Everett, T. H. 1981. The New York Botanical Garden Illustrated Encyclopedia of Horticulture. Garland Publ., New York, 10 vols. Froeschner, R. C. 1949. Contributions to a synopsis of the Hemiptera of Missouri, Pt. IV 172 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Hebridae, Mesoveliidae, Cimicidae, Anthocoridae, Cryptostemmatidae, Isometopidae, Miridae. Am. Midi. Nat. 42:123-188. Henry, T. J. 1982. New synonymies and a new combination in the North American Miridae (Hemiptera). Proc. Entomol. Soc. Wash. 84:337-341. Henry, T. J., J. W. Neal, Jr. and K. M. Gott. 1986. Stethoconus japonicus (Heteroptera: Miridae): a predator of Stephanitis lace bugs newly discovered in the United States, promising in the biocontrol of azalea lace bug (Heteroptera: Tingidae). Proc. Entomol. Soc. Wash. 88:722-730. Henry, T. J. and C. L. Smith. 1979. An annotated list of the Miridae of Georgia (Hemiptera- Heteroptera). J. Ga. Entomol. Soc. 14:212-220. Henry, T. J. and A. G. Wheeler, Jr. 1979. Palearctic Miridae in North America: records of newly discovered and little-known species (Hemiptera: Heteroptera). Proc. Entomol. Soc. Wash. 81:257-268. Henry, T. J. and A. G. Wheeler, Jr. 1988. Family Miridae Hahn, 1 833 (=Capsidae Burmeister, 1 835). The plant bugs. Pages 25 1-507 in: T. J. Henry and R. C. Froeschner (eds.). Catalog of the Heteroptera, or True Bugs, of Canada and the Continental United States. E. J. Brill, Leiden and New York. Hsiao, T. Y. 1942. A list of Chinese Miridae (Hemiptera) with keys to subfamilies, tribes, genera and species. Iowa State Coll. J. Sci. 16:241-269. Knight, H. H. 1939. Three new species of Miridae from North America (Hemiptera). Bull. Brooklyn Entomol. Soc. 34:21-23. Knight, H. H. 1941. The plant bugs, or Miridae, of Illinois. 111. Nat. Hist. Serv. Bull. 22(1): 1-234. Lee, C. E. 1971. Heteroptera of Korea. Illustrated Encyclopedia of Fauna & Flora of Korea. Vol. 12 (Insecta IV), pp. 99-448, 475-601, 1051-1059. Schaffner, J. C. 1 974. Texocoris secludis, a new genus and species of Orthotylinae from Texas (Heteroptera: Miridae). J. Kans. Entomol. Soc. 47:281-284. Schwartz, M. D. 1989. New records of Palearctic Heteroptera in New York State: Micro- physidae and Miridae. J. New York Entomol. Soc. 97:1 1 1-1 14. Wheeler, A. G., Jr. and J. L. Herring. 1979. A potential insect pest of azaleas. Q. Bull. Am. Rhododendron Soc. 33:12-14, 34. Received September 26, 1988; accepted December 1, 1988. J. New YorkEntomol. Soc. 97(2): 173-1 86, 1989 TANYSTOMA DIABOLICA NEW SPECIES (COLEOPTERA: CARABIDAE: PLATYNINI) FROM BAJA CALIFORNIA AND ITS BIOGEOGRAPHIC SIGNIFICANCE James K. Liebherr Department of Entomology, Comstock Hall, Cornell University, Ithaca, New York 14853-0999 Abstract. — Tanystoma diabolica, new species, is described from Baja California, and a key to the five species of Tanystoma is presented. A cladistic analysis of the species permits elu- cidation of past vicariant events leading to the present-day species. The long-time disjunction of the California fauna by vicariance at the Salinas Valley is supported by the area cladogram derived for this group. The genus Tanystoma Motschulsky comprises species found along the Pacific Coast of North America from Baja California to Oregon. Liebherr (1985) resurrected the generic name, recognizing four species. More recent studies of other genera phylo- genetically close to Tanystoma (Liebherr, 1989) allowed assignment of an unde- scribed species from Baja California to Tanystoma. In this paper, I describe this new species, and present a cladistic analysis of the five species of Tanystoma. As four of the five species of Tanystoma are allopatric, or only narrowly parapatric, analysis of their geographic distributions in light of their cladistic relationships allows elucidation of portions of the pattern of past vicariant events leading to the present-day species. MATERIALS AND METHODS Specimens were examined by light microscopy using techniques of Liebherr ( 1 985). Scanning electron micrographs (SEM) were made on gold/pallidium-coated speci- mens using an Amray 1000 scanning electron microscope. Gold-coated body parts were mounted on points beneath the dissected specimens. The cladistic analysis utilized the Phylogenetic Analysis Using Parsimony (PAUP) algorithm of Swofford (1985). For an initial analysis, character states were coded using out-group comparison. Where character-state polarities were ambiguous due to out-group heterogeneity, the primitive state was estimated using two criteria. Where the character transformation could be interpreted as the reduction of a complex character, Dollo’s Rule of Reduction was used to establish the polarity. For other ambiguous characters, the primitive state was considered to be the more general state throughout Platynini. Two equally parsimonious cladograms were derived using these rules. In a second analysis, characters ambiguously polarized after out-group comparison and application of Dollo’s Rule were coded as missing data in the hypothetical ancestor entered in the PAUP data file. In this way, ambiguous character polarizations were free to change based on overall in-group parsimony. This analysis found one cladogram shorter than those found by the initial analysis. This shortest, and thus preferred, cladogram was used to establish likely character states in the ancestor of 174 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Tanystoma, allowing a choice between Paranchodemus and Rhadine as the sister group of Tanystoma. The geographic distributions of the species can be placed on the preferred cladogram to produce an area cladogram. This area cladogram is used as the basis for a scenario explaining speciation in the group. In this analysis, the widespread distribution of T. maculicolle is assumed to have arisen via dispersal subsequent to origin of its sympatric species. The area cladograms derived from the cladistic analysis do not include information derived from the widespread range of T. maculicolle. CHARACTERIZATION AND AFFINITIES OF TANYSTOMA MOTSCHULSKY The genus Tanystoma is a member of the tribe Platynini, subtribe Platyni. Species of the genus can be recognized by their pelage of sparse setae over the body surface, best observed on the basal 3 antennomeres; a mentum with anteromedial setae close together, set adjacent to the anterior marginal bead (Fig. 9); flight wings either di- morphic in the species or vestigial; and a female reproductive tract containing a short, tubular, angulate spermatheca (Fig. 14). The gonocoxae bear two lateral and one dorsal ensiform setae, (Figs. 6, 14), and 6 or 9 furrow pegs in the apical depression (Figs. 1-5). Spermathecal configuration is shared with species of Rhadine, prompting Liebherr (1985) to propose sister group status for the two genera. Species of Rhadine differ from Tanystoma in the placement of the antero-medial mentum setae, which are further apart and set further from the anterior margin in Rhadine (Fig. 10). Rhadine is also characterized by the presence of only 2 furrow pegs in the apical depression of the female gonocoxae (Figs. 7, 8). If the most common number of furrow pegs observed in Platynini to date— 4 — is judged the primitive state, setal number has decreased during the evolution of Rhadine and increased in the phyletic line leading to Tanystoma. Several Rhadine species exhibit short, sec- ondary setae on the pedicel and third antennomere, this trait best developed in the cave species (Barr, 1983; Rhadine longicollis Benedict, Rhadine perlevis Casey, Rha- dine koepkei Barr, Rhadine babcocki Barr, and Rhadine subterranea Van Dyke of my experience). These species lack secondary setae on the scape as observed in Tanystoma (see Liebherr, 1985:1 189, Fig. 16), suggesting that setal development on the antennae is of parallel derivation in the two groups. The rarity of these setational characters in Platynini argues against an interpretation of shared-primitive status. Liebherr (1989) noted that the Sino-Japanese genus Paranchodemus Habu also exhibits an elevated gonocoxal furrow peg number of 9, a condition shared with several species of Tanystoma. Females of Paranchodemus possess short tubular sper- mathecae with a broad duct similar to those of Rhadine and Tanystoma. Parancho- demus species lack the basal pronotal seta, an absence observed in many Rhadine, though not Tanystoma. The basal 3 antennomeres of Paranchodemus are glabrous, unlike those of Tanystoma, but like those of some Rhadine. The anteromedial setae Figs. 1-4. Apical depression of female gonocoxa, showing short furrow pegs and long ne- matiform setae. 1. T. maculicolle, with 9 furrow pegs, 3,050 x. 2. T. cuyama, with 6 furrow pegs, holes in membrane due to clearing of dissection, 2,561 x. 3. T. sulcata, with 6 furrow pegs, 3,698 X . 4. T. striata, with 9 furrow pegs, 2,323 x . 1989 TANYSTOMA DIABOLICA 175 176 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) 1989 TANYSTOMA DIABOLICA 177 Figs. 9-10. Mentum ventral view. 9. T. diabolica. 10. Rhadine caudata. Fig. 1 1. Pronotum of T. diabolica, holotype. of the Paranchodemus mentum are positioned as observed in Rhadine. Finally, the fourth metatarsus of members of all 3 genera lacks the subapical setae seen in many other platynine groups (Liebherr, 1989, Figs. 28-30). For these reasons, Rhadine and Paranchodemus were both considered potential out-groups for the initial deter- mination of character polarities within Tanystoma. In the context of Platynini possessing a short, tubular angulate spermatheca, the monophyly of Tanystoma is based on the following derived traits: 1) antero-medial mentum setae close together and adjacent to anterior bead; 2) basal 3 antennomeres and body surface with sparse pelage of very short setae; 3) female gonocoxae with 6 or 9 furrow pegs in the apical depression. The value of the last character awaits a full survey of this trait across more Rhadine species and other Platynini. KEY TO ADULTS OF TANYSTOMA This key may be used to identify the 5 species of Tanystoma, and supercedes that of Liebherr (1985). In that work, an unfortunate mistake in the micrometer conver- sion factor resulted in incorrectly reported body lengths. The correct range of body lengths is given parenthetically in the key below, that length determined as the sum of distances from: 1) apex of left mandible to cervical collar, 2) from median pronotal apex to median pronotal base, plus 3) from median pronotal base to elytral Figs. 5-6. T. diabolica. 5. Apical depression of female gonocoxa, 6 furrow pegs, two ne- matiform setae removed, 4,376 x. 6. Left gonocoxa, ventral view, apical lateral ensiform seta broken off, 422 x. Figs. 7-8. Apical depression of female gonocoxa. 7. Rhadine umbra Casey, 3,218 X. 8. Rhadine caudata Leconte, 1,636 x. 178 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) apex. All measurements were made horizontally in the mieroscope held of vision with an ocular micrometer. Measurements were conhrmed with a manual slide mi- crometer. 1 . Body with lateral edges testaceous, disc of pronotum and elytra contrastingly darker, piceous to rufous 2 Body color more uniform, little contrast between disc and edges of pronotum and elytra 3 2(1). Elytral disc with dark cloud expanded laterally to 6th stria medially, expanded to 4th stria in basal V4 and apical % of length, expansion from 4th to 6th stria abrupt; flight wings dimorphic, either fully developed or vestigial (length 8.2-12.6 mm) . . T. maculicolle (Dejean) - Elytral disc with central cloud evenly bordered, intervals 8 and 9 progressively more testaceous approaching lateral margin; brachypterous (length 7. 8-9. 8 mm) T. cuyama Liebherr 3(1). Pronotum colored as the elytral disc, brunneous to piceous; brachypterous 4 - Pronotum rufous, paler than the rufo-piceous head and elytra; flight wings dimorphic, either fully developed or vestigial (length 7.6-10.0 mm) T. diabolica, n. sp. 4(3). Pronotum with hind margin strongly expanded posteriorly, hind angle evident, lateral margin sinuate before basal seta, notch often present in marginal bead laterad basal seta (length 9.3-1 1.1 mm) T. sulcata (Dejean) - Pronotum with hind margin moderately expanded posteriorly, hind angle rounded, lateral margin convex to straight before basal seta, marginal bead without notch near basal seta (length 7.8-1 1.3 mm) T. striata (Dejean) Tatty stoma diabolica, new species Diagnosis. Head and center of elytra brunneous; pronotum, antennae, legs and elytral margins paler, rufous; pronotum narrowed hastily, lateral margins sinuate, hind angles rounded (Fig. 11); flight wings dimorphic, fully developed or flap-like stubs, internal sac of male aedeagus with 3 distinct patches of spines (Figs. 12, 13). Description. HEAD. Eyes moderately convex, temple expanded slightly to meet hind margin of eye; mentum with narrowly rounded well-developed median tooth (Fig. 9); foveolar pits of mentum small, shallowly sloping outside central dimple; scape, pedicel and third antennomere covered with short erect setae in addition to longer apical setae; frons and clypeus brunneous, clypeus rufo-brunneous, maxillary palps and antennal scape paler rufo-testaceous, antennal pedicel and flagellum brun- neous. PROTHORAX. Pronotum constricted basally, laterobasal margins weakly sinuate before obtuse-rounded hind angles (Fig. 1 1); lateral marginal depression narrow from front angle to basal Vs of length, gradually widening and more reflexed basally to hind seta; laterobasal depression shallow, bordered laterally by elevated margins; posterior pronotal margin convexly lobate just inside hind seta, the margin without a defined bead; median basal margin with obscure broad bead, weak longitudinal wrinkles anterad basal bead; median longitudinal impression finely engraved, meeting shallow anterior transverse impressions; anterior margin with weak bead medially, a narrow well-defined bead laterally; front angles narrowly rounded. ELYTRA. Humeri weak even in fully-winged individuals, elytra moderately con- vex, sides evenly rounded, subapical sinuation weak; apex of each elytron narrowly 1989 TANYSTOMA DIABOLICA 179 Figs. 12-14. T. diabolica. 12. Median lobe of male aedeagus, internal sac everted, dextro- ventral view, spicular fields numbered 1 and 2, holotype. 13. Apex of median lobe and sac, laevo-dorsal view, spicular fields numbered 2 and 3, holotype. 14. Female reproductive tract, ventral view. 180 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) rounded; basal groove depressed relative to disc of elytra, evenly recurved at humerus; lateral margin narrow throughout; elytral intervals slightly convex, the striae fine but continuous; 5 or occasionally 6 setae in or adjacent to the third elytral interval; 1 5 or 1 6 setae laterad eighth elytral stria; sutural interval, scutellum and elytral margins rufous, disc darker, brunneous to piceous. METATHORAX AND FLIGHT WINGS. Metepisternum relatively elongate, ra- tio of medial length to width (c/a ratio of Liebherr, 1985, p. 1 188) ranging from 1.34 to 1.50 (5 specimens); wings either with reflexed apex and fully developed (2 spec- imens) or reduced to vestigial flaps not extending beyond the metanotum (3 speci- mens). MICROSCULPTURE AND PELAGE. Frons and vertex with well-developed iso- diametric microsculpture, slightly stretched transversely in places; pronotal disc with regular transverse mesh microsculpture, base and laterobasal depressions with stron- ger, more isodiametric meshes; elytral intervals with strong isodiametric microsculp- ture, appearing weakly granulate; body surface covered with a pelage of sparse, small setae ranging from 0.02-0.05 mm long, these more evident on body edges and recessed areas such as the sternites posterad the metacoxae. MALE GENITALIA. Median lobe of aedeagus evenly recurved, the apex short, narrowly rounded (Fig. 1 2); the aedeagal internal sac bearing 3 areas of stout spicules, two (positions 1 and 2) eudorsally near the base of the sac (Figs. 12, 13) and the third near the gonopore on the eudorsal side of the sac; fine, lightly sclerotized spicules covering the remainder of the sac. FEMALE REPRODUCTIVE TRACT. Basal gonocoxite with apical fringe of 12- 1 3 setae (Fig. 6); apical gonocoxite with 2 lateral and 1 dorsal ensiform setae (Figs. 6, 14); 6 furrow pegs and 2 nematiform setae in apical pit of gonocoxite (Fig. 5); bursa copulatrix short, broad, without evident microtrichia; spermatheca tubular, angulate at base and at apical '/s; spermathecal gland duct entering at base of sper- matheca. LENGTH. 7.6-10.0 mm. Holotype. Male: MEXICO: Baja California Norte: 40 mi. E El Rosario, 0.5 mi. S Hwy. 1, 4-III-1981, 1,500' elev., P. Lesica (CUIC). Paratypes. MEXICO: Baja California Norte: Miller’s Landing, 29-III-1973, sea level, J. Doyen and S. L. Szerlip, white light along rd. (13, 19, CISC). Baja California Sun San Bartolome Bay, 2-VI-1925, H. H. Keifer (13, 19, CAS). Both paratypical localities are indicated in Michelbacher and Ross (1942, p. 15). Etymology. This species is named both to honor my friend and botanical collector, Peter Lesica, through the latinization of his eolloquial name, and for the difficulty I had in placing the species to genus. Distribution and Habits. This species is known only from the Pacific Coast of central Baja California (Fig. 15B). The holotype was taken under a sleeping bag after Fig. 15. Distribution of Tanystoma species. Stippled areas on maps indicate upland pine forest. A. T. maculicolle. B. T. diabolica (area a), with localities indicated by dots; T. cuyama (area b); T. striata (areas c + d); T. sulcata (areas d + e). 1989 TANYSTOMA DIABOLICA 181 182 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Table 1 . Matrix of primitive (0) and derived ( 1 ) character states for five Tanystoma species. Species Characters \* 2 3 4 5 6 7 8 9 10 T. maculicolle 0/1 0 0 1 0 0 0 0 1 1 T. diabolica 1/0 1 0 0 0 0 0 0 0 1 T. cuyama 1/0 1 1 1 0 1 0 0 1 1 T. striata 1/0 0 1 0 1 1 1 1 0 0 T. sulcata 1/0 0 1 0 1 1 1 1 0 0 * Initial character values to left of slash were reversed to values at right of slash in construction of most parsimonious cladogram. Text details reasons and methods for final character-state polarization. a nightly camp in desert habitat. Keifer (in Michelbacher and Ross, 1942) made note of “excessive aridity” at his stop at San Bartolome Bay. The two specimens taken at light include a brachypterous female and a winged male. The San Bartolome Bay specimens also comprise a brachypterous female and a winged male. The holotype male is brachypterous. CHARACTERS, CLADISTICS, AND BIOGEOGRAPHY Character Analysis The cladistic analysis of the five species of Tanystoma is based on 10 binary characters (Table 1). The initial estimates of transformation series polarity were based on out-group comparison using Rhadine and Paranchodemus and Dollo’s Rule of Reduction. A description of the character states and criteria for the initial polarity decisions follow. Character 1. In the primitive state the aedeagal internal sac lacks definite spinose fields, and is covered by lightly sclerotized microtrichia. In the derived state, spinose setal fields are situated on the sac. This coding is ambiguous because Paranchodemus have an aspinose sac, but Rhadine generally have one or more sclerotized fields at various positions on the sac. Character 2. In the primitive state, a spinose field near the basal dorsum of the aedeagal internal sac is comprised of less than 20 spines (Fig. 12, position 2). The derived state exhibits more than 20 spines in this field. This character is unambig- uously polarized because neither out-group possesses a spinose patch of configuration or position as seen in Tanystoma. Character 3. In the primitive state, slender spines occur in the spinose field at position 2 (Fig. 1 2), whereas in the derived state, stout spines occur in this spicular field, as seen in T. cuyama (Liebherr 1985, Fig. 22). The restriction of the spicular field at position 2 to Tanystoma allows unambiguous coding of this character. Character 4. The pronotal base is laterally sinuate in the primitive state, and convexly rounded in the derived state. Both out-groups possess cordate prothoraces, allowing unambiguous coding of this character. Character 5. In the primitive state, the pronotal hind angle is rounded, either broadly rounded, or tightly rounded between nearly perpendicular basal and lateral margins (Fig. 1 1). In the derived state, the hind angle is more angulate. This character 1989 TANYSTOMA DIABOLICA 183 is ambiguously coded as Rhadine may have either rounded or very sharp pronotal hind angles. Paranchodemus possess sharp pronotal hind angles. Character 6. In the primitive state, at lease some members of the species possess fully-developed flight wings. In the derived state, all members are brachypterous. Based on out-group comparison this would be ambiguously coded, as Rhadine are all brachypterous whereas Paranchodemus are winged. But, flight wing loss is con- sidered the loss of a complex structure. Thus, this character is unambiguously po- larized using Dollo’s Rule of Reduction. Character 7. In the primitive state, the ratio of median length to width of the metepisternum (Liebherr, 1985: 1188, c/a ratio) ranges from 1.0- 1.7 within a species. The derived state is given to species in which members possess further reduced metepisterna with c/a ratios ranging from 0.94-1.12. This character coding would be ambiguously supported by out-group comparison for the same reasons as for character 6. However, just as with flight wing reduction, metepisternal reduction is considered to be an irreversible specialization. Character 8. In the primitive state, the integument is colored anywhere from a pallid testaceous to rufo-brunneous. In the derived state, the cuticle is more heavily sclerotized and is rufo-brunneous to piceous in color. This character is ambiguously coded as Rhadine is heterogeneous, and Paranchodemus has a metallic-blue piceous integument. Character 9. Primitively the integument is unicolorous, although appendages are usually paler than the body. In the derived state, the body dorsum is darker medially, with pallid testaceous margins. This coding is unambiguous, as all Rhadine and Paranchodemus are unicolorous. Character 10. The primitive state is considered to be the possession of 3-5 dorsal elytral setae, and the derived state is the presence of more than 5 setae in at least some members of the species. This character is unambiguously coded as individuals of both Paranchodemus and Rhadine possess 5 or less dorsal setae. Cladistic Analysis The cladistic analysis based on predetermined out-group polarization results in two cladograms, both of length 14 (Fig. 16 A, B). In one cladogram, parallelisms are proposed for characters 2, 4, and 9 (Fig. 1 6 A), and in the second, parallelisms are proposed for characters 2, 3, and 6 (Fig. 16B). Each cladogram includes one reversal, either character 10 (Fig. 16 A) or character 1 (Fig. 16B). These two cladograms are similar in topology to the two equally parsimonious cladograms resulting from the analysis of four Tanystoma species presented by Lieb- herr (1985). T. diabolica is added to this cladogram near the base, with derived states in characters 1,2, and 10. In the prior analysis, character weighting was used to choose a preferred cladogram, that similar to topology A (Fig. 1 6). In this analysis an alternate, more objective procedure is made possible by the second cladistic analysis, that done with ambiguously polarized characters considered unordered. When eharacters 1,5, and 8 are considered unordered, a cladogram of the topology of Figure 16B is produced, with a total length of 13. This one-step reduction is brought about by the reversed polarity in character 1 . By this interpretation, a spinose aedeagal internal sac is considered the primitive state, with the aspinose sac of T. 184 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Fig. 16. Cladograms derived in the two cladistic analyses; slashes indicate character-state advances, x’s indicate reversals; distributional areas of species indicated parenthetically after species name. A. Cladogram of 14 steps derived by first analysis with character polarities determined by out-group comparison. B. Second cladogram of 14 steps derived as in A, which converts to 1 3-step cladogram when the primitive state for character 1 is changed to maximize in-group parsimony. In 13-step cladogram, character 1 occurs once, as an advance in the stem below T. maculicolle. macuUcolle the result of derived secondary loss of spinose fields. By the criterion of parsimony, the topology of Figure 16B is thus considered the preferred topology. Its acceptance results in preference of Rhadine as the closest out-group to Tanystoma, as they share the state of a spinose aedeagal sac (character 1). Paranchodemus would be placed as the second closest out-group, based on a sac lacking spines, lack of pubescence on the body and basal 3 antennomeres, and a slightly different sper- mathecal configuration. The acceptance of Rhadine as the sister group of Tanystoma supports the inter- pretation of parallel increase in gonocoxal furrow peg number in Tanystoma and Paranchodemus. If the numbers of furrow pegs in each Tanystoma species are overlaid onto the shortest cladogram, the basal state would be 6 furrow pegs, as observed in T. diabolica, T. cuyama, and T. sulcata (Figs. 2, 3, 5), with parallel increases to 9 furrow pegs in T. maculicolle and T. striata (Figs. 1, 4). Biogeographic Analysis If the two cladogram topologies of the cladistic analysis (Fig. 16 A, B) are converted to area cladograms representing the areas of endemism of the four geographically restricted Tanystoma species, two fundamentally different cladograms result. The preferred cladogram (Fig. 1 6B) supports an earliest vicariant event between California north of the Salinas Valley, and the Southern Coast ranges plus portions of Baja California. In the T. maculicolle-T. cuyama-T. diabolica clade, vicariance between area a of coastal Baja California (Fig. 15B) and areas to the north follows the initial vicariance at the Salinas Valley. The second vicariant event within this clade sepa- rated T. cuyama, restricted to the southern Coast Range, from the widespread T. maculicolle. The divergence of T. cuyama can be interpreted as the speciation of peripheral populations of a proto- 7". maculicolle. T. contains populations 1989 TANYSTOMA DIABOLICA 185 comprised of over 50% brachypterous individuals in the area now inhabited by T. cuyama, and essentially 100% brachypterous individuals on the Channel Islands (Liebherr and Hajek, 1986). During the Pliocene and Lower Pleistocene, the San Joaquin embayment connected with the Pacific Ocean via a strait at the present Santa Maria river valley, running east-west just north of Santa Barbara (Peabody and Savage, 1958). Such a barrier would have split the current range of T. cuyama, and may have isolated brachypterous populations of the common ancestor of T. macu- licolle and T. cuyama in peripheral island habitats similar to those of today’s Channel Islands. Based on the preferred cladogram, the current range of T. maculicolle has resulted from range expansion into at least parts of area b after the origin of T. cuyama, and into areas c, d, and e after the earliest vicariance within Tanystoma at the Salinas Valley. Colonization of the Central Valley of California by T. maculicolle would have occurred last, during the late Pleistocene drying of the valley floor (Wahr- haftig and Birman, 1965). The T. striata- T. sulcata clade comprises two brachypterous species, resulting from vicariance somewhere within area d, with subsequent limited dispersal establishing their current area of sympatry. Although Liebherr (1985) favored a pectinate cladogram equivalent to the current 14-step pectinate cladogram (Fig. 16 A) minus T. diabolica, the area relationships considered representative of a general pattern in that earlier paper are represented in this study only by the preferred cladogram (Fig. 1 6B). The inclusion of T. diabolica above T. maculicolle on a pectinate cladogram (Fig. 1 6 A) supports an oldest vicariant event between the Oregon + Northern to Southern California Coast Ranges, and the Sierra Columbia + Sierra Vizcaino of Baja California. The depauperate mesic-adapt- ed fauna of Baja California limits the number of taxa likely to have sister groups including the rest of the Pacific Coast Ranges north to Oregon. But, based on past studies, sister-group vicariance between central Baja and the rest of California has not been considered a general pattern (Truxal, 1960). In summary, preference of a taxonomic cladogram topology in this study different from the taxonomic cladogram topology preferred by Liebherr (1985), results in support for the same area cladogram, with the additional recognition of the Sierra Columbia + Sierra Vizcaino of Baja California as the sister area to the Southern Coast Range of California. The area relationships can be expressed as {a + b) -\- (c + e), using the area designations of Figure 15B. ACKNOWLEDGMENTS Specimens were obtained through the courtesy of John A. Chemsak, California Insect Survey, University of California, Berkeley (CISC), David H. Kavanaugh, California Academy of Sci- ences, San Francisco (CAS), and the Cornell University Insect Collection (CUIC). I thank Dave Kavanaugh for his insight and help in recognizing this species. This research was supported by NSF grant BSR-86 14628. LITERATURE CITED Barr, T. C., Jr. 1983. The cavernicolous anchomenine beetles of Mexico (Coleoptera: Carab- idae: Agonini). Assoc. Mexican Cave Stud. Bull. 8: 16 1-1 92/Texas Mem. Mus. Bull. 28; 161-192. 186 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Liebherr, J. K. 1985. Revision of the platynine carabid genus Tanystoma Motschulsky (Co- leoptera). J. New York Entomol. Soc. 93:1182-1211. Liebherr, J. K. 1989. Review of the Palaearctic genus Paranchodemus Habu (Coleoptera: Carabidae: Platynini). Pan-Pac. Entomol. Liebherr, J. K. and A. E. Hajek. 1986. Geographic variation in flight wing development and body size of the tule beetle, Tanystoma maculicolle (Coleoptera: Carabidae). Pan-Pac. Entomol. 62:13-22. Michelbacher, A. E. and E. S. Ross. 1942. Contributions toward a knowledge of the insect fauna of Lower California. Proc. Cal. Acad. Sci. 4. Ser. 24:1-20. Peabody, F. E. and J. M. Savage. 1958. Evolution of a coast range corridor in California and its effect on the origin and dispersal of living amphibians and reptiles. Pages 159-198 in: C. L. Hubbs (ed.). Zoogeography. Publ. No. 51, Amer. Assoc. Adv. Sci., Washing- ton, DC. Swoflbrd, D. L. 1985. PAUP, Phylogenetic Analysis Using Parsimony, Version 2.4.1 Illinois Nat. Hist. Sur., Champaign Truxal, F. S. 1960. The entomofauna with special reference to its origins and affinities. Syst. Zool. 9:165-170. Wahrhaftig, C. and J. H. Birman. 1965. The Quaternary of the Pacific mountain system in California. Pages 229-340 in: H. E. Wright, Jr. and D. G. Frey (eds.). The Quaternary of the United States. Princeton Univ. Press, Princeton, New Jersey. Received November 4, 1988; accepted January 12, 1989. J. New YorkEntomol. Soc. 97(2): 187-191, 1989 A NEW MICROCADDISFLY GENUS (TRICHOPTERA: HYDROPTILIDAE) FROM THE INTERIOR HIGHLANDS OF ARKANSAS, U.S.A. Michael L. Mathis and David E. Bowles Department of Zoology and Department of Entomology, University of Arkansas, Fayetteville, Arkansas 72701 Abstract.— Paucicalcaria, a new hydroptilid genus, and P. ozarkensis, the type species, are described from the Magazine Mountain area of Arkansas. The genus is distinguished by its unique tarsal spur count (0.1.2), the absence of ocelli, and a greatly reduced forewing venation. The new genus appears to be most closely related to the hydroptiline genus Hydroptila based on the similarities of the genitalia and thoracic nota and the mutual absence of ocelli. During a recent survey of the caddisflies of mountainous Arkansas, we encountered two male specimens representing a new genus of Hydroptilidae. The specimens were taken in a UV- light trap sample collected at Gutter Rock Creek near Green Bench, Logan County, Arkansas. Gutter Rock Creek is a small, intermittent stream origi- nating on the northwest corner of Magazine Mountain, the highest point between the Appalachian and Rocky Mountains, and has a gradient of 208.6 m/km between the origin and the type locality. This high gradient, along with the coarse substrate and relatively pristine riparian corridor, make the stream a rare habitat type for the central United States. Morphological terminology follows that of Marshall (1979). Paucicalcaria, new genus Figs. 1-8 Description. Male. Forewings narrow, pointed at apex; venation reduced, indistinct; costal fringe long. Hindwings narrow, pointed at apex; venation reduced. Head un- modified; tentorium complete; ocelli absent; antennae simple, 24-segmented, shorter than forewing. Mesoscutellum subtriangular with convex anterior margin; postero- dorsal edge touching posterior margin of notum on meson; transverse suture lacking. Metascutellum semicircular; not extending to lateral margin of notum but joined to it by narrow strap-like piece. Tarsal spur formula 0.1.2; hind tarsus with one spur each in apical and subapical positions. Setose lateral process of abdominal segment V present. Segment VII with ventral process. Segment VIII small; unmodified. Seg- ment IX fused; flexed dorsally; with antero-lateral apodeme and postero-lateral pro- cess. Inferior appendages simple, robust, long, and straight; extending posteriorly well beyond apex of tergite X. Tergite X short; lightly sclerotized; highly modified. Subgenital appendages simple. Aedeagus relatively long and slender; with distinct proximal and distal regions separated by a constriction; distal region sclerotized; intromittent organ present, demarcated basally by suture; spiral titillator present. Female and immatures unknown. Type species. Paucicalcaria ozarkensis n. sp. 188 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Diagnosis. Paucicalcaria can be distinguished from all other members of the family Hydroptilidae on the basis of its unique spur count (0.1.2), especially considering the arrangement of the spurs on the hind leg. Discussion. Most characteristics are consistent between the two specimens, but there is some variation in the shape of the metascutellum. It is semicircular in the holotype and pentagonal in the paratype. More specimens must be collected before we can establish which shape, if either, is more common within the genus. Based on the nature of the forewings and the absence of the mesokatepisternal suture, the new genus is a member of the subfamily Hydroptilinae. The form of the genitalia and the absence of a mesoscutellar transverse suture and bracteoles suggest a relationship to members of the tribe Hydroptilini. Within the tribe, Paucicalcaria appears to be most closely related to the genus Hydroptila as evidenced by the following features of the new genus that are shared by all or a number of species of Hydroptila: 1 ) the absence of ocelli, 2) the form of the genitalia (aedeagus relatively long and slender, consisting of distal and proximal ejaculatory ducts and a well demarcated intromittent organ, with distinct proximal and distal regions separated by a constriction, regions not differing greatly in maximal widths, distal portion entirely or at least with main axis sclerotized, with spiral titillator; claspers simple, long, and straight; subgenital appendages present; subgenital plate absent), and 3) the conhguration of the thoracic nota and terminal abdominal segments (pronotal warts round, appressed along midline; mesoscutellum subtriangular with convex anterior margin, extending to or nearly to posterior of notum; metascutellum pen- tagonal to triangular, joined to lateral notal margin by strap-like piece, abdominal segment VII with a ventral process; segment VIII unmodified, lacking a ventral process; segment IX annular, with antero-lateral apodeme and postero-lateral pro- cess). Although the genitalia of both genera exhibit the same basic theme, the claspers of Paucicalcaria are more robust and extend further posteriorly relative to tergite X than in Hydroptila. Possibly the two genera diverged from a common ancestor that had a complete tentorium, the distinctive genitalia and thoracic nota, a 0.2.4 spur count, typical postoccipital warts, and that lacked ocelli. The lineage leading to Hydroptila could be achieved through reduction of the tentorium, modification of the postoccipital warts into scent caps, and slight changes in the genitalia. The other lineage could lead to Paucicalcaria through loss of tarsal spurs on the two posterior pairs of legs, modification of the forewings, and small alterations of the genitalia. Although these conclusions are well supported by the available data, they are some- what tenuous because of the lack of supporting evidence that might be provided by the female and immatures and the monotypic nature of the genus. Etymology. Pauci-, Latin, meaning few; calcar, Latin, meaning spur; referring to the low tarsal spur count. Gender: feminine. Paucicalcaria ozarkensis, new species Figs. 1-8 Description. Forewings 1 .9-2. 1 mm long; base color light brown in alcohol; trichia darker brown; with erect black setae scattered on dorsal surface. Hindwings 1.6-1. 8 mm long; coloration as in forewings; lacking erect setae. Head and thorax light brown in alcohol. Postoccipital warts subelliptical, not hinged; antennae 24-segmented, 1989 NEW GENUS OF HYDROPTILIDAE 189 Figs. 1-4. Paucicalcaria ozarkensis. 1. Head, dorsal. 2. Thorax, dorsal. 3. Forewing. 4. Hindwing. Scale bar (Figs. 1,2) = 0.2 mm. Scale bar (Figs. 3, 4) = 0.5 mm. 190 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Figs. 5-8. Paucicalcaria ozarkensis, male genitalia. 5. Dorsal view. 6. Ventral view. 7. Lateral view. 8. Aedeagus, lateral and dorsal views. Ae, aedeagus; I A, inferior appendage; SgA, Subgen- ital appendage. Scale bar = 0.1 mm. 1989 NEW GENUS OF HYDROPTILIDAE 191 short; fifth segment of maxillary palpi longest, third and fourth subequal to fifth, first and second short. Pronotal warts circular, appressed at midline; mesonotum with few scattered setae; metanotum lacking setation. Abdomen creamy white in alcohol. Sternal gland with short lateral process; one long and one or two short setae at apex; pit not sculptured. Segment VII with long ventral process; process and midline of sternite with row of uniform setae. Segment VIII short, with 2 pairs of long setae dorsally, numerous lateral and ventral setae. Segment IX fused; apodeme extending into segment VII and opening anteriorly; posterolateral process with two long setae at apex. Tergite X deeply cleft dorsally, fused ventrally; forming a pair of hood- shaped lobes from dorsal view; ovoid in lateral aspect. Subgenital appendages strap- like; with small spine distally; sword-shaped in lateral view. Inferior appendages elongate, diverging apically; disto-lateral surface rugose. Aedeagus with distinct prox- imal and distal regions; basal portion widest proximally, tapering distally; titillator spiralling anteriorly one revolution before giving rise to short, slender posteriorly- directed process; distal portion of aedeagus tapering throughout length before ex- panding at apex. Holotype. 6. ARKANSAS, Logan Co., Gutter Rock Creek at low- water bridge on road to Green Bench (35°1 1'46''N, 93°39'46"W; elev. 396 m), 1 May 1987, R. A. B. Leschen, black light. Paratype. $. Same data as holotype. The holotype and paratype are deposited in the United States National Museum of Natural History, Smithsonian Institution, Washington, D.C. ACKNOWLEDGMENTS We thank Mr. Richard A. B. Leschen for collecting the specimens, Drs. Oliver S. Flint and Steven C. Harris for reviewing the manuscript and illustrations, and Drs. Mark D. Schram, Eugene H. Schmitz, Jacob R. Phillips, Robert T. Allen, and Mr. Christopher M. Carlton for reviewing the manuscript. Published with the approval of the Director, Arkansas Agricultural Experimental Station, Fayetteville, Arkansas 72701, U.S.A. LITERATURE CITED Marshall, J. E. 1979. A review of the genera of the Hydroptilidae (Trichoptera). Bull. British Mus. (Nat. Hist.), Entomol. 39:135-239. Received August 22, 1988; accepted November 4, 1988. J. New York Entomol. Soc. 97(2): 192-2 17, 1989 RELATIONSHIPS AMONG HOLARCTIC GENERA IN THE CYRTOGASTER-G^OW WITH A REVIEW OF THE SPECIES OF NORTH AMERICA NORTH OF MEXICO (HYMENOPTERA: PTEROMALIDAE) Steven L. Heydon Department of Entomology, NHB, Mail Stop 165, Smithsonian Institution, Washington, D.C. 20560 Abstract.— Cyrtogaster-group, which includes the genera Cyrtogaster Walker, Novitz- kyanus Boucek, Tricyclomischus Graham, and Callicarolynia, n.g., is defined as containing those miscogasterine pteromalid genera that possess three symmetrically arranged clypeal den- ticles and a weakly sculptured petiole with a distinct median and pairs of sublateral and lateral carinae. A phylogenetic analysis of relationships among these genera and their relationships to other miscogasterine genera is presented. Polycystus Westwood is herein synonymized with Cyrtogaster Walker. Seven Nearctic Cyrtogaster species are treated: C. anapodisis, n. sp., C. britteni Askew, C capitanea, n. sp., C. clavicornis Walker, C. reburra, n. sp., C tryphera (Walker), and C vulgaris Walker. Keys to the Nearctic Cyrtogaster species and a summary of the biology of the genus and each species are presented. A new Hawaiian species, C. annectens, n. sp., is also described. Polycystus clypeatus Girault is transferred to Thinodytes Graham; Polycystus propinquus Waterston, Polycystus nigriscapus Howard, and Polycystus luteipes How- ard are transferred to Halticoptera Spinola; and Polycystus nigritus Howard belongs in a new genus near Thinodytes Graham. A new genus Callicarolynia is described to accommodate a single Nearctic species, C. eruga, n. sp. While sorting museum collections during the author’s continuing study of the Nearctic miscogasterine Pteromalidae, three new Nearctic Cyrtogaster species were found (C. anapodisis, n. sp., C. capitanea, n. sp., and C. reburra, n. sp.), two Palearctic Cyrtogaster species were found to occur in the Nearctic region (C. britteni Askew and C. clavicornis Walker), and a new Hawaiian Cyrtogaster species morphologically intermediate between Cyrtogaster ?ind Polycystus Westwood was also found. Material belonging to a new Nearctic genus similar to Cyrtogaster was found {Callicarolynia, n. g.). In this paper, Polycystus is synonymized with Cyrtogaster, the four new Nearctic Cyrtogaster species are described, the two new Nearctic records are reported, and the new genus, Callicarolynia, is described containing its type species, C. eruga, n. sp. Study of Palearctic miscogasterine genera at the BMNH revealed that Novitzkyanus Boucek and Tricyclomischus Graham were phenetically similar to Cyrtogaster and Callicarolynia. Cladistic analysis by the author, reported herein, supports a hypothesis that Cyrtogaster, Novitzkyanus, Callicarolynia, and Tricyclomischus form a single monophyletic generic cluster defined by the following two synapomorphies: 1) three symmetrically arranged teeth or lobes on the anterior margin of the clypeus (Fig. 2) and 2) a weakly sculptured petiole with a distinct median carina and pairs of diverging submedian and lateral carinae extending its entire length (Fig. 3). Prior to this study, Cyrtogaster, Novitzkyanus, and Tricyclomischus were not thought to be closely related 1989 CYRTOGASTER-GKO\J^ RELATIONSHIPS 193 genera. Graham (1969) placed Tricyclomischus in the Miscogasterini and Cyrtogaster and Novitzkyanus in the Sphegigasterini. The genera of the Cyrtogaster-grouTp can be differentiated by the characters given in Table 1. Two related genera, Syntomopus Walker and Nodisoplata Graham, with three symmetrically arranged anterior clypeal denticles were not included in this generic group because other characters show that their petiole structure differs from that given above and because their closest affinities lie in other generic groups. Syntomopus belongs with those genera similar to Halticoptera Spinola because of similarities in the structure of the propodeal carinae and because the hind margin of the first gastral tergum is sinuous laterally and emarginate medially. Nodisoplata is morphologically identical to Seladerma Walker except for the symmetrical arrangement of the clypeal denticles. The genus Novitzkyanus contains two described species, N. cryptogaster Boucek (1961) from Europe and N. tridentatus Delucchi (1962a) from Morocco. Tricyclo- mischus is monotypic, with only T. celticus Graham (1956) from the United Kingdom. The genus Callicarolynia is also monotypic, containing only C eruga from the United States and Canada. Cyrtogaster contains nine described species: C. annectens from the Hawaiian Islands; C britteni Askew (1965), C clavicornis Walker (1833), and C. vulgaris Walker (1833) from the Holarctic region; C. mallorcensis Askew (1975) from the Palearctic region; C. nigra (Risbec, 1955) from the Ethiopian region; C. javensis Girault (1915) from the Oriental region; C. fluitantis De Santis (1972) and C santaclarae De Santis (1964) from the Neotropical region; and C. anapodisis, C. capitanea, C. reburra, C. tryphera (Walker, 1843) from the Nearctic region. Based on examination of their type specimens in the USNM and the BMNH, Polycystus clypeatus Girault (1918) is transferred to Thinodytes Graham; Polycystus propinquus Waterston (1915), Polycystus nigriscapus Howard (1897), and Polycystus luteipes Howard (1897) are transferred to Halticoptera Spinola; and Polycystus ni- gritus Howard (1897) belongs in a new genus near Thinodytes Graham. Species of Polycystus for which generic placement is uncertain include P. ivondroi Risbec (1952), P. madagascariensis Risbec (1959), and P. pauliani Risbec (1959) from the Ethiopian region. Terminology in this paper generally follows that of Graham (1969), except that genal concavity is used instead of genal hollow and club is used instead of clava. In addition, the gastral terga are numbered 1-7 beginning with the first tergite after the petiole. The following abbreviations are used: the median ocellar diameter is MOD, ocel-ocular distance is OOL, posterior ocellar distance is POL, lateral ocellar distance is LOL, multiporous plate sensillae are MPP sensillae, lower ocular line is LOcL, antennal funicular segments are FI through F6, and the gastral terga are T1 through T7. The units of measurement given in the descriptions can be converted to milli- meters by multiplying by 0.02. The author’s concept of Tricyclomischus celticus is based on a pair of specimens on loan from the BMNH that were collected at Awbridge, Romsey, Hampshire, England in September 1980 by C. Vardy, and determined by Z. Boucek. The concept of Novitzkyanus cryptogaster is based on a female from Moncada, Spain, collected 14 September 1978 from a calliphorid in a snail by M. J. Verdue, and a male from St. Chamas (near Marseilles), France, collected 9 June 1973 by Z. Boucek. These two specimens were determined by Z. Boucek, and were on loan to the author from 194 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Table 1. Character comparisons among the genera Thcyclomischus Graham, Cyrtogaster Walker, Novitzkyanus Boucek, and Callicarolynia n.g. Character Tricylomischus Cyrtogaster Novitzkyanus Callicarolynia Body color Clypeal margin Genal hollow Male maxillary palps Antennal inser- tion First funicular segment Male flagellum Pronotal collar Scutellar setae Basal cell Postmarginal vein length Petiole propor- tions Lateral patch of setae on T1 Hind margin ofTl Length of T 1 dark green projecting lobes round- ed absent slender below LOcL small, nearly annelliform similar to that of females acarinate 4 pairs setate longer than marginal vein transverse extensive but sparsely se- tate weakly con- cave < ’/2 gastral length blue-green to dark green teeth sharp extending ~V3 genal length globularly en- larged at or below LOcL same size as second similar to that of females carinate 4-5 pairs setate to bare shorter than marginal vein elongate to transverse poorly devel- oped concave < Vi gastral length bluish gray teeth sharp extending <‘/4 genal length slender above LOcL same size as second pedunculate w. erect setae carinate 2-3 pairs bare shorter than marginal vein elongate poorly devel- oped straight covering entire gaster dark green only middle tooth sharp absent weakly expanded below LOcL same size as sec- ond similar to that of females acarinate 4-many pairs setate shorter than marginal vein transverse well developed straight covering entire gaster the BMNH. Two female and one male N. cryptogaster specimens from Valencia, Spain, collected in November 1916 by E. A. Beck from a snail infested by Diptera (USNM) were also seen. The authors concept of Cyrtogaster is based on the Cyrto- gaster species treated in this paper. PHYLOGENETIC ANALYSIS OF THE CYRTOGASTER -GKOVtV GENERA For the phylogenetic analysis of the genera of the Cyrtogaster-gyouy), a character matrix was constructed by scoring the four genera of the Cyrtogaster-groxxp and five outgroup genera for 2 1 characters. The five outgroup taxa were chosen to represent four of the generic groups within the tribes Miscogasterini and Sphegigasterini (sensu 1989 CYRTOGASTER-GKOW RELATIONSHIPS 195 Graham, 1969) as defined in Heydon (1988). Lamprotatus Westwood and Misco- gaster Walker represent the Lamprotatus-group’, Merismus Walker, the Merismus- group; Halticoptera Spinola, the Halticoptera-gxo\xp\ and Sphegigaster Spinola, the Sphegigaster-group. The polarity of the characters was determined with respect to Lamprotatus, the most primitive of the outgroup taxa used. The 21 characters and their states are listed below. 1. Body Color: green to blue-green (0); bluish gray (1). 2. Anterior Margin of Clypeus: three asymmetrically arranged denticles (0), three symmetrically arranged denticles (1), two denticles (2). 3. Denticle Shape: denticles pointed (0), denticles rounded lobes (1). 4. Genal Concavity: absent (0), slight (1), profound (2). 5. Male Maxillary Palp: slender (0), green and globularly expanded (1), yellow and lamellately expanded (2). 6. Antennal Insertion: above a line between lower orbits (0), at or below a line between the lower orbits (1). 7. First Funicular Segment: about as large as second funicular segment (0), nearly annelliform (1). 8. Male Flagellum: similar in appearance to female flagellum (0), with strong sexual dimorphism (1). 9. Pronotal Collar: pronotum declining immediately from mesoscutum (0), hor- izontal collar present (1). 10. Anterior Margin of Collar: lacking anterior transverse carina (0), transverse Carina present (1). 1 1. Mesoscutal sculpture: imbricate (0), reticulate (1). 12. Scutellar Setae: four to many pairs present (0), three or fewer pairs present (1). 13. Basal Cell of Fore Wing: setate (0), bare (1). 14. Postmarginal Vein: longer than marginal vein (0), shorter than marginal vein (1). 15. Petiole Proportions: transverse (0), as long as or longer than wide (1). 16. Petiolar Carinae: carinae absent or short (0), median carina present (1), di- verging sublateral and lateral carinae present in addition to median carina (2). 17. Petiolar Base: acarinate ventrally at base (0), with ventral flange at base (1). 18. Lateral Setal Patch on Tl: absent or poorly developed (0), extensive (1). 19. Hind Margin of Tl: straight (0), broadly concave (1). 20. Mesal Hind Margin of Tl: straight (0), emarginate medially (1). 21. Length of Tl (females): < half the length of gaster (0), nearly equal to length of gaster (1). The character matrix (Table 2) was run on the Phylogenetic Analysis Using Par- simony (PAUP) program. Version 2.4, written by D. S. Swoffbrd (Illinois Natural History Survey). Because the data matrix was small enough, the branch and bound option was selected, thereby guaranteeing that all most parsimonious trees would be found. Three multistate characters (2, 5, and 16) were run as unordered characters because no plausible a priori hypothesis of character state transitions among the three character states could be made; however, the same tree was produced when character 1 6 was run as an ordered character. Both character states occurred in characters 1 3 196 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Table 2. Character matrix for phylogenetic analysis of the Cyrtogaster-%roup. Character state Genus 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 Lamprotatus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Miscogaster 0 0 0 2 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 Merismus 0 0 0 2 0 0 0 0 1 1 1 1 0 0 1 0 1 1 0 0 0 Halticoptera 0 2 0 2 2 1 0 0 1 0 1 1 1 1 1 1 1 1 0 1 0 Sphegigaster 0 2 0 2 0 0 0 0 1 0 1 0 1 1 1 0 1 1 1 0 0 Cyrtogaster 0 1 0 2 1 1 0 0 1 1 1 0 7 1 7 2 1 1 1 0 0 Novitzkyanus 1 1 0 1 0 0 0 1 1 1 1 1 1 1 1 2 1 1 0 0 1 Callicarolynia Tricyclomis- 0 1 0 0 2 1 0 0 1 0 1 0 0 1 0 2 1 0 0 0 1 chus 0 1 1 0 0 1 1 0 1 1 1 0 0 0 0 2 1 0 0 0 0 and 1 5 in Cyrtogaster. These characters were coded as missing (?), thereby freeing the PAUP algorithm to select either state during tree building since no a priori assumptions about the primitive or advanced states of these characters within the genus were made. Although C. anapodisis lacks the globularly enlarged male maxillary palps characteristic of other Cyrtogaster species, several apomorphic characters sup- port a relationship between it and C. vulgaris, the most apomorphic of the described Cyrtogaster species. This suggests that character 5 has reverted to the primitive state in C. anapodisis. Therefore, Cyrtogaster was coded as having state 1 for character 5 even though the genus as a whole was polymorphic for this character. A tree of 40 steps with a consistency index of 0.625 was produced (Fig. 1, Tree A). The Cyrtogaster-gYoup is defined on the tree by the symmetrical arrangement of the three clypeal denticles, the transverse carina on the pronotum, and the pattern of carinae on the petiole. The carinate pronotum is also characteristic of genera in the Merismus-gronp. Thus, either it gives evidence of a relationship between genera of the Merismus-group and genera of the Cyrtogaster-groxxp or it has arisen inde- pendently in each group. The data set used in this analysis favors an independent origin of this character in each group. The following two synapomorphies define Cyrtogaster: 1 ) The hind margin of T 1 is broadly concave. This apomorphic character state also occurs in Sphegigaster. 2) The male maxillary palps are globularly enlarged, at least primitively. This apo- morphic character state is also found in Haliplogeton De Santis (not seen) and some species of Sphaeripalpus Forster. Sphaeripalpus has other characters that place it among the genera of the Lamprotatus-group. Autapomorphies defining Novitzkyanus are the bluish gray body color and a strong sexual dimorphism in the antennal flagellum. The antennae of female Novitzkyanus resemble those of other species in the Cyrtogaster-group (as in Figs. 8, 19); the male funicular segments, however, are pedunculate, cylindrical, 2-3 times as long as wide, and are covered with long and semierect setae. Two other apomorphies set Novitz- kyanus off from other genera of the Cyrtogaster-group: the antennae are inserted in the middle of the face (a reversion to an ancestral condition), and there is a reduction in the number of pairs of scutellar setae (a common apomorphic character state). 1989 CYRTOGASTER-GKO\5V RELATIONSHIPS 197 Callicarolynia, although fairly distinct in appearance, is rather poorly defined cla- distically. The two apomorphies defining this genus are the expanded maxillary palps of the males and the loss of the transverse carina on the pronotum. This latter condition is a reversion to a common plesiomorphic state. Besides occurring only in the males, the lamellately expanded maxillary palps are characteristic of Halticoptem, and therefore not unique to Callicarolynia. Autapomorphies defining the genus Tricyclomischus are the rounded clypeal teeth, the reduced first funicular segment of the female antenna, and the lengthening of the postmarginal vein. This latter is a reversion to a common plesiomorphic state. The behavior of character 2 1 depends on the type of optimization employed. Farris optimization would have T 1 becoming elongate in the common ancestor of Novitz- kyanus, Callicarolynia, and Tricyclomischus and then reverting to the ancestral state in Tricyclomischus. Another equally parsimonious change would be the independent evolution of the elongate T1 in both Novitzkyanus and Callicarolynia. I judge this latter occurrence to be more probable because a greatly elongate T1 has arisen in- dependently in a number of other miscogasterine genera such as Cryptoprymna Forster, Notoglyptus Masi, and Toxeuma Walker. One anomalous result of this analysis is the grouping of the overall rather plesio- morphic genus Tricyclomischus with the apomorphic Callicarolynia. The four char- acter state transitions in characters 4, 13, 15, and 1 8, which define the branch uniting these two taxa, are all reversions to the primitive state. In addition, because of the apomorphic placement of Callicarolynia, character 14, the relative length of the postmarginal vein, undergoes a reversion apomorphy on the branch between Tri- cyclomischus and the common ancestor of Callicarolynia and Tricyclomischus. This character is rather constant within genera in the Miscogasterinae, which indicates it may not be readily subject to reversion. Moving Tricyclomischus to a more intuitively correct position at the base of the Cyrtogaster-%rowp might resolve this difficulty as well as the need for a reversion to the primitive state in character 2 1 . Tricyclomischus was moved to the base of the Cyrtogaster-%voup using the define tree option with PAUP. However, rerooting Tricyclomischus did not resolve these difficulties satis- factorily because the new analysis produced significantly longer trees. The new trees were a minimum of four steps longer than Tree A (Fig. 1, Tree B). One extra step is saved in character 21, but a step is added in characters 13, 15, and 18 and two steps are added in character 4. The reversion apomorphy in character 1 4 still occurs because the postmarginal vein is shortened in Halticoptera and Sphegigaster, genera ancestral to the Cyrtogaster-growp, and a majority of the genera in the Cyrtogaster-gvoup. Cyrtogaster Walker Cyrtogaster'^^WiQY, 1833:371, 381. Type species: Cyrtogaster rufipesVsfdAkQv, 1833. Westwood, 1839:68; Forster, 1856:52, 53, 54-55 (synonymy); Thomson, 1878:18, 25; Ashmead, 1904:331, 332 (key); Schmiedeknecht, 1909:375, 376, 382-383; Viereck, 1916:468; Nikol’skaya, 1952:248 (key); Peck, 1963:14, 626; Peck et al., 1964:38 (key); Askew, 1965: 179-195; Graham, 1969:124, 141-144; Dzhanokmen, 1978:77 (key); Burks, 1979:787. Polycystus Westwood, 1839:68. Type species: Polycystus matthewsii Westwood, 1839. Thomson, 1878: 18, 26; Ashmead, 1904:331, 332; Schmiedeknecht, 1909:375,376, 198 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) 380-381; Graham, 1956:261; Peck etal., 1964:38; A.skew, 1965:183-184; Graham, 1969:124, 144; Dzhanokmen, 1978:77 (key). New Synonym. Diconnus Forster, 1840:38. Type species: Dicormus aquisgranensis Forster, 1840. Forster, 1856:55. Hatia Risbec, 1955:248. Type species: Hatia nigra Risbec, 1955. Boucek, 1976:14. Description. Color: Head, mesosoma, coxae, petiole metallic blue-green to dark green; gaster dark metallic green to black. Female. Head transversely oval to more pentagonal, often weakly protuberant at level of antennal insertions; antennal scrobes deep so head length at inner orbits nearly half again its median length; clypeus usually nearly smooth with three sym- metrically arranged sharp marginal denticles; gena with concavity above base of mandibles, concavity sometimes weak; eye often setate; vertex with low transverse ridge behind ocelli in C. vulgaris and C. anapodisis\ occiput acarinate. Antenna inserted near level of ventral edge of eyes; formula 1:1:2:6:3; scape slender, length approximately 9 x width; flagellum strongly clavate (width of club often 2 x width of FI), often appearing granulate; MPP sensillae coarse, prominent, in single trans- verse row on each flagellar segment; club lacking terminal spine or large patch of micropilosity (though a small patch is present on apical segment in C. vulgaris), blunt apically. Mandibles four-toothed. Mesosoma flattened dorsally in C. britteni; prono- tum with collar separated from neck by transverse carina, collar smooth posterior of carina; notauli usually complete but indistinct posteriorly in C. capitanea and C. vulgaris-, scutellum as long as wide, with four to many pairs of lateral setae, frenal sulcus complete and distinct; propodeum rugulose but median carina and plicae usually traceable, spiracle circular to shortly ovate, callus with setae relatively dense, nucha undeveloped; mesopleuron with upper epimeron smooth. Larger species with two hind tibial spurs, smaller species with one. Fore wing with relative lengths of veins as follows: submarginal > marginal > postmarginal > stigmal; basal cell and speculum varying from completely bare (C. capitanea) to completely setate (C. clav- icornis). Petiole varying from transverse to longer than wide; nearly smooth except for median, diverging sublateral, and lateral carinae (carinae poorly developed in C. vulgaris and C. anapodisis)’, basal ventral flange present; one pair of lateral setae present in C. capitanea. Gaster ovate; T1 and T2 covering nearly its entire dorsal surface; hind margin of T 1 broadly concave; remainder with hind margin straight. Male. Similar to female except flagellum parallel-sided and MPP sensillae less prominent. Maxillary palps may be totally slender (C. anapodisis)-, with segment three metallic green and globularly expanded and the fourth segment present (Fig. 5) {Cyrtogaster sensu authors); or with segment three metallic green and globularly enlarged, the terminal segment lost, and an additional globular lobe off the stipites (Fig. 7) {Polycystus sensu authors). Discussion. Both Askew (1965:184) and Graham (1969:144) noted that the genus Polycystus is very similar to Cyrtogaster and might be reduced to a subgenus of Cyrtogaster. Graham (1969) separates the two genera on the basis of the pattern of setae on the fore wing and the structure of the male maxillary palp. Species in Cyrtogaster have a well developed speculum and at least the proximal third of the basal cell bare; the type species of Polycystus, P. clavicornis Walker, has the basal cell setate and the speculum absent. Male Cyrtogaster have the penultimate segment 1989 CYRTOGASTER-GKOU^ RELATIONSHIPS 199 of the maxillary palps globularly expanded and the stipites unmodihed (Fig. 5), male P. clavicornis have the terminal segment expanded and another globular lobe from the stipites (Fig. 7). A new Cyrtogaster species from the Hawaiian Islands, C an- nectens, n. sp., has the fore wing setal pattern of Cyrtogaster and the male maxillary palpal structure of Polycystus. In addition, males of the new Nearctic species C. anapodisis, n. sp. have maxillary palps that are slender and totally lack globular enlargement. Because of the variability in the two characters previously used to separate these two genera, the wing setation pattern and the structure of the male maxillary palps, I am synonymizing Polycystus Westwood with Cyrtogaster Walker. Cyrtogaster is here redefined as containing those species in the Cyrtogaster-group that have the hind margin of the first gastral tergum concave and otherwise agree with the characters given for Cyrtogaster in Table 1. BIOLOGY OF CYRTOGASTER Known hosts and the habits and plant associations of those hosts are presented in Table 3. Species of this genus are pupal parasitoids of a number of different leaf-, stem-, or seed-mining Diptera or other small dipterous pupae that might be adhering to plant material, since nonleaf-mining flies of the genera Lonchoptera (Lonchopteridae), Drosophila (Drosophilidae), and Brachydeutera (Ephydridae) are also recorded as hosts (Graham, 1969; Burks, 1979). An old record of Cyrtogaster vulgaris reared from an aphid by Haliday was questioned by Reinhard (1859), but now, after almost 130 years, Haliday may be vindicated. There is a specimen of C. vulgaris in the USNM which was reared from a mummy of the aphid Macrosiphum euphorbiae (Thomas). Such mummified aphids may sufficiently resemble fly pupae that they are occasionally parasitized by this species. The number of instars passed by species of Cyrtogaster is uncertain. Cameron (1939) reported that C. vulgaris has a life-history “in most respects like that of Sphegigaster flavicornis" [=S. pallicornis (Spinola)], which he gave as having five larval instars. Simmonds (1952) found that C. tryphera had three larval instars and a resting (prepupal) stage. Mating behavior in Cyrtogaster is typical for chalcidoids in general, and details for C. tryphera were given by Simmonds (1952). Simmonds (1952), also reported host-feeding by adult female C. tryphera. Askew (1965) reported C. vulgaris females can be taken on Salix catkins in the spring, so adults probably take nectar as well. Adult female Cyrtogaster vulgaris overwinter in various types of tufted or loose plant material such as evergreen foliage, Carex tufts, moss, galls, haystacks, bird nests, etc. (Askew, 1965 and VanderSar, 1978). Adults of Cyrtogaster britteni (Graham, 1969) and C. tryphera have been taken in February and March so overwintering by adults may be common in this genus. Cyrtogaster clavicornis, however, overwinters inside its host (Burghele, 1959). KEY TO THE SPECIES OF CYRTOGASTER WALKER 1. Vertex with transverse ridge immediately behind posterior ocelli (Fig. 6) 2 - Vertex smoothly rounded behind posterior ocelli (Fig. 4) 3 2. Both sexes with scutellum alveolate. Notauli obscure posteriorly. Males with middle 200 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Table 3. Biology of Cyrtogaster species. Cyrtogaster sp. Host Host biology/Plant host C. anapodisis unknown C. annectens Tephritidae \xv)k.no\yn/ Bidens cosmoides (A. Gray) C. britteni unknown C. capitanea unknown C. clavicornis' Hydrellia ghseola (Fallen) (Ephydri- dae) Leaf miner/primarily Gramineae H. nasturtii (Collin) Stem mmQv/ Nasturtium officinale L. Leaf miner/several vegetables and Pegomya hyoscyami Panzer (Antho- myiidae) Agromyza sp. (Agromyzidae) Lonchoptem sp. (Lonchopteridae) weeds C. reburra unknown C. tryphera- Brachydeutera argentata (Walker) Feeds on organic debris in tempo- (Ephydridae) rary puddles Oscine/la frit (Linneaus) (Chloropi- Stem and seed miner/grasses and dae) small grains Paregle cinerella (Fallen) (Antho- myiidae) Drosophila sp. (Drosophilidae) Associated with bovine dung C. vulgaris^ Phytomyza crassiseta Zett. (Agro- Leaf minQv/ Vernonia officinalis myzidae) Linneaus P. ilicis Curtis Leaf miner///px aquifolium Lin- neaus Chromatomyia horticola (Goureau) (Agromyzidae) Leaf minQT/ Aster spp. C nigra (Meigen) Leaf mmer/ Chrysanthemum and Gazania Cerodontha caricicola (Hering) (Agromyzidae) Leaf mmQv/Carex spp. Agromyza sp. (Agromyzidae) /Lupinus sp. Opomyza florum (F.) (Opomyzidae) Shoot and stem miner/grasses and small grains Oscinella frit (Linneaus) (Cecido- Stem and seed miner/grasses and myiidae) small grains Lonchoptera sp. (Lonchopteridae) Chloropidae sp. (Chloropidae) Associated with decaying plant material Biological references. 1. Askew (1965), Graham (1969), Fulmek (1962), Henriksen (1919). 2. Burks (1979), Blume (1986), Simmonds (1952). 3. Askew (1965), Graham (1969), Henriksen (1919), Nikol’skaya (1952), Peck (1963), von Rosen (1964). 1989 CYRTOGASTER-GRO\]P RELATIONSHIPS 201 tibia and tarsus black, pretarsus expanded (Fig. 1 6). Maxillary palps globularly enlarged vulgaris Walker - Males with scutellum coriaceous. Notauli distinct to hind margin of mesoscutum. Mid tibia and tarsus concolorous with fore and hind tibiae and tarsi, middle pretarsus as wide as tarsal segments 1-4 (as in C. tryphera. Fig. 1 5). Maxillary palps slender, segments cylindrical. (Female unknown.) anapodisis, n. sp. 3. Dorsum of mesosoma flat, scutellum flat between pairs of lateral setae . . . britteni Askew - Dorsum of mesosoma convex, particularly the part of the scutellum between the pairs of lateral setae 4 4. Hind margin of T1 with parabolic emargination extending half length of T1 (Fig. 9). Basal cell and vein glabrous. Large species (length 2. 2-3.0 mm capitanea, n. sp. - Hind margin of T1 straight mesally or with concavity extending less than a third maximum length ofTl (Fig. 13). Basal vein and cell setate. Smaller species (< 2.2 mm in length) 5 5. Fore wing lacking speculum. Male maxilla with terminal segment of palp lost and with an additional globular expansion off stipites (Fig. 7) clavicornis Walker - Fore wing with speculum (Figs. 10, 12, and 14). Male maxilla with the slender terminal segment of palp visible extending from globularly expanded third segment (Fig. 5) . . . 6 6. Fore wing with basal cell entirely setate (Fig. 12). Eye with numerous short erect setae (Fig. 11) reburra, n. sp. - Fore wing with basal half of basal cell bare (Fig. 14). Eye bare tryphera Walker Cyrtogaster anapodisis, new species Description. Holotype Male. Color: Head, mesosoma, petiole, gaster blue-green, frons more green, occiput and neck dark blue. Antenna with scape brownish yellow, pedicel yellowish brown, flagellum brown. Legs with coxae brown and only weakly metallic, pretarsi brown, remainder of leg brownish yellow. Wing veins smoky yellow. Sculpture: Clypeus smooth, head weakly alveolate, mesoscutum alveolate, scutel- lum coriaceous, frenum weakly alveolate, dorsellum with some irregular sculpturing medially, propodeum rugose and subalveolate between rugae, gaster smooth. Structure: Body length 1.3 mm. Head broadly ovate in anterior view, width 1.2 x height (26:2 1 ), 2.0 x length (26: 1 3); genal concavity extending about V3 malar distance (2. 0:5. 5); eye bare, height 1.3 x length (12:9), 2.2 x malar distance (12.0:5.5), length 3.0 X temple length (9:3); vertex with transverse carina one ocellar diameter behind lateral ocelli, occiput dropping abruptly behind carina; ratio of MOD, OOL, POL, LOL as 2:4:6:3; torulus on LOcL. Antenna with length of pedicel plus flagellum 1.6 x head width (41:26); ratio of lengths of scape, pedicel, annelli, Fl-6, club as 13.0:4.0: 1.0:4.5:4.5:4.0:4.5:4.5:3.5:10.5; ratio of widths of FI, F6, club as 2:2:2; annelli an- nular, transverse; MPP sensillae sparse, two or three visible at a time on each funicular segment. Maxillary palp slender, unexpanded. Mesosoma length 1.8 x width (35:20); notauli distinct to hind margin of mesoscutum; propodeum with spiracles circular, one diameter from anterior margin of propodeum. Wing length 2.4 x width (70:29); ratio of lengths of submarginal, marginal, postmarginal, stigmal veins as 24:14:15: 8; basal vein with single row of five setae with one on the cubital vein; basal cell with two setae; speculum open posteriorly. Legs with middle pretarsus slender, cy- lindrical. Petiole length 1.2 x width (5:4); lacking lateral setae. Gaster ovate, truncate posteriorly; length 1.6 x width (24:15); T1 shallowly concave, median length 0.92 x maximum length (16.5:18.0). 202 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Lamprotatus 4:0->2 15:0->1 Miscog aster 10 :0->l 12:0->1 9:0->l 17:0->1 18:0->1 ■ Merismus 6:0->l 16:0->2 5:0->2 12:0->1 16:2->1 20:0->l 13:0->1 14:0->1 Tree A Halticoptera 2:2->l 10:0->1 5:0->l 19:0->1 Cyrtogaster 1:0->1 6:l->0 4:2->l 21:0->1 Sphegigaster Novitzkyanus 4 :l->0 13:l->0 15:l->0 Lamprotatus Callicarolynia 3:0->l 7;0->l 14:l->0 4 :0->2 15:0->1 Miscogaster Tricyclomischus 10 :0->l 12:0->1. 9:0->l 11 : 0->l 17:0->1 18:0->1 • Merismus 6:0->l 16:0->2 5:0->2 12 :0->l 16:2->1 20:0->l 2:0->2 13:0->1 14;0->1 Tree B Halticoptera 4 :l->2 5:0->l 4 : l->0 10:0->1 13:1- 15:1- ■Sphegigaster Cyrtogaster 1:0->1 12:0->1 6:l->0 13:0->1 8:0->l 15:0->1 4 :l->0 5:0->2 10:l->0 Novitzkyanus Callicarolynia 3:0->l 4:l->0 7:0->l 14 : l->0 21:l->0 ■Tricyclomischus Fig. 1. Cladograms. Tree A. PAUP cladogram derived from character matrix in Table 2. Tree B. Intuitive cladogram input into PAUP showing the optimized character states that resulted. [Character number : state in immediate ancestor ^ state in immediate descendent.] Female unknown. Variation. The body color of the paratype male is green with bronzy reflections on the dorsum of the mesosoma. Its propodeum is finely but distinctly alveolate between the rugae. Its body length is 1 .5 mm. The length of the pedicel plus flagellum is 1.5 X the head width. The median length of T1 is % the maximum length (15:18). Discussion. Cyrtogaster anapodisis differs from other known species of Cyrtogaster because it lacks the globularly enlarged male maxillary palps. In my opinion, this 1989 CYRTOGASTER-GKOU^ RELATIONSHIPS 203 Figs. 2-7. Cyrtogaster tryphera. 2. Male clypeus. 3. Male petiole. 4. Male vertex. 5. Male maxillary palps. Cyrtogaster vulgaris. 6. Male vertex. Cyrtogaster clavicornis. 7. Male maxillary palps. characteristic is a reversion to the ancestral state because this species otherwise resembles C. vulgaris, the most morphologically advanced species in the genus. Both species have a carinate vertex and similar sculpturing on the propodeum and petiole, but can be distinguished from each other by the charaeters given in the key. In addition, the flagellum is brown in C. anapodisis, and yellow in C. vulgaris. Further, in males of C. anapodisis, the length of the pedicel plus flagellum is 1.5-1. 6 times 204 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) the head width, in males of C. vulgaris, it is about as long as the head width. Though the female of C anapodisis is unknown, it is likely to have a carinate vertex like that of the males and therefore would run out to couplet 2 in the key. The same differences in the development of the notauli and in the sculpturing of the scutellum given there for distinguishing the males of these two species are likely to distinguish the females as well. Etymology. The species name, from the Greek word anapodisis meaning a going back, refers to the reversion to slender maxillary palps in males. Type material. The holotype male and one paratype male (both INHS) were col- lected in Colorado at Green Mountain Falls (Teller County) on 17 July 1938 by H. H. Ross. Cyrtogaster britteni Askew Cyrtogaster britteni Askew, 1965:180-182, 184; Graham, 1969:142, 143; Askew, 1975:16; Dzhanokmen, 1978:81. The type of C. britteni was not in the collection at Oxford as reported in Askew (1975). The Nearctic specimens compare rather exactly with the description by As- kew. Discussion. This species is easily recognized by the flattened dorsum of the meso- soma. A tiny straight edge laid along the median longitudinal line of the mesosoma would contact the following structures simultaneously: the posterior portion of the mesoscutum, the entire scutellum, the dorsellum, and sometimes the transverse Carina along the anterior margin of the propodeum. In C. reburra, the scutellum is fairly flat between the rows of lateral setae but the mesosoma is continuously curved anterioposteriorly so that the surface of the dor- sellum is not coplanar with that portion of the scutellum anterior to the frenal sulcus. In addition, the female antenna of C. reburra has FI distinctly transverse, and the length of the pedicel and flagellum is 0.75 and 0.80 times the head width in the two known female specimens. The FI in female C. britteni is about as long as wide, and the combined length of the pedicel plus the flagellum varies between 0.87 and 0.92 times the head width in the Nearctic female specimens. The distributions of the two species are also highly disparate, with C. britteni known from northeastern Canada, and C. reburra from west-central United States. Distribution. Cyrtogaster britteni was described from England (Askew, 1965) but has not yet been reported from continental Europe. In the Nearctic region, it has been collected from two sites around Hudson Bay and one around Ungava Bay (all CNC). Canada. MANITOBA: Churchill, 5-VIII-1952, 19; Warkworth Creek (near Churchill), 21 -VI- 1952, 19. ONTARIO: Moose Factory, 27-VIII-I959, 19. QUEBEC: [Fort] Chimo, 17- 18-VIII-1959, 39. Biology. Nothing is known of the host(s) of this species. Cyrtogaster capitanea, new species Figs. 8-10 Description. Holotype Female. Color: Head, mesosoma, coxae, petiole dark green with strong purplish reflections; gaster black with greenish reflections. Antenna with scape yellow-brown, pedicel brown with weak metallic green reflections, remainder 1989 CYRTOGASTER-GROIJP RELATIONSHIPS 205 of flagellum black. Mandible yellow, teeth reddish. Legs yellow-brown; pretarsi brown. Wing veins translucent brown, stigma distinctly darker. Sculpture: Head with clypeus smooth; face coriaceous; frons alveolate medially, more imbricate dorsolaterally; mesoscutum alveolate, side lobes weakly alveolate; scutellum alveolate; dorsellum rugulose; propodeum smooth, median panels with rugae; petiole subreticulate with numerous longitudinal carinae laterally; gaster smooth. Structure: Body length 2.3 mm. Head subtriangular in anterior view, width 1.2 x height (36:31), 1.8 x length (36:20); genal concavity extending almost V3 malar dis- tance (4: 1 1); eye with scattered short erect setae, eye height 1.4 X length (19:14), 1.7 x malar distance (19:11), length 2.5 x temple length (14.0:5.5); ratio of MOD, OOL, POL, LOL as 2. 0:6. 0:7. 0:3. 5; vertex acarinate, rounding smoothly into occiput; tor- ulus with upper edge on LOcL. Antenna with length of pedicel plus flagellum 1 . 1 x head width (38:36); ratio of lengths of scape, pedicel, annelli, Fl-6, club as 21.0:6.0: 3.0:4.0:3.5:4.0:4.0:4.0:3. 5:7.0; ratio of widths of FI, F6, club as 3.5:5.0:5.0; both annelli quadrate; three to seven MPP sensillae visible at a time on each funicular segment. Mesosoma length 1.7 x width (49:29); notauli obscure posteriorly, demar- cated as line of distinct texture and color; scutellum with frenum obscure; propodeum with spiracles oval, placed on anterior edge of propodeum. Fore wing length 2.6 x width (79:30); ratio of lengths of submarginal, marginal, postmarginal, stigmal veins as 33:15:10:7; basal cell and vein bare; speculum open posteriorly. Petiole length 1.3 X width (9. 5:7. 5); one pair of lateral setae anteriorly. Gaster ovate, length 1.6 x width (45:28); T1 with parabolic emargination reaching half way to base, median length ofTl 0.53 x maximum length (18:34). Allotype male. Color: Body color green with weak coppery reflections along dorsum; antennal flagellum brown. Structure: Body length 2.3 mm. Antenna with flagellum 1.6 X as long as head width (43:27); ratio of lengths of scape, pedicel, annelli, Fl-6, club as 23.0:5.0:4.0:4.5:4.5:4.0:4.0:3.5:3.5:10.5; widths of FI, F6, club as 3.5:4.0: 4.0. Maxillary palp metallic green, third segment globularly enlarged, terminal seg- ment needlelike. Legs with middle pretarsus cylindrical. Petiole length 1.7 x width (12:7). Gaster ovate, length 1.4 x width (36:25). Variation. Female body length varies between 2.5 and 2.7 mm, and male body length varies between 2.2 and 2.9 mm. Specimens from the Rocky Mountain region often have a diffuse brown patch of pigment in the fore wing just behind the stigma. The scutellum is sometimes more reticulate and the dorsellum smooth. The color of the males varies from yellow-green to blue-green, and their flagellum is sometimes brown. Discussion. This species is easily recognized by its relatively large body size. The smallest male examined was 2.2 mm long, a length attained among the other species only by the largest C. vulgaris females. Other unique characters of this species are the glabrous basal cell and vein and the hind margin of T 1 , which is emarginate half way to its base. This species and C. vulgaris both have elongate petioles and notauli that are shallow posteriorly. Etymology. The species name, from the Latin capitaneus meaning chief in size or large, refers to the relatively large size of the individuals of this species compared with individuals of other species in the genus. Type material. The holotype female (USNM) is from O’Sullivan Dam, Grant County, Washington, and was collected on 22 July 1 954 by M. T. James. The allotype male (USNM) is from Midland County, Michigan, and was collected on 28 June 206 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Figs. 8-16. Cyrtogaster capitanea. 8. Female head, lateral view. 9. Female gaster, dorsal view. 10. Female fore wing. Cyrtogaster reburra. 1 1. Female head, lateral view. 12. Female fore wing. 13. Female gaster, dorsal view. Cyrtogaster tryphera. 14. Female fore wing. 15. Male middle leg. Cyrtogaster vulgaris. 16. Male middle leg. 1953 by R. R. Driesbach. Sixteen paratypes are distributed as follows (CNC, CSU, INHS, SEC, USNM, UW): Canada. NORTHWEST TERRITORIES: Yellowknife, 5-VI-1953, 19. ONTARIO: James Bay, 23-28-VIII-1976, 2$. United States. COL- ORADO: Fort Collins, 17-VII-1985, U. IDAHO: Parma, 14-XI-1935, 19; Tuttle Co., 1 -VI I- 1949 (on Salsola pestifer^Q\son), 19. ILLINOIS: Algonquin, 15-VII-1984, 19. 1989 CYRTOGASTER-GKOUV RELATIONSHIPS 207 INDIANA: \6. MICHIGAN: Calhoun Co., 4-IX-1958, 1<5; Cheboygen Co., 13-VIII- 1942, \6, Midland Co. 28-VI-1958, \6. NEW MEXICO: La Joya Wildlife Preserve, 20 mi. N Socorro, 15 -25- VII- 1976, 19; Ruidoso, 26-VI-1940, 19; Rio Grande, 6-VII- 1953, 19. VIRGINIA: Winchester, 1 3-VI- 1 964, 19. WISCONSIN: Dane Co., VI- 1 946, M; Dunn Co., 29-VII-I949 (on Ulmus americanus), 19. Biology. Nothing is known of the host(s) of this species. Cyrtogaster clavicornis Walker Fig. 7 Cyrtogaster clavicornis Walker, 1833:383; Delucchi, 1955:175; Graham, 1956:261; Graham, 1969:144. Cyrtogaster obscura Walker, 1833:383; Delucchi, 1955:175; Graham, 1956:261; Gra- ham, 1969:144. Polycystus matt hewsii Westv/ood, 1839:68; Graham, 1956:261; Graham, 1969:144. Polycystus scapularis Thomson, 1878:26; Henriksen, 1919:164-165; Graham, 1956: 261; Burghele, 1959:124. Polycystus clavicornis (Walker): Graham, 1956:261; Fulmek, 1962:33, 44; Askew, 1965:183-184; Graham, 1969:144; Dzhanokmen, 1978:80; Graham, 1979:276. The lectotype of C. clavicornis (BMNH Hym. Type No. 5.1816) was examined. The rest of the synonymy is accepted as given by Graham (1969). Discussion. In the USNM collection, there is a single male specimen, collected 22 June 1953 from Cherry Creek in Pinos Altos, New Mexico, which differs in no immediately discernible way from males of Cyrtogaster clavicornis from Burnham Beeches Park, England, in my own collection. Perhaps as more Nearctic specimens are collected, especially females, differences with the Palearctic material will be dis- covered. Biology. Known hosts of C. clavicornis in Europe are listed in Table 3. The biology of this species should make a very interesting study because of the strong aquatic associations of its known hosts. Two of its hosts, ephydrids of the genus Hydrellia (Graham, 1969), are miners of plants in aquatic habitats. Hydrellia griseola (Fallen) larvae form blotch mines either above or below the surface of the water in over 40 genera of Graminaceae and 20 non-graminaceous plants (Deonier, 1971). Burghele (1959) reared Cyrtogaster clavicornis in large numbers from Hydrellia griseola pupae collected in mid-winter from puddles formed in the cracks in the bottoms of shallow ponds. He observed the wasps emerging from the puparia and moving in the water until they found a hold on which to crawl out of the water. Another host, H. nasturtii Collin, the watercress stem miner, is a pest of watercress {Nasturtium officinale L.) in England (Taylor, 1928). Watercress is a sprawling perennial that grows in shall water. Its stems are often partly submerged. The developmental stage in which Cyrtogaster clavicornis attacks its hosts and whether it can attack them below the surface of the water are unknown. Cyrtogaster reburra, new species Figs. 11-13 Description. Holotype Female. Color: Head, mesosoma, coxae, petiole dark green; gaster black; remainder of legs brown, lighter at “knees” and near apex of middle tibia. Antenna with scape, pedicel dark green; flagellum dark brown. Wing veins pale 208 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) brown, stigma darker, basal half of submarginal vein lighter, apical tip of submarginal vein just before junction with marginal vein colorless. Sculpture: Head, mesonotum subalveolate except clypeus smooth and face cori- aceous; scutellum coriaceous, cells of frenum longer than wide; propodeum granulate- rugose; petiole granulate; gaster smooth, granulate basolaterally. Structure: Body length 1.5 mm. Head ovate in anterior view, width 1.2 x height (23.5:20.5), 2.0 x length (23.5: 12.5); genal concavity extending V4 malar distance (1.5: 6.5); eye setate, height 1.3 x length (11.0:8.5), 1.7 x malar length (11.0:6.5), length 3.4 X temple length (8. 5:2. 5); vertex acarinate, rounding smoothly into occiput; ratio of MOD, OOL, POL, LOL as 1 .5:3. 5:6. 0:3.0; torulus located just below LOcL. Antenna with length of pedicel plus flagellum 0.81 x head width (19.0:23.5); ratio of lengths of scape, pedicel, annelli, Fl-6, club as 11.0:4.0:1.0:1.5:1.5:1.5:1.5:1.5: 1.5: 5.0, ratio of widths of FI, F6, club as 2:4:4; annelli annulate, transverse; MPP sensillae sparse, only one or two visible at one time on Fl-4. Mesosoma length 1.6 x width (34:21); notauli groovelike; propodeum with spiracles circular, located one inner diameter from anterior margin of propodeum. Fore wing length 2.5 x width (64.0:25.5); ratio of lengths of submarginal, marginal, postmarginal, stigmal veins as 24.5:1 1.5:7. 0:6.0; basal cell completely setate; speculum reduced, closed by several rows of setae posteriorly. Petiole length 0.5 x width (4:8), with no lateral setae. Gaster ovate; length 1.5 x width (32:22); median length of T1 0.84 x its maximum length (12:14). Allotype male. Color is similar to holotype except scape yellow-brown, flagellum brown, legs with color pattern like female but paler. Sculpture like that of holotype. Structure: Body length 1.4 mm. Antenna with length of pedicel plus flagellum 0.96 x head width (24:25); ratio of lengths of scape, pedicel, annelli, Fl-6, club as 13.5:4.5: 1.0:2.0:2.0:2.0:2.0:2.0:2.0:7.0, widths of FI, F6, club as 2.0:2.5:3.0. Maxillary palps with third segment metallic green and globularly enlarged, terminal segment fusiform, slender. Legs with middle pretarsus cylindrical. Petiole length 0.5 x width (4:8). Gaster parabolic in shape, length 0.83 x width (19:23). Variation. Body length measured 1.5 and 1.6 mm for females; all males were 1.4 mm in length. The five known specimens are from the same rearing, and little morphological variation is found among them. Discussion. Cyrtogaster reburra is readily distinguished by its small body size, a female antenna that is short (length of pedicel and flagellum of females measured 0.75 and 0.80 x head width) and clavate (FI is half the width of F6), distinctly setate eyes, and a completely setate basal cell. Etymology. The species name, from the Latin reburrus meaning one with bristling hair, refers to the short, erect setae on the eyes that are characteristic of this species. Type material. The holotype female, allotype male, and 1 female and 2 male paratypes (all USNM) were collected on an animal carcass on the Bear River, Box Elder County, Utah, on 20-VII-1982 by S. W. Skinner. Biology. Nothing is known of the hosts of this species, but the type series was taken on an animal carcass. Cyrtogaster tryphera (Walker) Figs. 2-5, 14, 15 Lamprotatus trypherus Walker, 1 843: 1 58 (Lectotype, BMNH Hym. Type No. 5.82 1 ; examined); Walker, 1846:32; Burks, 1975:161. 1989 CYRTOGASTER-GKO\JV RELATIONSHIPS 209 Cyrtogaster glasgowi Crawford, 1914:36-37 (Holotype, USNM Type No. 18241; examined); Simmonds, 1952:525-528 (biology); Peck, 1963:626; Askew, 1965:179, 182-183, 184, 186; Graham, 1969:142, 143-144; Askew, 1975:16. Cyrtogaster trypherus: Burks, 1975:161-162; Burks, 1979:787; Blume, 1986:216- 217 (biology). Discussion. Cyrtogaster tryphera is rather difficult to characterize since it lacks the apomorphic characteristics that make each of the other Cyrtogaster species distinc- tive. The redescription of C tryphera (as C glasgowi) by Askew (1965) was based on a single specimen and should be corrected or supplemented as follows. The body length of females varies between 2. 1 and 1.3 mm. The body size may depend on the size of the host since a series of nine females reared on Oscinella frit (L.) are all close to 1.8 mm in length. Body color is generally very dark green, not black as stated. Head height 1.2 ± (SE=)0.01x width (N = 10), width 1.9 ± 0.00 x length; eye glabrous, height 1.7 ± 0.04 x malar distance; ratio of OOL, POL, LOL as 6.2 ± 0.19:3.8 ± 0.08:3.4 ± 0.15; torulus located just below LOcL. Antenna with length of pedicel plus flagellum 1 .0 ± 0.02 x head width; ratio of lengths of scape, pedicel, Fl-6, club as 12 ± 0.28:3.7 ± 0.20:1.6 ± 0.12:1.9 ± 0.11:2.0 ± 0.14:2.1 ± 0.11: 2.1 ± 0.12:1.9 ± 0.12:5.8 ± 0.16; ratio of widths of FI, F6, club as 2.1 ± 0.12:3.4 ± 0.13:3.7 ± 0.14. Propodeal sculpture variable, ranging from generally cobbled in appearance with weak rugae to nearly smooth with strong rugae. Fore wing length 2.4 ± 0.03 X width; ratio of lengths of marginal, postmarginal, stigmal veins as 13 ± 0.60:6.6 ± 0.34:5.6 ± 0.22; speculum open posteriorly. Petiole length 0.88 ± 0.034 X width; no lateral setae present. Gaster 1.8 ± 0.05 x as long as wide; T1 with median emargination extending about a quarter of its median length. Males generally paler in color than females, green to dark green with legs brownish yellow. Body length of males varies between 2.0 and 0.92 mm. Antenna with length of flagellum plus pedicel 1.1 ± 0.01 x head width (N = 5); ratio of lengths of scape, pedicel, Fl-6, club as 8.6 ± 1. 1:2.8 ± 0.49:1.5 ± 0.16:1.6 ± 0.19:1.6 ± 0.32:1.6 ± 0.20:1.6 ± 0.29:1.4 ± 0.25:5.2 ± 0.76; widths of FI, F6, club as 1.5 ± 0.16:2.0 ± 0.17:2.1 ± 0.17. Maxillary palp with only third segment globularly expanded, ter- minal segment very slender, fusiform. Distribution. Cyrtogaster tryphera is by far the most common Nearctic Cyrtogaster species and probably occurs throughout North America. It has been collected from the following U.S. states and Canadian provinces and territories (CNC, CU, INKS, SFC, UCR, USNM, UW): Arizona, California, Delaware, Florida, Georgia, Illinois, Kansas, Kentucky, Michigan, Minnesota, Missouri, New Jersey, New York, South Dakota, Texas, Virginia, Wisconsin, British Columbia, Northwest Territories, On- tario, Quebec, and Yukon Territory. Biology. Simmonds (1952) studied the biology of C tryphera with specimens obtained from wheat stems heavily infested by Oscinella frit, but also infested by some chloropids and the Hessian fly. He has found that specimens could be readily reared on O. frit puparia in the laboratory. Simmonds gave a description of the egg that is very similar to one given by Cameron (1939) for Cyrtogaster vulgaris. Sim- monds reported that the egg is usually laid in a furrow on the pupa and that only one egg is laid per host. An individual female may lay as many as 150 eggs during her lifetime. At 24°C, eggs take 24 hours to hatch. The first instar lasts 2-3 days, the second and third instar each take 2 days, a prepupal stage lasts one day, and the 210 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) pupal instar lasts six days. Females mate only once, and in laboratory situations, mating often occurs immediately upon emergence of the female. Females may live more than a month; males live only about 10 days. Recorded hosts are listed in Table 3. Adult C. tryphera have been taken in the months of February and March, a finding that suggests adults overwinter. I have collected adults by sweeping in Illinois from 19 May through 1 November. Cyrtogaster vulgaris Walker Figs. 6, 16 Cyrtogaster vulgaris Walker, 1833:382; Reinhard, 1859:192; C. G. Thomson, 1878: 25;Ashmead, 1894:55-56; Dalla Torre, 1898:168; Lameere, 1907:233; Schmiede- knecht, 1909:383; Grimshaw, 1915:349; Henriksen, 1919:164; Baird, 1938:87, 132, 140; Baird, 1939:106, 120, 124; Cameron, 1939:178-180, 197-199; Baird, 1940:99, 116, 1 2 1 ; Downes and Andison, 1940:948; McLeod, 1951:33; Simmonds, 1952:525 (biology); Nikol’skaya, 1952:252; Baird and McLeod, 1953:234; Mc- Leod, 1954:22-23; Delucchi, 1955:174, 175 (synonymy); W. R. Thomson, 1958: 591 (biology); Delucchi, 1962b: 14; Peck, 1963:627; Peck etal., 1964:38; von Rosen, 1964:[not seen, cited in Askew (1965)]; Askew, 1965:179-180, 181, 183, 184, 185- 186 (biology, synonymy); Graham, 1969:142-143 (synonymy); Askew, 1975:14, 15-16; Jones, 1976:91, 99, 100; Boucek, 1977:34; Dzhanokmen, 1978:80; Nor- lander, 1978:89-90; Graham, 1979:276 (distribution); Hedqvist, 1983:167. Cyrtogaster scotia Walker, 1833:382-383; Askew, 1965:179-180 (synonymy); [But see Graham, 1969:143, 202.] Cyrtogaster thoracica Walker, 1833:382; Delucchi, 1955:174; Graham, 1969:142. Cyrtogaster rufipes Walker, 1833:383; Westwood, 1839:68; Grimshaw, 1915:349; Delucchi, 1955:174; Graham, 1969:142. Cyrtogaster tenuis 1833:384; Askew, 1965:179-180; Graham, 1969:143. Cyrtogaster cingulipesV^2^kQr, 1833:384; Delucchi, 1955:174; Graham, 1969:143. Dicormus aquisgranensisVovsiQY, 1840:38; Delucchi, 1955:174-175; Graham, 1969: 143. Cyrtogaster poesosV^2L\\iQY, 1848:107, 164; Graham, 1969:142, 143. Lamprotatus acarnas Walker, 1848:111, 168; Graham, 1969:142, 143. Cyrtogaster biglobus Forster, 1861:33. Delucchi, 1955:174; Graham, 1969:142, 143. Sphegigaster deneger Walker, 1872:117; Graham, 1969:142, 143. The lectotypes of C vulgaris, C. rufipes, and C. thoracica (in the main collection of the BMNH) were examined. The remainder of the synonymy is accepted as given by Graham (1969). Discussion. Cyrtogaster vulgaris is easily recognizable because it and C. anapodisis are the only Nearctic species with a transverse carina on the vertex just posterior to the ocelli. The two species are readily separated by the characters given in the dis- cussion section for C. anapodisis. The notauli are rather weak posteriorly in C. vulgaris and C. capitanea. The males are readily distinguished from those of other Nearctic species by having the terminal segment of the maxillary palp clavate, the middle tarsi much darker than the fore and hind tarsi, and the middle basitarsus expanded and subcordiform. Distribution. Cyrtogaster vulgaris is widespread throughout the Palearctic (Graham, 1969) and occurs as far south as Morocco (Delucchi, 1962b). Although said to be 1989 CYRTOGASTER-GKOUV RELATIONSHIPS 211 introduced into British Columbia in 1937 (Baird, 1938), it was already present in the Nearctic. A female in the CNC was collected in 1931 from Vernon, British Columbia, and a male from South Bristol, Maine, was collected in 1933. This species may have been accidentally introduced into North America earlier because it has been intercepted at ports of entry into the United States on at least five separate occasions. Cyrtogaster vulgaris has been collected from the following U.S. states and Canadian provinces and territories (CMNH, CNC, INHS, UBC, UCD, UCR, UW, USNM): Alaska, California, Illinois, Maine, Michigan, Oregon, Pennsylvania, Wis- consin, British Columbia, Labrador, New Brunswick, Newfoundland, Nova Scotia, Ontario, Quebec, and Yukon Territory. Biology. Known hosts of this species are given in Table 3. In addition, this species has been collected on or associated with the following plants in North America: Malus pumila Mill, Pisum sativum L., Senecio jacobaea L., Vida angustifolia L., Vida sp., and leaf miners on Brassica, Papaver, Picea, and Primula vulgaris Hudson. Cameron (1939) reported that C. vulgaris is a solitary primary parasite on the pupal stage of the holly leaf miner, Phytomyza ilids, and that the biology of C. vulgaris closely resembles that of Sphegigaster flavicornis [=pallicornis (Spinola)], which he said is an external parasite within the puparium of the fly and goes through five larval instars. He also noted that adult C. vulgaris have a singularly effective habit of feigning death. Females of this species overwinter in a variety of coarse or tufted plant material (Askew, 1965; Graham, 1969; VanderSar, 1978). These females have no eggs in their ovaries during the winter, but are fertile and can resume egg laying the following spring (Askew, 1965; Cameron, 1939). Cyrtogaster annectens, new species Fig. 17 Description. Holotype Male. Color: Head, mesosoma, coxae, petiole, gaster dark green, frons with strong yellowish reflections. Antenna with basal third of scape yellowish brown; remainder, pedicel brownish green; flagellum reddish brown. Legs brownish yellow; pretarsi black. Wings with submarginal vein and stigma pale brown; marginal, postmarginal, stigmal veins pale yellow-brown. Sculpture: Clypeus smooth; head coriaceous, more coarse around torulus; meso- scutum coriaceous, weakly alveolate medially; scutellum coriaceous; propodeum smooth, rugose medially; petiole smooth; gaster smooth. Structure: Body length 1.6 mm. Head ovate in anterior view, width 1.3 x height (28:22), 1 .8 X length (28: 1 6); genal concavity extending half malar distance (3:6); eye glabrous, height 1.3 x length (13:10), 2.2 x malar distance (13:6), length 2.9 x temple length (10:3.5); vertex acarinate, ratio of MOD, OOL, POL, LOL as 2:5:8:3.5; torulus located 1 x own diameter above LOcL. Antenna with length of flagellum plus pedicel 1.2 X head width (33:28); ratio of lengths of scape, pedicel, annelli, Fl-6, club as 14.0:4.0:1.5:2.5:3.5:3.5:3.5:3.0:3.0:10.0; width of FI, F6, club as 2:2:2; annelli an- nulate, transverse; 1-2 MPP sensillae visible at a time on each funicular segment. Maxillary palp with terminal segment and stipites enlarged. Mesosoma length 1.7 x width (38:23); notauli complete as row of elongate punctures; dorsellum carinate anteriorly; propodeum with spiracles circular, placed on anterior margin of propo- deum. Fore wing length 2.4 x width (72:30); ratio of lengths of submarginal, marginal, postmarginal, stigmal veins as 27:15:9:7; basal cell with anterior row of setae, 3 212 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Figs. 17-19. Cyrtogaster annectens. 17. Male fore wing. Callicarolynia eruga. 18. Female head, anterior view. 19. Female habitus, dorsolateral view. irregular rows down basal vein; cubital vein setate; speculum reduced, closed pos- teriorly. Legs with pretarsi cylindrical. Petiole length 0.75 width (4. 5:6.0); with no lateral setae. Caster ovate, truncate apically; length 1.1 x width (22:20); T1 with median length 0.81 x maximum length (12.5:15.5). Allotype female. Color similar to holotype except head blue laterally. Sculpture similar to holotype. Structure: Body length 2.5 mm. Antenna with length of pedicel plus flagellum 0.96 x head width (34.0:35.5); ratio of lengths of scape, pedicel, annelli, Fl-6, club as 16.5:5.0:2.0:3.0:3.5:3.5:3.0:3.0:3.0:8.5; width of FI, F6, club as 2:3:4. Petiole length 0.7 x width (5.0:7. 5). Caster cordate, length 1.8 x width (50.0:28.5). Discussion. Cyrtogaster annectens is annectant between Cyrtogaster dind Polycystus. It has the speculum on the fore wing characteristic of Cyrtogaster and the male maxillary palp structure characteristic of Polycystus. Principally on the basis of these character distributions, I synonymize Polycystus with Cyrtogaster because females of the two genera are indistinguishable. Aside from the male palpal structure, this species resembles C. tryphera. The females can be distinguished by the position of the toruli. In C. annectens, the toruli are just above the LOcL; in C. tryphera, they are at or below this line. Etymology. The species name is from the Latin word annectens, meaning linking or joining, and refers to the morphologically intermediate structure of this species between the genera Cyrtogaster and Polycystus. Type material. The holotype male (USNM) is from Pu’u Hapapa, 4 km SW Wahi- awa, O’ahu, Hawaii, and was collected 1 6 January 1 927 by O. H. Swezey. The allotype 1989 CYRT0GASTER-GK015V RELATIONSHIPS 213 female (USNM) is from Kumuweia Ridge, 14 km SW Ha’ena, Kauai, Hawaii, and was reared from a tephritid on Bidens cosmoides (A. Gray) Sherlf on 19 June 1932 by O. H. Swezey. Biology. The allotype female was reared from a tephritid on Bidens cosmoides (A. Gray) Sherlf. Callicarolynia, new genus Type species: Callicarolynia eruga, n. sp. Description. Color: Head, mesosoma, coxae, petiole metallic dark green; gaster black. Female. Head pentagonal in anterior aspect, antennal scrobes deep; clypeus nearly smooth, with three symmetrically arranged marginal denticles; gena without hollow above base of mandible; eye setate; occiput acarinate. Antenna inserted just below LOcL; formula 1:1:2:6:3; scape slender, length approximately 9x width; flagellum strongly clavate (width of club about 2 x width of FI), finely granulate; MPP sensillae coarse, prominent, in single dense transverse row; club blunt apically, lacking terminal spine or large patch of micropilosity. Mandibles four-toothed. Mesosoma with pro- notal collar rounding smoothly over into neck; notaulus complete as deep septate groove; scutellum as long as wide, with four to many pairs of lateral setae, frenal sulcus distinct; propodeum rugose with sculpture between rugae irregular, spiracle shortly ovate, callus with setae relatively dense, nucha undeveloped; mesopleuron with upper epimeron smooth. Legs with two hind tibial spurs. Fore wing with relative lengths of veins as follows: submarginal > marginal > postmarginal > stigmal; basal cell setate; speculum nearly closed posteriorly. Petiole transverse, rather smooth except for median, diverging sublateral, and lateral carinae; basal ventral flange present; without lateral setae. Gaster ovate; T1 covering nearly its entire dorsal surface, hind margin straight. Male. Similar to female except flagellum parallel-sided and MPP sensillae less prominent. Maxillary palp with terminal segment somewhat enlarged as a weak- walled yellow sack. Discussion. Callicarolynia is distinguished from the other genera in the Cyrtogaster- group by the characters given in Table 1 . Callicarolynia would key out to Syntomopus Walker in Graham (1969), but can be distinguished from that genus by having the dorsum of the mesosoma arched, T 1 nearly covering the entire gaster, and the male maxillary palps lamellately expanded. Etymology. The generic name is derived from the Latin calli-, meaning beautiful, and the name of my wife, Carolyn. The gender is feminine. Callicarolynia eruga, new species Figs. 18, 19 Description. Holotype Female. Color: Head, mesosoma, coxae, petiole olive green; face, scape, pedicel green; flagellum, gaster black; legs yellow-brown with some me- tallic coloration on femora, tarsi paler; mandible reddish brown; maxillary palp brown. Sculpture: Clypeus smooth; head, scape, mesoscutum alveolate; scutellum, axilla coriaceous; gaster polished. Structure: Body length 2.0 mm. Head width 1 .2 x height (34:28), 2. 1 x length (34.0: 214 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) 16.5); eye with scattered short erect setae, height 1.4 x length (16.0: 1 1.5), 1.6 x malar length (16:10), length 2.9 x temple length (1 1. 5:4.0); ratio of MOD, OOL, POL, LOL as 2. 0:5. 0:7. 0:3. 5; torulus located just below LOcL. Antenna with length of flagellum plus pedicel 0.94 x head width (32:34); ratio of lengths of scape, pedicel, annelli, FI- 6, club as 18.0:5.0:1.5:3.5:3.5:3.5:3.5:3.5:3.0:8.0, ratio of widths of FI, F6, club as 2. 5:4. 0:4.0. Mesosoma length 1 .7 x width (46:27); scutellum with four pairs of lateral setae and one more medial pair between first two pairs; propodeal callus with long dense setae; spiracle length 2x width, 1.5 x outside diameter from anterior margin of propodeum. Fore wing length 2.7 x width (77:29); ratio of lengths of submarginal, marginal, postmarginal, stigmal veins as 32: 1 5: 1 2: 1 1; costal cell with single complete row of setae; basal cell with distal % setate; speculum open posteriorly. Petiole length 0.6 X width (4:7). Gaster length 1.4x width (41.5:28.5); dense hair patches present laterally on T 1 . Allotype male. Color similar to holotype female except head, mesosoma yellow- green with side lobes and hind margin of mid lobe of mesoscutum, scutellum with strong purplish reflections; maxillary palp yellow-brown; T1 dark green basally; legs lacking any metallic coloration, middle tibial spur black. Sculpture. Face imbricate, otherwise similar to the holotype. Structure. Body length 2.0 mm. Antenna inserted just above LOcL; length of flagellum plus pedicel 1.0 x head width (34:34); ratio of lengths of scape, pedicel, annelli, Fl-6, club as 18.0:4.5:1.5:3.0:3.5:3.5:3.5:3.5:3.0: 9.0; widths of FI, F6, club as 3. 0:3. 5:4.0. Terminal segment of maxillary palps lamellately expanded, length 2.3 x width (7:3). Legs with middle tibia slightly ex- panded, ventral edge sharp. Petiole length 0.64 x width (4. 5:7.0). Gaster length 1.3 x width (31:23). Variation. The body length of the female types varies from 1.8-2. 3 mm. Body color varies from blue-green to olive-green. The base color of the legs varies from pale brownish yellow to pale reddish brown. The dark bands on the femora may or may not have metallic reflections. Etymology. The specific name is derived from the Latin word erugo, meaning clear of wrinkles or smooth, and refers to the rounded pronotal collar. Type material. The holotype was collected in Mackinac County, Michigan, on 30 August 1959 by R. and K. Driesbach (USNM). The allotype was collected 7 km SW Carleton Place, Ontario, on 1 1-17 July 1980 by S. J. Miller (CNC). Seven paratypes were collected as follows (CNC, SEC, USNM): Canada. ALBERTA: Flatbush, 12- IV- 1960, 2$ (ex pupae of Odontomyia pubescens Day). MANITOBA: 4 mi. N White- water, 30-VII- 1 958, 19. ONTARIO: Bells Comers, 22-VII- 1941, 19; Ottawa, 19. United States. UTAH: Neola, 29-VI-1954, 19. WYOMING: South Pass City, 29-VII- 1954, 19. Biology. The two specimens from Flatbush, Alberta, were reared from pupae of Odontomyia pubscens (Diptera: Stratiomyidae). ACKNOWLEDGMENTS I thank the following persons for the loan of material: Dr. G. E. Wallace, Carnegie Museum of Natural History (CMNH), Pittsburgh, PA; Dr. G. P. Gibson, Canadian National Collection (CNC), Ottawa, ON; Dr. B. C. Kontratieff, Colorado State Museum (CSU), Fort Collins, CO; Dr. L. L. Pechumen, Cornell University (CU), Ithaca, NY; Dr. G. W. Byers, Snow Entomological Collection (SEC), Lawrence, KS; Dr. S. G. Cannings, University of British Columbia (UBC), 1989 CYRTOGASTER-GKOUV RELATIONSHIPS 215 Vancouver, BC; R. O. Schuster, University of California (UCD), Davis, CA; J. Hall, University of California (UCR), Riverside, CA; Dr. E. E. Grissell, United States National Museum (USNM), Washington, D.C.; and Dr. B. J. Harrington, University of Wisconsin (UW), Madison, WI. I would like to express my appreciation to Audrey Hodgins and Drs. L. I. Nevling, W. E. LaBerge, and George Godfrey of the Illinois Natural History Survey (INHS), Champaign, IL, and an anonymous reader for reviewing this paper. I would like to thank my advisor. Dr. W. E. LaBerge, and Dr. E. E. Grissell for help and encouragement, J. Sherrod (INHS) for assistance with the illustrations, J. Noyes (BMNH) for use of his card file, the staff at the Center of Electron Microscopy at the University of Illinois at Urbana-Champaign for use of the facilities, and Molly Scott (INHS) for preparing the plates. LITERATURE CITED Ashmead, W. H. 1984. Descriptions of new parasitic Hymenoptera. Trans. Amer. Entomol. Soc. 21:318-344. Ashmead, W. H. 1904. Classification of the chalcid flies or the superfamily Chalcidoidea, with descriptions of new species in the Carnegie Museum, collected in South America by Herbert H. Smith. Mem. Cameg. Mus. l:i-ix, 225-551. Askew, R. R. 1965. The Holarctic species of Cyrtogaster Walker and Polycystus Westwood (Hym., Pteromalidae) including the description of a new species of Cyrtogaster from Britain. Entomophaga 10:179-195. Askew, R. R. 1975. Some Chalcidoidea from Majorca, with descriptions of two new species. Eos Rev. Esp. Entomol. 49:13-27. Baird, A. B. 1938. Summary of insect parasites and predators liberated in Canada (to December 31, 1937). Can. Insect Pest Rev. 16(1) Suppl.:77-154. Baird, A. B. 1939. Summary of parasite liberations in Canada during the year 1938. Can. Insect Pest Rev. 17:102-128. Baird, A. B. 1 940. Summary of insect parasites and predators liberated in Canada and New- foundland 1939. Can. Insect Pest Rev. 18:94-126. Baird, A. B. and J. H. McLeod. 1953. 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McLeod, J. H. 1954. Statuses of some introduced parasites and their hosts in British Columbia. Proc. Entomol. Soc. B. C. 50:19-27. 1989 CYRTOGASTER-GROU^ RELATIONSHIPS 217 Nikol’skaya, M. N. 1 952. [The Chalcid fauna of the USSR (Chalcidoidea)]. English translation- 1963. Israel Program for Scientific Translations, Jerusalem. Nordlander, G. 1978. Parasitoids of the frit fly, Oscinella frit (L.) on oats. Norw. J. Entomol. 25:89-90. Peck, O. 1963. A catalogue of the Nearctic Chalcidoidea (Insecta, Hymenoptera). Can. Ento- mol. Suppl. 30:1-1092. Peck, O., Z. Boucek and A. Hofler. 1 964. Keys to the Chalcidoidea of Czechoslovakia (Insecta: Hymenoptera). Mem. Entomol. Soc. Can. 34:1-120. Reinhard, H. 1859. Die Blattlausen lebenden Pteromalinen. Entomol. Zeitg. 20:191-197. Risbec, J. 1952. Contribution a I’etude des Chalcidoides de Madagascar. Mem. Inst. Scient. Madagascar (Serie E) 2:1-448. Risbec, J. 1955. Hymenopteres parasites du Cameroon. Bull. Inst. Fr. Afr. Noire 17:191- 266. Risbec, J. 1959. Pteromalidae de Madagascar (Hymenoptera Chalcidoidea). Mem. Inst. Scient. Madagascar (Serie E) 11:1 29-1 7 1 . Rosen, H. von. 1964. Untersuchungen fiber die Verbreitung und Biologi von zwie Pterom- aliden in Rapsschoten (Hym., Chalcidoidea). Medd. Vaxtskyddsanst. 12:453-465. Schmiedeknecht, O. 1909. Hymenoptera family Chalcididae. In: P. Wytsman (ed.). Genera Insectorum, 97:1-550. M. P. Wytsman, Bruxelles. Simmonds, F. J. 1952. Parasites of the frit-fly, Oscinella frit (L.) in eastern North America. Bull. Entomol. Res. 43:503-542. Taylor, T. H. 1928. The watercress stem-miner. Entomol. Mon. Mag. 64:126-128. Thomson, C. G. 1878. Hymenoptera Scandinaviae. 5. Pteromalus (Svederus) continuatio. Lund. Thomson, W. R. 1958. Catalogue of parasites and predators of insect pests. Section 2. Host parasite catalogue. Part 5. Hosts of the Hymenoptera (Miscogasteridae to Trigonalidae), Lepidoptera and Strepsiptera. Commonwealth Institute of Biological Control, Ottawa. VanderSar, T. J. D. 1978. Emergence of predator and parasites of the white pine weevil, Pissodes strobi (Coleoptera: Curculionidae) from Engelmann spruce. J. Entomol. Soc. B. C. 75:14-18. Viereck, H. L. 1916. The Hymenoptera, or wasp-like insects, of Connecticut. In: W. E. Britton (ed.). Guide to the Insects of Connecticut. Part 3. Conn. St. Geol. Nat. Hist. Sur. Bull. 22:1-824. Walker, F. 1833. Monographia Chalcidum. Entomol. Mag. 1:367-384. Walker, F. 1843. Description des Chalcidites trouvees au Bluff de Saint-Jean, dans la Floride orientale, par MM. E. Doubleday et R. Forster. Annls. Soc. Entomol. Fr. (2)1:145-162. Walker, F. 1846. List of the specimens of Hymenopterous insects in the collection of the British Museum. Part L— Chalcidites. London. Walker, F. 1848. List of the specimens of Hymenopterous insects in the collection of the British Museum. Part II.— Chalcidites. Additional species. London. Walker, F. 1872. Notes on Chalcidoidea. Part 7. E. W. Janson, London. Waterston, J. 1915. New species of Chalcidoidea from Ceylon. Bull. Entomol. Res. 5:325- 342. Westwood, J. O. 1839. Synopsis of the genera of British insects. Pages 1-158. [Bound with, J. O. Westwood. An introduction to the modem classification of Insects; founded on the natural habits and corresponding organization of the different families, Vol. 2.] Long- man, Orme, Brown, Green, and Longman, London. Received October 21, 1987; accepted November 10, 1988. J. New YorkEntomol. Soc. 97(2):2 18-231, 1989 STINGING BEHAVIOR AND RESIDUAL VALUE OF WORKER HONEY BEES {APIS MELLIFERA) Steven A. Kolmes' and Linda A. Fergusson-Kolmes^ ^Department of Biology, Hobart and William Smith Colleges, Geneva, New York 14456 “Department of Entomology, New York State Agricultural Experiment Station, Geneva, New York 14456 Abstract. — 'Novktr honey bees of two subspecies {Apis mellifera mellifera andyl. m. ligustica) were bioassayed in the laboratory to determine their willingness to sting at various ages. Foragers returning to field colonies of both subspecies were captured, bioassayed for stinging behavior, and the sugar concentration and volumes of their nectar loads were measured. Residual value theory, which has previously been demonstrated to be a good predictor of the intensity of risky nest defense by parental birds, was used to interpret these data. Workers of the more highly defensive A. m. mellifera displayed residual value sensitivity in their stinging behavior, while the less defensive A. m. ligustica did not. The stinging behavior of worker honey bees is the final action in a suicidal form of colony defense. A bee that stings a vertebrate typically has its barbed stinging structures catch in the vertebrate’s epidermis and pull out of the bee’s abdomen, killing the bee. Queen honey bees have a weakly barbed stinger (see plate 1.43, Erickson, Carlson and Garment, 1986) used only to dispatch any rival queens in a colony. No model has previously been proposed to explain whether worker bees should sting more readily at specific ages, or be equally ready to sting throughout their adult lives. Collins, Rinderer, Tucker, Sylvester and Lackett (1980) proposed a model of colony defense that incorporated genetic and environmental variables, but not worker bee age. Kolmes (1985a, b, c) and Jeanne (1986) discussed the behavioral transition from hive to field duties in terms of the hazardous nature of foraging activities, but neither addressed stinging as an especially hazardous task. Worker bees are nonreproductive members of insect societies, capable of perform- ing various tasks which allow their society as a whole to produce new workers and reproductives. The evolution of this system is intimately tied to the haplodiploid nature of hymenopteran genetics, and the greater degree of genetic relatedness between a worker and her full-sisters than between a worker and her own hypothetical offspring (Hamilton, 1 964). Although worker bees are not themselves reproductive, their efforts on behalf of their colony can all be thought of as helping to produce future generations of closely related individuals (Oster and Wilson, 1978; Kolmes 1986). When worker bees are considered in the fashion just described, another body of literature exists that may help us model their stinging behavior. The intensity of nest defense by song birds has been studied in terms of the residual reproductive value of the defending parents (Curio, 1987; Curio, Regelmann and Zimmermann, 1985, 1984; Pianka and Parker, 1975; Curio, Klump and Regelmann, 1983; Windt and Curio, 1986; Regelmann and Curio, 1986, 1983). Parental birds have proven to be 1989 STINGING BEHAVIOR 219 most likely to engage in vigorous and hazardous defensive behavior when: there were more young to defend; as time in the breeding season progressed; as age of young increased; as number of young in second broods increased; in association with sex- specific differences in residual reproductive value; and as the expected number of neighboring mobbers increased (Curio, 1987; Regelmann and Curio, 1986, 1983; Curio, Regelmann and Zimmermann, 1985, 1984; Windt and Curio, 1986; Curio, Klump and Regelmann, 1983). When a parental bird had a lower future expectation of reproductive success, in either the current or future breeding seasons, it would tend to defend its present brood more vigorously. Present risk and future expectation of success seemed to be inversely related. Parallels exist between parental birds and sterile insect workers defending nests in which they are helping to produce closely related siblings. In both instances the possibility exists of hazardous defensive be- havior being related to an individual’s residual value. At a moment of crisis containing the potential for self-sacrificial defensive behavior, the balance between the benefit of stinging and a worker bee’s future value to the colony may govern individual behavior. Residual value theory predicts that a worker bee should be more likely to sting when its anticipated future value to its colony is lower. We predict therefore that (a) bees should sting more readily as they age and possess shorter future life expectancies. We can also predict that bees should sting more readily when they are able to perform less work for their colony. It is difficult to quantify the work a bee performs within the hive, but a forager that is harvesting a richer floral resource should be of greater value to a colony than a forager harvesting a poor resource. Worker bees typically develop constancies to a single type of flower. Honey bees are known to shift from one source of forage to another due to experimental manip- ulations of sucrose concentrations in feeders (Seeley, 1986) or natural declines in previously exploited nectar sources (Ribbands, 1 949). However, a variety of evidence points to a typical flower constancy in an individual forager unless environmental conditions change considerably during its relatively brief foraging life. Workers that develop an initial attraction to one variety of apple blossom continue to forage at that variety so long as it retains its relative attractiveness, despite the presence nearby of other varieties representing more abundant floral resources (Free, 1 966). Individual flowers within patches in the temperate zone have a great enough longevity in spring or early summer (ca. 5 to 7 days, Prirnack, 1985) that the retention of attractiveness of a species is a likely condition. Learning curves of bees trained to feeders are unaffected by wide ranges of concentrations or availabilities of sugar solutions during the first few rewards (Menzel, Erber and Masuhr, 1974; Menzel and Erber, 1972). Bees trained to artificial floral arrays develop constancy to wavelengths of floral reflectance (Jones, Scannell, Kramer and Sawyer, 1986). Constancy of foraging bees to specific flower colors has been identified as strong enough to serve as a pre- pollination isolating mechanism between species of Cercidium. The average “mistake frequency” of honey bees was 5.63% when two different UV floral patterns were presented, despite nearly equal caloric rewards presented by different species of Cer- cidium (Jones, 1978). Other experiments have shown that food odor is entrained to even more strongly by honey bees than flower color (Free, 1970). The actual foraging area returned to by individual worker bees has been shown to be quite small (Rib- bands, 1949; Free, 1966; Levin, 1966). Bees are most constant to floral sources that 220 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) supply both nectar and pollen (Ribbands, 1 949). Most foragers are constant to one type of pollen (Sekiguchi and Sakagami, 1966; Free, 1963). Pollen constancy tends to be retained even when a colony is moved to a new location, and under circum- stances where different pollen sources become available at various times of day (Free, 1963). Other data providing evidence of flower constancy that are too numerous to cite are presented in Grant (1950), and Wells and Wells (1983). The preceding material points to a great potential for flower constancy in the foraging behavior of most honey bees. Workers tend to remain constant to the resource they initially begin to exploit, and bees develop preferences for floral re- sources within a wide range of acceptability. The variability of floral resource value, and the likelihood that a bee will continue to exploit one resource unless it deteriorates considerably, leads to our second prediction about honey bee stinging behavior and residual value. We predict that (b) foragers returning to their colony with less valuable nectar loads might be expected to sting more readily than more “valuable” foragers. The experiments were performed to test predictions (a) and (b). In one, we indi- vidually measured the threshold stimulus required to elicit stinging for worker bees of known ages using a laboratory bioassay. This was done for workers from colonies of two subspecies of honey bees, British bees {Apis mellifera melliferd) and Italian- derived bees {A. m. ligustica). In a second series of experiments we captured foragers as they returned to colonies, and measured their threshold stimulus to elicit stinging as well as the volume and sugar concentration of nectar in their honey stomachs. The latter experiments allowed us to ask whether or not foragers were modifying their stinging behavior on the basis of the value of the nectar load they carried. MATERIALS AND METHODS This research was conducted at the Bee Research Unit of University College, Cardiff, Wales. Two colonies of honey bees were used in the experiment, one of dark British honey bees {Apis mellifera mellifera) and the other light Italian honey bees {A. m. ligustica). The Italian bee colony had been established using a mated queen imported from New Zealand. The colonies were each possessed of the distinctive behavioral traits typical of their respective subspecies. The Italian bees were “gentle” and the “highly defensive” British workers were much more prone to emerge rapidly from the hive when disturbed, and to attack and sting in the apiary more readily and persistently than the Italian colony. When the defensiveness of British and Italian colonies in our apiary was measured by bouncing a black suede ball at their entrances (Free, 1961) the British colonies had a shorter latency to delivery of the first sting than the Italian bees (avg. of 3.5 sec vs. 17 sec), the British colonies delivered more stings to the suede ball in the first minute of the test (avg. of 39.5 stings vs. 2 stings), and the British bees continued to attack the suede ball when it was withdrawn further than did the Italian bees (avg. 6.3 m vs. 2.5 m) (Echazarreta and Paxton, unpublished data). Each colony fully occupied one deep British standard hive body and filled most of one shallow hive super above it for honey stores, and contained healthy levels of eggs and brood throughout the experiment. Experiment I. Frames of capped brood ready to emerge were removed from both colonies and placed overnight in an incubator at 32-34°C. The newly emerged workers were collected the following day, marked, lightly sprayed with sugar syrup, and reintroduced into their colonies of origin. The Italian colony received 842 marked 1989 STINGING BEHAVIOR 221 workers, and the British colony received 504 marked workers. All 1,346 bees were returned to their natal colonies on the same day in order to prevent environmental variables that fluctuate from day to day from affecting the experimental bees differ- ently. One hundred of the workers placed in each colony were individually marked on their thoraxes with colored and numbered tags. The other marked bees were color- coded with ink on their thoraxes as an age cohort. Workers were removed from the colonies at intervals to test for their willingness to sting. The intervals were irregular because the colonies were not sampled during rainy periods, and because of an intentionally long period at the end of the experiment so that bees well into their foraging lives could be tested. An initial 4 day period was given for workers to be accepted and acclimated to their colonies. Twenty British and 20 Italian bees were then removed from their colonies at 5, 7, 10, 12, 14, 18, 20, 21, 24 and 31 days of age. Workers were tested for their willingness to sting by being gently removed from their colony with forceps, placed individually between the lid of a small plastic petri plate (57 mm diameter) and a 7.6 cm x 12.7 cm index card, and allowed to remain there for several minutes until there was no apparent disturbance among them. Each petri plate assembly and bee in turn was then gently transferred to the test apparatus, and the index card was slid out from beneath the bee. The new surface that the bee stood on consisted of a grid of parallel steel wires of approximately 2 mm diameter and 3.5 mm interwire spacing. Under the wires was a floor consisting of a prewashed black suede target. Each suede target was used for testing one bee, removed, and washed and aired for at least a day before being used on subsequent trials. Testing bees individually allowed us to avoid the type of group-size effect known in metabolic responses to alarm pheromones (Southwick and Moritz, 1985; Moritz, Southwick and Breh, 1985; Moritz and Burgin, 1987). The wires were arranged so that they alternated positive and negative polarity. A bee standing on any two adjacent wires completed a circuit. The positive and negative connections ran to a Coutant 200.2 DC power supply. A potential difference between the positive and negative wires beginning at 0 volts was increased by 1 volt per second whenever a bee was in contact with two adjacent wires. One observer watched the bee to determine when contact was being made, and a second observer slowly increased the voltage whenever it was appropriate to do so. The maximum current flow through the system was preset at 50 milliamps. The individual bees were free to climb onto the wall or ceiling of the petri plate lid, groom themselves, and generally to behave naturally within the limits of their confinement. This ability of the bees to move off of the flooring lengthened the time required to test each bee considerably, but it was felt to be compensated for by the more natural behavior that it allowed. At some point a threshold voltage was reached and the test bee suddenly stung downwards into the suede target with a characteristic flexion of its abdomen and a braced stance. This stinging posture was identical to the typical stinging behavior familiar to anyone who has worked with honey bees. A total of 400 known-aged bees (200 British and 200 Italian) were tested using this procedure. Worker bees sting more readily in defense of their hive than at flowers, and they are attracted to sting by dark colors and coarse surface textures (Free, 1961). In these regards, presenting a black suede target to a bee collected at its colony provides a good representation of the situation present when a bee defends its hive from a (typically) dark-colored 222 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Fig. 1 . Threshold stimuli required to elicit stinging behavior in the British subspecies, Apis mellifera mellifem. Values are expressed as means with standard deviation bars. A total of 200 bees were bioassayed at the various ages indicated. mammalian threat. See Kolmes and Fergusson-Kolmes (in press) for further descrip- tion and discussion of this method. The age at which workers began foraging was determined by observing the hive entrances. Observations were carried out at worker ages of 3, 5, 7, 10, 12, 14, 18, 20, 21, 24, 27, 28, and 31 days. Irregular sampling intervals were due to variable weather, with observations not being carried out on rainy days in order to avoid humidity effects (Collins, 1981). Entrances were blocked with a wire screen for 15 min and then observed for an additional 15 min while the entrances remained blocked. The first foraging trip was considered to be the date that an individually marked worker was initially seen outside the hive during entrance observations. Although a small number of these flights may have been orientation flights rather than foraging flights, previous studies have indicated that error due to this factor is not significant (Winston and Katz, 1982; Winston and Punnett, 1982; Winston and Fergusson, 1985, 1986; Fergusson and Winston, 1988). Data were analyzed using Wilcoxon matched-pairs signed-ranks tests. Chi-squared 2x2 contingency tests corrected for continuity, Mann- Whitney U tests, Kruskal- Wallis one-way ANOVAs, and Chi-squared tests for two independent samples (Sie- gel, 1956). All of these are nonparametric statistics that do not make the assumptions of normal distributions or homogeneity of variance among the data sets. Experiment II. Workers returning to the Italian (48 bees) and British (48 bees) colonies were captured with forceps, and individually isolated. Only foragers return- ing to their colony without pollen loads were collected, because of the considerably increased difficulty in evaluating the worth of simultaneous pollen and nectar loads. The bees were tested for their threshold voltages required to elicit stinging as already described. After this procedure, bees were rendered unconscious with carbon dioxide, and the contents of their honey stomachs were drawn from their mouths into a micropipette by pressure on their abdomen (Gary and Lorenzen, 1976) followed by 1989 STINGING BEHAVIOR 223 dissection to ensure that all of the contents had been gathered. The difference between capturing these workers as they alighted at the hive entrance, and removing workers from frames within the hive in Experiment I, precluded direct comparisons of thresh- old voltages measured in the two experiments. The volume of each honey stomach’s contents was measured by determining what portion of a micropipette’s length was filled (4 cm = 20 microliters). Sugar concen- tration of each honey stomach’s contents was measured in sucrose equivalents with a refractometer. Data were analyzed in order to examine the relationship between the value of the nectar load being brought back to the colony and the threshold stimulus required to elicit stinging. The data were analyzed 3 ways, as threshold stimulus vs. (a) microliters being carried, (b) micrograms of sucrose equivalents per microliter in the honey stomach contents, and (c) total micrograms of sucrose equivalents in the nectar load (concentration x volume). A fourth analysis compared the volume of nectar loads to their concentration in sucrose equivalents, to see whether bees were discriminating in their foraging as might be expected (Schmid-Hempel, Kacelnik and Houston, 1985; Wells and Giacchino, 1968; Seeley, 1986). Bees with empty honey stomachs were often foragers marked visibly with Impatiens pollen, but with no nectar load at the moment of capture. These empty bees were excluded from analyses involving nectar concentrations. Kendall Rank Correlation Coefficients (Siegel, 1956) were used in data analysis. RESULTS Experiment I. Stinging behavior measurements for individual workers of the two subspecies of honey bees differed in a fashion consistent with the dark bouncing suede ball test results already mentioned. British bees stung at lower threshold stim- ulus levels than Italian-derived bees, with very significant differences between them when data from both colonies at every age date were compared with a Wilcoxon matched-pairs signed-ranks test (sum of less frequent ranks = 0, P < .005) (Figs. 1, 2). British bees were, overall, more defensive. The stinging behavior of each subspecies of bee varied with age. Italian bees had threshold stimuli required to elicit stinging ranging from 10.88 ± 2.38 volts (mean ± standard deviation) at 5 days of age to 7.60 ± 1.87 volts at 18 days of age (Fig. 2). British bees had threshold stimuli required to elicit stinging range from 9.23 ± 3.50 volts (mean ± standard deviation) at 5 days of age to 6.80 ± 1.70 volts at 20 days of age (Fig. 1). The overall picture for both subspecies is of an initially declining threshold stimulus required to elicit stinging, to a minimum value around 18 or 20 days of age. British bees demonstrated only a slight increase in threshold stimuli required to elicit stinging after this age, up to 7.23 ± 2.25 volts at 31 days of age. Italian bees showed a considerably greater rebound in their threshold stimuli required to elicit stinging, reaching 10.43 ±3.16 volts at 31 days of age. Both British and Italian bees therefore were less likely to sting when young, and became more likely to sting at intermediate ages. Only the Italian bees subsequently became considerably less likely to sting as they aged further. The age at which worker bees demonstrated their greatest willingness to sting coincided with the period immediately prior to their average age of first foraging. British bees had an average age of first foraging of 20.52 ± 6.39 days (mean ± 224 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) AGE IN DAYS Fig. 2. Threshold stimuli required to elicit stinging behavior in the Italian subspecies, Apis mellifera ligustica. Values are expressed as means with standard deviation bars. A total of 200 bees were bioassayed at the various ages indicated. standard deviation) while Italian bees began to forage at 20.03 ± 5.60 days of age. When the first foraging ages for the 56 bees for which data was collected were categorized according to their subspecies and compared using a Mann- Whitney U test, the ages of first foraging for representatives of the two subspecies did not differ significantly from one another (U = 315, P = .3734, 2-tailed test). In order to determine whether the age-related variability of threshold stimuli required to elicit stinging in both British and Italian bees was significant, data for each subspecies were analyzed separately using a Kruskal- Wallis one-way analysis of variance. This approach asked whether the differences between the samples (thresh- old data for different ages) for each subspecies were great enough to indicate that they were drawn from different populations (i.e., if bees of the same subspecies at different ages demonstrated significantly different threshold stimuli for stinging). For both British and Italian bees, the age-related differences proved to be significant. The one-way ANOVA results for British bees were H (adjusted for ties) = 17.1 1 {P < .05, 9 degrees of freedom). The one-way ANOVA results for Italian bees were H (adjusted for ties) = 46.03 {P < .001, 9 degrees of freedom). Both British and Italian bees did vary significantly in their likelihood of stinging according to their age. Experiment II. The British and Italian bees proved to be similar in two regards in the relationships of their stinging responses to the characteristics of the nectar loads they were bearing back to their colonies. First, neither type of forager dem- onstrated a significant association between their threshold stimulus to elicit stinging and the concentration of their nectar load in micrograms of sucrose equivalents per microliter (Fig. 3) ((a) British bees: Kendall Rank Correlation Coefficient = .1530, P=.1131,N = 31, 1 -tailed test; (b) Italian bees: Kendall Rank Correlation Coefficient = -.1010, P = .2266, N = 28, 1-tailed test). Second, both types of forager demon- strated a significant positive association between the volume of their nectar load in microliters and the nectar concentration in micrograms of sucrose equivalents per 1989 STINGING BEHAVIOR 225 VOLTAGE VS. MICROGRAMS/ MICROLITER QC UJ O QC O (0 □ BRIT, UG/UL ♦ ITAL, UG/UL Fig. 3. Threshold stimuli required to elicit stinging behavior for two subspecies of foragers, plotted with respect to the concentration of nectar in their honey sacs expressed as micrograms/ microliter of sucrose equivalents. microliter (Fig. 4) ((a) British bees: Kendall Rank Correlation Coefficient = .2776, P = .0 1 43, N = 3 1 , 1 -tailed test; (b) Italian bees: Kendall Rank Correlation Coefficient = .3184, P = .0087, N = 28, 1 -tailed test). The foragers did not modify their stinging behavior due to differences in micrograms of sucrose equivalents per microliter of nectar borne by them. The foragers did gather a greater volume of more concentrated nectar, demonstrating the expected discrimination. The British bees displayed stinging behavior that varied with their nectar loads in two significant fashions. There was a significant positive association between thresh- old stimuli required to elicit stinging and total micrograms of sucrose equivalents carried (Fig. 5) (Kendall Rank Correlation Coefficient = .1919, P = .0274, N = 48, 1 -tailed test). There was also a significant positive association between threshold stimuli required to elicit stinging and microliters of nectar carried (Fig. 6) (Kendall Rank Correlation Coefficient = .1895, P = .0287, N = 48, 1 -tailed test). The Italian bees differed by not demonstrating a significant association between threshold voltage required to elicit stinging and either total micrograms of sucrose equivalents carried (Fig. 5) (Kendall Rank Correlation Coefficient = .0833, P = .2005, N = 48, 1 -tailed test) or microliters of nectar carried (Fig. 6) (Kendall Rank Cor- relation Coefficient = .0520, P = .3015, N = 48, 1 -tailed test). British bees therefore appeared to modify their stinging behavior in terms of their foraging suceess in significant ways that Italian bees did not. DISCUSSION Residual value theory has proven to be a useful model to predict changes in the stinging behavior of some honey bees. Workers from the colony of British bees {Apis 226 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) MICROLITERS VS MICROGRAM/MICROLITER CL o cc o (f) cc LU O CC o MICROGRAMS SUCROSE EQUIV PER MICROLITER BRIT, UL ITAL, UL Fig. 4. The volume in microliters of nectar loads carried in the honey sacs of two subspecies of foragers, plotted with respect to the concentration of nectar in their loads expressed as micrograms/microliter sucrose equivalents. mellifera mellifera) appeared to sting in a residual-value sensitive fashion in two ways: (a) they sting more readily when carrying fewer microliters of nectar, and (b) they sting more readily when older than approximately 10 days of age (Figs. 1, 6). The significant positive association between threshold stimuli to elicit stinging and total micrograms of sucrose equivalents carried (Fig. 5) is probably a function of the relationship already mentioned between threshold stimuli and microliters carried, because total micrograms of sucrose equivalents are calculated in part on the basis of microliters carried. There was no independent evidence that workers of this sub- species modified their stinging behavior on the basis of the concentration of their nectar loads alone (Fig. 3). In contrast, Italian bees {Apis mellifera ligustica) displayed none of the modifica- tions of their readiness to sting that are consistent with residual value theory. They possess a high threshold stimulus to elicit stinging at an advanced age (Fig. 2) and have no significant associations between stinging behavior and either nectar load size, concentration, or total net load (Figs. 3, 5, 6). The workers of this subspecies were both less willing to sting overall and appeared to be less residual value sensitive than the workers of A. m. mellifera. Increased locomotion and wing flicking among groups of caged bees presented with alarm pheromones (Collins, 1980), electroantennagram responses to alarm phero- mones (Allan, Slessor, Winston and King, 1987), and increases in oxygen consump- tion among groups of caged workers exposed to alarm pheromones (Moritz, South- wick and Breh, 1985) have all been used to examine how groups of bees of various ages react to chemicals released when a hive is threatened. In all instances the bees reacted more strongly at intermediate ages than immediately after emergence, and 1989 STINGING BEHAVIOR 227 VOLTAGE VS. MICROGRAMS BRIT, MICROGR ITAL, MICROGR Fig. 5. Threshold stimuli required to elicit stinging behavior for two subspecies of foragers, plotted with respect to the total amount of carbohydrate in their honey sacs expressed as micrograms of sucrose equivalents. reactivity either reached a plateau or subsequently declined. These group data are consistent with the sorts of stinging response measures reported in our data. Both the British and Italian workers were significantly more likely to collect larger nectar loads when foraging at more concentrated floral resources (Fig. 4). This is consistent with what we know about the sensitivity of honey bees in general to varying environmental resources (Schmid-Hempel, Kacelnik and Houston, 1985; Seeley, 1986). Workers of the two subspecies also began to forage at the same age, which is consistent with data suggesting that foraging onset is controlled in part by sensitivity to environmental circumstances (Kolmes, 1985a). Apparently British and Italian bees differ from one another in their stinging behavior, but not in their foraging behavior in the manner of European vs. Africanized bees (Winston and Katz, 1982). The significant positive association between the size and concentration of nectar loads for bees foraging on natural floral resources (Fig. 4) is consistent with reports by von Frisch (1934, 1965, 1 97 1) that workers collect larger nectar loads from artificial feeders when the sucrose solution is more concentrated. Both the data collected from foragers on natural floral sources (Fig. 4) and those of von Frisch (1934, 1965, 1971) are contradictory to those reported by Wells and Giacchino (1968) which showed no relationship between nectar load volume and concentration. The short distance (50 m) between hive and feeders in Wells and Giacchino’s study (1968) may have resulted in more typical discriminatory foraging behavior breaking down. To further examine honey bee stinging behavior in terms of residual value theory, a number of questions must be answered. These include: (a) How variable are different colonies of both subspecies in their stinging behavior and in its modifiability? Stinging behavior may be related to the annual colony life cycle of honey bees, especially as individual workers may be of greater value to newly founded colonies than to older 228 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) VOLTAGE VS. MICROLITERS Q. o a: o o cc o □ BRIT. MICROL ♦ ITAL, MICROL Fig. 6. Threshold stimuli required to elicit stinging behavior for two subspecies of foragers, plotted with respect to the volume carried in their honey sacs expressed in microliters. populous colonies, (b) Do these two subspecies differ in their ages of guard duties? Although both subspecies have similar minimum threshold stimuli to elicit stinging immediately prior to their average age of first foraging, the British bees maintain approximately this level of stinging threshold while the Italian bees have a declining readiness to sting at a later age. This could be reflected by a broader age-range of guarding behavior among A. m. mellifera workers, if threshold stimuli required to elicit stinging are motivationally related to guarding behavior, (c) Is there any influ- ence of the value of a floral resource in terms of pollen on stinging behavior? The data presented in this paper dealt only with nectar foragers because of the difhculties inherent in quantifying the values of pollen loads or of mixed pollen and nectar loads, (d) Would data that included more information about foraging workers (e.g., distance to floral resources; numbers of round trips per hour; precise ages of workers; amount of competition for resources with workers from other colonies; worker life expectancy) demonstrate a stronger residual value sensitivity for workers of either subspecies? (e) Are workers of additional subspecies of honey bees, such as African- ized bees, sensitive to their residual value in their stinging behavior? (f ) Do workers of other social insects possess residual value sensitivity in their defensive behavior? Finally, residual value sensitivity in stinging behavior may be important to in- corporate into general models of colony defense (Collins, Rinderer, Tucker, Sylvester and Lackett, 1980) for some if not all honey bee subspecies. It may be that variable behavioral responsiveness to stimuli eliciting stinging are related to sensory changes with age (Allan, Slessor, Winston and King, 1987). The relationship between readiness to sting of workers at various ages and guarding behavior (Moore, Breed and Moor, 1987) also remains to be elucidated. This first examination of stinging behavior by worker honey bees in terms of residual value theory is a small beginning compared to the more fully developed empirical evidence in the ornithological literature (Curio, 1989 STINGING BEHAVIOR 229 1987; Curio, Regelmann and Zimmermann, 1984, 1985; Pianka and Parker, 1975; Curio, Klump and Regelmann, 1983; Windt and Curio, 1986; Regelmann and Curio, 1986, 1983) but it may provide a useful approach to studying and predicting the behavior of the social insects. ACKNOWLEDGMENTS This research was conducted at the Bee Research Unit, Department of Zoology, University College, Cardiff, Wales CFl IXL, UK. We thank Professor R. S. Pickard and the Bee Research Unit, of which he is Head, for so generously acting as hosts during the course of this research, R. J. Paxton and other inhabitants of the Black Box, too numerous to individually name, and Professor J. B. Free provided us with help and stimulating conversation. R. J. Paxton and C. Echazarreta supplied us with the “bouncing ball” data for the Cardiff apiary. This work was supported through a Junior Leave for SAK and other research funds provided by the Hewlett and Mellon Foundations and Hobart and William Smith Colleges. The staff of the International Bee Research Association head- quarters in Cardiff kindly assisted with library resources. We thank two anonymous reviewers for their helpful comments. LITERATURE CITED Allan, S. A., K. N. Slessor, M. L. Winston and G. G. S. King. 1987. The influence of age and task specialization on the production and perception of honey bee pheromones. J. Insect Physiol. 33:917-922. Collins, A. M. 1980. Effect of age on the response to alarm pheromones by caged honey bees. Ann. Entomol. Soc. Am. 73:307-309. Collins, A. M. 1981. Effects of temperature and humidity on honeybee response to alarm pheromones. J. Apic. Res. 20:13-18. Collins, A. M., T. E. Rinderer, K. W. Tucker, H. A. Sylvester and J. J. Lackett. 1980. A model of honeybee defensive behaviour. J. Apic. Res. 19:224-231. Curio, E. 1987. Brood defence in the great tit: the influence of age, number and quality of young. Ardea 75:35-42. Curio, E., G. Klump and K. Regelmann. 1983. An anti-predator response in the great tit (Parus major): is it tuned to predator risk? Oecologia (Berl.) 60:83-88. Curio, E., K. Regelmann and U. Zimmermann. 1984. The defense of first and second broods by great tit {Parus major) parents: a test of predictive sociobiology. Z. Tierpsychol. 66: 101-127. Curio, E., K. Regelmann and U. Zimmermann. 1985. Brood defense in the great tit {Parus major): the influence of life history and habitat. Behav. Ecol. Sociobiol. 16:273-283. Erickson, E. H., Jr., S. D. Carlson and M. Garment. 1986. A Scanning Electron Microscope Atlas of the Honey Bee. Iowa State University Press, Ames. Fergusson, L. A. and M. L. Winston. 1988. The influence of wax deprivation on temporal polyethism in honey bee colonies. Can. J. Zool. 66:1997-2001. Free, J. B. 1961. The stimuli releasing the stinging response of honeybees. Anim. Behav. 9: 193-196. Free, J. B. 1963. The flower constancy of honeybees. J. Anim. Ecol. 32:1 19-132. Free, J. B. 1966. The foraging areas of honeybees in an orchard of standard apple trees. J. Appl. Ecol. 3:261-268. Free, J. B. 1970. Effect of flower shapes and nectar guides on the behaviour of foraging honeybees. Behaviour 7:269-285. Frisch, K. von. 1934. Uber den Geschmacksinn der Bienen. Z. vergl. Physiol. 21:1-156. Frisch, K. von. 1965. Tanzsprache und Orientierung der Bienen. Springer- Verlag, Berlin. 230 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) Frisch, K. von. 1971. Bees, Their Vision, Chemical Senses, and Language, Revised Edition. Cornell University Press, Ithaca. Gary, N. E. and K. Lorenzen. 1976. A method for collecting the honey-sac contents from honeybees. J. Apic. Res. 15:73-79. Grant, V. 1950. The flower constancy of bees. Bot. Rev. 16:379-398. Hamilton, W. D. 1 964. The genetical theory of social behavior I and II. J. Theor. Biol. 7:1-52. Jeanne, R. L. 1986. The evolution of the organization of work in social insects. Monitore Zool. Ital. (N.S.) 20:119-133. Jones, C. E. 1978. Pollinator constancy as a pre-pollination isolating mechanism between sympatric species of Cercidium. Evolution 32:189-198. Jones, C. E., C. L. Scannell, K. J. Kramer and W. E. Sawyer. 1986. Honeybee constancy to ultraviolet floral reflectance. J. Apic. Res. 25:220-226. Kolmes, S. A. 1 985a. A quantitative study of the division of labor among worker honey bees. Z. Tierpsychol. 68:287-302. Kolmes, S. A. 1985b. An information-theory analysis of task specialization among worker honeybees performing hive duties. Anim. Behav. 33:181-187. Kolmes, S. A. 1985c. An ergonomic study of Apis mellifera (Hymenoptera: Apidae). J. Kans. Entomol. Soc. 58:413-421. Kolmes, S. A. 1986. Have hymenopteran societies evolved to be ergonomically efficient? J. New York Entomol. Soc. 94:447-457. Kolmes, S. A. and L. A. Fergusson-Kolmes. In press. Measurements of stinging behaviour in individual worker honeybees {Apis mellifera). J. Apic. Res. Levin, M. D. 1966. Orientation of honeybees in alfalfa with respect to landmarks. J. Apic. Res. 5:121-125. Menzel, R. and J. Erber. 1972. The influence of the quantity of reward on the learning performance in honeybees. Behaviour 41:27-42. Menzel, R., J. Erber and T. Masuhr. 1974. Learning and memory in the honeybee. Pages 195-2 1 7 in: L. Barton Browne (ed.). Experimental Analysis of Insect Behaviour. Springer- Verlag, New York, 366 pp. Moore, A. J., M. D. Breed and M. J. Moor. 1987. The guard honey bee: ontogeny and behavioural variability of workers performing a specialized task. Anim. Behav. 35:11 59- 1167. Moritz, R. F. A. and H. Burgin. 1987. Group response to alarm pheromones in social wasps and the honey bee. Ethology 76:15-26. Moritz, R. F. A., E. E. Southwick and M. Breh. 1985. A metabolic test for the quantitative analysis of alarm behavior of honeybees {Apis mellifera L.). J. Exp. Zool. 235:1-5. Oster, G. F. and E. O. Wilson. 1978. Caste and Ecology in the Social Insects. Princeton University Press, Princeton. Pianka, E. R. and W. S. Parker. 1975. Age-specific reproductive tactics. Am. Nat. 109:456- 464. Primack, R. B. 1985. Longevity of individual flowers. Ann. Rev. Ecol. Syst. 16:15-37. Regelmann, K. and E. Curio. 1983. Determinants of brood defence in the great tit Parus major h. Behav. Ecol. Sociobiol. 13:131-145. Regelmann, K. and E. Curio. 1986. Why do great tit {Parus major) males defend their brood more than females do? Anim. Behav. 34:1206-1214. Ribbands, C. R. 1949. Theforagingmethodofindividualhoney-bees.J. Anim. Ecol. 18:47-66. Schmid-Hempel, P., A. Kacelnik and A. I. Houston. 1985. Honeybees maximize efficiency by not filling their crop. Behav. Ecol. Sociobiol. 17:61-66. Sekiguchi, K. and S. F. Sakagami. 1 966. Structure of foraging population and related problems in the honeybee, with considerations on the division of labour in bee colonies. Hokkaido Agricultural Experiment Station Rep. 69:1-65. 1989 STINGING BEHAVIOR 231 Seeley, T. D. 1986. Social foraging by honeybees: how colonies allocate foragers among patches of flowers. Behav. Ecol. Sociobiol. 19:343-354. Siegel, S. 1956. Nonparametric Statistics for the Behavioral Sciences. McGraw-Hill, New York. Southwick, E. E. and R. F. A. Moritz. 1985. Metabolic response to alarm pheromone in honey bees. J. Insect Physiol. 31:389-392. Wells, H. and P. H. Wells. 1983. Honey bee foraging ecology: optimal diet, minimal uncer- tainty, or individual constancy? J. Anim. Ecol. 52:829-836. Wells, P. H. and J. Giacchino. 1968. Relationships between the volume and the sugar con- centration of loads carried by honeybees. J. Apic. Res. 7:77-82. Windt, W. and E. Curio. 1 986. Clutch defense in great tit {Pams major) pairs and the Concorde fallacy. Ethology 72:236-242. Winston, M. L. and L. A. Fergusson. 1985. The effects of worker loss on temporal caste structure in colonies of the honeybee {Apis mellifera L.). Can. J. Zool. 63:777-780. Winston, M. L. and L. A. Fergusson. 1986. The influence of the amount of eggs and larvae in honeybee colonies on temporal division of labor. J. Apic. Res. 25:238-241. Winston, M. L. and S. J. Katz. 1982. Foraging differences between cross-fostered honeybee workers {Apis mellifera) of European and Africanized races. Behav. Ecol. Sociobiol. 10: 125-129. Winston, M. L. and E. N. Punnett. 1982. Factors determining temporal division of labor in honeybees. Can. J. Zool. 60:2947-2952. Received July 29, 1988; accepted November 5, 1988. NOTES AND COMMENTS J. New YorkEntomol. Soc. 97(2):232-233, 1989 EVALUATION OF THE SPIDER STEATODA TRIANGULOSA (ARANEAE: THERIDIIDAE) AS A PREDATOR OF THE RED IMPORTED FIRE ANT (HYMENOPTERA: FORMICIDAE) Spiders have been shown to be important predators of ants (MacKay, 1 982; Porter and Eastmond, 1982). Seventy-hve percent of the prey of the black widow spider, Latrodectus mactans (F.), consists of the fire ant, Solenopsis invicta Buren, in cotton fields in Texas (Nyffeler et ah, 1988). An additional 16 species of spiders prey on the fire ant (Nyffeler et ah, 1988). The spider genus Steatoda is especially important, consisting of a group of specialized ant predators (Holldobler, 1970; MacKay, 1982). We are currently evaluating the impact of imported fire ants {Solenopsis invicta) on electrical equipment. We have shown an attraction of fire ants to such equipment (MacKay et ah, unpubh). Ants cause considerable damage in such equipment by shorting of circuitry and damaging electrical components (MacKay, 1989; Vinson and MacKay, 1989). A number of species of spiders occur in such equipment and are obviously predators of fire ants as ants occur in their webs. We have observed spiders killing fire ants on two occasions. Steatoda triangulosa (Keyserling) is the most common species; other species include Scytodes sp. (Scytodidae) and Crosso- priza stridulans Millot (Pholcidae, introduced from Madagascar). We visited traffic control signal cabinets in the cities of College Station and Bryan, Texas throughout 1987 and 1988 at monthly intervals. We estimated the numbers of ants in each of fifteen cabinets and counted the numbers of spiders in each cabinet. We tested the hypotheses that at lower densities of fire ants (50 or less/cabinet), the spider population density would be inversely correlated with the fire ant population density due to predation on the ants. At higher densities (over 50 ants/cabinet) the spider population would be positively correlated with the fire ant population due to immigration of spiders to these cabinets. Such movement of spiders to areas of higher ant density have been shown to occur (MacKay, 1982). We found no correlation between spider and ant population densities either at low ant density (Pearson correlation coefficient: = —0.20, P = 0.16, N = 36 for Bryan; = — 0.0 1 , P = 0.94, N = 62 for College Station) or at higher densities (Pearson correlation coefficient = 0.16, P = 0.50, N = 19, data lumped for both cities), indicating that neither population had any statistically significant effect on the other. We conclude that spiders, especially S. triangulosa prey on fire ants in electrical equipment, although the effect of this predation is not statistically significant. Spiders were removed or disturbed each time a cabinet was checked, which may account to some extent for the lack of impact of spiders on ants. We suggest that whatever practice is employed to control ants in such equipment, the spider population should 1989 NOTES AND COMMENTS 233 be protected, as spiders do kill ants, and they do little or no harm to the equipment. — William P. MacKay and S. Bradleigh Vinson, Dept, of Entomology, Texas A&M University, College Station, Texas 77843. ACKNOWLEDGMENT We thank the Texas State Department of Highways and Public Transportation for supporting the research. Allen Dean identified the spiders and critically reviewed the manuscript. Approved as number TA-24344 by the Texas Agricultural Experiment Station. LITERATURE CITED Holldobler, B. 1 970. Steatoda fulva (Theridiidae), a spider that feeds on harvester ants. Psyche 77:202-208. MacKay, W. P. 1 982. The effect of predation of western widow spiders (Araneae: Theridiidae) on harvester ants (Hymenoptera: Formicidae). Oecologia 53:406-41 1. MacKay, W. P. 1989. The impact of the red imported fire ant on electrical equipment. In: S. B. Vinson (ed.). The Economic Impact of the Red Imported Fire Ant. Governer’s Fire Ant Conference. Sportsman Conservationists of Texas, Austin, Texas, in press. Nyffeler, M., D. A. Dean and W. L. Sterling. 1988. The southern black widow spider, Latro- dectus mactans (Araneae, Theridiidae), as a predator of the red imported fire ant, Sole- nopsis invicta (Hymenoptera, Formicidae), in Texas cotton fields. J. Appl. Entomol. 106: 52-57. Porter, S. and D. Eastmond. 1982. Euryopis coki (Theridiidae), a spider that preys on Po- gonomyrmex ants. J. Arachnol. 10:275-277. Vinson, S. B. and W. P. MacKay. 1989. Effects of the fire ant, Solenopsis invicta, on electrical circuits and equipment. In: K. Jaffe and R. Vander Meer (eds.). The Current Status of Pest Ants. Westview Press, Boulder, Colorado, in press. BOOK REVIEWS /. New YorkEntomol. Soc. 97(2):234-241, 1989 CLADISTICS IN THE FAST LANE Hennig86. Version 1.5.— J. S. Farris. 41 Admiral Street, Port Jefferson Station, New York 1 1776. $50. The advent of cladistic philosophy and methods has given systematics a more active, one might even say fundamental, role in the general framework of comparative biology. With emphasis on establishing only monophyletic taxonomic groups on the basis of synapomorphy and reflecting relationships in the form of cladograms, cla- distics necessitates explicit formulation of hypotheses and results. Concurrent with the widespread acceptance of these tenets has been the development of a more critical protocol for character and character state elucidation, coding, and analysis. This shift toward greater empiricism, coupled with parsimony as the ultimate arbiter in clado- gram selection, has certainly placed a burden on the systematist by requiring clado- grams to represent character state distributions as accurately as possible. Systematists are, however, constrained in their ability to construct cladograms by hand since the possible number of equally parsimonious cladograms rises dramati- cally with the addition of taxa and/or characters. Even a general attempt at manual construction will be precluded by the sheer number of characters traditionally rec- ognized, for example, in arthropod or vertebrate groups. The simultaneous evolution of computers and cladistics packages has seen a trend from relatively inaccessible mainframe programs to the large-scale distribution of microcomputer versions, such as PHYLIP and PAUP, developed by Joseph Felsen- stein and David Swofford, respectively. Recently, a series of empirical comparisons have been reported for various mainframe (e.g., Luckow and Pimentel, 1985) and PC versions (e.g., Fink, 1986; Platnick, 1987, 1988, in press; see also Coddington, 1987). With such publicity, coupled with rumors of “this new version” or “that new program” about to be released, one gets the impression that we are in the midst of an event which will be of great benefit to systematists: a programming race to produce faster algorithms for finding all minimum-length trees, yet which are compatible on a variety of PC’s. The latest contender in this race is Hennig86, version 1.5, developed by J. S. Farris (1988) for MS-DOS, IBM-compatible PC’s. This review is not intended to make empirical comparisons of features or results from Hennig86 with those, for example, from PHYLIP or PAUP. The most current information on these aspects has been prepared by Platnick (in press) as an update of his earlier analyses (Platnick, 1987). Hennig86 is a surprisingly compact program (49K), yet extremely powerful and impressive in its tree-building capabilities, speed, and extensive tree/data-manipu- lating commands. Further, it does not require a math co-processor. The distribution disk contains three files: 1) ssxom, which is Hennig86 per se; 2) dox, a command help file which can be accessed while working within the program; and 3) peg, a sample character state matrix. About 5 1 2K of RAM are required for its operation. 1989 BOOK REVIEWS 235 Hennig86 is fully interactive, providing a variety of cladogram and character editing features. Cladograms are generated by strict parsimony analysis. There is, however, no facility for implementing any form of Dollo or Camin-Sokal parsimony (ala PAUP; Swolford, 1985). Workers familiar with the PHYSYS mainframe program, developed by J. S. Farris and M. F. Mickevich (Mickevich and Farris, 1984), will hnd much of the operation of Hennig86 very familiar. Those more familiar with any of the other available programs will probably come to fine the ease, logic, and (best of all) speed of Hennig86 to be extremely gratifying. The documentation for Hennig86 might seem sparse, comprising only 15 text pages divided into 23 sections. This and its rather terse wording will likely be intim- idating to some. Initial apprehensions aside, the user should find all instructions and examples quite comprehensible. Most importantly, the documentation is arranged such that descriptions of commands and internal program files in one section will usually have a direct bearing on how one or a series of commands in a later section can be successfully initiated. With each section dependent information-wise upon earlier sections, it is best to initially proceed through the documentation in sequence, sparing one the frustration of having to continually backtrack to determine why a command will not work. Also, working through each of the documentation’s examples with the sample data matrix, peg, is quite helpful. Indeed, the documentation’s idiosyncratic style is especially effective in prodding the user into experimenting with various commands and options, reducing the degree to which one might be inclined to simply view Hennig86 as just another “black box” program. This makes Hennig86 a must, not only for the established systematist, but for students as well. Several sections, however, should probably be expanded to include additional examples as well as further details about applications, interpreting results, and avoiding pitfalls. Data matrix files can be entered into Hennig86 as DOS or ASCII word-processing files. While the documentation describes the format for setting up a data matrix, users should examine the contents of peg for a good example. As noted earlier, peg is also a good sample data matrix with which to explore all available options. Hennig86 can accept from 1 to 999 characters and 4 to 180 taxa. Character states must be integer coded. In the edition of 1.5 reviewed here (obtained in August 1988), the number of states for a character is limited to the range 0-9. A future update of 1.5 will extend this range to about 36 states (Farris, pers. comm.). Missing or unknown data are allowed, coded as “?” or Characters can be differentially weighted, with weights ranging from 0 to 100 (the default value is 1 ; also see discussion below on successive approximations weighting). Multistate characters may be treated as additive (“ordered”) or non-additive (“unordered”). In the additive form, transfor- mation series can only be arranged linearly. Input of user-defined branching character state trees will be available in a later version for those who wish to explore this option. In the meantime, one can achieve the same end through a variety of additive coding methods (e.g., Pimentel and Riggins, 1987; O’Grady and Deets, 1987). Poly- morphic characters per se cannot be designated. The ccode command enables the user to delete, weight, or change the additivity of characters for a given analysis. Unfortunately, there is no similar command for temporarily deleting taxa. The documentation does not explicitly forewarn users of one minor detail which could cause initial problems. Hennig86 always numbers input taxa and characters, internal program files, and all output listings in a consecutive manner starting with 236 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) 0. Thus, for example, if one decides to input a data matrix with the fifth character weighted higher than the default value, this character must be referred to as character 4 in the weighting command. Hennig86 offers a wide range of cladogram-calculating commands based on “exact” and “approximate” algorithms. When multiple minimum-length cladograms are gen- erated, only unique topologies are retained, i.e., redundant cladograms with unsup- ported or zero-length branches are collapsed to show all polytomies. Cladograms must always be rooted by at least one outgroup taxon and there is the option of designating any number of additional taxa as secondary outgroups. A command for rerooting cladograms is also available (reroot, described below). The “exact” algorithm, ie (“implicit enumeration”), will find all minimum-length cladogram(s), but its success may be dependent upon the number of final cladograms that are saved by a particular option and/or the amount of available memory. Based upon the particular option selected with ie, the number of final cladograms retained can be limited to 1, may go up to 100, or all available memory may be used. Because of the exhaustive search strategy performed by the ie command, its use may be prohibitive timewise due to size of the data set, amount of homoplasy, and/or size and speed of the computer’s microprocessor. This is a matter of how long the user wants to tie up the machine, especially if it is not multi-tasking. The bottom line is that Hennig86 does not discriminate on the basis of data size when it comes to how extensively one wants a search to be executed. But, even on my Toshiba lap- top, with a 9.54 MHz 80C86-1 microprocessor, I have been able to run relatively large data sets very quickly. There are two “approximate” algorithms, each with several options. The least effective of these, hennig, makes a single pass through the data, constructing one cladogram, which may not be of minimum length. Limited branch-swapping can be applied to this cladogram, but again, only a single cladogram is retained. The com- mand, mhennig, constructs several initial cladograms, each by a single pass, but adds taxa in several different combinations, saving all minimum-length cladograms. Lim- ited branch-swapping can be performed on these cladograms with mhennig*. For very large or messy data sets, the only feasible approach to obtaining optimal or near-optimal results in a timely fashion is mhennig* in combination with bb or bb*. The bb command performs extended branch-swapping on all cladograms generated from mhennig*, saving all cladograms it can find up to a limit of 100. The number of cladograms retained can be upgraded to the limit of available memory by using bb* . The efficiency of mhennig* with bb* to find all most parsimonious cladograms appears to be quite good (Platnick, 1988, pers. comm.). If I were to order the search strategies from best to worst from the standpoint of finding as many minimum-length cladograms as possible, I would suggest the fol- lowing: 1) ie*, 2) ie, 3) ie- with bb or bb*, 4) mhennig* with bb or bb*. Similar suggestions are made by Farris (1988; see also Platnick, 1988 for comparisons of results). Again, the trade-off is the possibility of not finding all minimum-length cladograms for the benefit of a shorter run time. Cladograms can be output in the form of branching diagrams or in parenthetical notation, and can be examined as output directly from the monitor and/or saved to a disk file. Upon obtaining results, one can select, with the tchoose command, a particular group of cladograms which can be further examined on the monitor or 1989 BOOK REVIEWS 237 saved to a disk file using the tsave command. A particularly nice option when printing out cladograms is that branches can be displayed using extended ASCII symbols (the default), which could conceivably make diagrams of publishable quality. Moreover, cladograms are diagrammed such that taxa are placed on consecutive, single-spaced lines, considerably reducing the amount of printout. Cladograms can be diagnosed to varying degrees with the xsteps command in conjunction with a series of specified options. Diagnoses can include, for example, length, consistency (Cl) and retention indices (r^) of each cladogram, Cl and of each character, number of steps required for each character on each cladogram, best and worst fits (Cl and r,) of each character for a set of cladograms, and all possible states at the nodes of each cladogram (i.e., hypothetical ancestral states). The retention index, developed by Farris for Hennig86, is a measure of the ability of a character to function as a synapomorphy relative to the overall consistency of that character. The index is not described in the documentation and a formal de- scription has not yet been published. The index is defined as = (hj-Si)/(hi-li), where h, is the largest number of steps possible for character i on any cladogram topology for a given set of taxa, 1, is the smallest number of possible steps, and S; is the observed number of steps for the actual cladogram (Farris, pers. comm.). An r; of 1 denotes a character which is completely consistent on the cladogram and with at least one state acting as a synapomorphy, whereas a value of 0 indicates unique character state changes limited only to terminal taxa. Values less than 1 and greater than 0 indicate some degree of homoplasy or reversal. Since the is sensitive to the number of states acting as synapomorphies it will not always correspond to the Cl. In the event there are different equal-length transformation series for a particular character, including equal possibilities of reversal or parallelism, Hennig86 will au- tomatically list all possible states that that character can manifest at a given node. This is comparable to output from the CSPOSS command in PAUP (Swofford, 1985) but is presented in a more concise manner in Hennig86. A graphic represen- tation of character state ambiguity is provided by the tree editor, Dos Equis (see below). All diagnostics are printed out as very compact tables which take up as little space as possible. This is not only a convenience when having to examine reams of printout, but, like the cladograms, makes for ease of examination directly from a monitor. My only complaint is that there is no comparable listing for unique state changes (e.g., a reversal or loss) occurring in terminal taxa. Autapomorphies in terminal taxa can be detected by comparing the listing of total step changes for a character with that character’s total number of steps at the nodes. A discrepancy indicates that terminal-taxon changes have occurred. One will then have to examine the original data matrix to find which taxa have uniquely derived the condition, then map the change on the cladogram(s). This is actually not much of an inconvenience, but should be watched when mapping character states onto cladograms. Another method of searching for terminal taxon changes is with the tree editor, Dos Equis (see below). Farris (pers. comm.) is in the process of making it easier to account for autapomor- phies with the xsteps command. The command, nelsen, calculates a Nelson consensus tree (Nelson, 1979; Nelson and Platnick, 1981) of all cladograms from a given tree file. Hennig86 also allows for a successive approximations weighting procedure (Farris, 238 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) 1969; see also Carpenter, 1988), which is effective in reducing the number of equal- length cladograms by an iterative series of a posteriori weightings. This has the effect of choosing the cladogram(s) with the most consistent (i.e., cladistically reliable) characters. Successive approximations weighting affords one the opportunity to re- duce the number of cladograms that must be inspected. This might be useful if one is more concerned with getting patterns of relationship based on as few cladograms as possible without sacrificing character support, or deleting cladograms on the basis of a priori assumptions. In this procedure, each character in the initial set of cladograms is assigned a weight, scaled between 0 and 10. Weights are calculated by the xsteps command with the w option as products of the highest Cl and r, values as determined from the best fits statistics. The data matrix is rerun with characters weighted accordingly. New weights are then calculated from the new cladogram(s), applied again to the original data, and rerun. The procedure is terminated at the point in which weights no longer change with each iteration, indicating cladogram topologies are not changing from one run to the next. Unlike PHYSYS, there is no command loop available to automatically switch weights and carry out each run to termination (Carpenter, 1 988). The weighting command in Hennig86, however, makes the task so easy that this is hardly an inconvenience. The weighting function in Hennig86 differs from that in PHYSYS in that weights in the latter are calculated as the mean Cl of each character (Carpenter, 1988). Since mean Cl takes all values into consideration from all cladograms generated, it is probably a stronger weighting function than that in Hennig86. Carpenter (1988) reiterated the suggestion made by Farris ( 1 969) that integer-coded, additive multistate characters should be recoded in additive binary form to avoid uneven weighting and effects of character state dependency. Additive binary coding effectively treats each state independently. This alternative is not feasible in the case of integer-coded, non- additive multistate characters since conversion to a non-additive binary form pre- cludes determination of nodal conditions or transformation series (Farris, pers. comm.) Hennig86 accepts the input of user-defined cladograms by use of the tread com- mand. This has the utility, for example, of taking a published cladogram on which no character support has been shown (which is quite common) and optimizing char- acters from the original data matrix onto this topology. Subsequently, by using the tree editor, Dos Equis (described below), one may interactively examine, edit, and/ or save results of any further manipulations. Another approach to the same problem, however, is to simply edit the cladogram(s) generated from the data matrix by Hennig86, using Dos Equis, to conform to the published topology. Discrepancies between published results and those found by Hennig86 can be readily determined from the diagnostics output from xsteps or Dos Equis. User-defined cladograms to be input into Hennig86 (using the tread command) must be expressed in some form of parenthetical notation. Unresolved groupings are allowed and terminal taxa may be referred to by number or name. There is no need to balance all parentheses, especially in complex asymmetrical topologies. For ex- ample, the expression (((1 2)(3 4))(5 6)) can also be input as 1 2)(3 4))(5 6. Hennig86 is very liberal in the types of symbols that are allowed to delimit groups, i.e., ( ), \ /, [ ], { }, and comma. When used with asymmetrical topologies, each symbol has a given priority level relative to all others such that one symbol will force symbols 1989 BOOK REVIEWS 239 of equal or lower priority to balance out when read. The symbols shown above are ordered from lowest to highest priority. The amount of time and effort needed to input topologies is considerably decreased. Thus, the notation (0[(1 2) \ 3(4(5 6(7 8 is the same as 0((1 2)(3(4(5 6(7 8 and (0((1 2)(3(4(5 6(7 8)))))). One of the examples provided in the Hennig86 documentation (Section 7: Tree Input) is in error; 0[1(2(3 4/6(7(8 9 is said to be equivalent to 0(( 1(2(3 4)))(6(7(8 9)))). The first expression will not designate a sister-group relationship between (1 2 3 4) and (6 7 8 9). The abbre- viated notation should actually be something like 0[ 1(2(3 4][6(7(8 9. Hypothetical (nodal) ancestors can be specified in notation as a number preceded by a period (e.g., .0 or .2), with descent from an ancestor denoted by the connection sign “ — Large or complex clades can then be split apart by use of the comma as a delimiter (see above). This has the utility of making potentially unwieldy expressions easier to handle. For example, the expression .0\.l, .2, .0-0\l 2, .l-3\4 5, .2-6\7, states initially that ancestors .1 and .2 are descended from .0, establishing the sister group (3 4 5)(6 7). The entire grouping is therefore the same as 0\1 2,[3\4 5]\6 7 or (0(1 2)(3(4 5)(6 7))). The process of compressing (or collapsing) zero-length or unsupported branches to polytomies during cladogram construction in Hennig86 can also be applied to a set of user-defined cladograms using the xsteps command with the u option. Say, for example, the topologies (0(1 (2(3 4)))) and (0(1 (3(2 4)))) are input. If there is no character support for either (3 4) or (2 4), these are compressed to a polytomy resulting in a single unique cladogram: (0(1(2 3 4))). Since Hennig86 does not generate redun- dant cladograms with unsupported branches, when would one need to worry about compressing cladograms? Both PHYLIP and PAUP (prior to version 3.0) will gen- erate only fully dichotomous cladograms, including all possible (and redundant) fully resolved topologies for polytomous conditions. For the purposes of comparing only unique cladograms produced by either program, all cladogram topologies can be input into Hennig86, compressed, and examined. Often this will substantially reduce the number of cladograms that must be examined and affords easy comparison with cladogram(s) generated from the same data set by Hennig86. A cladogram or set of cladograms can be rerooted using a different outgroup by the reroot command. If a new outgroup has been designated, invoking the reroot command will produce all new and unique cladograms based on this new outgroup. The new cladogram(s) can then be diagnosed with the xsteps command. Similarly, cladograms can be rerooted, examined, and diagnosed from the tree editor, Dos Equis, described next. Hennig86 provides a very nice interactive tree editor, Dos Equis, which has ca- pabilities very similar to those seen in the program MacClade, developed by Wayne and David Maddison for Macintosh computers. Cladograms generated by Hennig86 or other user-defined cladograms can be used. Dos Equis is entered, not surprisingly, with the command, xx. A single cladogram from the program’s current internal tree file is displayed until another cladogram is chosen. The states for a particular character are indicated for each terminal taxon and at all nodes. As mentioned earlier, in the event that several equal-length transformation series are possible for a given character, all possible nodal conditions are presented. Additional diagnostic data are shown below the cladogram and include total cladogram length, which character is being displayed, current weight of that character, if the character is additive or non-additive. 240 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) if it has been used in cladogram construction (active) or not (non-active), and the number of steps required by that character to fit on the cladogram. Editing with Dos Equis is very straight-forward, with onscreen editing controls displayed in a concise manner. As is typical of Hennig86, a minimal number of key strokes are required to initiate commands. Cladogram topology modifications include moving terminal branches or clades, and deleting nonterminal branches (clades). Upon making such changes the user is updated as to total cladogram and character length. Unfortunately, a single terminal taxon cannot be deleted. Possible character modifications include changing weights, activity, and additivity. Again, the user is updated on the changes incurred with these editing procedures with regard to» total character length. All desired changes can then be saved, or else the user can exit directly from Dos Equis. While one can move or rotate branches or taxa, and see the effects directly, the same immediate results are not achieved with character modifications. For example, if a character is made inactive (i.e., essentially deleted for purposes of cladogram construction), this is indicated on the update, but the character is still shown on the cladogram as though it were active. In order to see what effect this change actually has on the cladogram topology, one must save the modified cladogram and character settings and rerun this new data set. In all, Hennig86 meets virtually all the criteria one would expect in a cladistics program. Its small size, low cost, and compatibility make it readily accessible to a wide audience. The few problems pointed out here are certainly minuscule compared to the overall benefits provided. Obviously, one’s acceptance and use of a particular program is an indication that it meets, at least minimally, the user’s expectations, which might include 1) ease of interaction, 2) ability to handle a variety of data sets of different sizes, 3) relatively good speed in analyzing data, 4) a variety of search strategies, 5) receiving concise and accurate results, and 6) being able to easily handle and interpret output. No doubt with the introduction of new programs, and revisions of old ones, users will begin to weigh differences and similarities based on their own expectations. Differences of opinion will probably develop mainly as a function of these expec- tations, as well as due to theoretical and research proclivities, and associated ad hoc assumptions deemed allowable. Biases aside, systematists should definitely take the time to assess for themselves what they perceive to be the strengths and weaknesses of Hennig86 as it pertains to their own research and teaching.— A'/r/c Fitzhugh, De- partment of Invertebrates, American Museum of Natural History, New York, New York 10024. LITERATURE CITED Carpenter, J. M. 1 988. Choosing among multiple equally parsimonious cladograms. Cladistics 4:291-296. Coddington, J. A. 1987. The sixth annual meeting of the Willi Hennig Society. Cladistics 3: 178-184. Farris, J. S. 1969. A successive approximations approach to character weighting. Syst. Zool. 18:374-385. Farris, J. S. 1988. Hennig86 reference. Documentation for version 1.5. Fink, W. L. 1986. Microcomputers and phylogenetic analysis. Science 234:1 135-1 139. 1989 BOOK REVIEWS 241 Luckow, M. and R. M. Pimentel. 1985. An empirical comparison of numerical Wagner computer programs. Cladistics 1:47-66. Mickevich, M. F. and J. S. Farris. 1984. PHYSYS documentation. Nelson, G. 1 979. Cladistic analysis and synthesis: principles and definitions, with a historical note on Adanson’s “Families des plantes” (1763-1764). Syst. Zool. 28:1-21. Nelson, G. and N. I. Platnick. 1981. Systematics and Biogeography; Cladistics and Vicariance. Columbia University Press, New York. O’Grady, R. T. and G. B. Deets. 1987. Coding multistate characters, with special reference to the use of parasites as characters of their hosts. Syst. Zool. 36:268-279. Pimentel, R. M. and R. Riggins. 1987. The nature of cladistic data. Cladistics 3:201-209. Platnick, N. I. 1987. An empirical comparison of microcomputer parsimony programs. Cla- distics 3:121-144. Platnick, N. I. 1988. Programs for quicker relationships. Nature 335:310. Platnick, N. I. In press. An empirical comparison of microcomputer parsimony programs, II. Cladistics. Swofford, D. L. 1985. PAUP (phylogenetic analysis using parsimony). Documentation for version 2.4. Illinois Natural History Survey, Champaign. J. New YorkEntomol Soc. 97(2):24 1-242, 1989 TWO NEW TRUE BUG CATALOGS Catalog and Bibliography of the Leptopodomorpha (Heteroptera).— R. T. Schuh, B. Galil, and J. T. Polhemus. 1987. Bulletin of the American Museum of Natural History 185:243-406. $10.65. For most biologists, and especially for museum curators, taxonomists, and bio- geographers, the most important source of reference is a worldwide catalogue. Un- fortunately, few people want to undertake the tedious and time-consuming work involved in making such a catalogue. The present volume is therefore received with great enthusiasm. For many higher groups of Heteroptera or true bugs, the only worldwide catalogue is still that of L. Lethierry and G. Severin (1 893-1 896). Needless to say, this catalogue is hopelessly outdated. The “General Catalogue of the Hemiptera,” initiated in 1927 (Editors G. Horvath and H. M. Parshley) was never completed as far as the Heter- optera is concerned. In fact, only two heteropteran families were ever treated, the Mesoveliidae by G. Horvath and the Pyrrhocoridae by R. F. Hussey (both in 1929). Fortunately, other worldwide catalogues have appeared, foremost among them the impressive catalogues on the Miridae by J. C. M. Carvalho (1957-1960), on the Lygaeidae by J. A. Slater (1 964), and on the Tingidae by C. J. Drake and F. A. Ruhoff (1965). Nevertheless, most families of the Heteroptera, including such large and important groups as the Reduviidae, Coreidae, and Pentatomidae, have not been adequately catalogued. The present “Catalog and Bibliography of the Leptopodomorpha” covers one of the smallest of the heteropteran infraorders with less than 300 described species. The infraorder comprises the shore bugs and allied groups. Most species inhabit damp soil close to water, either fresh or saline. A few species are intertidal marine. The 242 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) taxonomy of leptopodomorphans is generally well known and the phytogeny above the genus-level has been analysed quite recently (Schuh and Polhemus, 1980). The authors apply the most recent classification of the Leptopodomorpha which divides the infraorder into four families, Leptopodidae (7 genera, 28 species), Oma- niidae (2, 4), Aepophilidae (1, 1), and Saldidae (25, 264). The odd genus Leotichius, which previously had its own family, is now included in the Leptopodidae. The remarkable marine bug, Aepophilus bonnairei Signoret, of western Europe is the single member of the family Aepophilidae. The catalogue has an excellent introductory chapter. The section describing the format is necessary for the use of the catalogue and at the same time provides a layout which may well be followed by future cataloguers. The organization of the catalog is alphabetically within each level of the hierarchy. Species are catalogued by genera while subgeneric placements are ignored although recorded. Since subgeneric placements often are controversial, it is understandable why the authors have adopted this procedure. Subspecies are not catalogued separately but listed under the species they belong to. This is a wise decision which evades the problems caused by incon- sistent usages of “forms,” “varieties,” and “subspecies” by previous authors. The typography used is clear and consistent and makes it quite easy for the reader to find his way around. The section for each species is headed by the species name and the name of the author. It is misleading, however, that author names are cited without the use of parentheses to distinguish between original and subsequent com- binations with generic names. It is correct that Article 5 1 (c) of the Code only provides that the name of the author is enclosed in parentheses if a species name is combined with a generic name other than the original one. However, in this catalogue the reader must find out for himself what is the correct form. For each species, references and distributional records are listed chronologically. The authors seem to have made an exhaustive search for literature covering both taxonomical and faunistical references, as well as works on biology, ecology, etc. However, it is difficult to understand how the authors missed the catalogue of the Heteroptera of Sweden by Coulianos and Ossiannilsson (1976), one of the most reliable faunal lists published in any European country. The bibliography is divided in two parts. The first part includes only those ref- erences which are cited in the catalogue. The second part contains additional but uncatalogued references. Each reference is followed by a short but very useful note stating its key contents. The choice of references for both parts seems relevant and exhaustive, making this new catalogue extremely useful in itself and an example to follow by future cataloguers.— M oiler Andersen, Zoological Museum, Universi- tetsparken 15, DK-2100 Copenhagen, Denmark. LITERATURE CITED Coulianos, C.-C. and F. Ossiannilsson. 1976. Catalogus Insectorum Sueciae. VII. Hemiptera- Heteroptera, 2nd Edition. Ent. Tidskr. 97:135-173. Schuh, R. T. and J. T. Polhemus. 1980. Analysis of taxonomic congruence among morpho- logical, ecological and biogeographic data sets for the Leptopodomorpha (Hemiptera). Syst. Zool. 29:1-26. 1989 BOOK REVIEWS 243 /. New YorkEntomol. Soc. 97(2):243-245, 1989 Catalog of the Heteroptera, or True Bugs, of Canada and the Continental United States.-Thomas J. Henry and Richard C. Froeschner (eds.). 1988. E. J. Brill, Leiden, New York, Kobenhaven, Koln, 958 pp. $58.50. There was a time when the Catalogue of the Hemiptera of America North of Mexico, by E. P. Van Duzee (1917), was considered the Bible of the group in the Nearctic. Unfortunately, that time passed at least two decades ago, and in the meantime, for many groups. North American heteropterists have had to rely on species lists of their own making, or worse yet, with no comprehensive listings at all. In the Lygaeidae, Miridae, and Tingidae the problem was alleviated by World catalogs in those groups (respectively: Slater, 1964; Carvalho, 1957-1960; Drake and Ruholf, 1965), but for all other groups Van Duzee was still the only source. Now, things have changed. Not only have World compilations appeared for the Leptopodomorpha (see review on page 241) and the Aradidae (Kormilev and Froeschner, 1987), but finally we have available what might be considered a sequel to Van Duzee— an up-to-date catalog of the North American Heteroptera fauna. Editors Henry and Froeschner, who are also authors of a large number of chapters in the volume, will certainly gain the gratitude of their colleagues for the devotion of time and energy required to bring this work to fruition (Other contributors are: the late P. D. Achlock, J. D. Lattin, D. A. Polhemus, J. T. Polhemus, A. Slater, and C. L. Smith.). In this age of the global economy and overnight travel to anywhere on the face of the earth, one might ask, “Why prepare a regional catalog?” The answer is that such catalogs are extremely valuable to great numbers of entomologists and other biologists whose work is regional in character. But possibly more important, knowledge in many groups of insects is not sufficiently well organized or developed to allow for preparation of truly effective World catalogs. In many cases there are no workers who are capable or inclined to prepare such a volume for every family— something that is certainly true for the Heteroptera at the present. In nearly all groups, development of knowledge of faunas has always presaged the preparation of World revisions and catalogs. With the appearance of this Catalog we might expect to see an increase in the quantity and quality of work on the North American fauna, simply because a complete enumeration of the taxa for the region is now available at an affordable price. Each chapter, which falls in alphabetical order by family name and which presents all contained taxa in alphabetical order as well, begins by offering a brief overview of the habits and taxonomic history of a family as well as some habitus illustrations. The catalog is laid out in an attractive and easy-to-read manner, orienting the reader’s eye to the various subdivision in what seems a natural fashion. Rather than being comprehensive, the catalog lists references to the original description for each genus and species as well as citations for synonymies and misspellings— the last seeming like a poor use of space and bibliographic effort to this reader. For many higher groups, genera, and species, notes are provided, leading users to important additional references on classification, synonymy, and biology. The type locality (or broader distribution) is indicated with the reference to the original description for each species. 244 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) and the bibliographic references for each species are followed by a summary listing of occurrences drawn from a broader sampling of the literature than that presented in the catalog. The editors argue passionately for the use of the term Heteroptera at the ordinal level. The idea cannot be faulted with regard to recognition of a monophyletic group, although the absolute rank of the taxon will probably continue to be a subject of continued debate. At the very minimum, this broad-based work will almost certainly influence North American workers in the future to adopt the term Heteroptera, a usage more in line with that of students of the true bugs from other parts of the World. The Catalog presents an up-to-date classification for many groups, e.g., the Miridae and Saldidae, taxa with which I have some familiarity. In other cases, the presentation is what I would call idiosyncratic, e.g., treating the Phymatinae at family rank, whereas this group was long since recognized as a subgrouping within the Reduviidae (Car- ayon, Usinger, and Wygodzinsky, 1958), and has been treated as such by many subsequent authors. The Literature Cited section (comprising 1 3 1) pages will by itself serve as a valuable reference on many occasions. Not since the publication of Parshley’s (1925) “A Bibliography of North American Hemiptera-Heteroptera” have heteropterists had available such an extensive listing of literature on the North American fauna. The index will be no less important for use of the catalog. It will allow the user to locate nearly every name (and its variant spellings) ever used for the North American Heteroptera. Although the catalog would otherwise be worthwhile, the index will make it a truly valuable research resource. There seems to be a tradition— which has been followed by Henry and Froeschner— in the preparation of catalogs dealing with the Nearctic fauna, be it of Hemiptera, Coleoptera, Diptera, or Hymenoptera, to draw the line for inclusion of taxa at the border between Mexico and the United States. It may be that the language and customs of these two countries seem very different to many, but the Neartic fauna is certainly not restricted to Canada and the continental United States. Thus, except for those groups for which World catalogs are available, knowledge of the large com- ponent of the Nearctic fauna occurring on the Mexican plateau— with many groups reaching well into Central America at higher elevations— will continue to require recourse to the original literature. How many times in the past have I wished that I had a catalog just like this one? Although it will not resolve all questions one might have about a group, it will provide a starting point— and at least for a while— will be right up to date. As a reference, it will be a must for all students of the Heteroptera, as well as for all libraries in North America that deal with Qniomo\o%y Randall T. Schuh, Depart- ment of Entomology, American Museum of Natural History, New York, New York 10024. LITERATURE CITED Carayon, J., R. L. Usinger and P. Wygodzinsky. 1958. Notes on the higher classification of the Reduviidae, with the description of a new tribe of Phymatinae (Hemiptera-Heter- optera). Rev. Zool. Bot. Africaines 57:256-281. 1989 BOOK REVIEWS 245 Carvalho, J. C. M. 1957-1960. Catalogue Miridae of the World. Arq. Mus. Nac., Rio de Janeiro, 5 vols. Drake, C. J. and F. A. Ruhoff. 1965. Lacebugs of the World; a catalog (Hemiptera: Tingidae). Bull. U.S. Nat. Mus. 243:i-viii + 634 pp. Kormilev, N. A. and R. C. Froeschner. 1987. Flat bugs of the World. A synonymic list (Heteroptera: Aradidae). Entomography 5:1-246. Parshley, H. M. 1925. A Bibliography of the North American Hemiptera-Heteroptera. Smith College, Northampton, Massachusetts, 252 pp. Slater, J. A. 1964. A Catalogue of the Lygaeidae of the World. Univ. Connecticut, Storrs, 2 vols. Van Duzee, E. P. 1917. Catalogue of the Hemiptera of America North of Mexico. Univ. California Pubs., Tech. Bull., Entomology 2:i-xiv + 902 pp. J. New YorkEntomol. Soc. 97(2);245-248, 1989 LIFE CYCLES AND DIAPAUSE Insect Development: Photoperiod and Temperature Control. — Victor A. Zaslavski. 1988. Springer- Verlag, Berlin, xi + 187 pp. Hardbound $99.95. The topic of this book is much narrower than the general title of “Insect Devel- opment: Photoperiodic and Temperature Control” might indicate. Make no mistake about it, this is a book on diapause. The broader issue of the role that temperature and photoperiod play in other developmental processes is dismissed, on page 1 1 , by the statement, “Mathematical expression of these dependencies can be found in ecological manuals.” Although this is a book about diapause, don’t expect deep physiological insights into the processes involved. The approach taken is what has been described in the physical sciences as phenomenological. Within these limits, however, I believe this is an important contribution to the literature on diapause, following the honorable tradition of contributions that phenomenological models have made in areas such as physics. Zaslavski has organized his book into three chapters. The first is primarily intro- ductory in nature. The second begins to develop the underlying theme of the phe- nomenological model, and the third states the form of the model and applies it to examples introduced in the first chapter. My review will be structured according to his organization. The objective of the first chapter is to lay the empirical groundwork for the sub- sequent model by defining terms, providing a basic classification scheme for pho- toperiodic reactions, and to illustrate the diversity and complexity of the photope- riodic response by describing numerous experimental results. The abundance of examples is valuable if for no other reason than providing an introduction to the rich literature on diapause, and in particular, the Russian literature that might not be familiar to Western readers. There is, however, a major problem with the first chapter. It is difficult going, almost to the point of brutality. I suspect that many readers, even those with serious interests in diapause, will become frustrated and 246 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) give up. This problem arises from several sources. Diapause, and its control by the interaction of photoperiod and temperature, is an inherently complex subject. Con- founding this complexity is Zaslavski’s (or translator Vasilyev’s) choice of words used for his dehnitions. The relationship between the common use of a word and the phenomenon described is occasionally obscure and sometimes misleading. A glossary of terms would have been extremely useful, and I advise readers to make one as they progress through this chapter. Some of the problems with dehnitions undoubtedly resulted in the process of translation from the Russian. Vasilyev’s translation some- times leads to archaic and/or arcane usage. Throughout the book, but particularly in the hrst chapter, Zaslavski’s hgures are often confusing, although I must admit that with persistence they invariably led to a more complete understanding of his narrative. Additionally, hgures often augment text discussion that may be several pages removed from placement of the hgure, and I caution the reader to avoid the temptation of trying to understand a hgure before it is discussed in the text. Most hgure captions are not self-contained. Finally, the sheer weight of examples in the hrst chapter is formidable. Through use of a wide diversity of empirical examples, Zaslavski intends to make a point, but it may be overkill. The dominating theme of the hrst chapter is the complexity of insect diapause and seasonality. By the time the reader has hnished the hrst chapter, it is patently obvious that a conceptual model of diapause is an absolute necessity to avoid the morass of a seemingly inhnite variation on the theme of seasonality. This is, of course, the reaction that Zaslavski anticipated. He has, in a sense, set the reader up for what follows in the remaining two chapters. In the hrst 3 pages of Chapter 2, Zaslavski makes an eloquent argument for a unihed physiological basis for diapause control in the insects. He goes on to present many experimental examples. However, as opposed to the examples in Chapter 1, the underlying theme of Chapter 2 is one of unihcation. Zaslavski bases his unihcation on a two phase process (a “dual control mechanism”). His historical review of the development of a duality concept of diapause is quite good (pp. 114-117) and logically results from the preceding empirical data. Duality, leading to a “profound relationship between inductive and spontaneous processes in seasonal development of insects,” is reinforced by discussion in the remainder of this chapter. Actual formulation of Zaslavski’s two phase model is the subject of Chapter 3. This formulation is composed of two parts. The hrst, on pages 129-131, sets out the quantitative basis for the expression of photoperiod. The second, on pages 133-136, dehnes the components of the model and describes their properties and intercon- nections. A physiological basis underlies both discussions. As previously noted, Zas- lavski’s approach is descriptive rather than mechanistic. For example, the basic shape of the photoperiodic threshold curve (a 2 hump curve, or perhaps more descriptively, a valley and a hill) as presented on page 137 is absolutely necessary for proper functioning of the model (e.g.. Figs. 84, 86, 87, 88, and 89). This specihc shape results from integrating the area under two simple curves describing the photoperiod effect on enzyme synthesis. The temporal relationship between these two curves in turn results in the appropriate shape for the photoperiodic response. The result of all this is a simple model that integrates the effects of photoperiod and temperature and also provides a unihed description for a complex array of observed photoperiodic 1989 BOOK REVIEWS 247 responses. These results are important enough that I, for one, plan to discuss the physiological basis of the model in more detail with a qualified specialist. Although Zaslavski does indeed provide a synthetic model for diapause, I was personally disappointed that he does not provide a formal mathematical statement of that model. A mathematical statement of the model would have made it easier for me to understand the arguments that Zaslavski uses to develop his model and to convince myself that his consequences logically followed from his assumptions. More importantly, mathematics provides a formal mechanism to test (validate) a model. Zaslavski makes an attempt at validation through a process he terms “pre- diction.” I disagree with his use of the term. Zaslavski has used his model to describe various patterns of observed photoperiodic response and seasonality (see pages 137- 1 58). Clearly, his model is flexible and can describe a wide array of empirical diapause patterns. However, just as clearly, this is not prediction in the prospective sense of the word. I have a hunch that Zaslavski’ s model as stated is overparameterized, in other words, a model that is flexible in descriptive capabilities but lacks predictive power. Use of his model for prediction will require constraints that may be implicitly included in his discussion but that were not explicitly stated. The topic of Zaslavski’s book is an important one. Appropriate timing of life history events is arguably the single most important adaptation of insects living in seasonal environments. Additionally, the appropriate modeling of diapause assumes economic importance because its termination initiates subsequent phenology. Ac- curate prediction of insect phenology is often critical to effective control. Given this importance, Zaslavski makes a significant contribution to the literature on insect ecology. Almost 30% of the 442 articles cited are in Russian. A significant proportion of the remaining articles are in non-English language journals. By using this literature to build the empirical basis for his model, Zaslavski provides insights into current trends in diapause research in the USSR. Considering that both the US and the USSR are notoriously xenophobic in their scientific outlook (e.g., Garfield, 1988), books such as Zaslavski’s can do much to promote a scientific glasnost. The second significant contribution is the major reason Zaslavski undertook this project, and that is to present a dual phase model that unifies photoperiodic and temperature control of diapause. Zaslavski is largely successful in this presentation. As previously stated, I am personally disappointed in the lack of mathematical rigor in his formulation of the model. In my opinion, the rigorous, unambiguous math- ematical statement of his model would make an excellent Chapter 4, and the appli- cation of the model for parameter estimation would make an excellent Chapter 5. Perhaps all this is just to say that Zaslavski’s is not the last word on the subject, and there is work yet to be done for the rest of us. Zaslavski’s work is particularly useful for establishing priorities in diapause re- search, and he does so on many occasions. For example, on page 139 regarding the interrelationship between temperature and photoperiod, he states, “Temperature affects the commanding centers of poikilotherm organisms directly and thus inev- itably. In contrast, to perceive the daylength a mechanism of photoperiodic clock is necessary, the presence or functional activity of which seems to be nonobligatory, since photoperiodically neutral species exist. Therefore, the temperature reaction should be considered both separately and in combination with the photoperiodic one.” Zaslavski also makes a serious attempt to place diapause within the total eco- 248 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) logical matrix of factors impinging on the success of a species. By doing this, his book is motivation for collaboration between insect physiologists, insect ecologists, and mathematical modelers. It is only through such collaboration that we can expect to make progress toward a more complete understanding of diapause. A final note on the cost of this book is probably in order. Translations of technical books with limited distribution are notoriously expensive. At a price of $100 for a book of 187 pages (including References and Index) “Insect Development” is no exception to this rule. I for one, however, am glad that Springer- Verlag is willing to undertake such ventures, and if such prices are necessary, then so be it. I anticipate that I will refer often to this book in my work involving diapause and insect sea- sonality. I also anticipate significant insights will be gained into procedural methods for modeling such processes. My advice to serious researchers in the area of diapause and insect ecology is to buy the book and then make it widely available to students and others who might find it prohibitively expensive.— /I. Logan, Department of Entomology and Department of Forestry, Virginia Polytechnic Institute and State University, Blacksburg, Virginia 24061. LITERATURE CITED Garfield, E. 1988. Is the American scientific press provincial? Current Contents 19(34):3-7. /. New YorkEntomol. Soc. 97(2):248-249, 1989 The Evolution of Insect Life Cycles. — F. Taylor and R. Karban (eds.). 1986. Springer- Verlag, New York, 287 pp. $64.00. This book is a collection of sixteen papers on the evolution of life cycles in insects. The papers are modified from a symposium at the XVII International Congress of Entomology in 1984. The stated goal of the volume is “. . . to provide a compre- hensive view of current research on insect life cycles . . . .” The book is organized into four sections: Geographical Patterns in Insect Life Cycles (5 papers). Diversity of Life Cycle Patterns (6 papers). Mechanisms of Insect Life Cycle Evolution (4 papers) and Concluding Remarks (1 paper); however, many of the papers are ap- propriate for more than one section. The book addresses several important issues in the study of insect life histories, among them the reality of environmental uncertainties which can produce consid- erable year to year variation in selection pressures. Several chapters discuss this point in both theoretical and empirical terms, as well as the consequences for genetic variability of life history traits. Several other papers consider exceptions to gener- alizations about life histories. For example, A. Shapiro reports that r- and K-type traits do not consistently occur at a taxonomic level; however, they do correspond to what is known of the ecology of the organisms and the presumed selection pressures. Several of the chapters discuss genetic aspects of life histories. This is a crucial issue in the study of the evolution of life histories and I felt a general weakness of some of the papers was implicit assumptions about genetic systems. There seems to be a tendency for authors to infer past selection from present patterns. For example. 1989 BOOK REVIEWS 249 although few empirical studies have been able to demonstrate natural selection, in chapter 1 3 the ability of natural selection to affect genetic systems is given consid- erable weight. Not only are traits selected but their inheritance (polygenic vs. single locus heritability) is also selected. A second example is chapter 2 in which it is assumed that a correlation between ovipositor length and habitat moisture demonstrates se- lection balance. I feel that correlation does not imply causation, the role of selection needing to be rigorously demonstrated, and thus these are areas where empirical work is needed. Several of the chapters were enjoyable and thought provoking reading, including the chapter by D. Neumann on a slow growing intertidal midge (38-95 days) in which the adults are short lived (30 minutes to a few hours); the chapter by Valarie Brown which includes an amazing amount of data on the effect of plant succession on insect life cycle strategies; the review of variation in diapause induction by critical photoperiod as a function of latitude by Taylor and Spaulding; and the chapter by Lounibos and Machado-Allison describing parental care in mosquitos and demon- strating that rainfall is a selective pressure for maternal egg brooding. The volume summarizes a broad range of research in insect life history evolution. I feel its strength is that it not only suggests areas for future research but I came away with many ideas of how such research needs to be conducted to provide contributions to this area. For this reason, I recommend it to researchers of insect life cycles.— Lori Stevens, Department of Zoology, University of Vermont, Burlington, Vermont 05405-0086. J. New York Entomol. Soc. 97(2):249-250, 1989 DARWIN’S INSECTS Darwin’s Insects. Charles Darwin’s Entomological Notes. — K. G. V. Smith (ed.). 1987. Bulletin of the British Museum (Natural History) 14(1): 1-143. Natural His- tory Museum Publications, Cromwell Road, London SW7 5BD, U.K. £25. This publication is a careful, painstakingly prepared, account (essentially a cata- logue) of Charles Darwin’s insect collections of those of his specimens that have been located in the present-day entomological holdings or various institutions with the field notes made during the voyage of the British Navy’s H.M.S. .§^<2^/^(1831-1835), on which vessel he served as naturalist. The text is organized in 10 sections, of which hve record material housed in different institutions, all in various parts of The United Kingdom, most specimens being in the collections of the British Museum (Natural History). Almost one-half of the volume (67 pages) is taken up with Darwin’s en- tomological notes from the Beagle voyage and their annotation. A list of scientific names is given that are formed from the surname Darwin, and used in the Insecta. Two indices are included: one, of geographical place names, and names of institutions, ships and persons cited in the text; and one of names of taxa. Illustrations provided are maps showing positions of collecting localities in South America, reproductions of plates illustrating specimens of new taxa that were based on Darwin-collected 250 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(2) material, and photographs of store-boxes, in several institutions, of Darwin’s insect specimens. Although Darwin has an excellent reputation for his contributions to entomology, the accolades are based on his ecological observations and hypotheses (significance of wing loss from adults; social behavior; pollination; other insect-plant relationships). He did not work on insect taxonomy or classification, and thus did not require or prepare extensive, working, entomological collections. The insect material gathered during his one great expedition, the voyage of the Beagle, was turned over to others for curation and study. This material was documented by perfunctory field notes recorded by Syms Covington, Darwin’s assistant on the Beagle, who did much of the collecting. These notes were turned over to G. R. Waterhouse by Darwin, and, presently, the original manuscript is in the Entomological Library of the British Museum (Natural History) under the title “Copy of Darwin’s notes in reference to Insects collected by him.” The bulk of the Darwin collection is a random accumulation of insects encountered during biotic and geologic explorations by the naturalist of the Beagle and his assis- tants. The collection was made solely for faunal documentation, in the least restrictive sense of that term. The collection was not used in any significant way by Darwin in developing his ideas about evolution. The value of the collection is in its individual components: the numerous new taxa based on this material, that were described by Darwin’s contemporaries at his behest and with his encouragement. From this material, entomologists gained early insights about the nature of portions of the insect fauna of the Southern Hemisphere visited by the Beagle. Few new insights emerge from this publication because the volume is intended for documentation,, only. The Editor did not offer any explicit goals or formal justification for preparing the publication. I was depressed, however, by the seedy appearance of the Darwin material illustrated by the photographs of the store-boxes in which spec- imens are housed. One would think that these institutions, especially Cambridge University, would be more mindful of such material, considering the eminence of the collector. The justification for preparing this document resides, in fact, in the eminence of Charles Darwin. His outstanding contributions to biology make it worthwhile that all facets of his scientific life be documented as fully as possible. Kenneth Smith’s publication elucidates one part of Darwin’s scientific life. As a result of this work, future taxonomists who desire to locate specific parts of Darwin’s collection will have a relatively easy task. Many taxa, however, were not accounted for. Perhaps this publication will provide the information and impetus necessary for curators with unrecognized Darwin material in their collections to recognize it and to make known their discoveries. This publication should be in the better biological libraries of the world. However, I doubt that it has sufficient general value to be sought out by entomologists other than systematists, and only those in the latter group who are particularly interested in entomological history or in the entomological aspects of Charles Darwin. — E. Ball, Department of Entomology, University of Alberta. {Continued from back cover) LIFE CYCLES AND DIAPAUSE Insect Development: Photoperiod and Temperature Control The Evolution of Insect Life Cycles Jesse A. Logan 245-248 Lori Stevens 248-249 DARWIN’S INSECTS Darwin’s Insects. Charles Darwin’s Entomological Notes George E. Ball 249-250 INSTRUCTIONS TO AUTHORS The Journal of the New York Entomological Society is devoted to the advancement and dissemination of knowledge of insects and related taxa. The costs of publishing the Journal are paid by subscriptions, membership dues, page charges, and the proceeds from an endowment established with bequests from the late C. P. Alexander and Patricia Vaurie. The Journal will consider for publication manuscripts of any length dealing with original research in entomology. Longer papers will be printed as articles, shorter ones as “scientific notes.” Book reviews will be solicited by the Book Review Editor. Manuscripts should be submitted in duplicate to: Dr. Randall T. Schuh, Editor, Journal of the New York Entomological Society, c/o Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024. All material must be double-spaced (including references, tables, captions, etc.) and prepared in the format of a recent issue of the Journal. Longer manuscripts intended for submission as articles should be accompanied by a brief abstract. Footnotes should be avoided. Tables should be prepared as separate pages; they should be kept to a minimum because of the high cost of typesetting, but may be submitted as photographically reproducible material (see below). The list of references is headed “Literature Cited” and should follow the format indicated in the CBE Style Manual or as found in a recent issue of the Journal. Illustrations (originals whenever possible) should be submitted flat, never rolled or folded, with the manuscript. Size of illustrations should be kept manageable and suitable for shipment through the US mail without being damaged. Proportions of illustrations should conform to single column format, actual page size 1 1.5 x 17.5 cm. Please assemble illustrations in a com- pact fashion in order to use journal space effectively. Mount line drawings and halftones on separate plates. Figures should be numbered consecutively. Figure captions should be double- spaced, grouped together and placed at the end of the manuscript. If tables are submitted for photographic reproduction, they should be neat and clean and conform to journal format proportions. Authors will receive page proofs, with the original manuscripts, directly from the printer. Corrected proofs should be returned promptly to the Editor to avoid publication delays. Re- visions in proof should be kept to the absolute minimum. Excessive alterations in proof will be charged to the author at the rate of $2.00 per revision line. Society members will be charged a fee of $20.00 per printed page and $5.00 per plate of figures. Non-members will be charged $45.00 per printed page and $5.00 per plate of figures. Member authors who do not have institutional funds may petition to the Society for waiver of page charges for no more than eight pages on a once a year basis. Because of limited funds, all such requests will be handled on a first-come first-serve basis. Authors will receive a reprint order blank with the proofs. Reprints are ordered directly from the printer with no benefit accruing to the Society. Journal of the New York Entomological Society VOLUME 97 APRIL 1989 NO. 2 CONTENTS A fossil solpugid, Haplodontus proterus, new genus, new speeies (Arachnida: Sol- pugida) from Dominican Amber George O. Poinar and Jorge A. Santiago-Blay 125-132 On the abundance and ecology of Ricinulei (Arachnida) from Central Amazonia, Brazil Joachim U. Adis, Norman I. Platnick, Jose W. de Morais, and Jose M. Gomes Rodriguez 133-140 Two new species of Eosentomon from Chickasaw State Park, Tennessee (Protura, Eosentomidae) William A. Outten and Robert T. Allen 141-150 New species rediscriptions, and cladistics of the genus Pseudocentroptiloides (Ephemeroptera: Baetidae) R. D. Waltz and W. P. McCafferty 151-158 Review of Daleapidea Knight (Heteroptera: Miridae: Orthotylinae: Orhtotylini) Randall T. Schuh 159-166 Texocoris nigrellus: Distribution and hosts of an enigmatic plant bug (Heteroptera: Miridae: Orthotylinae) A. G. Wheeler, Jr. 167-172 Tantysoma diabolica new species (Coleoptera: Carabidae: Platynini) from Baja California and its Biogeographic Significance James K. Liebherr 173-186 A new microcadddisfly genus (Trichoptera: Hydroptilidae) from the interior high- lands of Arkansas, U.S.A. Michael L. Mathis and David E. Bowles 187-191 Relationships among Holarctic genera in the Cyrtogaster-growp with a review of the species of North American north of Mexico (Hymenoptera: Pteromalidae) Steven L. Heydon 192-217 Stinging behavior and residual value of worker honey bees {Apis mellifera) Steven A. Kolmes and Linda A. Eergusson-Kolmes 2 1 8-23 1 Notes and Comments Evaluation of the Spider Steatoda triangulosa (Araneae: Theridiidae) as a Predator of the Red Imported Fire Ant (Hymenoptera: Formicidae) William P. MacKay 232-233 Book Reviews CLADISTICS IN THE FAST LANE Hennig86. Version 1.5 Kirk Fit zhugh 234-241 TWO NEW TRUE BUG CATALOGS Catalog and Bibliography of the Leptodomorpha (Heteroptera) Nils Moller A ndersen 24 1 -242 Catalog of the Heteroptera, or Ture Bugs, of Canada and the Continental United States Randall T. Schuh 243-245 {Continued on inside back cover) JULY 1989 No. 3 Vol. 97 Journal of the New York Entomological Society (ISSN 0028-7199) i j Devoted to Entomology in General JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Editor: Randall T. Schuh, Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024 Book Review Editor: David A. Grimaldi, Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024 Publications Committee: Louis Trombetta, St. Johns University, Queens, New York, Chairman; Alfred G. Wheeler, Jr., Pennsylvania State Department of Agriculture, Harrisburg; Joseph M. Cerreta, St. Johns University, Queens, New York. The New York Entomological Society Incorporating The Brooklyn Entomological Society President: Dennis J. Joslyn, Department of Biology, Rutgers University, Camden, New Jersey 08102 Vice President: Durland Fish, Medical Entomology Laboratory, New York Medical College, Armonk, New York 10504 Secretary: Richard Falco, Westchester County Health Department, White Plains, New York 10601 Treasurer: Louis Sorkin, Department of Entomology, American Museum of Natural History, New York, New York 10024 Trustees: Class of 7955— Henry M. Knizeski, Jr., Mercy College, Dobbs Ferry, New York; Michael D. Schwartz, American Museum of Natural History, New York, New York; Class of 7959— Christine Falco, West- chester County Health Department, White Plains, New York; James S. Miller, Department of Entomology, American Museum of Natural History, New York, New York. Annual dues are $23.00 for established professionals with journal, $10.00 without journal, $15.00 for students with journal, $5.00 without journal. Sustaining memberships are $53.00 per year, institutional memberships are $125.00 per year, and life memberships are $300.00. Subscriptions are $40.00 per year domestic and $45.00 foreign. All payments should be made to the Treasurer. Back issues of the Journal of the New York Entomological Society, the Bulletin of the Brooklyn Entomological Society, Entomologica Americana, the Torre-Bueno Glossary of Entomology and other Society publications can be purchased from Lubrecht and Cramer, RD 1, Box 244, Forestburgh, New York 12777. Meetings of the Society are held on the third Tuesday of each month (except June through September) at 7 p.m. in the American Museum of Natural History, Central Park West at 79th Street, New York, New York. Mailed August 22, 1989 The Journal of the New York Entomological Society (ISSN 0028-7199) is published 4 times per year (January, April, July, October) for the Society by Allen Press, Inc., 1041 New Hampshire, Lawrence, Kansas 66044. Second class postage paid at New York, New York and at additional mailing office. Postmaster: Send address changes to the New York Entomological Society, % American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024-5192. Known office of publication: American Museum of Natural History, New York, New York 10024. Journal of the New York Entomological Society, total copies printed 700, paid circulation 602, mail subscription 602, free distribution by mail 19, total distribution 621, 79 copies left over each quarter. THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER. J. New York Entomol. Soc. 97(3):25 1-260, 1989 ANNOTATED CHECKLIST OF THE THYSANOPTERA OF BERMUDA SuEO Nakahara* and Daniel J. Hilburn^ ‘Systematic Entomology Laboratory, Agricultural Research Service, USD A, Beltsville, Maryland 20705 and ^Department of Agriculture and Fisheries, P.O. Box HM834, Hamilton HMCX, Bermuda Abstract.— Thirty -QighX introduced species of thrips representing 28 genera in three families are known from Bermuda. The species were apparently introduced from eastern North America or the Caribbean region, except for three species. Hosts, collection data and world distribution are presented for each species and the economic importance of several species is mentioned. Bermuda is a small archipelago in the North Atlantic located at 32°18'N latitude, 64°46'W longitude. Seven main islands and numerous smaller ones lie in a hsh hook- shaped cluster. The closest land is Cape Hatteras, North Carolina, U.S.A., 1,040 km to the west northwest. Bermuda’s total land area is approximately 53 km2 with a maximum elevation of only 74 m. The climate is mild and frost free due to the nearby Gulf Stream. Rainfall is distributed fairly evenly throughout the year and averages 1 ,420 mm annually. Vegetation is lush and subtropical. The island is densely populated with approximately 60,000 residents. Due to its small size and isolation, Bermuda’s native terrestrial flora and fauna are depauperate. There are few endemic species. Most species originated from south- eastern North America or the Caribbean region and were introduced accidentally by human activity. Kevan (1981) presented a historical review of the terrestrial arthropods in Bermuda, from the earliest mention of insects in 1603 through 1900. Ogilvie (1928) published a list of Bermuda’s insects, which contained six species of Thysanoptera (thrips). Eleven additional species were reported by Waterston (1940, 1941), and one species each by Wilson (1975), Bennett et al. (1985) and Nakahara (1985). The Bermuda Department of Agriculture and Fisheries (BDAF) has maintained an insect collection since 1928. Specimens collected by Lawrence Ogilvie between 1923 and 1927 formed the nucleus of this collection. Since then, additions to the collection have been sporadic. Major contributions were made by Idwal Hughes between 1954 and 1975 and Francis Monkman between 1 964 and 1 97 1 . A permanent position for an entomologist with the official title of Plant Protection Officer was created in 1983. Kevin Monkman, the first to hold this post, made additions between 1983 and 1986. In 1987 a project was initiated by the junior author to upgrade the collection. Specialists in various insect groups were invited to study the collection and to survey the island for additional material. Since then, many unreported species have been found and the collection has been enlarged by addition of hundreds of specimens collected in the surveys. The thrips in the BDAF collection were examined by the senior author in June 252 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) 1988 in conjunction with a joint survey of the islands by the authors. Twelve species were found for the first time in Bermuda during the survey. Thirty-eight introduced species representing 28 genera in three families are now known from Bermuda. The species were apparently introduced from eastern North America or the Caribbean region, except ^OY Asprothrips seminigricornis (Girault), Dorcadothrips caespitis Pries- ner and Hercinothrips bicinctus Bagnall. In addition to the species collected in the survey, the following list includes species and host plants reported by Ogilvie (1928), Waterston (1940, 1941), Wilson (1975), Bennett et al. (1985) and Nakahara (1985); species collected by various collectors but not reported in the literature; and material deposited in the United States Museum of Natural History (USNM). The 20 species reported for the first time for Bermuda are indicated with asterisks. Species and host- plant records reported by Waterston (1940, 1941), which were based mainly on quarantine interceptions at United States (US) ports of entry on cutflowers and plants from Bermuda, were not verified. Voucher specimens are deposited in the Bermuda Natural History Museum, which now houses the main portion of the BDAF collec- tion, and the Thysanoptera collection of the USNM located at Beltsville, Maryland. AEOLOTHRIPIDAE Aeolothrips fasciatus (L.)* ‘Waterville,’ Paget, Chrysanthemum frutescens, 1 9-VI-88, S. Nakahara. Ferry Point Park, St. George’s, Cakile lanceolata, 24- VI-8 8, S. Nakahara. Botanical Gardens, Paget, sweeping, 26-VI-88, D. Hilbum. Spittal Pond Nature Reserve, Smith’s, sweep- ing, 27-VI-88, D. Hilbum. Preys on thrips, aphids, beetles and other insects (Lewis, 1973:299); species known from Europe, Canada and US. Franklinothrips vespiformis (D. L. Crawford)* Gilbert Nature Reserve, Somerset, Citharexylum spinosum, 20-VI-88, D. Hilbum & S. Nakahara. Shelly Bay Park, Hamilton, Schinus terebinthifolius, 22-VI-88, S. Nakahara. Botanical Gardens, Paget, Araceae, Vitis sp., 23-VI-88, D. Hilbum & S. Nakahara; Rosa sp., 23-VI-88, D. Hilbum; sweeping, 26-VI-88, D. Hilbum. A predator in tropical and subtropical areas of the New World that feeds on thrips, red spider mites, whiteflies and leafhoppers (Stannard, 1952:16). PHLAEOTHRIPIDAE Aleurodothrips fasciapennis Franklin Waterston, 1940:6, 1941:51. Paget Marsh, Paget, Sabal blackburniana, 24-1-33, T. A. Russell. A predator of armored scale insects (Lewis, 1 973:299) occurring in US from Florida to Texas, Caribbean Islands and Central America. Eury thrips modes tus (Bagnall)* Spittal Pond, Smith’s, sweeping, 27-VI-88, D. Hilbum. Reported from Cuba, St. Vincent, Trinidad, Panama, and Brazil. 1989 THYSANOPTERA OF BERMUDA 253 Gynaikothrips ficorum (Marchal) Bennett et al., 1985:120. A common species causing leaf distortion and curling of Ficus microcarpa. Ex- tremely abundant during May and June and a nuisance to residents. First collected in Bermuda, 3-1-50, by E. C. G. Bedford. A widespread species known from the tropics and subtropics on Ficus. Haplothrips gowdeyi (Franklin) Haplothrips dozieri Watson: Waterston, 1940:7, 1941:16. Haplothrips gowdeyi (Franklin): Waterston, 1940:7, 1941:16, 28, 35, 42. Stoke’s Point Nature Reserve, St. George’s, Bidens pilosa, 25-III-88, M. B. Stoetzel & D. Hilburn. ‘Waterville’ area, Paget, Carpobotrus edulis, 19-VI-88, S. Nakahara. Spittal Pond Nature Reserve, Smith’s, unknown composite, 21 -VI-88, D. Hilbum; Bidens sp., 21 -VI-88, S. Nakahara & P. Stevens. ‘Locust Hall,’ Devonshire, grass, 21 - VI-88, P. Stevens, D. Hilbum & S. Nakahara. Shelly Bay Park, Hamilton, grass, 22- VI-88, P. Stevens & D. Hilbum. Botanical Gardens, Paget, Citharexylum spi- nosum, 23-VI-88, D. Hilbum & S. Nakahara. Ferry Point Park, St. George’s, grass, 24-VI-88; D. Hilbum. According to Waterston, H. dozieri was intercepted in the US on Chrysanthemum from Bermuda in 1923, and H. gowdeyi was intercepted in the US in 1936, 1938 and 1939 on Chrysanthemum sp., Freesia refracta, Lathyrus odoratus and Nerium oleander from Bermuda. Quarantine interceptions made at New York and Boston from Bermuda on Gerbera sp., Lilium longiflorum and pampas grass are deposited in the USNM. A cosmopolitan species in the tropics and subtropics. Haplothrips kurdjumovi Kamy Nakahara, 1985:894. Intercepted at predeparture quarantine inspection at Bermuda: Passijlora sp. flow- ers, 13-VII-68, A. Kaplan & J. Fons; Lonicera sp., l-VII-69, G. Holt; cutflowers, 21 -VI-70, J. Fons; Passijlora sp. flower, 17-VII-70, J. Fons. Hamilton, Pembroke, Ipomoea villosa, 23-III-88, M. Mello. ‘Waterville’ area, Paget, Chrysanthemum fru- tescens, Carpobotrus edulis, Durant a repens, Ipomoea villosa, Lonicera sp.. Nastur- tium officinale, Pimenta officinale, 19-VI-88, S. Nakahara. Admiralty House Park, Pembroke, Solidago sp., 20- VI-8 8, S. Nakahara. Gilbert Nature Reserve, Sandy’s, Carissa grandijlora, 20-VI-88, D. Hilbum & S. Nakahara. ‘Garthowen Estate,’ Dev- onshire, Persea americana, 22- VI-8 8, D. Hilbum; Diant hus caryophyllus, Sambucus sp., S. Nakahara. Shelly Bay Park, Hamilton, Tamarix sp., 22- VI-8 8, S. Nakahara. ‘Windy Bank Farm,’ Smith’s, Bromus sp., 22- VI-8 8, P. Stevens, D. Hilbum & S. Nakahara. Jubilee Road, Devonshire, Clerodendron sp., 22-VI-88, P. Stevens, D. Hilburn & S. Nakahara. Botanical Gardens, Paget, pan trap, 27-III-88, M. B. Stoetzel; Citharexylum spinosum, Punica granatum, Rosa sp., Vitis sp., 23-VI-88, D. Hilbum & S. Nakahara. Ferry Point Park, St. George’s, Jasminum sp., 24-VI-88, S. Nakahara. Paget, Phaseolus vulgaris, 24- VI-8 8, D. Hilburn & S. Nakahara. A predator of mites and moth eggs in Europe, Canada and eastern United States. 254 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Hoplandrothrips sp.* Shelly Bay Park, Hamilton, Tamarix sp., 22-VI-88, S. Nakahara. Hoplothrips marginalis (Hood & Williams) Trichothrips marginalis Hood & Williams: Waterston, 1941:6. Quarantine interception in the US in 1933 on Antirrhinum majus; known from southeastern United States. Karnyothrips flavipes (Jones)* Shelly Bay Park, Hamilton, grass, 22- VI-8 8, D. Hilbum & P. Stevens. A cosmopolitan predator of scale insects, whiteflies and mites (Pitkin, 1976:263) in tropical and subtropical regions. Karnyothrips longiceps (Hood)* Spittal Pond Nature Reserve, Smith’s, sweeping, 27-VI-88, D. Hilbum. Known from United States and Caribbean area. Karnyothrips melaleucus (Bagnall)* Shelly Bay Park, Hamilton, grass, 22- VI-8 8, D. Hilbum & P. Stevens. Spittal Pond Nature Reserve, Smith’s, sweeping, 27-VI-88, D. Hilbum. Species reported preying on a soft scale (Pitkin, 1976:263). Known from various areas of the tropics. Nesothrips lativentris (Kamy)* ‘Windy Bank Farm,’ Smith’s, Bromus sp., 22-VI-88, D. Hilbum, P. Stevens & S. Nakahara. Horseshoe Bay, Southampton, sweeping grasses, 24-VI-88, D. Hilbum. ‘St. Mark’s Church,’ Smith’s, Gladiolus sp., 24- VI-8 8, D. Hilbum & S. Nakahara. Botanical Gardens, Paget, sweeping, 26-VI-88, D. Hilbum. A widespread species feeding on fungal spores in the tropics. Stephanothrips occidentalis Hood & Williams Waterston, 1940:7, 1941:51. On leaves of Sabal bermudana in 1933. Species known from Caribbean Islands, Mexico, India, South Africa, Australia and US (Florida, Hawaii). THRIPIDAE Anisopilothrips venustulus (Priesner) Wilson, 1975:32. Quarantine interception at New York City; Durant a repens, 1-II-36, Field & Kostal. Gilbert Nature Reserve, Sandy’s, leaves of unknown tree, 20- VI-8 8, D. Hilbum & S. Nakahara. Species known from the US (Florida), West Indies, Surinam, Fiji and Taiwan. 1989 THYSANOPTERA OF BERMUDA 255 Asprothrips seminigricornis (Girault)* Gilbert Nature Reserve, Sandy’s, leaves of unknown tree, 20- VI-88, D. Hilbum & S. Nakahara. A species with disjunct distribution and unknown origin; known from Australia, Hawaii, and New York state in greenhouse. Caliothrips insularis (Hood)* Botanical Gardens (in greenhouse), Paget, Stenotaphrum secundatum var.floratum, 16-XII-86, D. Hilbum; sweeping, 26-VI-88, D. Hilbum. Species known from the Caribbean area, Panama and Brazil. Chirothrips spiniceps Hood* Ferry Point Park, St. George’s, grass, 24- VI-8 8, D. Hilbum. An endemic species in the conterminous United States. Dendrothripoides innoxius (Kamy)* Bermuda, Ipomoea sp. leaves, 28-VIII-50, F. D. Bennett. Gilbert Nature Reserve, Sandy’s, grass, 20- VI-8 8, D. Hilbum & S. Nakahara. Reported causing moderate damage to lettuce and yams (Dioscorea sp.) and mod- erate to considerable damage to sweetpotato shoots in Hawaii (Wilson 1975:104). Species known also from Central and South America, West Indies, India and Aus- tralia. Dorcadothrips caespitis Priesner* ‘Windy Bank Farm,’ Smith’s, Bromus sp., 22- VI-8 8, D. Hilbum, P. Stevens & S. Nakahara. Shelly Bay Park, Hamilton, grass, 22- VI-8 8, D. Hilbum & P. Stevens. A species with disjunct distribution; known from Hawaii, Egypt, Sudan, Uganda and India. Echinothrips americanus Morgan Ogilvie, 1928:29. Bermuda, greenhouse plants, VII-27, L. Ogilvie. Paget, Phaseolus vulgaris, 24-VI- 88, D. Hilbum & S. Nakahara. Botanical Gardens, Paget, sweeping, 26-VI-88, D. Hilbum. Endemic species in conterminous United States. Frankliniella bispinosa (Morgan)* Agricultural Station, Gladiolus sp., 1 5-VI-32, T. A. Russell. ‘Waterville’ area, Paget, Lonicera sp., Pimenta officinale, 19-VI-88; S. Nakahara. Par La Ville Park, Hamilton, Pembroke, Agapanthus africana, Brunfelsea americana, 19-VI-88, S. Nakahara. Ad- miralty House Park, Pembroke, Borrichia arborescens, 20-VI-88, S. Nakahara. ‘Lo- cust Hall,’ Devonshire, Antirrhinum majus, Brassica sp., 21 -VI-88, P. Stevens, D. Hilbum & S. Nakahara. Ferry Point Park, St. George’s, Cakile lanceolata, 24-VI- 88, S. Nakahara. 256 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Quarantine interceptions at New York City from 1935 to 1947 on Antirrhinum majus, asters, Dianthus caryophyllus, Freesia sp., Gerbera sp., lily, Lilium harissi, Hibiscus sp., Melia azedarach, Nerium oleander, Rosa sp., Tagetes sp.; specimens deposited in the USNM. Species known from southeastern United States, and Ba- hama Islands. Frankliniella cephalica (Crawford) Waterston, 1940:6, 1941:6, 23, 28, 31, 35-36, 49, 56. Stoke’s Point Nature Reserve, St. George’s, Bidens pilosa, 25-III-88, M. B. Stoetzel & D. Hilbum. ‘Waterville’ area, Paget, Bidens sp., 19-VI-88, S. Nakahara. Spittal Pond Nature Reserve, Smith’s, Bidens sp., Pluchea sp., unknown composite, 2 1 -VI- 88, P. Stevens, D. Hilbum & S. Nakahara. According to Waterston (1940, 1941), this species was intercepted in the US in 1936 and 1937 on Antirrhinum majus, Dianthus sp., Freesia refracta, Gerbera ja- mesoni. Gladiolus sp.. Hibiscus rosa-sinensis, Lathyrus odoratus, Lilium longiflorum var. eximium, Nerium oleander, Rosa spp., Tagetes spp. Some of these records apparently are misidentifications of F. bispinosa. Species known from southern US, Caribbean Islands and Central America. Frankliniella cubensis Hood Waterston, 1941:28. Intercepted in the United States in 1939 on Gerbera jamesoni. Species known from Cuba and several islands in the Caribbean. Frankliniella hemerocallis Crawford* Botanical Gardens, Paget, Hemerocallis sp., 23-VI-88, D. Hilbum, S. Nakahara & P. Stevens. Species is recorded from Japan and US (Florida, Maryland, New York and Wis- consin) on daylily. Frankliniella insularis (Franklin) Ogilvie, 1928:29; Waterston, 1941:6, 10, 20, 23, 28, 31, 35-36, 38, 42, 44, 49, 57. Experiment Station, hibiscus flowers, 13-XII-24, L. Ogilvie. Hamilton, Pembroke, Ipomoea villosa, 23-III-88, M. J. Mello. Botanical Gardens, Paget, pan trap, 27-III- 88, M. Stoetzel. ‘Waterville’ area, Paget, Carpobotrus edulis. Chrysanthemum fru- tescens, Duranta repens, Ipomoea villosa, Lonicera sp., Malvaviscus arboreus, Pi- menta officinale, 19-VI-88, S. Nakahara. Pomander Road, Paget, Hibiscus rosa- sinensis, 19-VI-88, S. Nakahara. ‘Garthowen Estate,’ Devonshire, Dianthus cary- ophyllus, 22-VI-88, S. Nakahara. Jubilee Road, Devonshire, Clerodendron sp., 22- VI-88, D. Hilbum, P. Stevens & S. Nakahara. Botanical Gardens, Paget, Canna sp., Lagerstroemia indica. Magnolia grandiflora, Punica granatum, Rosa sp., 23-VI-88, D. Hilbum & S. Nakahara. According to Waterston (1941), this species was intercepted in the US during 1 932, 1989 THYSANOPTERA OF BERMUDA 257 1933, 1935-1936, and 1938-39, or collected in Bermuda on Antirrhinum majus, Bougainvillea spectabilis, Codiaeum variegatum, Dianthus sp., Gerbera jamesoni. Gladiolus sp.. Hibiscus rosa-sinensis, Lathyrus odoratus, Lilium longiflorum var. eximium, Magnolia grandijlora, Nerium oleander, Passiflora incarnata, Rosa sp., and Vicia faba. Species known from tropical and subtropical areas of the New World. Heliothrips haemorrhoidalis (Bouche) Ogilvie, 1928:28; Waterston, 1941:20. Bermuda Agriculture Station, Codiaeum sp., l-IX-34, T. A. Russell. Bermuda, Acalypha sp., lO-VIII-53, 1. W. Acalypha wilkesiana, 6-X-50, F. D. Bennett; Persea americana, 23-VII-82, K. D. Monkman. ‘Waterville’ area, Paget, Viburnum sp., 19-VI-88, S. Nakahara. Admiralty House Park, Pembroke, Acalypha wilkesiana, Cyrtomium falcatum, 20- VI-8 8, D. Hilbum. Spittal Pond Nature Reserve, Smith’s, Schinus terebinthifolius, 21 -VI-88, S. Nakahara & P. Stevens. ‘Garthowen Estate,’ Devonshire, Persea americana, 21 -VI-88, D. Hilbum. Botanical Gardens, Paget, Rosa sp., 23-VI-88, P. Stevens & S. Nakahara. According to Waterston (1941) this species was intercepted in the United States in 1923 on Codiaeum variegatum. A widespread pest of various fruit and ornamental plants in the tropics and subtropics, and a pest in greenhouses in temperate areas. Hercinothrips bicinctus Bagnall Waterston, 1940:7, 1941:51. Bermuda, banana fruit, 28-VI-50, F. D. Bennett. Intercepted in predeparture quar- antine inspection at Bermuda: Passiflora sp. flower, 1 l-IV-69, J. Fons. Spittal Pond Nature Reserve, Smith’s, Ipomoea villosa, 21 -VI-88; P. Stevens & S. Nakahara. Botanical Gardens, Paget, Araceae, Citharexylum spinosum, Scindapsus aureus, 23- VI-88, D. Hilbum & S. Nakahara; sweeping, 26- VI-8 8, D. Hilbum. Waterston reported that this species was collected in 1933 on leaves of Sabal bermudana. Known from tropical areas of Africa, Asia and Australia, H. bicinctus is a banana pest in the tropics, and a minor pest in greenhouses in temperate areas (Wilson, 1975:156). Hercinothrips femoralis (Reuter) Heliothrips femoralis Reuter: Ogilvie, 1928:28; Waterston, 1941:32, 40. Bermuda, greenhouse plant, VII-27, L. Ogilvie; banana fruit, 28-VI-50, F. D. Bennett. Botanical Gardens, Paget, pan trap, 27-III-88, M. B. Stoetzel; Araceae, 23- VI-88, D. Hilbum & S. Nakahara. Gilbert Nature Reserve, Sandy’s, Citharexylum spinosum, grass, leaves of unknown tree, 20- VI-8 8, D. Hilbum & S. Nakahara. Seymour Pond, Southampton, Allium cepa leaves, 20- VI-88, D. Hilbum & S. Na- kahara. Waterston reports it from Ipomoea cathartica and Musa cavendishii. Widespread species in the tropics and subtropics and in greenhouses in Europe and the United States. 258 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Microcephalothrips abdominalis (D. L. Crawford)* Quarantine interception at New York City; Zinnia sp., 2-IX-49, Chapman & Trafford. Pomander Road, Paget, Wedelia trilobata, I9-VI-88, S. Nakahara. ‘Water- ville’ area, Paget, Bidens sp., 19-VI-88, S. Nakahara. Spittal Pond Nature Reserve, Smith’s, Bidens sp., Borrichia arborescens, Pluchea odorata, 2 1 -VI-88, P. Stevens & S. Nakahara. Botanical Gardens, Paget, Tagetes sp.. Salvia farinacea, 23-VI-88, S. Nakahara. Horseshoe Bay, Southampton, Solidago sp., 24- VI-8 8, S. Nakahara. Species known from the New World, Hawaii and the Orient. Neohydatothrips portoricensis (Morgan)* Seymour Pond, Southampton, Allium cepa, 20-VI-88, D. Hilbum & S. Nakahara. ‘Locust Hall,’ Devonshire, Ipomoea batatas, 21 -VI-88, P. Stevens, D. Hilbum & S. Nakahara. Botanical Gardens, Paget, sweeping, 26-VI-88, D. Hilbum. Known from Puerto Rico, Lesser Antilles and South America. Plesiothrips perplexus (Beach)* Paget, Phaseolus vulgaris, 24-VI-88, D. Hilbum & S. Nakahara. Spittal Pond Nature Reserve, Smith’s, sweeping, 27-VI-88, D. Hilbum. A New World species known also from Hawaii, Guam and several South Pacific Islands. Rhamphothrips pandens Sakimura* ‘Garthowen Estate,’ Devonshire, Sambucus sp., 22-VI-88; S. Nakahara. A species with disjunct distribution; known from Brazil, Jamaica, Florida, Hawaii, and several South Pacific Islands. Selenothrips rubrocinctus (Giard) Waterston, 1940:7, 1941:10. ‘Waterville’ area, Paget, Viburnum sp., 19-VI-88, S. Nakahara. Spittal Pond Nature Reserve, Smith’s, Coccoloba uvifera, Schinus terebinthifolius, 21 -VI-88, P. Stevens & S. Nakahara. Shelly Bay Park, Hamilton, Tamarix sp., Schinus terebinthifolius, 22-VI-88, D. Hilbum, P. Stevens & S. Nakahara. Botanical Gardens, Paget, Cith- arexylum spinosum, Scindapsus aureus, Schinus terebinthifolius, 23-VI-88, D. Hil- bum & S. Nakahara. Ferry Point Park, St. George’s, Schinus terebinthifolius, 24- VI- 88, S. Nakahara. According to Waterston, this species was intercepted in the United States on Bougainvillea spectabilis in 1932 from Burmuda. A serious pest of cacao, cashew and mango plants (Wilson 1975:234). It occurs in tropical and subtropical areas of the New World, Southeast Asia and Africa. Taeniothrips eucharii (Whetzel) Physothrips eucharii 1923:30; Ogilvie, 1928:29; Waterston, 1941:25; Tae- niothrips eucharii (Whetzel): O’Neill, 1962:400. 1989 THYSANOPTERA OF BERMUDA 259 Paget, East Bermuda, Eucharis (lily), 11-22, H. H. Whetzel. Waterston (1941) reports it from Eucharis grandiflora. Quarantine interceptions at United States ports of entry from Bermuda on Amaryllis, carnations, cutflowers, easter lily, Ereesia, Hibiscus and Lilium deposited in the USNM. An Oriental species known from Japan, China and southeastern United States, infesting mainly bulbs, leaves and flowers of Amaryllidaceae. Thrips hawaiiensis (Morgan)* Botanical Gardens, Paget, pan trap, 27-III-88, M. B. Stoetzel. ‘Waterville’ area, Paget, Chrysanthemum frutescens, Carpobotrus edulis, Duranta repens, Ipomoea vil- losa, Lonicera sp.. Nasturtium officinale, Pimenta officinale, 19-VI-88, S. Nakahara. Par La Ville Park, Hamilton, Pembroke, Agapanthus africana, Brunfelsea americana, 19-VI-88, S. Nakahara. Admiralty House Park, Pembroke, Leucaena glauca, Soli- dago sp., 20-VI-88, S. Nakahara. Gilbert Nature Reserve, Sandy’s, Carissa grandi- flora, 20- VI-88, D. Hilbum & S. Nakahara. ‘Locust Hall,’ Devonshire, Antirrhinum majus, grass, Lantana sp., 21 -VI-88, D. Hilbum, P. Stevens, & S. Nakahara. Spittal Pond Nature Reserve, Smith’s, Bidens sp., Borrichia arborescens, Carissa grandiflora, 21 -VI-88, P. Stevens & S. Nakahara. ‘Garthowen Estate,’ Devonshire, Dianthus caryophyllus, Sambucus sp., 22-VI-88, S. Nakahara. Jubilee Road, Devonshire, Cle- rodendron sp., 22-VI-88, P. Stevens, D. Hilbum & S. Nakahara. Botanical Gardens, Paget, Abelia sp., Citharexylum spinosum, Hemerocallis sp., Lagerstroemia indica, Magnolia grandiflora, Rosa sp.. Salvia farinacea, 23-VI-88, P. Stevens, D. Hilbum & S. Nakahara. Ferry Point Park, St. George’s, Jasminum sp., 24- VI-88, S. Nakahara. Paget, Phaseolus vulgaris, 24- VI-8 8, D. Flilbum & S. Nakahara. Species known from the Orient, Pacific Islands and southern United States. Dam- ages coffee, mango, citrus, apple, pear, passion fruit, rose and banana, and is a beneficial pollinator of certain flowers in India (Palmer and Wetton, 1987:397). Thrips simplex (Morison) Taeniothrips simplex Movi^orv: Waterston, 1940:7, 1941:29, 31. Devonshire, 26-V-32, T. A. Russell; Gladiolus, 14-11-33, T. A. Russell; Gladiolus, 24-11-33, T. A. Russell. Bermuda Agriculture Station, Gladiolus, 15-VI-32, T. A. Russell; 22-VI-32, T. A. Russell. Paget Marsh, Paget, Sabal bermudana, 24-1-33, T. A. Russell. Waterston (1941) lists Gladiolus sp. and Hippeastrum vittatum as hosts. A cos- mopolitan pest of gladiolus. Thrips t abaci Lindeman Ogilvie, 1928:29; Waterston, 1940:7, 1941:4. Bermuda Agriculture Station, Brassica oleracea, 24-11-33, T. A. Russell. Triming- ham Hill, Paget, Brassica oleracea, 24-III-88, M. B. Stoetzel, D. Hilbum & M. J. Mello. ‘Waterville’ area, Paget, Chrysanthemum frutescens, Pimenta officinale, 19- VI-88, S. Nakahara. Par La Ville Park, Hamilton, Pembroke, Agapanthus africana, 19-VI-88, S. Nakahara. Admiralty House Park, Pembroke, Solidago sp., 20-VI-88, 260 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) S. Nakahara. Seymour Pond, Southampton, Allium cepa, 20- VI-8 8; D. Hilbum & S. Nakahara. Spittal Pond Nature Reserve, Smith’s, Carissa grandijlora, 2 1 -VI-88, P. Stevens & S. Nakahara. ‘Locust Hall,’ Devonshire, Brassica sp., grass, 21 -VI-88, P. Stevens, D. Hilbum & S. Nakahara. ‘Garthowen Estate,’ Devonshire, Allium cepa, 22-VI-88, D. Hilbum; Dianthus caryophyllus, Sambucus sp., 22-VI-88, S. Nakahara. Shelly Bay Park, Hamilton, Foeniculum vulgare, 22- VI-8 8, S. Nakahara. Botanical Gardens, Paget, Canna sp., 23-VI-88, S. Nakahara & D. Hilbum. Ferry Point Park, St. George’s, Cakile lanceolata, 24-VI-88, S. Nakahara. Reported by Waterston (1941) on Allium cepa, A. porrum and Brassica oleracea. A cosmopolitan pest of onions and other plants and a vector of tomato spotted wilt virus. ACKNOWLEDGMENTS We thank the following reviewers of the manuscript for their comments and suggestions: R. J. Beshear, University of Georgia, Experiment; D. C. Ferguson, Systematic Entomology Lab- oratory, Washington, D.C.; M. B. Stoetzel, same laboratory, Beltsville, Maryland; W. Sterrer, Bermuda Natural History Museum, Flatts; and R. Dow and K. D. Monkman, Bermuda Dept. of Agriculture and Fisheries, Paget. We are grateful to R. Dow for checking the scientific names of the host plants. LITERATURE CITED Bennett, F. D., 1. W. Hughes, F. J. Simmonds, and M. Yaseen. 1985. Biological control in Bermuda. In: M. J. W. Cock (ed.), A review of biological control of pests in the Com- monwealth Caribbean and Bermuda up to 1982. Commonwealth Inst. Biol. Cont. Tech. Com. No. 9:119-142. Kevan, D. K. McE. 198L The terrestrial arthopods of the Bermudas; an historical review of our knowledge. Archives Nat. Hist. 10(1): 1-29. Lewis, T. 1973. Thrips, Their Biology, Ecology and Economic Importance. Academic Press, London and New York, 349 pp. Nakahara, S. 1985. Haplothrips kurdjumovi Kamy in North America with a new junior synonym (Thysanoptera: Phlaeothripidae). Proc. Entomol. Soc. Washington 87(4):894- 895. Ogilvie, L. 1928. The insects of Bermuda. Bermuda Dept. Agric. Pub., 52 pp. O’Neill, K. 1962. An Oriental Taeniothrips (Thysanoptera: Thripidae) infesting certain Ama- ryllidaceae. Ann. Entomol. Soc. America 56:399-401. Palmer, J. M. and M. N. Wetton. 1987. A morphometric analysis of the Thrips hawaiiensis (Morgan) species-group (Thysanoptera: Thripidae). Bull. Entomol. Res. 77:397-406. Pitkin, B. R. 1976. A revision of the Indian species of Haplothrips and related genera (Thysa- noptera: Phlaeothripidae). Bull. Brit. Mus. (Nat. Hist.) Entomol. 34(4):223-280. Stannard, L. J. 1952. Phylogenetic studies of Franklinothrips (Thysanoptera: Aeolothripidae). J. Washington Acad. Sci. 42(1): 14-23. Waterston, J. M. 1940. Supplementary list of Bermuda insects. Bermuda Dept. Agric. Pub., 10 pp. Waterston, J. M. 1941. A list of food-plants of some Bermuda insects. Bermuda Dept, of Agric. Pub., 63 pp. Whetzel, H. H. 1923. Report of the plant pathologist for the period January 1st to May 31st 1922. [Bermuda] Repts. Board and Dept. Agric. for 1922:30-37. Wilson, T. H. 1975. A monograph of the subfamily Panchaetothripinae (Thysanoptera: Thripi- dae). Mem. American Entomol. Inst. No. 23, 354 pp. Received December 12, 1988; accepted January 25, 1989. J. New York Entomol. Soc. 97(3):26 1-264, 1989 ANNOTATED CHECKLIST OF THE WHITEFLIES (HOMOPTERA: ALEYRODIDAE) OF BERMUDA SuEO Nakahara' and Daniel J. Hilburn^ ‘Systematic Entomology Laboratory, Agricultural Research Service, USD A, Beltsville, Maryland 20705 and ^Department of Agriculture and Fisheries, P.O. Box HM834, Hamilton HMCX, Bermuda Abstract.— Ttn species of whiteflies (Aleyrodidae) representing seven genera are reported from Bermuda. One species, Aleurodicus cocois (Curtis), has not been found since its original collection and apparently is not established in Bermuda. Nine species were introduced from eastern United States or the Caribbean region, and one species, Aleyrodes proletella (L.), was apparently introduced from Great Britain. Collection data, hosts, and general distribution are presented for each species, and comments are made about the economic importance of several species. A historical account of the insects of Bermuda was provided by Nakahara and Hilbum (1989). As part of a project initiated by the junior author to upgrade the insect collection in the Bermuda Department of Agriculture and Fisheries (BDAF) and to update the knowledge of Bermuda’s insect fauna, we surveyed the islands for whiteflies (Aleyrodidae) in June, 1988. Four species, Aleyrodes proletella (L.), Paraley- rodes sp., Trialeurodes floridensis (Quaintance), and T. notata Russell, were found for the first time in Bermuda during this survey. Two other species, Paraleyrodes naranjae (Dozier) and Bemisia tabaci (Gennadius), found in Bermuda prior to and after the survey have not been reported in the literature. Of the ten species in seven genera reported here from Bermuda, only Aleurodicus cocois (Curtis) apparently is not established. Some of the native plants were examined for endemic whiteflies, but only introduced species were found. Nine of the species were introduced either from eastern United States or from the Caribbean region, and one species, Aleyrodes proletella (L.), was introduced apparently from Great Britain. In addition to the new species records, the following list includes species and host plants reported by Ogilvie (1928), Waterston (1941) and Mound and Halsey (1978). Six species reported for the first time from Bermuda are indicated with asterisks. Voucher specimens are deposited in the Bermuda National History Museum, which houses the main portion of the BDAF collection, and/or the whitefly collection of the United States National Museum of Natural History, Beltsville, Maryland. Aleurodicus cocois (Curtis) Ogilvie, 1928:23. According to Ogilvie, this species was intercepted on coconut palms from Trinidad, in February 1928. Since then, it has not been collected in Bermuda. Aleurodicus cocois is reported from the Caribbean Islands, Central America, and several countries in South America. 262 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Aleyrodes proletella (L.)* ‘Waterville,’ Paget, Bmssica sp., 19-VI-88; S. Nakahara. Botanical Gardens, Paget, Brassica sp., 23-VI-88, D. Hilbum & S. Nakahara. Reported by Mound & Halsey (1978:99) from Europe, USSR, Canary Islands, Egypt, Morocco, Kenya, Angola, Mozambique, Brazil and New Zealand. A poly- phagous species; according to Mound (1966:404), an occasional pest of brussels- sprouts and cabbages in England. Bemisia tabaci (Gennadius)* Pomander Road, Paget, Euphorbia pulcherrima, 27-X-88, K. D. Monkman. A cosmopolitan species and serious pest of various agricultural crops in tropical and subtropical regions of the world, and a vector of various viral diseases. In recent years, it has been found in greenhouses in colder regions on ornamental plants, especially poinsettias. Dialeurodes citrifolii (Morgan) Ogilvie, 1928:23; Waterston, 1941:18. Devonshire, Citrus sinensis, 22-VI-88, D. Hilbum & S. Nakahara. ‘Windy Bank Farm,’ Smith’s, Citrus sinensis, 22-VI-88, D. Hilbum, P. Stevens & S. Nakahara. Botanical Gardens, Paget, Citrus sp., 23-VI-88, S. Nakahara. Recorded from the Orient and New World, and a pest of Citrus. Metaleurodicus cardini (Back) Mound and Halsey, 1978:244; Bennett et al., 1985:122. Gilbert Nature Reserve, Sandy’s, Citharexylum spinosum, 20- VI-88, D. Hilbum & S. Nakahara. Devonshire, Citrus sp., 22-VI-88, S. Nakahara & D. J. Hilbum. Botanical Gardens, Paget, Psidium cattleianum, 23-VI-88, S. Nakahara. A common species in Bermuda on fiddlewood, Citharexylum spinosum-, known from several Caribbean Islands and United States (Florida). Paraleyrodes naranjae (Dozier) (as currently recognized)* Devonshire, Citrus sp., 13-X-86, D. Hilbum. ‘Windy Bank Farm,’ Smith’s, Citrus sinensis, 22- VI-8 8, D. Hilbum, P. Stevens & S. Nakahara. Botanical Gardens, Paget, Citrus sp., 23-VI-88, S. Nakahara. Recorded from the West Indies and US (Florida and Hawaii). Paraleyrodes sp.* Botanical Gardens, Paget, Citrus sp., 23-VI-88, S. Nakahara. This is apparently the same undescribed species known from Mexico and United States (California, Texas, Florida) on Citrus, Chamaedorea, and other hosts. 1989 WHITEFLIES OF BERMUDA 263 Trialeurodes floridensis (Quaintance)* ‘Waterville’ area, Paget, Persea americana, 19 & 23-VI-88, S. Nakahara. This species is known from the US (Arizona, Florida), Caribbean Islands, Mexico, and Central America. Trialeurodes not at a Russell* Spittal Pond Nature Reserve, Smith’s, Ipomoea villosa, 21 -VI-88, S. Nakahara. Russell (1963:151) records this species in US from District of Columbia, Illinois, Kansas, Maryland, Missouri, New York, Pennsylvania and Virginia. Trialeurodes vaporariorum (Westwood) Ogilvie, 1928:23; Waterston, 1941:4, 16, 17, 21, 22, 26, 30, 50, 54, 56, 59. ‘Waterville’ area, Paget, Lonicera sp.. Nasturtium officinale, Sonchus oleraceus, 19-VI-88, S. Nakahara. Spittal Pond Nature Reserve, Smith’s, Euphorbia sp., 21- VI-88, S. Nakahara. ‘Garthowen Estate,’ Devonshire, rosea, Helianthus annuus, 22-VI-88, S. Nakahara & D. Hilbum. Other recorded host plants are Ageratum houstonianum, Citharexylum spinosum, Citrullus lanatus, Cucumis melo, C. sativus, Cucurbita maxima. Euphorbia peplus, Helichrysum bracteatum, Lycopersicon esculentum, Psidium guajava, Ruta graveo- lens, Solanum tuberosum, Tithonia rotundiflora. Verbena sp. and Zinnia elegans (Ogilvie, 1928; Waterston, 1941). A cosmopolitan species out-of-doors and in greenhouses. A serious pest of various agricultural crops and reported as a vector of several viral diseases in greenhouses in Europe and Japan. ACKNOWLEDGMENTS We thank the following reviewers of the manuscript for their comments and suggestions: R. J. Gill, California Dept, of Food and Agriculture, Sacramento; A. S. Menke, Systematic Ento- mology Laboratory, Washington, D.C.; R. L. Smiley, same laboratory, Beltsville, Maryland; and R. L. Dow and K. D. Monkman, Bermuda Dept, of Agriculture and Fisheries, Paget. LITERATURE CITED Bennett, F. D., M. J. W. Cock, I. W. Hughes, F. J. S. Simmonds, and M. Yaseen. 1985. Biological control in Bermuda. In: M. J. W. Cock (ed.), A review of biological control of pests in the Commonwealth Caribbean and Bermuda up to 1982. Commonwealth Inst. Biol. Contr. Tech. Com. No. 9:119-142. Mound, L. A. 1966. A revision of the British Aleyrodidae (Hemiptera: Homoptera). Bull. Brit. Mus. (Nat. Hist.) Entomol. 17(9):399-428. Mound, L. A. and S. H. Halsey. 1978. Whitefly of the World. A Systematic Catalogue of the Aleyrodidae (Homoptera) with Host Plant and Natural Enemy Data. British Museum (Nat. Hist.) and John Wiley and Sons, 340 pp. Nakahara, S. and D. J. Hilbum. 1989. Annotated checklist of the Thysanoptera of Bermuda. J. New York Entomol. Soc. (in press). Ogilvie, L. 1928. The insects of Bermuda. Dept. Agric. Bermuda Pub., 52 pp. 264 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Russell, L. M. 1963. Hosts and distribution of five species of Trialeurodes (Homoptera: Aleyrodidae). Ann. Entomol. Soc. Am. 56(2): 149-1 53. Waterston, J. M. 1941 . A list of food-plants of some Bermuda insects. Dept. Agric. Bermuda Pub., 63 pp. Received February 6, 1989; accepted March 28, 1989. J. New York Entomol. Soc. 97(3):265-270, 1989 BLISSUS BREVIUSCULUS: NEW DISTRIBUTION RECORDS OF A LITTLE-KNOWN CHINCH BUG (HETEROPTERA: LYGAEIDAE) A. G. Wheeler, Jr.* and Jonathan E. Fetter^ •Bureau of Plant Industry, Pennsylvania Department of Agriculture, Harrisburg, Pennsylvania 17110 ^Department of Entomology, University of Wisconsin, Madison, Wisconsin 53706 Abstract.— Blissus breviusculus Barber is a poorly known blissine lygaeid that is rare in col- lections because of its cryptic habits; nymphs and adults live deep within crowns of little bluestem, Schizachrium scoparium (Michx.) Nash (Graminiae). Known previously from Con- necticut, Maine, and Massachusetts, B. breviusculus is recorded form a serpentine barren in southeastern Pennsylvania. A suction-sampling machine (D-Vac) was used to extract specimens from crowns of its host plant. A New York record, based on a specimen collected prior to Barber’s original description and housed in the U.S. National Museum of Natural History collection, is given; two additional Massachusetts localities, based on USNM specimens, are also cited. Parshley’s (1917) record oi"" Blissus hirtus Montandon” from Maine is referred to B. breviusculus. Blissus breviusculus Barber belongs to the lygaeid subfamily Blissinae, an Old and New World group restricted to monocotyledonous plants, particularly grasses. Rather than feeding on seeds like most other Lygaeidae, a great many of which live in litter, blissines extract sap from leaves, stems, and roots of host plants (Slater, 1976). As currently recognized, the genus is limited to the Western Hemisphere (Slater, 1979). Its members include the infamous chinch bug, B. leucopterus leucopterus (Say), a grain pest that threatened early North American agriculture; B. leucopterus hirtus, a major pest of turfgrasses in New England and the mid- Atlantic states; and B. insularis Barber, a well-known lawn pest in the southern United States (Leonard, 1966; Tashi- ro, 1987). Other species, especially those associated with native grasses in the western states and midwestern prairies, or living along coastal dunes, are seldom collected. Herein, we give new distribution records for a little-known eastern species, B. breviusculus. Listed previously only from three localities in New England, this blissine is recorded from a serpentine barren in southeastern Pennsylvania. An unpublished New York record, based on a museum specimen collected prior to Barber’s original description, is given along with two additional Massachusetts records based on mu- seum specimens. We also have determined that Parshley’s (1917) record of Blissus hirtus Montandon” from Maine should be referred to B. breviusculus. Voucher spec- imens from Pennsylvania have been deposited in the collections of Cornell University (CUIC), Pennsylvania Department of Agriculture (PDA), and U.S. National Museum of Natural History (USNM). 266 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Blissus breviusculus Barber Barber (1937) described this small (about 2.4 mm), black, densely pilose bug (Fig. 1) from four specimens from Wareham, Massachusetts, and one from Harbor, Maine. ^ The Maine specimen would appear to be the one Parshley (1917) cited as B. hirtus Montandon from the same locality; month, day, and collector also are identical, and the year, 1912, could have been misread by Barber as 1915. The only other published record of this poorly known lygaeid is New Haven, Connecticut, where Leonard (1968) collected nymphs (instars I-V) and adults from crowns of little bluestem, Schizachrium scoparium (Michx.) Nash, a perennial bunch grass formerly placed in the genus Andropogon. Leonard (1968) erroneously credited Barber (1937) with saying that this species “. . . was collected on a lake shore, by shaking out clumps of grass.” In the original description, however, the Massachusetts specimens were said to have been taken under stones by C. A. Frost. We have not located a published source of the statement Leonard attributed to Barber, but four specimens in the USNM collection, taken by Frost at Framingham, Massachusetts, subsequent to the original description (30 May 1947), bear labels indicating they were collected on the shore of a farm pond from tufts of grass growing in sand. Frost collected another specimen near this pond on 30 Sept. 1949. The USNM collection also contains a specimen collected by Frost at Berlin, Massachusetts, 30 July 1937, by sweeping grass, and one from Indian Lake, Sabael, New York, 19 Aug. 1921 (collector unknown). The latter specimen, bearing H. G. Barber’s undated determination label, undoubtedly was identified after he described this species. As Leonard (1968) noted, B. breviusculus lives deep within crowns of S. scoparium and is difficult to collect. Our discovery of this lygaeid in Pennsylvania was fortuitous. We found two adults among arthropod material extracted with a Berlese funnel from little bluestem plants that had been dug on 21 June 1988 from the New Texas barren south of Wakefield (Lancaster Co.) near the Maryland state line (Fig. 2). This ser- pentine community of about 103 ha (255 acres) is characterized by large open areas of little bluestem and other grasses (Wheeler, 1988). We obtained additional material by using a D-Vac machine (Dietrick, 1961) to suck nymphs and adults from crowns of S. scoparium. On 18 August, we collected 4 third-instar nymphs and 3 fourth instars; 6 fifth instars and 22 adults were taken on 16 September. All adults collected were brachypterous. Sweet (1964) discussed wing polymor- phism in lygaeids, emphasizing that brachyptery usually is associated with relatively permanent habitats. The serpentine barrens are such a habitat, having had minimal human disturbance. Blissus breviusculus does occur in more temporary habitats (e.g., in grass along the shore of farm ponds), and macropterous individuals are known from New England populations of this typically brachypterous species (Leonard, 1968). Slater (1977) noted that in addition to habitat permanency, the “ecological- time” stability of the bug’s specific habitat (niche) is important in the development of flightlessness in the Lygaeidae. He concluded that wing reduction is apparently ^ “Harbor, Maine” perhaps should refer to Bar Harbor, Northeast Harbor, Seal Harbor, Southwest Harbor, or other locality in the Mount Desert region of Hancock County that has “Harbor” as part of its name. 1989 BLISS US BRE VIUSCUL US 267 Fig. 1. Blissus breviusculus, adult habitus. restricted to litter-inhabiting species and to monocot sheath-living species (lamina- philes). For B. breviusculus, leaf sheaths of little bluestem in serpentine barrens would seem to provide a specific habitat or niche of ecological-time stability (sensu Slater, 1977). Although we found B. breviusculus in different areas of the New Texas barren, we were unable to collect it in two other state line barrens— Goat Hill and Nottingham County Park (Wheeler, 1988)— even when a D-Vac machine was used. This lygaeid 268 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Fig. 2. Known distribution of Blissus breviusculus. Open circles represent previously pub- lished records; closed circles represent new records: New York (and additional Massachusetts localities) based on USNM specimens, and Pennsylvania, based on our recent collecting. appears to be abundant and generally distributed at New Texas but may not be present in all nearby barrens. Its absence from similar habitats might be expected. Serpentine barrens have been described as relatively undisturbed ecological islands within the eastern deciduous forest. The sparse, usually stunted vegetation of these nutrient-poor soils, which contrasts with the surrounding forest or farmland, is char- acterized by endemic or near-endemic species, morphological variants, and plants of disjunct distribution (Proctor and Woodell, 1975; Miller, 1977; Mansberg and Wentworth, 1984; Wheeler, 1988). Diabrotica cristata, a chrysomelid characteristic of midwestern prairies, recently was found in five eastern serpentine barrens but could not be collected in a sixth, nearby barren (Wheeler, 1988). Blissus breviusculus is now known from habitats ranging from rather temporary 1989 BLISSUS BREVIUSCULUS 269 (shore of a farm pond) to more permanent (serpentine barrens); almost certainly it is more widely distributed than current records reflect (Fig. 2). Additional information on distribution of this and other seldom-collected species of the genus might be useful in establishing biogeographic patterns and, in a broad sense, contribute to phyloge- netic analyses of the group. Blissus breviusculus and the midwestem B. iowensis, another rarely collected species, not only are morphologically similar but are univoltine lygaeids that may specialize on little bluestem. They may have had a common ancestor that was widely distributed across the country during xeric phases of the Pleistocene and, when climate became cooler and more humid, became isolated by the return of forests in the East. In contrast, economically important Blissus spp. are multivoltine (Leonard, 1 966, 1 968) and tend to have a broader host range (Slater, 1976). The poorly known members of the genus should not be ignored. A better understanding of their habits might help elucidate the evolution of host relationships in Blissus and help explain why certain species have become pests (e.g., with selection away from the brachypterous condition in midwestem B. 1. leucopterus), while B. breviusculus and others remain obscure. ACKNOWLEDGMENTS We are grateful to J. F. Stimmel (BPI, PDA) for assistance in the field, the photograph used in Figure 1 , and comments on the manuscript; K. Valley (BPI, PDA) for reading the manuscript; T. J. Henry (Systematic Entomology Laboratory, USDA, % U.S. National Museum of Natural History, Washington, D.C.) for confirming the identification of B. breviusculus and providing label data from USNM specimens; and J. A. Slater (Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs) for valuable discussion and for reviewing the manu- script. LITERATURE CITED Barber, H. G. 1937. Descriptions of six new species of Blissus (Hemiptera-Heteroptera: Ly- gaeidae). Proc. Entomol. Soc. Wash. 39:81-86. Dietrick, E. J. 1961. An improved backpack motor fan for suction sampling of insect pop- ulations. J. Econ. Entomol. 54:394-395. Leonard, D. E. 1966. Biosystematics of the ^"leucopterus complex” of the genus Blissus (Het- eroptera: Lygaeidae). Conn. Agric. Exp. Stn. Bull. 611 , 47 pp. Leonard, D. E. 1 968. A revision of the genus Blissus (Heteroptera: Lygaeidae) in eastern North America. Ann. Entomol. Soc. Am. 61:239-250. Mansberg, L. and T. R. Wentworth. 1984. Vegetation and soils of a serpentine barren in western North Carolina. Bull. Torrey Bot. Club 1 1 1:273-286. Miller, G. L. 1977. An ecological study of the serpentine barrens in Lancaster County, Penn- sylvania. Proc. Pa. Acad. Sci. 51:169-176. Parshley, H. M. 1917. Fauna of New England. 14. List of the Hemiptera-Heteroptera. Occas. Pap. Boston Soc. Nat. Hist. 7:1-125. Proctor, J. and S. R. J. Woodell. 1975. The ecology of serpentine soils. Adv. Ecol. Res. 9: 255-366. Slater, J. A. 1976. Monocots and chinch bugs: a study of host plant relationships in the lygaeid subfamily Blissinae (Hemiptera: Lygaeidae). Biotropica 8:143-165. Slater, J. A. 1977. The incidence and evolutionary significance of wing polymorphism in lygaeid bugs with particular reference to those of South Africa. Biotropica 9:217-229. Slater, J. A. 1979. The systematics, phylogeny, and zoogeography of the Blissinae of the world (Hemiptera, Lygaeidae). Bull. Am. Mus. Nat. Hist. 165:1-180. 270 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Sweet, M. H. 1964. The biology and ecology of the Rhyparochrominae of New England (Heteroptera: Lygaeidae). Parts I, II. Entomol. Am. 43:1-124; 1-201. Tashiro, H. 1987. Turfgrass Insects of the United States and Canada. Cornell University Press, Ithaca and London, 39 1 pp. Wheeler, A. G., Jr. 1988. Diabrotica cristata, a chrysomelid (Coleoptera) of relict midwestem prairies discovered in eastern serpentine barrens. Entomol. News 99:134-142. Received December 14, 1988; accepted February 6, 1989. J. New York Entomol. Soc. 97(3):27 1-276, 1989 THREE NEW SPECIES OF UNCUS (HEMIPTERA: PENTATOMIDAE) FROM PALMS L. H. Rolston Department of Entomology, Louisiana Agricultural Experiment Station, Louisiana State University Agricultural Center, Baton Rouge, LA 70803 Abstract.— ThrQQ new ochlerine pentatomid species, Lincus hebes, L. spurcus and L. male- volus, are described from specimens collected in Peru on various palms. Of the 30 species treated in my revision of Lincus, only two specimens were accompanied by data that presumably indicate host plants (Rolston, 1983). Both specimens were L. vandoesburgi Rolston from Surinam, one taken “on roots [of] Liberian coffee” and the other on “oliepalm” (oil palm). Since then three species of Lincus have been reported from palms, L. lethifer Dolling, L. apollo Dolling and L. croupius Rolston (Dolling, 1984). L. lethifer is known to transmit “marchitez” of oil palm and the other two species have been associated with “heartrot” of coconut, both diseases caused by the flagellate Phytomonas staheli. The three new species described here, all from Peru, were also collected on palm. Lincus hebes, new species Figs. 1-5 Description. Juga projecting little if any beyond apex of tylus, their lateral margins reflexed for entire length, somewhat constricted above antennifers (Fig. 4). Eyes small, each 0.24-0.30 of interocular width; distance across ocelli 0.92-1.00 of interocular width. Vertex of head moderately convex. Anterolateral margins of pronotum sin- uous, reflexed, most strongly reflexed along convexity immediately behind pronotal lobes; pronotal lobes depressed, curving dorsad, truncate apically, extending ante- riorly beyond middle of eyes and laterally beyond eyes about 1.0- 1.5 width of eye; emargination behind each lobe relatively shallow, distance across pronotum at this point about 1 .4 times width of head across eyes. Metastemum slightly concave, without Carina. Proctiger protruding far past posterior margin of pygophore, curving well above last tergite (Fig. 3); apex narrowly rounded from dorsal and caudal views (Figs. 1, 2). Pygophoral emargination U-shaped but narrowly rounded ventrally, thinly rimmed, with broad, shallowly concave border interrupted mesoventrally by confluence of rims into narrow, mesial carina (Fig. 2). Genital cup a simple concavity without setal tufts or conspicuous carinae, but rugose posteriorly. Genital plates as in Figure 5; posterolateral border of basal plates impressed; second gonocoxae polished. Dorsum fuscous with few to several small, pale, interstitial macules scattered on hemelytra and usually on scutellum and pronotum; lateral margins of tylus pale bordered toward apex; pale, marginal macule often present on each of some or all connexival sclerites. Pronotum and scutellum appreciably rugose, particularly toward base of each, with punctures arranged in irregular transverse lines. 272 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Figs. 1-5. L. hebes. 1. Pygophore, dorsal view. 2. Same, caudoventral view. 3. Same, lateral view. 4. Head and anterior portion of pronotum. 5. Genital plates, caudoventral view. Symbols: bp, basal plate; gx2, second gonocoxae; pr, proctiger. Venter fuscous, partially pale bordered on each side of pronotum and base of hemelytron, and sometimes on abdominal stemites. Three basal segments of antennae fuscous, apical two segments lighter in color with basal half of last segment and sometimes apical half of segment 4 cream; rostrum, trochanters, sides of tibiae, tarsi and trichobothrial tubercles usually light brown or yellowish brown. Punctuation on abdominal venter shallow, moderately dense. Measurements (mm). Width of head 1 .75-1 .90, length 1 .85-2. 10; interocular width 1.10-1.20; distance across ocelli 1.05-1.15, between ocelli 0.80-0.95, from each ocellus to nearest eye 0.25-0.30. Length of segments 1-5 of antennae 0.60-0.70, 0.60-0.90, 0.90-1.00, 0.95-1.10, 1.2. Length of segments 1-4 of labium 1.10-1.30, 2.60-2.85, 2.00-2.30, 2.15-2.30. Pronotum 4. 7-5. 7 wide, 2. 2-2. 7 long mesially. Scutellum 2. 7-3. 6 wide at base, 3. 7-4.4 long. Body length, excluding proctiger of males, 9.2-1 1.3. Distribution. Peru (Madre de Dios). 1989 NEW SPECIES OF UNCUS 273 Holotype. Male, labeled “Perou-Mazuco, (Madre de Dios), 27 sept. 1987, F. Kahn & J. Llosa coll.” and “sur Astrocaryum sp. aff. A. macrocalyx Burrett. ref. FK2094.” Deposited in the Museum National d’Histoire Naturelle, Paris. Paratypes. 3 females and 7 males with same labeling as holotype, and 6 males labeled “Perou, Puerto Maldonado, (Madre de Dios), 1 novem. 1987, F. Kahn & J. Llosa coll.” and “sur Astrocaryum sp. aff A. macrocalyx Burrett, ref FK2147.” Comments. This species belongs among the “little eyed” group of species in which the width of each eye is less than one-half of the interocular width. The only other species of this group in which the proctiger is known to project well beyond the posterior pygophoral margin is varius Rolston, although armigera Breddin and lev- iventris Rolston are known only from the female holotypes. In my key to Lincus species (Rolston, 1983), this species runs to armigera, but the two species apparently differ significantly in the form and size of the pronotal lobes. Lincus spurcus, new species Figs. 9-11, 15, 16 Description. Juga projecting slightly past tylus, their lateral margins reflexed along preocular concavity, parallel between this concavity and apical convexity. Width of eye 0.41-0.46 of interocular width; distance across ocelli 0.95-1.00 of interocular width. Ridge on each side of head running along ventral surface from base of head to antennifer weak, not differentially colored. Vertex of head moderately convex. Anterolateral pronotal margins sinuous, somewhat reflexed; pronotal lobes tapering to narrowly rounded apex, extending anteriorly to or near imaginary, transverse line at posterior limit of eyes and laterally beyond eyes by 0.4-0. 7 width of eye; emar- gination behind each lobe shallow, distance across pronotum at this point about 1.1- 1.2 width of head across eyes. Metastemum flat, hirsute, with weak, thin mesial Carina failing to reach posterior margin. Proctiger contained entirely within genital cup, its apex greatly expanded (Fig. 9). Shallow depression within genital cup on each side of superior ridge bearing tuft of setae. Pygophoral emargination somewhat lyre shaped from caudo ventral view; rim on each side of emargination joining mesoventrally into broad, flat carina (Fig. 10). Deep, ovoid, mesial impression present at base of inferior ridge. Apices of parameres visible from caudoventral view. Profile of pygophore slightly sinuous (Fig. 1 1). Gen- ital plates as in Figure 15; basal plates convex, posterolateral border faintly impressed at most; second gonocoxae dull. Spermathecal bulb with three diverticula (Fig. 16). Dorsum usually black, occasional specimen fuscous to light brown, with small, pale macule on disk of each corium and on each humeral angle; pale markings on connexival segments irregularly and variably shaped, inconspicuous. Pronotum and basal disk of scutellum rugosely punctate, most punctures arranged in irregular, transverse rows. Venter fuscous to black with anterolateral margins of propleura and interstices between punctures on abdomen, excepting broad, mesial vitta, yellowish brown. Three basal segments of each antenna black or brown, fourth usually lighter, fifth pale. Rostrum and tarsi yellowish brown. Punctation on abdomen black, moderately strong and irregularly spaced where interstices pale. Measurements (mm). Width of head 1 .95-2. 1 5, length 1 .80-1 .95; interocular width 274 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Figs. 6-17. 6-8. L. varius. 6. Pygophore, dorsal view. 7. Same, caudoventral view. 8. same, lateral view. 9-1 1, 15, 16. L. spurcus. 9. Pygophore, dorsal view. 10. Same, caudoventral view. 11. Same, lateral view. 15. Genital plates, caudoventral view. 16. Spermathecal bulb. 12-14, 17. L. malevolus. 12. Pygophore, dorsal view. 13. Same, caudoventral view. 14. Same, lateral view. 1 7. Spermathecal bulb. Symbols: i, impression; ir, inferior ridge; p, paramere; pr, proctiger; s, setal tuft; sr, superior ridge. 1989 NEW SPECIES OF UNCUS 275 1.05-1.15; distance across ocelli 1.05-1.15, between each ocellus and nearest eye 0.20-0.25. Length of segments 1-5 of antennae 0.55-0.70, 0.65-0.75, 1.05-1.20, 1.25-1.40, 1.65-1.80. Length of segments 1-4 of labium 1.10-1.25, 2.20-2.30, 1.80- 1.95, 1.75-1.90. Pronotum 4. 9-5. 6 wide, 2. 2-2.4 long mesially. Scutellum 3. 0-3. 6 wide at base, 3.6-4. 3 long. Body length 9.1-10.8. Distribution. Peru (San Martin). Holotype. Male, labeled “Perou, Tocache, XL 1987. sur palmier. R. Huguenot” and “Plantation de palmesa 7196.” Deposited in the Museum National d’Histoire Naturelle, Paris. Paratypes. 1 male and 2 females with same labeling as holotype; 1 male and 2 females labeled “Perou, Sector Cahuto Tocache, palmier a huile. 8 II 1986. Esmilda Arevalo”; 1 male labeled “C. Bolivar. Saposoa. Peru. Piscoyem. 6/IX/40”; 12 males and 9 females labeled “Perou-Uchiza, Plantat. Palmas del Espino, 20.8.1987. G. Couturier & F. Kahn Coll.” and “sur Astrocaryum sp. aff. A. murumuru Mart. (Palmae) ref. herbier F. Kahn 1933”; 9 males and 7 females labeled “Peru-San Martin, Endepalma-Uchiza, april 1988, Julio Llosa coll.” and “host plant Elaeis guineensis (Palmae). Comments. This species also belongs among the “little eyed” group of species in which the width of each eye is less than one-half of the interocular width. In my key to Linens species (Rolston, 1983), it most nearly fits the alternatives leading to varius Rolston. These two species differ in several particulars, especially the form of the proctiger and punctation of the abdominal venter. In varius the proctiger protrudes from the genital cup and the punctation of the abdominal venter is inconspicuous (Figs. 6-8). Lincus malevolus, new species Figs. 12-14, 17 Description. Similar to L. spurcus in coloration, punctation and somatic mor- phology, differing markedly in male genitalia. On average slightly larger than L. spurcus, eyes often a little larger, 0.41-0.50 of interocular width. Metastemum slightly tectiform, hirsute. Proctiger contained entirely within genital cup, apex moderately expanded (Fig. 12). Pygophoral emargination V-shaped from caudoventral view (Fig. 13); rim on each side of emargination joining mesoventrally into broad, flat carina. Deep, mesial impression present at base of inferior ridge. Apices of parameres visible from cau- doventral view. Profile of pygophore moderately sinuous from lateral view (Fig. 14). Genital plates as in L. spurcus. Spermathecal bulb with two diverticula (Fig. 1 7). Measurements (mm). Width of head 2.00-2.20, length 1 .80-2. 10; interocular width 1.05-1.15; distance across ocelli 1.05-1.15, between ocelli 0.75-0.85, from each ocellus to nearest eye 0.20-0.25. Length of segments 1-5 of antennae 0.60-0.70, 0.55-0.80, 1.00-1.20, 1.15-1.40, 1.50-1.67. Length of segments 1-4 of labium 1.15- 1.30, 2.10-2.35, 1.70-1.90, 1.70-1.90. Pronotum 4.8-5. 7 wide, 2.1-2.5 wide at in- cisions behind lobes, 2. 1-2.6 long mesially. Scutellum 3. 1-3.6 wide basally, 3. 7-4. 4 long. Body length 9.6-1 1.3. Distribution. Peru (Loreto). Holotype. Male, labeled “Perou— 4°55S 73°40W, Jenaro Herrara, 1.9.1987, G. 276 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Couturier & F. Kahn Coll.,” “sur Astrocaryum macrocalyx (Palmae)” and “El Capite (quebrada).” Deposited in the Museum National d’Histoire Naturelle, Paris. Paratypes. 1 2 males and 1 2 females with same labeling as holotype; 7 males and 10 females with same labeling as holotype except date 31.8.1987; 16 females with same labeling as holotype except lacking last label and dated 27.8.1987; 1 male and 1 4 females with same labeling as holotype except lacking last label and dated 29.8. 1 987; 3 females with same labeling as holotype except lacking last label and dated 4.9.1987; 2 males and 2 females with same labeling as holotype except lacking last label and dated 30.8.1987; 1 male and 6 females labeled “Peru-Loreto, Jenaro Herrera, June 1988, Julio Llosa col” and “host plant Elaeis oleifera (Palmae)”; 3 males and 16 females labeled “Peru-Loreto, Maniti, June 1988, Julio Lloso col.” and “host plant Astrocaryum sp.” Comments. This species does not fall clearly into any of the species groups of convenience. Those specimens with an eye width less than one half of the interocular width most nearly fit the alternatives in my key (Rolston, 1 983) that lead to L. varius. The remaining male specimens key to the couplet separating L. substyliger and L. subuliger but fit neither alternative. ACKNOWLEDGMENTS The specimens in this study were provided by G. Couturier of ORSTOM and J. M. Maldes of CIRAD. I thank J. B. Chapin and D. A. Rider for their critical review of the manuscript. The paper was approved for publication by the Director of the Louisiana Agricultural Exper- iment Station as manuscript number 88-17-2675. LITERATURE CITED Dolling, W. R. 1984. Pentatomid bugs (Hemiptera) that transmit a flagellate disease of cul- tivated palms in South America. Bull. Entomol. Res. 74:473-476. Rolston, L. H. 1983. A revision of the genus Linens StM (Hemiptera: Pentatomidae: Disco- cephalinae: Ochlerini). J. New York Entomol. Soc. 9 1(1): 1-47. Received November 15, 1988; accepted February 27, 1989. J. New YorkEntomol. Soc. 97(3):277-304, 1989 RECONSTITUTION OF COELOMETOPINI, TENEBRIONINI AND RELATED TRIBES OF AMERICA NORTH OF COLOMBIA (COLEOPTERA: TENEBRIONIDAE) John T. Doyen Department of Entomology and Parasitology, University of California, Berkeley, California 94720 Abstract.— tenebrionid subfamilies Tenebrioninae and Coelometopinae are diagnosed. The tenebrionine tribes Tenebrionini and Alphitobiini are defined, and most genera previously included in Tenebrionini are transferred to Coelometopini. Centronopini and Acropteronini are proposed as new tribes of Tenebrioninae. The coelometopine tribes Coelometopini, Stron- gyliini and Talanini are defined. Cnodalonini, Misolampini and Nodotelini have been based on superficial characters which primarily reflect loss of flying ability. Each of these groups consists of paraphyletic assemblages derived several times independently from Coelometopini, and they are placed as junior synonyms of that tribe. Keys are provided to the genera of these beetles for North and Central America. Since the work of Lacordaire (1859) members of two major lineages (tenebrionine and coelometopine lineages of Doyen and Tschinkel, 1982) of Tenebrionidae have been confounded and included in the tribe Tenebrionini. Lacordaire recognized the difficulty in defining his Tenebrionini and specifically addressed its apparent rela- tionship to Cyphaleini (= Heleini) and Cnodalonini (also to Pycnocerini). Heleini is very similar to Tenebrionini (Doyen et al., in press; Matthews and Doyen, in press), constituting part of the Tenebrionine lineage of Doyen and Tschinkel (1982), whereas Cnodalonini forms part of the Coelometopine lineage. These two lineages consistently differ in the configuration of the internal female reproductive tract, ovipositor, de- fensive glands and reservoirs, and other features (Tschinkel and Doyen, 1980). Other character differences are less constant (e.g., tarsal vestiture; aedeagal orientation; maxillary structure). The primary features used by Lacordaire for separating these groups (armature of maxillary lacinia; shape of mesostemum, etc.) belong to the second group of characters, and do not vary in concordance with tribal limits based on the female reproductive tract and other characters mentioned above. Lacordaire (1859:366) remarked upon the variability of these tribes, both in morphological and biological characteristics. Surprisingly he commented only briefly (p. 359) on their relationship to his Coelometopides, whose chief unifying feature is loss or great reduction of the wings. Subsequent to Lacordaire’s work, no formal definitions have been provided for any of these tribes. Genera have been assigned on the basis of external characters alone, resulting in some preposterous classifications. For example, as pointed out by Spilman (1962b) Zophobas, which is winged, is always included in Tenebrionini, while Rhinandrus, which is wingless, is placed in Coelometopini. All important characters indicate that they are sister genera within Tenebrionini. Similarly, Coe- locnemis differs from Iphthiminus primarily by loss of wings (Doyen, 1973), but this very feature places the two in different tribes. 278 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Based on characters of the internal female reproductive tract, ovipositor, defensive reservoirs and various external features, most genera now included in Tenebrionini are here transferred to Coelometopini, which is expanded to include Misolampini and Cnodalonini. In this sense Coelometopini is one of the larger and more diverse tribes of Tenebrionidae, consisting mostly of tropical and subtropical forms which are associated with decaying wood. Larvae of Strongyliini, the other major tribe of Coelometopinae, also inhabit rotten wood. The coelometopine lineage is formally defined below as the subfamily Coelometopinae. In contrast, Tenebrionini is reduced to a small, relatively uniform group, whose larvae, with the exception of Bius, are scavengers on animal or non-ligneous plant remains. Alphitobius (formerly in Triboliini) and Metaclisa marginalis (formerly in Cnodalonini are here placed in Alphitobiini, which differs from Tenebrionini in several external features, notably antennal sensory structures. The position of these taxa has long been uncertain, as indicated by Reitter’s original (1917) proposal of Alphitobiini and his (1922) placement of Metaclisa in Scaphidemini. Acropteronini is proposed for Acropteron, presently in Cnodalonini. Acropteron shows most of the diagnostic features of Tenebrioninae, but does not conform to any of the existing tribes. Its most notable characters are the presence of 1 0 elytral striae, internally open procoxal cavities (both primitive) and a highly derived ovipositor. Centronopus and Scotobaenus are superficially similar to certain Coelometopini (Lacordaire believed they resembled Menephilus), but lack the distinctive coelo- metopine female reproductive tract and ovipositor. They differ from other tribes of Tenebrioninae in defensive reservoir structure and other characters. Hence the tribe Centronopini is proposed. All of these tribes are placed in the subfamily Tenebrion- inae, which is formally defined. Detailed discussions of character interpretation and apparent relationships among the taxa addressed briefly above are given where appropriate below. Morphological terminology follows usage in Tschinkel and Doyen (1980) and Doyen and Tschinkel (1982). The geographic scope is America north of Colombia with occasional reference made to other areas. Table 1 provides a conspectus of the included taxa and taxonomic changes. Several genera which appear in catalogues are excluded from Table 1 because they have previously been moved into other tribes of Tenebrionidae or into other families. These are: 1) Adelonia Laporte, transferred into Belopini (Doyen and Tschinkel, 1982; Doyen, 1988); Merotemnus Horn and Rhacius Champion are junior synonyms of Adelonia (Spilman, 1961). 2) Alaephus Horn, transferred into Vacronini (Doyen and Lawrence, 1979). 3) Boros Herbst, separated as Boridae (Crowson, 1955). Eu- psophulus Cockerell {=Eupsophus Horn), transferred to Vacronini (Doyen and Law- rence, 1979). 4) Biomorphus Motschulsky has been placed in synonymy under Helops Fabricius (Aalbu et al., in press). In addition Maracia haagi Gebien is listed from Central America by Papp (1961). Gebien (1919:35) states, however, that the type locality is unknown, and Maracia has not been subsequently mentioned in the primary literature. It is not considered here. Reminius Casey was placed in synonymy under Strongylium by Spilman ( 1 959). Pteroglymmius Gebien is a synonym of Isaminas Champion (Doyen, 1987). Par- oeatus is listed by Papp (1961) as possibly from Central America. It is included in Table 1 and the keys even though I have seen specimens only from South America. 1989 NORTH AMERICAN TENEBRIONINI 279 Hesiobates, described from Dominican amber by Kaszab (1984) appears to belong in Coelometopini, and may be closely related to Hesiodus, Ilus and Choastes. It is not considered further in this work. Tenebrioninae Description. Adult. — Small to large (about 3 mm to 30 mm). Antennae filiform- serrate, incrassate or rarely capitate bearing only simple, setiform sensilla or occa- sionally with compound, stellate sensoria on apical five or six segments. Labrum transverse with basal membrane exposed or concealed. Mandible with mola striate or not. Maxilla with galea finely setose or with uncus of one or two teeth. Tentorium with bridge posterior, flat or arched. Procoxal cavities closed externally, open or closed internally. Mesocoxal cavities closed laterally by mesepimeron or sternum. Elytra with scutellary striole and 9 complete striae or estriate. Apical membrane comprising 25% of less of wing length; recurrent cell large to small or obsolete; subcubital fleck present or absent. Metendostemite usually with long stalk, long arms with subterminal muscle attachment flange; tendons inserted near midpoint or toward apex. Tarsi usually with ventral surface coarsely setose or spinose, occasionally with pads of pilose setae. Ovipositor usually with coxites clearly 4-lobed, occasionally with ovipositor shaft shortened and lobing reduced; lobes usually subequal in length; fourth lobe rarely free and digitate; paraprocts parallel to axis of ovipositor at rest. Internal female reproductive tract consisting of vagina, bursa copulatrix, long slender spermathecal accessory gland and spermatheca. Aedeagus with tegmen dorsal or rotated about 45° to 90° at rest, rarely inverted; median lobe freely extrusible or adnate to tegmen; sometimes with accessory lobes. Defensive reservoirs variable, often distinctive at tribal level (see Tschinkel and Doyen, 1980). Larva. — Variable in all important characters (see discussion below). Tenebrioninae corresponds to the combined tenebrionine, toxicine and opatrine lineages recognized by Doyen and Tschinkel (1982). This group includes those tribes in which the spermatheca is derived from the bursa copulatrix, and appears as a separate structure from the spermathecal accessory gland. (Alleculinae, which usually have this configuration, are recognized here as a separate subfamily.) Discussion. The ovipositors of this group are mostly similar to that of Tenebrio, with the paraproct subequal to the coxite and the coxite composed of four similar lobes. However, in Toxicini and Boletophagini, the fourth ovipositor lobe is free and digitate, as in Lagriini; in some Toxicini there are several bursa-derived spermathecae, again resembling Lagriinae. Among Alleculinae, which mostly conform to the te- nebrionine pattern, Lobopoda has both ovipositor and internal female reproductive structures similar to Lagriinae. These character distributions suggest that the tenebri- onine configurations may have been evolved independently more than a single time. There are also enormous differences in life style and habits of various tenebrionine tribes, however, which could indicate that some of these structural patterns (especially in ovipositor morphology) have arisen as secondary specializations from the basic tenebrionine configuration described above. For example, the ovipositors of Bole- tophagini and Toxicini, which both inhabit fruiting bodies of polypore fungi, are similar in structure to the ovipositors of many Diaperini (but not Diaperis), which also live on fungi. In contrast, the ovipositors of Eleodini and Opatrini (also Tenebri- 280 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Table 1. Tribal placements of North and Central American genera treated in text. Former placement indicated at right. Alphitobiini Alphitobius Stephens Triboliini Metaclisa J. DuVal Cnodalonini Tenebrionini Bins Motschulsky Tenebrionini Idiobates Casey Tenebrionini Neatus LeConte Tenebrionini Rhinandrus LeConte Coelometopini Tenebrio Linnaeus Tenebrionini Zophobas Blanchard Tenebrionini Centronopini Centronopus Sober Coelometopini, Tenebrionini Scotabaenus LeConte Coelometopini Tauwceras Hope Tenebrionini Acropteronini Acroptewn Perty Cnodalonini Coelometopini Alobates Motschulsky Coelometopini Apsida Lacordaire Diaperini Blapida Perty Cnodalonini Bothynocephalus Doyen Coelometopini Camaria Serville Cnodalonini Choastes Champion Tenebrionini Cibdelis Mannerheim Coelometopini Cnephalura Doyen Coelometopini Cnodalon Latreille Cnodalonini Coelocnemis Mannerheim Coelometopini Cyrtosoma Perty Cnodalonini Dinomus Breme' Misolampini Elomosda Bates Cnodalonini Epicalla Champion Cnodalonini Glyptotus LeConte Tenebrionini Gonospa Champion Diaperini Haplandrus LeConte Tenebrionini Hegemona Laporte Misolampini, Helopini Hesiodus Champion Tenebrionini Hicetaon Champion Tenebrionini Ilus Champion Tenebrionini Iphthiminus Spilman- Tenebrionini Isaminas Champion Misolampini Isicerdes Champion Tenebrionini Merinus LeConte Tenebrionini Mitys Champion Misolampini Moeon Champion Cnodalonini Mophon Champion Cnodalonini Mylaris Motschulsky^ Tenebrionini Nesocyrtosoma Marcuzzi"* Cnodalonini Nuptis Motschulsky Tenebrionini Oeatus Champion Tenebrionini Oenopion Champion Coelometopini Othryoneus Champion Cnodalonini 1989 NORTH AMERICAN TENEBRIONINI 281 Table 1. Continued. Oxidates Champion Misolampini Paroeatus Gebien Tenebrionini Polopinus Casey Coelometopini Polypleurus Eschscholtz Coelometopini Saziches Champion Misolampini Spinepicalla Pic' Cnodalonini Stenoboea Champion Tenebrionini Sycophantomorphus Pic' Cnodalonini Upis Fabriceus Tenebrionini Xenius Champion Cnodalonini Xylopinus LeConte Tenebrionini Strongyliini Cuphotes Champion Strongyliini Mentes Champion Helopini Otocerus Maklin Strongyliini Poecilesthus Blanchard Strongyliini Pseudotocerus Champion Strongyliini Strongylium Kirby Strongyliini Talanini Talanus Maklin Talanini ' Not examined or included in key. 2 Iphthiminus Spilman is replacement name for Iphthinus (=Iphthimus) of authors. See Spil- man (1973). ^ My laris Pallas = Nyct abates Guerin. See Spilman (1973). Nesocyrtosoma Marcuzzi 1976 (NEW STATUS), originally proposed as a subgenus of Cyr- tosoma, differs from Cyrtosoma s.s. in having the labroclypeal membrane concealed and in having a fossa in each elytron base in which the pronotal base rests. These characters are shared with Cnodalon which, like Nesocyrtosoma, is endemic to the Greater Antilles. oninae) are at least superficially similar to those of Phaleriini (subfamily Diaperinae). All of these beetles oviposit in loose, often sandy soil. Larvae of the tenebrionine tribes are as variable as adults. Body forms similar to that of Tenebrio, with a relatively strongly sclerotized body, slightly enlarged pro- thoracic legs with distinct combs of setae, much enlarged ninth abdominal tergite and annular spiracles are almost ubiquitous in soil-dwelling larvae. However all of these features vary greatly in Tenebrioninae which occupy other situations. For example in Boletophagini the body is grub-like; in Toxicini and Heleini the spiracles are surrounded by peripheral air tubes; in Ulomini and Lepispilus (Heleini) the ninth tergite is paraboloid, entirely covering the abdominal apex, with the anus concealed inside it (a similar shape occurs in Alleculinae); and in many Triboliini the setation of the legs is irregular and the forelegs are not enlarged. It has been no more obvious how to subdivide Tenebrioninae on the basis of larval than of adult features. In addition, larvae of many tribes are inadequately described or unknown. Splitting of Tenebrioninae into several subfamilies may eventually prove desirable (Watt [1974] recognized Toxicinae as a subfamily, for example), but is neither feasible nor practical in a work dealing with only the North and Central American fauna. Therefore, a generally conservative approach is taken here in accepting all the cur- rently recognized tribes without combining them into larger infrasubfamilial units. 282 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Only Tenebrionini, Centronopini, Alphitobiini and Acropteronini, all of which have been confounded with Coelometopini, are formally defined below. Tribe Tenebrionini, New Sense Tenebronides vrais Lacordaire, 1859 (in part) Tenebrionini, Reitter, 1920 Tenebrionini, various authors (in part) Description. Adult. — Small to large (about 6 mm to 30 mm). Eyes moderate in size, weakly emarginated to entirely divided by epistomal canthus; antennae serrate- filiform, weakly incrassate, bearing only simple, setiform sensilla; labrum about twice as broad as long, with basal membrane concealed or exposed; epipharynx asym- metrical (as in Fig. 1 1; Doyen and Tschinkel, 1982); mandibles with incisors bifid, molas striate or nonstriate; lacinia with or without (Rhinandrus) uncus, palp sub- cylindrical to broadly triangular; tentorium with bridge posterior, slender, not arched. Procoxal cavities closed externally, opep or closed internally; mesocoxal cavities closed laterally by mesepimeron; mesostemal apophysis developed as slender dorsal arm with or without anteroventral muscle attachment flange; elytra 9-striate with scutellary striole or estriate. Apical membrane comprising about one-fifth of wing length; recurrent cell large; subcubital fleck present (Bins) or absent. Metendostemite with stalk long to short {Rhinandrus), tendons inserted near midpoint of arms or close to apex; arms with subterminal muscle attachment flange (much enlarged in Rhinandrus). Tarsi clothed ventrally with spinose or pilose {Zophobas, Rhinandrus) setae. Ovipositor flexible with coxites and paraprocts subequal in length; coxites divided into four subequal lobes; fourth lobe not digitate; internal female repoductive tract with bursa reduced or absent, spermatheca tubular, coiled, long and slender to short, T-shaped and thick. Aedeagus with mediam lobe freely extrusible or adnate to tegumen {Zophobas, Rhinandrus), without accessory lobes. Defensive reservoir short, conical, eversible and with or without {Bius) common volume; reservoir walls without annulation, sometimes rigidified by cuticular strip from stemite 7 (see Ac- ropteronini, discussion); secretory ducts distributed over dorsal surface of reservoir, as basal line at neck of reservoir, or as few duct emptying at neck {Bius). Larva.— Cylindrical, moderately sclerotized and pigmented; ocelli present as weak pigment spots without lenses. Antenna with three segments; second segment subequal to basal, about 6-8 times as long as digitate third segment; sensorium single, arcuate around base of third segment, or multiple ellipses {Zophobas). Labrum two to two and one-half times as broad as long with anterior margin straight or weakly concave; epipharynx with pair of masticatory processes (right process usually larger), two central blunt spines and 6 annular sensilla (3 sensilla in Bius). Mandibles asymmet- rical, left with more promient retinaculum and mola; molas variably sculptured with coarse blunt teeth or ridges. Maxilla with mala entire, without uncus; spinose on mediodorsal surface. Hypopharyngeal sclerome with anterior comers prominent, middle straight or weakly bidentate. Prothorax with prestemum usually well defined; terga with anterior transverse carina well defined, especially on meso- and metatho- rax. Legs similar in size and configuration or anterior pair slightly larger, more coarsely spinose {Zophobas)', at least anterior pair bearing regular combs of spines on inner surface of femur and tibia. Ninth abdominal tergite expanded posteriorly, about two 1989 NORTH AMERICAN TENEBRIONINI 283 to three times as long as sternite, sometimes bearing short urogomphi; anus subter- minal, below tergite; pygopods moderately large, setose, with posterior surface weakly sclerotized. Spiracles simple ellipses. Discussion. Tenebrionini as conceived here is greatly reduced from present cata- logue listings, with most of the genera transferred to Coelometopini (Table 1). Major differences in ovipositor, internal female reproductive tract, and defensive gland and reservoir structure, as well as a number of other characters (type of antennal sensoria, structure of ninth segment of larvae) separate these groups. These characters are discussed in more detail below under Coelometopini, and most have been previously analyzed several times (Tschinkel and Doyen, 1980; Doyen and Tschinkel, 1982; Doyen et al., in press). The closest relatives of Tenebrionini are Triboliini and Alphitobiini, whose salient characters have been outlined previously (Doyen, 1985; Doyen et al., in press). As suggested in those publications, it may eventually prove desirable to recognize all three at the subtribal level. Alphitobiini is formally defined below. Tribe Alphitobiini Alphitobiini Reitter, 1917 Description. Adult. — Small (about 4 mm to 7 mm). Eyes emarginate but never divided by epistomal canthus; antennae incrassate, bearing stellate, compound sen- soria on apical six segments; labrum about two and one-half to three times broader than long, with basal membrane concealed; epipharynx asymmetrical; mandibles with incisors bifid, molas striate or nonstriate; lacinia with uncus; palp narrowly triangular; tentorium with bridge posterior, slender, not arched. Procoxal cavities closed externally and internally; mesocoxal cavities closed by epimeron or sterna; mesosternal apophyses with long, slender dorsal arm without anteroventral muscle flange; elytra 9-striate with scutellary striole. Apical membrane about one-fifth to one-third wing length; recurrent cell large; subcubital fleck present {Metaclisa) or absent. Metendosternite with long stalk, tendons inserted near apex of arms; arms with subterminal muscle attachment flange. Tarsi clothed ventrally with spinose setae. Ovipositor as in Tenebrionini; internal female reproductive tract with spermatheca long, slender and coiled {Metaclisa) or capsular, reniform. Aedeagus with median lobe adnate to tegmen, without accessory lobes. Defensive reservoirs short, conical, and with common volume or long, saccate, without common volume {Metaclisa)\ secretory ducts distributed over apical half of reservoir {Metaclisa) or as basal line on neck. Larva (based on Alphitobius). — Similar in nearly all features to larvae of Tenebrion- ini, differing as follows: sensorium on second antennal segment arcuate around base of third segment; mandibles with molas of subequal prominence; hypopharyngeal sclerome with anterior margin straight; prothorax without distinct prestemum; ab- dominal tergite nine terminating in single, short urogomphus. Discussion. Alphitobius adults and Old World Diaclina are similar in all diagnostic features. Metaclisa marginalis Horn is similar in most features except the defensive reservoirs, which are greatly enlarged and saccate. The secretory tissue drains through many ductules distributed over the dorsal surface of the reservoirs, as in most Te- nebrionini. Metaclisa is placed in Alphitobiini rather than Tenebrionini because the 284 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) antennae bear compound sensoria and because the median lobe of the aedeagus is adnate to the tegmen. I have not dissected other species of Metaclisa (all Old World), which may prove very different from marginalis, necessitating a new generic name for the latter. KEY TO GENERA OF TENEBRIONINI AND ALPHITOBIINI 1 . Antenna with compound, stellate sensoria on apical five segments; pronotal margin evenly curved from apex to base (Alphitobiini) 2 - Antenna with simple, setiform sensilla; pronotal margin recurved near base (Tene- brionini) 3 2(1). Epipleuron gradually narrowing to elytral apex; prostemal process prominent, sub- horizontal behind coxae; mesosternum acutely concave Alphitobius - Epipleuron abruptly narrowing at anterior margin of visible stemite five, not reaching elytral apex; prostemal process declivous, flattened behind coxae; mesosternum ob- tusely concave Metaclisa 3(1). Tarsi with ventral pads of dense, pilose, yellowish setae; labroclypeal membrane usually exposed, at least medially 4 Tarsi with stiff, sparse, usually dark colored setae ventrally; labroclypeal membrane concealed 5 4(3). Metastemum about twice length of mesocoxa Zophobas - Metastemum about as long as mesocoxa Rhinandrus 5(3). Eye not divided by epistomal canthus 6 - Eye divided by epistomal canthus into dorsal and ventral lobes Idiobates 6(5). Elytra with distinct striae 7 - Elytra with confused punctation, without striae Bins 7(6). Abdominal stemite five with very fine marginal groove Neat us - Abdominal stemite five without marginal groove Tenebrio Centronopini, new tribe Description. Adults. — Moderate to large (about 10 mm to 20 mm), elongate, flat- tened beetles. Eyes moderate in size, strongly emarginated by epistomal canthus; antennae incrassate; apical five or six segments bearing large, stellate sensoria, especially on inner apical margins; labrum about twice as broad as long, with basal membrane concealed; epipharynx symmetrical or nearly so; mandibles with incisors bifid, molas nonstriate; lacinia with uncus, palp weakly triangular; tentorium with bridge posterior, slender, not arched. Procoxal cavities closed externally and inter- nally; elytra 9-striate with short, sometimes poorly defined scutellary striole; meso- coxal cavities closed laterally by mesepimeron; mesostemal apophysis developed as large, anteriorly oriented muscle disk, without dorsal arm. Apical membrane com- prising about one-fifth of wing length; recurrent cell large; subcubital fleck absent. Metendosternite with long stalk, tendons inserted slightly beyond midpoint of arms; arms with subterminal muscle attachment flange. Tarsi clothed ventrally with pads of dense, fine, yellowish pubescence. Ovipositor (Fig. 1) strongly sclerotized, slightly compressed in lateral plane with lobing of coxites sometimes obscured and gonostyli papilliform; paraprocts about twice as long as coxites; internal female reproductive tract (Figs. 2, 3) with large bursa copulatrix, long slender accessory gland; spermatheca present or absent. Aedeagus rotated about 45°-60°; median lobe adnate to tegmen or nearly so. Defensive reservoirs (Fig. 4) elongate, without common 1989 NORTH AMERICAN TENEBRIONINI 285 Figs. 1, 2. Female genitalic characters of Centronopini. Scotobaenus parallelus LeConte. 1. Ovipositor, lateral. 2. Internal female reproductive tract. volume; mesal margins slightly expanded at about middle, reservoir walls without annulation; secretions drained by single large collecting duct opening at base of each reservoir. Larva. — Cylindrical, moderately sclerotized and pigmented; ocelli present as weak- ly developed pigment spots without lenses. Antenna with three segments; second segment about twice length of basal, bearing sinuate sensorium around base of digitate third segment. Labrum about twice as broad as long with evenly arcuate anterior margin; epipharynx with pair of large masticatory processes (right process larger), two central blunt spines and six annular sensilla. Mandibles asymmetrical, left with more prominent retinaculum and mola; incisors bilobed; molas with several coarse, transverse ridges. Maxilla with mala weakly indented at apex, without uncus; spinose on mediodorsal surface. Hypophryngeal sclerome with middle portion greatly elon- gate and usually apically bilobed, projecting far anterad of short ligula. Prothorax with distinct prestemum; thoracic terga lacking anterior transverse carina. Legs sim- ilar in size; bearing combs of spines on inner surfaces of femora and tibiae. Ninth abdominal tergite very large, produced as pair of stout, tapering, sharply pointed urogomphi; ninth sternite small, transverse, bearing short pygopods without spines; anus ventral, usually concealed with pygopods beneath ninth tergite. Spiracles simple annuli or ellipses. Discussion. Centronopini as defined here comprises Centronopus Solier (including Pyres Champion), Scotobaenus LeConte and Tauroceras Hope. Lacordaire (1859) included the first two genera in his Coelometopides, whereas LeConte 1862 and LeConte and Horn (1883), not recognizing Coelometopini, placed them in Tenebrion- ini. Recent catalogs (Gebien, 1942-1944; Backwelder, 1945; Papp, 1961) place Cen- tronopus and Scotobaenus in Coelometopini and Pyres in Tenebrionini. Spilman (1962a, b) clarified the generic nomenclature, and placed Pyres as a synonym of Centronopus. Immatures of Centronopus have been treated by St. George (1924) and Spilman (1979). Spilman (1963) described larvae which very likely represent Tauroceras. In their 286 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Figs. 3, 4. Internal characters of Centronopini. 3. Internal female reproductive tract of Centronopus suppressus (Say). 4. Defensive reservoirs (dorsal) of Scotobaenus pamllelus. morphological features these larvae were strikingly similar to those of Scotobaenus and Centronopus. The most important points of similarity include (1) the strongly sclerotized trunk; (2) the greatly developed middle lobe of the hypopharyngeal scler- ome; (3) the distinct combs of setae on the legs; (4) the configuration of the ninth abdominal tergite, with a pair of large urogomphi and several smaller, thom-like processes. The second and fourth may be considered as synapomorphies of Cen- tronopini, although in Centronopus there is a single pair of thorn-like processes on the ninth abdominal tergite and in Scotobaenus they are absent. Tschinkel and Doyen (1980; Appendix IV) noted that the internal female repro- ductive tract arrangement of Tauroceras is of the Tenebrionine type, with separate spermatheca and spermathecal accessory gland. This arrangement also occurs in Centronopus (see below). The ovipositor of Tauroceras is strongly sclerotized and very similar to that of Scotobaenus in the proportions of the coxite lobes. The defensive reservoirs of Tauroceras bear annular folds, and cuticular thickenings, whereas those of Centronopus and Scotobaenus are only irregularly annulate and lack the thickened rings. However, annulate defensive reservoirs have developed inde- pendently in several lineages of Tenebrionidae (Tschinkel and Doyen, 1980:332). Although the male secondary sexual characters produce a superficial dissimilarity, the balance of adult and larval features strongly supports the inclusion of Tauroceras in Centronopini. Past disagreements over taxonomic position will perhaps not be laid to rest here because Centronopini have features of both Tenebrioninae and Coelometopinae. Like Coelometopinae the adults have compound sensoria on the apical antennal segments and tarsal pads of fine, dense setae. However, neither of these features is diagnostic of Coelometopinae. Compound antennal sensoria occur also in Diaperinae (Doyen, 1984) and some Tenebrioninae (Amarygmini: Medvedev, 1977; Triboliini: Doyen, 1985; alphitobiini: Doyen et al., in press; also, see above discussion). Tarsal pads of fine, dense setae occur in many Tenebrionidae living on surfaces of logs or trees, including Heleini, Toxicinae, Nyctoporini and Tenebrionini (Rhinandrus). 1989 NORTH AMERICAN TENEBRIONINI 287 Moreover, features of the defensive glands, internal female reproductive tract and ovipositor preclude membership in Coelometopinae. Though enlarged, as in Coe- lometopini, and with glands drained by a single collecting duct, the reservoirs of Centronopini (Fig. 4) lack common volume and most lack annulation. The ovipos- itors of Centronopini (Fig. 1) are slightly compressed in the lateral plane, and quite strongly sclerotized, with reduced gonostyli. The basal coxite lobe is not elon- gate, as in Coelometopini, nor is the paraproct rotated. The primitive 4-lobed division of the coxite is clearly visible. In Hegemona and Saziches (placed here in Coelo- metopini), which have more highly modified, blade-like ovipositors, coxite lobation has been essentially eliminated (Doyen, 1987, fig. 3). The internal female reproductive tract is variable within Centronopini. In Centronopus (Fig. 2) and Tauroceras the large bursa copulatrix bears a long, slender accessory gland as well as a spermatheca, both attached to a slender, non-glandular duct leading to the vagina near the entrance of the common oviduct. Although differing in detail, this configuration is similar to that of various Tenebrioninae. In Scotobaenus (Fig. 3) the bursa copulatrix bears only an accessory gland, somewhat shorter and thicker than in Centronopus. This arrangement is like that of Hegemona and Saziches. The coelometopine tract is similar, except that the apex of the accessory gland forms an enlarged, nonglandular spermatheca. It seems clear that the configuration in Centropus and Tauroceras is plesiomorphic, that in Scotobaenus derived by loss of the spermatheca. In other adult and larval features Centronopini are so similar that it seems almost certain that they represent a monophyletic clade. Larval characteristics of Centronopini mostly suggest affinities with Tenebrioninae, rather than Coelometopinae. A general tenebrionine feature is the leathery, pigmented cuticle. Most Coelometopinae have more delicate, transparent cuticle. The epiphar- ynx is very similar to that of Tenebrio, with a single pair of large, subquadrate masticatory processes. In Coelometopinae, etc., the masticatory processes are usually elongate or dentate, but there is much variation. Centronopini and Tenebrionini also share the presence of a transverse prestemum (cervicostemum of Watt, 1970), which is lacking in Coelometopinae I have examined. The legs of larval Centronopini bear regular, longitudinal combs of spines on the femur and tibia, as in most Tenebrionini. In Coelometopinae the leg spines do not form regular combs. Finally, the ninth tergite of Centronopini is much more expanded ventrally than in Coelometopini. The ninth sternite is reduced to a narrow, transverse sclerite, and the ventral anus and pygopods may be concealed within the enlarged tergite. This configuration is very similar to that of Bassianus and most Heleini (Matthews and Doyen, in press). In Tenebrionini the ninth sternite is slightly larger, and in Coelometopinae the expansion of the ninth tergite is primarily dorsad and posteriad, so that the anus opens posteriorly and is never concealed within the tergite. The shape of the hypopharyngeal sclerome, with its very long anterior process, is similar to that of Ulomini and Alleculini, but both of these differ in numerous other features from Centronopini. The character state distributions discussed above show that Centronopus, Scoto- baenus and Tauroceras cannot be retained in Coelometopinae. While not entirely diagnostic, the female reproductive tract and especially the larval characters indicate placement in Tenebrioninae, close to Tenebrionini and Heleini. Apomorphic features distinguishing Centronopini include the sclerotized ovipositor and the anterior pro- cess of the hypopharyngeal sclerome. 288 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) In external characters Centronopini are very similar to typical Coelometopini (compound antennal sensoria; tarsal pads of fine, yellowish setae). For this reason they are included in the keys to that group (see below). Acropteronini, new tribe Description. Adult. — Moderate in size (about 6 mm to 20 mm), slender, elongate, subcylindrical beetles. Head deflexed, porrect; eyes large, bulging, anterior border weakly emarginate; antennae slender, basally filiform, becoming weakly serrate api- cally, bearing simple, hair-like sensilla; labrum about 4 times as broad as long with basal membrane exposed; epipharynx symmetrical or nearly so; mandible with incisor bluntly spatulate, undivided; mola finely, transversely striate; lacinia finely setose, without uncus; palp with apical segment triangular; tentorium with bridge posterior, stout, weakly arched above posterior arms. Procoxal cavities broadly closed exter- nally, open internally. Mesocoxal cavities closed laterally by mesepimeron; elytra with long scutellary striole and 10 complete striae; epipleuron gradually narrowing to apex. Metendostemite with long stalk, stout arms, with tendons located half distance to apex. Apical membrane comprising about one-sixth of wing length; re- current cell large, subcubital fleck present. Ovipositor with coxites strongly sclero- tized, compressed into vertical blade with papillate gonostyli (Fig. 5); paraprocts strongly sclerotized with base transversely expanded; not rotated at rest. Internal female reproductive tract with large bursa copulatrix, long, slender accessory gland and much shorter, thicker spermatheca (Fig. 6). Aedeagus rotated about 90° at rest; median lobe freely extrusible with apex slightly enlarged. Defensive reservoirs small, saccate, with little common volume and without annulation; lateral reservoir walls rigidified by cuticular strip from stemite 7 (Fig. 7); glandular tissue distributed over most of dorsal surface of reservoirs, emptying through diffuse tubules. Larva and biology. — Unknown. Discussion. Acropteronini includes only Acroptcron Perty, which has been included in the tribe Cnodalonini by all authors subsequent to Lacordaire (1859). Ischyomius Champion, originally placed in Cnodalonini near Acropteron is now included in Pythidae (Lawrence, 1982) or Trictenotomidae (Watt, 1987). As discussed below, Lacordaire’s Cnodalonini is based almost entirely on primitive or highly variable characters and cannot be differentiated from Coelometopini. More pertinent here are the important features by which Acropteron differs from all members of the Coelo- metopinae. 1) Only simple, setiform sensilla are present on the antennae of Acrop- teron. Compound sensilla are present in all Coelometopinae, at least on the apical segments. 2) The defensive reservoirs (Fig. 7) are short, saccate structures with the secretory tissue emptying through many tubules distributed diffusely over the dorsal reservoir wall. In most Coelometopinae the reservoirs are elongate, usually with annular foldings in the walls which allow for volumetric expansion. In all Coelo- metopinae the secretions are delivered through one or a few enlarged collecting tubules which empty near the neck of the reservoirs. In Strongyliini the reservoirs are short and saccate, but none of the other features are similar to Acropteron. 3) The internal female reproductive tract of Acropteron is of the type found in Tenebrioninae, with separate spermatheca and accessory gland (Fig. 6). In all Coelometopinae there is a single diverticulum from the bursa copulatrix, which is usually expanded apically as 1989 NORTH AMERICAN TENEBRIONINI 289 Figs. 5-7. Internal characters of Acropteronini. Acropteron sp. (Ex. Sta. Catarina, Brazil). 5. Ovipositor (lateral). 6. Internal female reproductive tract. 7. Defensive reservoirs (dorsal). the spermatheca. 4) In Coelometopinae the ovipositor is highly specialized, with the paraprocts rotated 1 80° at rest (see discussion under Coelometopini). In Acropteron the ovipositor (Fig. 5) is highly modified as a strongly sclerotized, blade-like organ. The parapfocts do not show any indication of the type of specialization found in Coelometopinae. As noted by Champion (1 887:268) and elaborated by Doyen (1987), the ovipositors of Hegemona and related genera are sclerotized in the form of two vertical blades. However, in Hegemona the coxites are curved strongly dorsad, where- as in Acropteron they are curved ventrad. In several other features Hegemona is similar to Coelometopinae (Doyen, 1987). Talanus also has a blade-like, sclerotized ovipositor (Tschinkel and Doyen, 1980, fig. 41), but is clearly coelometopine in all other important characters (see below). Thus, all the distinctive apomorphic features of coelometopinae are lacking in Acropteron. Two other unusual features displayed by Acropteron are clearly plesiomorphic and of no use in indicating cladistic relationship. 1) The internally open procoxal cavities occur also in other Tenebrioninae (Heleini, Toxicini and some Tenebrionini) and in Zolodininae. 2) Elytra with ten striae occur also in Lagriinae, Pimeliinae (=Tentyrii- nae), Zolodininae, and a partial or complete tenth stria occurs in Toxicini (Doyen and Tschinkel, 1982:137). All of these taxa show other apomorphic features not shared with Acropteron. Many of the characteristics discussed above are plesiomorphic and of limited use in determining cladistic position. None, however, disagree with placement in Te- nebrioninae. Tenebrioninae are defined to a large extent by lack of derived characters, and division into tribes is problematic. One feature of the defensive reservoirs suggests a possible relationship between various tribes of Tenebrioninae and may be primitive in this subfamily. In Acropteron the lateral walls of the defensive reservoirs are stiffened by a strip of cuticle from the seventh abdominal stemite (Fig. 7). A similar strip of cuticle occurs in all genera of Heleini, which have similar short saccate reservoirs. A less well developed strip of cuticle is present in a few Cyphaleini and is also present in Tenebrio. Similar strips of cuticle occur in some Opatrini {Blapstinus, Edylius, Pedinus, Ulus), and in Alleculinae the neck of the reservoir is sometimes noticeably sclerotized. Both of these latter groups also belong to the tenebrionoid lineage of Doyen and Tschinkel (1982) according to internal characters. 290 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Among primitive Tenebrioninae, Acropteron shares a general phenetic similarity with Titaenini (elongate, cylindrical body; deflexed head), but differs in ovipositor, defensive reservoirs and other characters. Lepispilus (Heleini) has a rigid, tube-like, sclerotized ovipositor which is somewhat similar to that of Acropteron, but shares no obvious synapomorphies. In addition, in Lepispilus, the paraproct bases rotate through a partial arc as the ovipositor is exserted or retracted, in a manner analogous to that in Coelometopinae (E. Matthews, pers. comm.). Finally, Acropteron differs from all other Tenebrioninae in its extremely broad labrum (about four times broader than long). For the reasons discussed above it is appropriate to recognize Acropter- onini as a distinct tribe of Tenebrioninae. SUBFAMILY COELOMETOPINAE Description. Adult. — Small to very large (about 5 mm to 45 mm) beetles of diverse shape and color. Antennae filiform, serrate, incrassate or weakly capitate; apical five to eight segments bearing stellate compound sensoria. Labrum transverse with basal membrane exposed or concealed. Mandible with mola finely striate, flat or occa- sionally coarsely ridged. Maxilla with galea finely setose or with uncus of one or two teeth. Tentorium with bridge posterior, flat or weakly arched. Procoxal cavities broad- ly closed both externally and internally. Mesocoxal cavities closed laterally by mes- epimeron; elytra with scutellary striole and 9 complete striae or estriate. Apical membrane usually comprising about 25% of wing length; recurrent cell moderate to large; subcubital fleck rarely present (e.g., Upis, Camaria). Metendostemite with stalk long or short and broad (wingless species), with long arms, usually with tendons inserted near midpoint and subterminal muscle attachment flanges. Tarsi with ventral pads of fine, dense pubescence or with sparser, coarser setae; tibiae frequently with setose inner apical margins. Ovipositor (Tschinkel and Doyen, 1980, figs. 22, 23, 39) with coxites clearly 4-lobed; basal lobe usually elongate (often longer than three apical lobes combined); paraprocts rotated about 145° about articulation with coxite at rest, or, rarely, rotated about 60° to 90° (e.g., Menephilus, not North America). Internal female reproductive tract (Tschinkel and Doyen, 1980, figs. 22, 23, 26) consisting of vagina, enlarged bursa copulatrix, and single appendant duct; duct glandular except at apex, which forms spermatheca, which is usually enlarged, subspherical. Aedeagus with tegmen dorsal at rest, rotated about 60° to 90°, or occasionally inverted (rotated 180°); median lobe usually adnate to tegmen, rarely freely extrusible. Defensive reservoirs saccate, with considerable common volume, and often with regular, an- nular foldings of the walls; defensive tissue draining through one to several enlarged collecting ducts, often with basal ampullae. Larva. — Elongate, cylindrical, usually with weakly sclerotized trunk segments. An- tennae with three segments, second about one to one and one-half times longer than basal, bearing semicircular or occasionally sinuate sensorium around base of digitate third segment. Labrum about as long as wide to twice as long as wide; epipharynx usually with two central spines subtending four to ten annular sensilla and elongate masticatory processes; tormae usually indistinct. Mandibles slightly to moderately asymmetrical, with either left or right mola more prominent; incisor bilobed, usually subtended by retinaculum, giving trilobed appearance. Maxilla with mala entire or weakly (occasionally moderately) indented at apex, sometimes produced medially 1989 NORTH AMERICAN TENEBRIONINI 291 Figs. 8-1 1. Tarsal structure in Coelometopinae. 8. Coelocnemis (Coelometopini). 9. Stwn- gylium (StTongyliini). 10. Talanus (Tsdanini). 11. (Coelometopini). Each illustration shows left mesotarsus from anteroventral aspect. near tip; inner surface spinose. Hypopharyngeal sclerome usually with anterior margin tridentate, occasionally with middle tooth absent or enlarged and produced anterad over ligula. Prothorax without prestemum; terga usually lacking anterior transverse Carina. Legs similar in size; femoral and tibial setae irregularly distributed, not ar- ranged in regular combs. Ninth abdominal tergite almost always produced as prom- inent pair of recurved urogomphi, sometimes with additional ridges, spines or cal- loses. Ninth sternite occupying ventral third of segment with anus subterminal and pygopods small or absent. Spiracles simple annuli or ellipses. Discussion. By far the most important defining characters of Coelometopinae are those of the ovipositor and internal female reproductive tract, in which intermediacy or exceptions are very uncommon. These features are discussed at greater length under Coelometopini, below. With very few exceptions (see Doyen, 1987, for ex- ample), this is one of the most clearly delimited higher taxa of Tenebrionidae, and its definition has been discussed at length in previous publications (Tschinkel and Doyen, 1980; Doyen and Tschinkel, 1982). Coelometopinae primarily occupy forest and woodland situations in the tropics and subtropics. Larvae inhabit rotten wood, usually when it has reached the punky stage of decay. Less frequently they are found in soil, beneath bark of more recently dead trees or in fruiting bodies of wood-rotting fungi. Adults are frequently found associated with various sorts of dead wood and are usually nocturnal. Loss of wings is common in this group, having led to convergence in body form among distantly related taxa (Doyen et al., in press and following discussion of Coelometopini). A substantial fauna of Coelometopinae inhabits the hardwood forests of eastern United States. This fauna is largely distinct from the larger Meso-American fauna at the generic level. The western North American fauna is depauperate, with two endemic genera {Coelocnemis, Cibdelis) and representatives of a few wide-ranging genera (Alobates, Iphthiminus, Strongylium). The following key includes Centronopini, which is similar to Coelometopini in external features. 292 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) KEY TO TRIBES OF COELOMETOPINAE AND CENTRONOPINI 1 . Antenna with stellate sensoria (visible at 50 x or higher) on apical five or six segments'; ventral surface of basal three or four tarsomeres covered by pads of dense, usually yellow pubescence; ventral surface of tarsomeres usually flattened (Fig. 8) 2 Antenna with stellate sensoria on apical seven or eight segments; ventral surface of tarsomeres covered by stiff, usually dark colored setae; tarsomeres cylindrical, at least on posterior two pairs of legs (Fig. 9) 3 2(1). Tarsomeres three and four subequal, each usually with pad of dense, yellowish pubescence (Fig. 9); body form variable Coelometopini and Centronopini (part) Fourth tarsomere much smaller than third bearing only a few long, ventral setae (Fig. 10); body form elongate, cylindrical Talanini 3(2). Prosternal process prominent, horizontal behind coxae; sharply acute and received in deep mesosternal fossa Coelometopini (part) Prosternal process declivous, flattened behind coxae; broadly rounded or truncate; mesosternal fossa very broad, shallow Strongyliini Tribe Coelometopini Coelometopini Lacordaire, 1859:358, Doyen et al., in press. Misolampini Lacordaire, 1859:440. Nodotelini Koch, 1950:67 (replacement name for Eutelini). Eutelini Sober, 1844:268 (not Walker, 1834; see Koch, 1950). Cnodalonini Lacordaire, 1859:414. Tenebrionini, various authors (in part). Hegemonini Reitter, 1922:5. Description. Adult. — Small to very large (about 5 to 45 mm) beetles of diverse body shape and color. Eyes reniform, moderate in size, separated dorsally by much more than width of single eye lobe. Antennae incrassate or weakly capitate, with stellate sensoria on apical five to six (rarely seven or eight) segments. Labral mem- brane exposed or concealed. Tarsi with ventral surface almost always flattened, bear- ing pads of yellowish, usually pilose setae; inner margins of tibiae frequently pilose, especially near apices. Ovipositor a flexible tube or rarely (Hegemona, Saziches) flattened, blade-like and strongly sclerotized; spermatheca swollen, spherical or rarely (Mylaris) isodiametric with accessory gland. Aedeagus with median lobe adnate to tegmen (freely extrusible in Tauroceras. Defensive reservoirs elongate, walls usually with annular folds (folds absent or rudimentary in Apsida, Camarid). Larva. — Moderately elongate, cylindrical or subcylindrical; ninth abdominal tergite usually with promient recurved urogomphi, rarely (Coelocnemis) with ring of acces- sory spines. Discussion. In catalogues and checklists most coelometopine genera are listed under tenebrionini, which are extremely different in several important internal features. 1) ‘ Stellate sensoria appear at 25 x to 50x as rounded, dome-shaped structures set in shallow pits on apical antennal segments. They are especially prevalent on the inner apical margins of the antennomeres. Above 50 x their stellate configuration may be discernible, particularly in larger specimens. 1989 NORTH AMERICAN TENEBRIONINI 293 Figs. 12-16. Antennal and leg variation in Coelometopinae. 12. Apsida. 13. Othryoneus. 14. Mophon, all from dorsal aspect. 15. Isicerdes. 16. Choastes, both from anterior aspect. In Tenebrionini the defensive reservoirs are short saccate with the secretion collecting ducts emptying diffusely through the dorsal surface (as in Fig. 7). The reservoir walls lack annular folds and are eversible. In Coelometopini the defensive reservoirs are elongate with one or a few collecting ducts emptying at the neck. The reservoirs are never eversible and their walls almost always have annular folds which allow ex- pansion (Tschinkel and Doyen, 1980, hg. 14e). 2) The ovipositor of Tenebrionini consists of a pair of basal paraprocts, subequal in length to the coxites (Doyen, 1966, figs. 71, 72). The coxites are subdivided into four subequal lobes. In Coelometopini the paraprocts are much shorter than the coxites, and at rest are rotated 1 80° so that the morphologically proximal ends lie distally beside the coxites (Tschinkel and Doyen, fig. 39). The proximal coxite lobe is longer than the distal lobes— often considerably longer than the three distal lobes combined. 3) In Tenebrionini the spermatheca and the spermathecal accessory gland are separate structures, either opening independently into the bursa copulatrix, or with the spermatheca emptying very near the base of the accessory gland (Tschinkel and Doyen, 1980, figs. 33, 34). In Coelometopini the apex of the accessory gland is nonglandular and functions as the spermatheca. In nearly all genera the apex is greatly dilated and spherical (Tschin- kel and Doyen, hg. 26). 4) The aedeagus of Tenebrionini has a connecting membrane between the median lobe and tegmen, allowing free extrusion of the median lobe. The median lobe extrudes below the tegmen. In Coelometopini the connecting mem- brane is almost always very short or absent, so that the position of the median lobe is hxed or nearly so relative to the tegmen. The aedeagus is usually rotated so that the median lobe is lateral or ventrolateral to the tegmen. 5) In Tenebrionini the antennae bear simple, setiform sensilla on all the segments. (Compound sensoria are present in closely related groups such as Alphitobiini and Triboliini, however.) In Coelometopini, in addition to setiform sensilla, there are compound, stellate sensoria on the apical hve to eight segments (see Medvedev, 1977, hgs. 31-55). These char- 294 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Figs. 17-19. Taxonomic characters of Coelometopini. 17. Elytral apex of Taurocems, left lateral aspect. 18. Head of Taurocems female, dorsal. 19. Elytral apex of Haplandrus fulvipes Herbst. acters are discussed in much greater detail in previous papers (Tschinkel and Doyen, 1980; Doyen and Tschinkel, 1982). Several larval differences between Coelometopini and Tenebrionini are discussed above under Centronopini. The genera assigned here to Coelometopini are split between Misolampini, Cno- dalonini and Coelometopini in catalogues. As pointed out earlier (Doyen et al., in press) Misolampini and Cnodalonini have not been redefined since Lacordaire (1859) originally proposed them on the basis of very superficial characters. Misolampini (and the Old World Nodotelini) simply comprise conglomerations of flightless Coe- lometopini, and it is now clearly evident that flightlessness has evolved independently dozens or even hundreds of times in Tenebrionidae. Certain of the “misolampine” genera appear to be closely related. Examples include Isaminas, Sphaerotus Kirby, Immedia Pascoe, Hemimedia Gebien and Parimmedia Gebien in the neotropics or Hegemona, Saziches and Promorphostenophanes Kaszab in the neotropics and ori- ental region respectively (Doyen, 1987). The latter group would correspond to He- gemonini of Reitter (1922). Other groups of genera, however, have certainly been derived from different parts of Coelometopini (e.g., Heliofugus Guerin, Myrmeco- dema Germain, Mitys Champion). Misolampus Latreille, from the Mediterranean region, differs from the most similar New World genera in several salient features, including the complete internalization of the scutellum, in the much shorter legs and partially exposed labral membrane, and in having the epipleural margins subparallel throughout, rather than broadened basally. It seems likely that Misolampus is more closely related to Coelometopus, which it resembles in general body form, than to any New World genus, but their exact derivation from winged forms is unclear. The genera discussed above all possess the complete inventory of diagnostic Coelometo- pine traits without significant variation. Diceroderes Sober, from Mexico, appears in catalogs under Eutelini (=Nodotelini). Properly, that genus, along with Ozolais and Calymmus, belongs in Toxicini (Doyen, 1988; Doyen et al., in press). In addition to the evidence cited above I have associated toxicine type larvae with Diceroderes. Cnodalonini Lacordaire is based on Cnodalon Latreille, which possesses all the important internal and external features of Coelometopini. Tschinkel and Doyen (1980) recognized cnodalonine categories for defensive reservoir configuration (res- 1989 NORTH AMERICAN TENEBRIONINI 295 Figs. 20-24. Taxonomic characters of Coelometopini. 20. Elytral apex of Hicetaon. 21. Head of Epicalla, dorsal. 22. Same, Merinus. 23. Mesothoracic leg of Alobates, anterior. 24. Same, Merinus, male. ervoir walls lacking annular folds), ovipositor (coxite lobes three and four fused), and internal female reproductive tract (spermatheca isodiametric with accessory gland. These distinctions were based on examination of other genera placed in Cnodalonini in catalogs. For example, defensive reservoirs of Camaria, Hapsida and several Old World genera lack annular folds. Likewise, Hapsida and several Old World “cno- dalonines” have coxite lobes three and four fused, and the latter do not have the spermatheca enlarged. Examination of additional taxa and characters shows that none of these character states is distributed in a recognizably systematic fashion. For example, while the defensive reservoirs completely lack annulation in Hapsida, they distinctly show basal annulation in Camaria and Blapida becoming nonannulate distally, especially in the latter. In Epicalla, phenetically similar to Camaria, the reservoirs are fully annulate. Again, examining the structure of the internal female reproductive tract, a non-enlarged spermatheca occurs in Apsida, Taphrosoma, Eu- cyrtus and Hemicera. Taphrosoma was previously classified in Tenebrionini, and is phenetically very different from the other genera. Hapsida contains highly specialized beetles previously classified as Diaperini (e.g., Triplehom, 1965, 1970), and the other two genera are Old World. Nor are the “cnodalonine” character states highly cor- related with one another: only Hapsida, Eucyrtus and Hemicera share all three “cno- dalonine” states. While Hemicera and Eucyrtus may be closely related, Hapsida is very different. In general it seems likely that all three of the “cnodalonine” characters are subject to convergence or else represent primitive retained characteristics. Thus, there is no morphological basis for recognizing a tribe Cnodalonini. Even if some of the more distinctive “cnodalonine” genera, such as Hapsida or the group related to Camaria, are eventually recognized, Cnodalonini could not be used no- 296 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) menclaturally since Cnodalon clearly belongs to the Coelometopini sensu stricto according to all diagnostic characters. Metaclisa marginalis, the only North American genus assigned to Cnodalonini, belongs to Tenebrioninae-Alphitobiini, discussed above. The genera included here in Coelometopini are listed in Table 1, where the former tribal association and pertinent references are also indicated. In this combined sense Coelometopini comprises one of the major groups of Tenebrionidae, and certainly one of the most variable, especially in tropical regions. Understanding its patterns of variation and producing a meaningful generic classihcation remains a major task. As in most large groups of organisms, not all genera of Coelometopini are equally easy to identify using keys. The following key is constructed so that several genera with intermediate or equivocal character states may be identified by following either alternative of the pertinent couplet. This is usually the reason that a taxon appears more than once in the key. In a few cases the genera as now conceived are polymorphic for the key characters, accounting for their multiple appearance in the key. This is the situation with Cibdelis, for example, where the epistomal border is emarginate in C bachei, often exposing the labro-clypeal membrane. In C. blaschkei the episto- mum is not emarginate and the membrane is concealed. KEY TO GENERA OF COELOMETOPINI AND CENTRONOPINI 1 . Labroclypeal membrane concealed beneath epistomum 7 Labrocly peal membrane broadly exposed just before epistomal margin 2 2(1). Tarsus with setal pad on penultimate segment much smaller than on preceding segments (Fig. 11), consisting of a narrow, apical fringe; lateral margins of pronotum dentate Cyrtosoma - Tarsus with setal pad on penultimate segment similar to those on preceding segments (as in Fig. 9); lateral margins of pronotum not dentate 3 3(2). Pronotum with base margined; antennae filiform, moniliform or gradually clavate (Figs. 12-14), with segments eight and nine subquadrate or longer than broad 4 Pronotum with base unmargined; antennae clavate (Fig. 12), with segments eight and nine about twice as broad as long Apsida 4(3). Prosternal process prominent, subhorizontal or horizontal behind coxae; mesosternum deeply excavate 5 - Prosternal process declivous immediately behind coxae; mesosternum very shallowly excavate, nearly flat Cibdelis 5(4). Antenna slender, filiform, much longer than head and prothorax combined (Fig. 14) 6 Antenna submoniliform (Fig. 1 3), shorter than head and prothorax combined Othryoneus 6(5). Eye with deep groove extending around apex of ventral lobe; epipleuron ter- minating abruptly at about anterior margin of fifth visible abdominal stemite Mophon - Eye with groove extending medially behind maxillary articulation from apex of ventral lobe of eye; epipleuron extending to elytral apex Elomosda 7(1). Elytra with apices conforming to shape of last abdominal sternite 9 Elytra with apices produced beyond last abdominal sternite as spine-like pro- cesses 8 1989 NORTH AMERICAN TENEBRIONINI 297 8(7). Foretibia strongly curved ventrad in apical third; metastemal length about equal to coxal diameter 41 - Foretibia nearly straight; metastemal length at least twice mesocoxal diameter Blapida 9(7). Metafemur abruptly emarginate on apical 'A to of inner surface (Figs. 15, 16); often with small spur or prominence just basad of emargination 10 - Metafemur with margins subparallel or gradually convergent toward apex, never abruptly emarginate 14 10(9). Body length greater than 3 cm; fifth visible abdominal stemite with marginal groove (some males of) Mylaris Body length less than 2 cm; fifth visible sternite without marginal groove ... 11 11(10). Metastemal length about 'A to Vi diameter of mesocoxa; all femora with spur on inner surface % distance to apex Cnephalura Metastemal length about 1.5 times diameter of mesocoxa; spurs present or absent from inner femoral surface 12 12(11). Metatibia arcuately curved (Fig. 15); prostemal process horizontal or sub- horizontal behind coxae Isicerdes, Ilus Metatibia straight except just before articulation with femur; (Fig. 16) proster- nal process declivous behind coxae 13 1 3( 1 2). Metafemora reaching anterior margin of fifth abdominal stemite; mesosternal fossa with margins strongly elevated, dentiform Choastes Metafemora reaching at most to anterior margin of fifth abdominal stemite; mesosternal fossa with margins slightly raised Hesiodus 14(9). Head with abrupt, arcuate escarpment across frons between eyes; eye with supertending groove, expanding into a large deep excavation behind eye . . . Bothynocephalus Head with frons uniformly curved or with epistomal region slightly depressed below level of frons; head sometimes with grooves along dorsal margin of eye, but never with large pits 15 15(14). Epipleuron abruptly broadened at fourth visible abdominal stemite then nar- rowed to elytral apex (Fig. 1 7); epistoma produced as sharp projection between eye and lateral epistomal suture; clypeus produced laterally much beyond epistomal suture (Fig. 18) (Centronopini) Tauroceras Epipleuron with margins subparallel or gradually converging posteriorly (sometimes abruptly narrowing at about anterior margin of fifth abdominal sternite) (Figs. 19, 20); epistoma evenly arcuate between eye and lateral epi- stomal suture; clypeus not produced laterally beyond epistomal suture 16 16(15). Dorsal lobe of eye with marginal groove around apex 17 - Dorsal lobe of eye without groove around apex 29 17(16). Metastemal length between coxae 1.5 to 2 times mesocoxal diameter 20 Metastemal length between coxae equal to or less than mesocoxal diameter 1 8 1 8(17). Tibiae with paired, narrow longitudinal lines of yellowish pubescence on apical V2 to % of inner surfaces 30 Tibiae without lines of pubescence, or with apical '/s to 'A with faint setal lines or patch 19 19(18). Legs long, metafemur reaching at least to base of fifth abdominal stemite, frequently extending beyond apex of abdomen 42 - Legs shorter, metafemur reaching no farther than third abdominal stemite Polypleurus 20(17). Epipleuron complete to apex of elytra, gradually narrowed posteriorly from fifth abdominal sternite (Fig. 20) 22 298 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) 21(20). 22(20). 23(22). 24(23). 25(22). 26(25). 27(25). 28(27). 29(16). 30(18, 29). 31(29). 32(31). 33(31). 34(26, 33). Epipleuron abruptly narrowing at about anterior margin of fifth abdominal sternite, disappearing before elytral apex (Fig. 19) 21 Mesostemal fossa deep, with lateral margins strongly raised; pronotal base with complete, raised margin Sthenoboea Mesostemal fossa broad, shallow, with lateral margins scarcely raised; pronotal base unmargined, at least medially Haplandrus Fifth abdominal sternite without marginal groove or line 25 Fifth abdominal sternite with marginal groove or impressed line 23 Frons behind epistomal suture much more coarsely punctate than posteriorly and around eyes Hicetaon Frons finely, evenly punctate 24 Fifth abdominal sternite with groove deeply excavate; mentum flat Oeatus Fifth sternite with fine, slightly impressed marginal line; mentum elevated as a prominent tubercle anteriorly Glyptotus Femora strongly clavate; profemur 2.5 to 4 times thicker in middle than at base 27 Femora subcylindrical; profemur no more than 2 times thicker in middle than at base 26 Epistomal suture faint in medial portion, shallowly impressed or obsolete ... 34 Epistomal suture very deeply incised in medial portion Paroeatus Pronotum finely and shallowly punctate (punctures much smaller than single eye facet); elytra punctate-striate 28 Pronotum coarsely and deeply punctate (punctures as large as several eye facets combined); elytra reticulately rugose Upis Epistomal suture strongly incised Nuptis Epistomal suture very fine, shallow, sometimes partially obsolete, never in- cised Merinus All tibiae with paired, narrow longitudinal lines of yellowish pubescence on apical V2 to % of inner surface 30 Tibiae without lines of pubescence or with apical Vs to 'A sometimes with single faint setal lines or with setal patch 31 Gena with deep, abrupt excavation at apex of ventral lobe of eye .... Oenopion Gena often coarsely rugose but never with distinct excavation at apex of ventral lobe of eye Coelocnemis Antenna gradually enlarged or serate-filliform (as in Figs. 13, 14); segments nine and ten no more than 1.5 times as broad as long 33 Antenna with apical five or six segments enlarged as more-or-less distinct club (Fig. 12); segments nine and ten about twice as broad as long 32 Elytral base broader than base of thorax; epipleuron expanded as prominent umbo at elytral base; elytra coarsely punctate-striate Cnodalon Elytral and thoracic bases equally broad; epipleuron not forming umbo; strial punctures of elytra extremely fine Gonospa Epipleuron gradually narrowing posteriorly, with subparallel margins through- out except near humerus and apex (Fig. 20); extending to elytral apex or nearly so 40 Epipleuron abruptly narrowed at about anterior margin of fifth abdominal sternite; not reaching elytral apex (Fig. 1 9) 34 Prosternal process horizontal, prominent behind coxae; mesostemum deeply excavate with raised lateral borders 35 Prosternal process declivous; mesostemum very shallowly excavate, lateral borders not raised 36 1989 NORTH AMERICAN TENEBRIONINI 299 35(34). 36(35). 37(34). 38(37). 39(37). 40(34). 41(8, 40). 42(19, 41). 43(42). 44(42). 45(44). 46(44). 47(40). 48(47). 49(48). Elytral interstices smooth; antenna extending posteriorly beyond eyltral base 36 Elytral interstices bearing large, shining tubercles; antenna not reaching base of prothorax Xenius Elytral length no more than 1.5 times width; pronotum with continuous raised margin around lateral and posterior borders Nesocyrtosoma Elytral length more than twice width; pronotum with submarginal groove along lateral border, becoming deeper posteriorly and interrupting raised mar- gin at corner Moen Femora clavate; profemur about twice as thick in middle as at base (Centronopini) 39 Femora subcylindrical; profemur less than 1.5 times thicker in middle than at base 38 Metastemum length between coxae 1.5 to 2 times mesocoxal diameter 50 Metasternum length between coxae less than mesocoxal diameter Cibdelis Fifth visible abdominal sternite with marginal groove Scotobaenus Fifth visible abdominal sternite without marginal groove Centronopus Metastemum length between coxae about 1.5 to 3 times mesocoxal diameter 47 Metastemum length between coxae equal to or less than coxal diameter .... 41 Legs long, metafemur reaching fifth abdominal sternite or beyond 42 Legs shorter, metafemur reaching third or fourth abdominal sternite Polopinus Metatarsus excluding claw at least % length of tibia 43 Metatarsus shorter, almost always about half length of tibia 44 Antenna with segments three and four subequal in length; segments six to ten about three times as long as wide; body more than 2 cm long Hegemona Antenna with segment three about 1 . 5 times longer than segment four; seg- ments six to ten about twice as long as wide; body less than 1 5 mm long . . Saziches Prostemal process horizontal, prominent behind coxae, fitting into deep meso- sternal fossa with strongly raised lateral margins 45 Prostemal process subhorizontal or declivous; mesostemum very shallowly concave or flat, with lateral margins not raised 46 Epistomal suture deeply impressed at angles anteromedial to eyes, usually forming distinct foveae Isaminas Epistomal suture shallow throughout its course, never deeper at angles Oxidates Mentum with anterior central portion elevated as a forward projecting tubercle Mitys Mentum almost flat, never with anterior elevation Cibdelis Head constricted abruptly just behind eyes, much narrower than before eyes (Fig. 21) 48 Head with lateral margins subparallel or gradually narrowed, not abruptly constricted behind eyes (Fig. 22) 50 Prostemal process horizontal or subhorizontal, prominent behind coxae; re- ceived in deep fossa with raised lateral borders in mesostemum 49 Prostemal process declivous behind coxae; received in shallow, obtuse fossa without raised lateral margins 53 Medial epistomal suture in broad, shallow depression with shallow fovea before each eye; head anterad of eyes shorter than width of dorsal eye lobe Epicalla 300 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) - Medial epistomal suture not depressed; no foveae or depressions before eyes; head anterad of eyes longer than width of dorsal eye lobe Camaria 50(38,47). Pronotal base with complete raised margin 51 Pronotum with base unmargined, at least medially Haplandrus Basal four tarsomeres with ventral surface entirely covered by pads of dense pubescence 52 Tarsomeres with two longitudinal rows of setae, separated by groove with few sparse setae Iphthiminus Elytra regularly striate (strial punctures sometimes very fine) 53 Elytra irregularly rugose; striae not discernible Upis Fifth abdominal stemite without marginal groove 55 Fifth abdominal stemite with marginal groove 54 Epistomal margin arcuately concave; head constricted behind eyes; more than 2.5 cm long My laris Epistomal margin nearly straight; head not constricted behind eyes; less than 2 cm long Xylopinus Meso- and metatibia arcuately curved (Fig. 23); pronotum with hind angles rounded Merinus Meso- and metatibia straight (Fig. 24); pronotum with hind angles sharp, right angled Alobates Tribe Strongyliini Strongyliides Lacordaire, 1859 and various authors. Strongyliini, various authors. Description. Adult. — Very small to moderate (about 3 mm to 20 mm) beetles, usually with elongate, subcylindrical body; color highly variable. Eyes large, usually separated dorsally by less than width of single eye lobe, frequently contiguous or nearly so, especially in males. Antennae usually serrate, sometimes weakly incrassate, rarely pectinate; apical seven or eight segments bearing stellate, compound sensoria. Labral membrane exposed. Tarsi nearly cylindrical, ventral surfaces usually covered by stiff, often dark colored setae; tibiae seldom bearing strips or patches of pile. Ovipositor a flexible tube; spermatheca swollen, spherical. Aedeagus with median lobe adnate to tegmen or only slightly extrusible. Defensive reservoirs short saccate, without annular folds. Larva. — Very elongate, cylindrical or nearly so; tergite nine rigidly sclerotized, formed into pair of large, apically bifid, recurved urogomphi, supertended anteriorly by a serrate, transverse ridge; tergite eight rigidly sclerotized, sometimes with trans- verse ridge or denticles; sternites eight and nine less rigidly sclerotized. Discussion. Many Strongyliini are brightly metallic or pastel, as in some Coelo- metopini. In the structure of the defensive glands there is no confusion between the tribes, at least in the New World fauna. All Strongyliini have stellate sensoria on the terminal seven or eight antennal segments. Among north and central American Coelometopini stellate sensoria occur on the terminal five or six segments, with the exception of Mophon, where they are borne on the apical eight segments. The form of the ninth abdominal segment of larval Strongyliini is superficially similar to that of some Coelometopini such as Coeloenemis, where a row of complex spines supertends the urogomphi. In Coeloenemis, however, tergite nine becomes weakly sclerotized anteriorly, and segment eight and stemite nine are no more strongly 51(50). 52(51). 53(52). 54(53). 55(54). 1989 NORTH AMERICAN TENEBRIONINI 301 Figs. 25, 26. Cranial structure in Strongyliini. 25. Oblique aspect of head of Strongylium. 26. Same, Merit es. sclerotized than the preceding segments. Triplehorn and Spilman (1973) described larvae of four species and pupae of two species of North American Strongylium. Mentes has been placed in the Helopini on the basis of Champion’s (1893) vague statement that it was probably a “degraded” member of that tribe. I have been able to dissect only males, whose antennae have stellate, compound sensoria on segments three to eleven, and whose defensive glands are of the short saccate type of Stron- gyliini, with much common volume and without annulation. In Helopinae compound sensoria are never present on the antennae, and the defensive reservoirs are elongate, medially expanded and without common volume (Tschinkel and Doyen, 1980). On the basis of these characters I am placing Mentes in Strongyliini, and I predict that when observed the ovipositor and internal female reproductive tract will support this transfer. Without a detailed study it is impossible to judge the cladistic relationship of Otocerus or Pseudotocerus to Strongylium. They are separated in the following key according to the characters used by Champion (1888). In this regard the other small genera of Strongyliini described by Pic and others must also be suspected as spe- cialized derivatives of Strongylium, whose New World species are unrevised since the work of Maklin (1862). The Chilean genus Homocyrtus Reitter, placed in Stron- gyliini in catalogues, is a member of Chalcodryidae (Watt, pers. comm.; Doyen, unpublished). KEY TO STRONGYLIINI 1 . Antenna with apical 5 or 6 segments much broader than long, forming a more or less distinct club 2 - Antenna with all segments longer than broad, filiform to serrate 3 2(1). Pronotum margined Poecilesthus - Pronotum not margined Cuphotes 3(1). Antenna inserted beneath strong epistomal canthus; canthus laterally elevated, emar- ginating eye posterior to antennal base (Fig. 25); antennae usually filiform 4 - Antenna inserted within emargination in eye; epistomal canthus very weak, narrow (Fig. 26); antenna serrate or pectinate, especially in males Mentes Champion 4(3). Antenna with segment three much shorter than segment four; medial apical angles of antennal segments four to ten usually produced, giving a serrate appearance 5 - Antenna with segment three as long or longer than segment four; segments four to ten usually filiform Strongylium 5(4). Hind femur reaching apex of elytra; basal segment of hind tarsus 2 to 3 times as long as apical segment Pseudotocerus - Hind femur shorter, not reaching apex of elytra; basal segment of hind tarsus 1 to 2 times as long as apical segment Otocerus 302 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Tribe Talanini Talanites Champion, 1887. Talanini, various authors. Dignamtini LeConte and Horn, 1883. Description. Adult. — Small to moderate (about 3 mm to 10 mm). Eyes moderate to large, bulging, but separated dorsally by much more than width of dorsal eye lobe. Antennae incrassate, bearing stellate, compound sensoria on apical five or six seg- ments. Labral membrane exposed. Tarsi with ventral surface flattened, with pads of yellowish, pilose setae; fourth tarsomere of first two pairs of legs much smaller than preceding tarsomeres and bearing only few long setae. Inner tibial surface setose or pilose. Ovipositor strongly sclerotized, laterally compressed, without apparent gono- styli or lobing of coxite (Tschinkel and Doyen, 1980, fig. 41); spermatheca swollen, spherical. Aedeagus with median lobe slightly extrusible. Defensive reservoirs large, saccate, constricted at neck, without annular folds. Larva. — Unknown. Discussion. Talanini contains only the Neotropical genus Talanus, which univer- sally possesses a highly modified, blade-like ovipositor, superficially similar to those in Hegemona, Saziches and Acropteron. In Talanus, however, the paraproct has the ability to rotate about its articulation with the coxite, as in nearly all Coelometopinae. In the others the paraproct cannot rotate, and they differ in numerous other important characters, as discussed above. Talanus lacks compound sensoria on antennal seg- ments four and five, and has the tarsi ventrally pilose, differentiating it from Stron- gyliini, near which it is usually placed in catalogues. The lack of annulation on the defensive reservoir walls is shared with Apsida and Camaria among the New World Coelometopini, and in Apsida the penultimate tarsomere is smaller than the anti- penultimate and has reduced pilosity, as in Talanus. Talanus differs from both of these in numerous other characters, however, and must be accorded an isolated position within Coelometopinae. Talanini has been long recognized, and unless more compelling evidence can be marshalled for derivation from some other tribe, it should be retained for these strongly apomorphic beetles. ACKNOWLEDGMENTS Many persons contributed ideas which were incorporated into the present work. Chief among these are K. W. Brown, (San Joaquin County Department of Agriculture, Stockton, CA), J. F. Lawrence (CSIRO, Canberra, Australia), E. G. Matthews (South Australian Museum, Adelaide), W. R. Tschinkel (Florida State University, Tallahassee) and C. S. Watt (DSIRO, Auckland, New Zealand). Needless to say they do not all agree on all the taxonomic aspects and are not in the least responsible for errors in fact or interpretation. R. Aalbu (California Department of Food and Agriculture, Sacramento) kindly made available information regarding the identity of Biomorphus tuberculatus. The drawings were done by Ms. C. Jordan (University of California, Berkeley). M. A. Ivie, Montana State University, read the manuscript and offered many helpful comments and ideas. LITERATURE CITED Aalbu, R. L., T. J. Spilman and K. W. Brown. 1989. The systematic status of Amblycyphrus asperatus, Threnus niger, Pycnomorpha californica, Emmenastus rugosus, and Biomor- phus tuberculatus Motschoulsky (Coleoptera: Tenebrionidae). Coleopt. Bull, (in press). 1989 NORTH AMERICAN TENEBRIONINI 303 Blackwelder, R. E. 1945. Checklist of the Coleopterous insects of Mexico, Central America, the West Indies, and South America. Part 3. Bull. U.S. Nat. Mus. 185:343-550. Champion, G. C. 1887. Biologia Centrali- Americana, Insecta, Coleoptera. 4(l):265-353. Champion, G. C. 1888. Biologia Centrali-Americana, Insecta, Coleoptera. 4(l):354-476. Champion, G. C. 1893. Biologia Centrali-Americana, Insecta, Coleoptera (Supplement) 4(1): 477-524. Crowson, R. A. 1955. The Natural Classification of the Families of Coleoptera. Lloyd and Company, London, 187 pp. Doyen, J. T. 1966. The skeletal anatomy of Tenebrio molitor (Coleoptera: Tenebrionidae). Misc. Publ. Entomol. Soc. Amer. 5:103-150. Doyen, J. T. 1973. Systematics of the genus Coelocnemis (Coleoptera: Tenebrionidae): a quantitative study of variation. Univ. Calif. Publ. Entomol. 73:1-110. Doyen, T. J. 1984. Reconstitution of the Diaperini of North America, with new species of Adelina and Sitophagus (Coleoptera: Tenebrionidae). Proc. Entomol. Soc. Wash. 86: 777-789. Doyen, J. T. 1985. Reconstitution of the tribes Ulomini and Triboliini for North and Central America (Tenebrionidae: Coleoptera). Proc. Entomol. Soc. Wash. 87:512-524. Doyen, J. T. 1987. New and little known Tenebrionidae from Mexico and Central America, with remarks on their classification. Pan-Pac. Entomol. 63:301-318. Doyen, J. T. 1988. Descriptions of some phylongenetically important larvae of Tenebrionidae (Coleoptera). Coleopt. Bull. 42:285-301. Doyen, J. T. and J. F. Lawrence. 1979. Relationships and higher classification of some Tenebrionidae and Zopheridae (Coleoptera). Syst. Entomol. 4:333-377. Doyen, J. T., E. G. Matthews and J. F. Lawrence. 1989. Classification and an annotated checklist of the Australian genera of Tenebrionidae (Coleoptera). Invert. Tax. 2 (in press). Doyen, J. T. and W. R. Tschinkel. 1982. Phenetic and cladistic relationships among tene- brionid beetles (Coleoptera). Syst. Entomol. 7:127-183. Gebien, H. 1919. Monographie der sudamerikanischen Camarien (Coleopt. Heterom.) nebst einer Ubersicht fiber die indischen Gattuugen der Camariinen. Archiv. f. Naturg. 83(1917): 25-168. Gebien, H. 1942-44. Katalog der Tenebrioniden. Teil III. Mitt. Mfinchener Entomol. Gesell. 32:729-760 [746-777], 33:339-430, 895-926 [778-841], 34:497-555 [842-899]. Kaszab, Z. 1 984. Fine neue Schwarzkafer— Gattung und— Art aus Domikanischem Bernstein (Coleoptera, Tenebrionidae). Stuttg. Beitr. Naturk. (B), 109:1-6. Koch, C. 1950. Proposed change of African generic names in the family of Tenebrionidae. Entomologist 83(1042):66-68. Lacordaire, T. 1859. Histoire naturelle des insects. Genera des Coleopteres. Tome 5, Roret, Paris, 750 pp. Lawrence, J. F. 1982. Coleoptera. Pages 482-553 In: Parker, S. P. (ed.). Synopsis and Clas- sification of Living Organisms. McGraw-Hill, Inc., New York. LeConte, J. L. 1862. Classification of the Coleoptera of North America. Prepared for the Smithsonian Institution. Smithson. Misc. Coll. 136:209-286. LeConte, J. L. and G. H. Horn. 1883. Classification of the Coleoptera of North America. Smithson. Misc. Coll. 507:38-567. Maklin, F. W. 1862. Die Arten der Gattung Acropteron Perty. Acta Soc. Sci. Fenn. 7:545- 554. Marcuzzi, G. 1976. New species of Neotropical Tenebrionidae (Coleoptera). Ann. Hist. -Nat. Mus. Nat. Hung. 68:117-140. Matthews, E. G. and J. T. Doyen. 1 989. A reassessment of the Australian species of Menephilus Mulsant (Coleoptera: Tenebrionidae) with descriptions of two new genera and a larva and pupa. Rec. S. Austr. Mus. (in press). Medvedev, G. S. 1977. Taksonomicheskoye znacheniye antennal’nikh sensill zhukovcher- 304 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) notelok (Coleoptera, Tenebrionidae). In: Akademia Nauk S.S.S.R., Trudi Vsesoyuznovo Entomologicheskovo Obschchestva 58:61-68. Morphologischeskie osnovi sistematiki nasekomikh. Akademia Nauk S.S.S.R., Leningrad. (The taxnomic significance of the antennal sensilla of the darkling beetles (Coleoptera: Tenebrionidae).) Papp, C. S. 1961. Checklist of the Tenebrionidae of America, north of the Panama Canal (Notes on North American Coleoptera, No. 14). Opusc. Entomol. 26:97-140. Reitter, E. 1917. Bestimmungstabellen europaischen Coleoptera. — 8 1 . Unterfamilien und tribus der palaeartikischen Tenebrionidae. Wien. Entomol. Zeit. 36:51-66. Reitter, E. 1 920. Bestimmungstabellen der Europaishen Coleopteren. Heft 87. Tenebrionidae. XV Teil. Edmund Reitter’s Nachfolger, Paskau (Mahren). Reitter, E. 1 922. Bestimmungstabellen der Europaishen Coleopteren. Heft 92. Tenebrionidae. XVI Teil. Unterfamilie Helopinae I. Wien. Entomol. Zeit. 39:1-44. Sober, A. J. 1844. Essai sur les collapterides de la tribu des Molurites. Mem. Accad. Sci. Torino (2)6:213-339. Spilman, T. J. 1959. Notes on Edrotes, Leichenum, Palorus, Eupsophulus, Adelium, and Strongylium (Tenebhonida.Q). Coleopt. Bull. 13:58-64. Spilman, T. J. 1961. Remarks on the classification and nomenclature of the American te- nebrionine genus Adelonia (Coleoptera: Tenebrionidae). Pan-Pac. Entomol. 37:49-51. Spilman, T. J. 1962a. The New World genus Centwnopus with new generic synonymy and a new species (Coleoptera: Tenebrionidae). Trans. Amer. Entomol. Soc. 88:1-19. Spilman, T. J. 1962b. A few rearrangements in the Tenebrionidae, with a key to the genera of the Ulomini and Tenebrionini of America, North of Mexico. Coleopt. Bull. 16:57-63. Spilman, T. J. 1963. On larvae, probably Tauroceras, from the Neotropics (Coleoptera: Te- nebrionidae). Coleopt. Bull. 17:58-64. Spilman, T. J. 1973. Nomenclatural problems in six genera of Tenebrionidae (Coleoptera). Proc. Entomol. Soc. Wash. 75:39-44. Spilman, T. J. 1 979. Larvae and pupae of Centwnopus calcaratus and Centwnopus suppressus (Coleoptera: Tenebrionidae) with an essay on wing development in pupae. Proc. Entomol. Soc. Wash. 81:513-521. St. George, R. A. 1924. Studies on the larvae of North American beetles of the subfamily Tenebrioninae with a description of the larva and pupa of Merinus laevis (Olivier). Proc. U.S. Nat. Mus. 65:1-22. Triplehom, C. A. 1965. Revision of Diaperini of America north of Mexico with notes on extralimital species (Coleoptera: Tenebrionidae). Proc. U.S. Nat. Mus. 117:349-458. Triplehom, C. A. 1970. A synopsis of the genus Apsida with description of a new species (Coleoptera: Tenebrionidae). Ann. Entomol. Soc. Amer. 63:567-572. Triplehom, C. A. and T. J. Spilman. 1973. A review of Stwngylium of America North of Mexico, with descriptions of two new species (Coleoptera, Tenebrionidae). Trans. Amer. Entomol. Soc. 99:1-27. Tschinkel, W. R. and J. T. Doyen. 1980. Comparative anatomy of the defensive glands, ovipositors and female genital tubes of tenebrionid beetles (Coleoptera). Intern. Jour. Ins. Morphol. & Embryol. 9:321-368. Watt, J. C. 1970. Coleoptera: Perimylopidae of South Georgia. Pac. Ins. Monogr. 23:243-253. Watt, J. C. 1974. A revised subfamily classification of Tenebrionidae (Coleoptera). New Zealand Jour. Zool. 1:381-452. Watt, J. C. 1 987. The family and subfamily classification and New Zealand genera of Pythidae and Scraptiidae (Coleoptera). Syst. Entomol. 12:111-136. Received November 30, 1988; accepted March 28, 1989. J. New York Entomol. Soc. 97(3):305-308, 1989 PITYOGENES BIDENTATUS (HERBST), A EUROPEAN BARK BEETLE NEW TO NORTH AMERICA (COLEOPTERA: SCOLYTIDAE) E. Richard Hoebeke Department of Entomology, Cornell University, Ithaca, New York 14853-0999 Abstract. —Pityogenes bidentatus, a widespread and abundant bark beetle in Europe, has been intercepted numerous times at U.S. ports of entry. It now has become established in the northeastern United States. A recent collection was made of this beetle from dying specimens of Bosnian pine, Pinus leucodermis, in a nursery plantation in Livingston County, New York, in April 1988. North American interception records, host plants, and habits of this bark beetle are summarized from the literature. An existing key to the North American species of Pityogenes is modified to include this newly detected immigrant, and scanning electron photomicrographs of diagnostic male and female structures are provided. During a routine nursery inspection (by J. P. Filkens, Horticultural Inspector, NYS Agric. & Markets) in early April 1988, several Bosnian pines {Pinus leucodermis Antoine)— native to the Balkans and one of Europe’s most common ornamental pines (Kriissmann, 1985)— were found dead or dying in a plantation in Lima, New York (Livingston County). The trees exhibited characteristic damage symptoms caused by bark beetles. Cut samples of these pines, with bark intact, were sent to the Insect Diagnostic Laboratory, Department of Entomology, Cornell University, for diag- nosis. By June, a number of adult beetles were reared, and identified as a European bark beetle, Pityogenes bidentatus (Herbst), not previously reported from North America. Because the infested Bosnian pines in the nursery plantation had been grown from seed and were a minimum of 10-15 years old (J. Filkens, pers. comm.), I am rea- sonably confident that the introduction of this European bark beetle into North America has not been a recent event. Inasmuch as the city of Rochester, only 33 km north of the detection site, is one of the major ports of entry in the Northeast, it is reasonable to assume that P. bidentatus was introduced on coniferous nursery stock at a much earlier date. For nearly a 40-year period, from 1948-1985, specimens of P. bidentatus were intercepted at major U.S. ports of entry on numerous occasions. There are at least 53 interception records documented in the “List of Intercepted Plant Pests,” compiled by the U.S. Department of Agriculture. Most specimens found during inspection were associated with wood {Pinus spp.), dunnage and wood crating, and pallets originating from various European countries (West Germany, Netherlands, Italy, Poland, Denmark, Finland, Sweden, Spain, France, Belgium and Portugal) and des- tined for a number of entry points in the United States (New York, Texas, Alabama, California, Delaware, Michigan, Louisiana, Georgia, Pennsylvania, Maryland, South Carolina, Illinois, Florida and Ohio). Pityogenes bidentatus, known in the European literature as the two-toothed bark 306 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Figs. 1-3. Pityogenes bidentatus. 1. Male elytral declivity. 2. Female elytral declivity. 3. Female head. beetle, is widely distributed and common throughout northern and central Europe, USSR, and Great Britain. Its principal host is Scotch pine [Pinus sylvestris L.], but it will also attack other pines [P. strobus L., P. mugo Turra., P. pinaster Ait., P. nigra Arnold, P. pithyusa Steven, P. pumila (Pall.) Regel, P. banksiana Lamb., P. cembra L., P. sibirica Du Tour, and P. sylvestris var. lapponica Fries], spruces [Picea abies (L.) Karst., P. obovata Ledeb., P. orientalis (L.) Link.], firs [Abies alba Mill., A. nordmanniana (Stev.) Spach], larch [Larix decidua Milk], Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco], cypress [Cupressus spp.], arborvitae [Thuja spp.], and false- cypress [Chamaecyparis lawsoniana (A. Murr.) Park] (Munro, 1926; Pavlovskii, 1955; Browne, 1968; Schwenke, 1974). Pityogenes bidentatus, a rather small (2.0-2. 5 mm) and variable bark beetle, usually breeds in slash and in cut or fallen limbs and branches. Occasionally it can be an 1989 SCOLYTIDAE 307 important secondary pest in young conifer plantations, attacking trees damaged by frost. The species is polygamous. Longitudinal egg galleries are cut by the female in the cambium region, and radiate from a central nuptial chamber. Numerous larval mines radiate away from the egg galleries and terminate in pupal cells. Pupation occurs in the cambium, and young adults emerge from individual exit holes in the bark (Browne, 1968). There are 2 generations recorded in Europe (Griine, 1979), with flight periods from May-June and July-August. Additional details of the biology and ecology of this species may be found in Eichhoff (1881), Escherich (1923), Balachowsky (1949), Chararas (1962) and Schwenke (1974). In North America, 6 species of Pityogenes are known to occur (Wood, 1982), and all breed in the twigs and thin-barked limbs and branches of pine. Although most species of the genus prefer to breed in slash, or attack trees weakened by drought, transplanting, ground hres, or mechanical means, young, healthy pines or vigorous trees in heavily infested areas occasionally are attacked (Baker, 1972). Adults of Pityogenes bidentatus differ from those of all other North American members of the genus by the following characters. In the male, the elytral declivity is armed by an upper pair of small, distinct spines adjacent to the suture and by a pair of large, hooklike teeth along the dorsolateral margin (Fig. 1). In the female, the frons lacks the large fossal excavation evident in the majority of the North American species, and is instead convex with a pair of small, bilateral indentations and a medial triangular area of dense, short pile (Fig. 3). Because of the unique combination of the armature of the male elytral declivity and the convex, unexcavated frons of the female, P. bidentatus does not trace to any species treated in the North American keys. To accommodate this newly detected European immigrant, couplets #1-4 of Wood’s (1982:650-65 1) key to the North and Central American species of Pityogenes should be modified, in part, and expanded as follows: 1 . Male elytral declivity armed by three pairs of coarse denticles; female declivity rather gradual, armed by three pairs of denticles of about equal size and equally spaced; female fossa large, undivided; in 5 -needle pines 2 - Male elytral declivity armed by at least one upper pair of large, hooklike teeth; female declivity rather steep, armed by three pairs of denticles with upper pair much smaller and much closer to second pair, or declivity without obvious pairs of denticles as in bidentatus-, female fossa divided by median septum, except undivided in carinulatus, and absent in meridianus and bidentatus 3 2. (Couplet #2 unchanged) 3. Female frons devoid of fossal excavation 4 Female frons with a conspicuous fossal excavation 5 4. Smaller species, 2. 0-3.0 mm; female frons flat to convex 4a Larger species, 3. 2-3. 4 mm; female frons along median line on upper half shallowly concave; Mexico; Pinus hartwegii 4. mexicanus Wood 4a. Male elytral declivity with one pair of small, but distinct spines adjacent to suture above pair of large, hooklike teeth (Fig. 1); female declivity without conspicuous pairs of denticles (Fig. 2); New York (introduced from Europe); Pinus leucodermis and other conifers; 2.0-2. 5 mm 8. bidentatus (Herbst) Male elytral declivity without pair of small spines above large, hooklike teeth; female declivity with three pairs of conspicuous denticles; North Carolina to Mississippi; Pinus echinata, P. taeda-, 2. 5-3.0 mm 3. meridianus Blackman 5. (Couplet #5 unchanged). 308 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) ACKNOWLEDGMENTS I thank D. M. Anderson (USDA, ARS, Systematic Entomology Laboratory, Washington, DC) and S. L. Wood (Department of Zoology, Brigham Young University, Provo, UT) for confirming the identification of the bark beetle, and J. P. Filkens (Horticultural Inspector, Prattsburg, NY) for bringing the bark beetle infestation to my attention. D. M. Anderson and A. G. Wheeler, Jr. (Penn. Dept. Agric., Harrisburg, PA) provided critical review of a draft of the manuscript. LITERATURE CITED Baker, W. L. 1972. Eastern forest insects. U.S. Dept. Agric., For. Serv., Misc. Publ. No. 1 175, 642 pp. Balachowsky, A. 1949. Coleopteres Scolytides. Faune de France 50, P. Lechevalier, Paris, 320 pp. Browne, F. G. 1968. Pests and diseases of forest plantation trees: an annotated list of the principal species occurring in the British Commonwealth. Clarendon Press, Oxford, 1330 pp. Chararas, C. 1962. Etude biologique des scolytides des coniferes. Ency. Entomol., Ser. A, Bd. 38, 556 pp. Eichhoff, W. L. 1881. Die Europaischen Borkenkafer. J. Springer, Berlin, 315 pp. Escherich, K. 1923. Die Forstinsekten Mitteleuropas. Bd. II. P. Parey, Berlin, 663 pp. Griine, S. 1979. Handbuch zur Bestimmung der europaischen Borkenkafer. M. & H. Schaper, Hannover, 182 pp. Kriissmann, G. 1985. Manual of cultivated conifers. Timber Press, Portland, Oregon, 361 pp. Munro, J. W. 1926. British bark-beetles. For. Comm. Bull. No. 8, H. M. Station. OIL, London, 77 pp. Pavlovskh, E. N. (Ed.) 1955. Forest pests. A handbook, II. Pages 425-1097. Moscow and Leningrad, Akad. Nauk SSSR (Zool. Inst.). [In Russian.] Schwenke, W. 1974. Die Forstschadlinge Europas, Bd. 2 P. Parey, Hamburg and Berlin, 500 pp. Wood, S. L. 1982. The bark and ambrosia beetles of North and Central America (Coleoptera: Scolytidae), a taxonomic monograph. Great Basin Nat. Mem. 6:1-1359. Received November 2, 1988; accepted February 6, 1989. J. New York Entomol. Soc. 97(3):309-316, 1989 NEW TRICHOPTERA EROM ALABAMA S. C. Harris Aquatic Biology Program, Department of Biology, University of Alabama, Tuscaloosa, Alabama 35487 Abstract. — Yxvq new species of Trichoptera, Rhyacophila carolae (Rhyacophilidae), Protoptila cahabensis (Glossosomatidae), Hydroptila micropotamis and Ochrotrichia weoka (Hydroptili- dae), and Ceraclea alabamae (Leptoceridae) are described and illustrated. In the course of a continuing survey of the caddisflies of Alabama, five undescribed species were collected using UV light traps. These new species, one in each of the genera Rhyacophila, Protoptila, Hydroptila, Ochrotrichia and Ceraclea, are described and diagnosed herein. Morphological terminology for Rhyacophila and Protoptila follows that of Schmid (1980); Hydroptila and Ochrotrichia that of Marshall (1979); and Ceraclea that of Morse (1975). Specimen length was measured from the tip of the head to the end of the wings. With more than one specimen, this length is given as a range for the species. Type material will be deposited at the National Museum of Natural History, Smithsonian Institution (NMNH), Illinois Natural History Survey (INHS), Florida State Collection of Arthropods (FSCA), University of Alabama Insect Collection (UA), and the personal collection of the author (SCH). Rhyacophila carolae, new species Figs. 1, 2 Diagnosis. This species, a member of the R. invaria species group of Schmid (1970), is most similar to R. kondratiejfi Parker and R. shenandoahensis Flint. These species have in common an elongate anal sclerite with small apical lobes and similarly shaped inferior appendages, but R. carolae is easily recognized by the short, emarginate apical lobe of segment IX and the lack of a concave segment X. As with females of the invaria group, the new species shares the posterodorsal incision and ventral median process of segment VIII and simple vaginal sclerites. With this new species, 1 1 Rhyacophila have been identified from Alabama. Description. Male: Length 7.8 mm. Head and thorax black, legs grayish. Wings black with scattered patches of yellow hair on forewings giving mottled appearance. Abdominal segment VII with small ventromesal projection. Segment IX narrow ventrally, thickened at line of articulation with inferior appendages; dorsal apical lobe short, barely extending beyond segment X, posterior margin with mesal notch. Segment X with posterior margins straight in lateral view; in caudal view rectangular with elongate setae at margins and short stout setae on posterior surface. Anal sclerite elongate in lateral view, narrow basal portion extending into posterior of segment IX, distal portion enlarged with apical lobes small and rounded, triangular laterally; in dorsal view, apical lobes narrowly separated and triangular in shape. Inferior appendages with ventral lobe narrow and rounded apically, dense patch of peg-like setae on dorsal surface; dorsal lobe approximately half length that of ventral lobe. 310 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Figs. 1, 2. Rhyacophila carolae, genitalia. 1. Male genitalia: A, lateral; B, apical lobe of segment IX, dorsal; C, segment X and anal sclerites, caudal; D, segment X and anal sclerites, dorsal; E, phallic apparatus, lateral. 2. Female genitalia: A, segment VIII, lateral; B, segment VIII, dorsal; C, vaginal sclerites, lateral; D, vaginal sclerites, ventral. slightly tapering to rounded apex, bearing several long setae. Phallic apparatus large with spiniform paramere, aedeagus lightly sclerotized and upturned at apex. Female: Length 8.7 mm. Overall appearance similar to male, with forewings less mottled. Abdominal segment VIII in lateral view with a narrow, posteroventral incision; dorsum with broad emargination posteriorly; venter with mesal projection, slightly emarginate on posterior margin. Vaginal sclerites brown; terminal sclerite elongate, in lateral view anterior portion generally rectangular with posterior portion 1989 NEW TRICHOPTERA FROM ALABAMA 311 3A Figs. 3, 4. Protoptila cahabenis, genitalia. 3. Male genitalia: A, lateral; B, sternum VIII, ventral. C, D, E, variation in rods of preanal appendages, lateral. 4. Female genitalia: A, ventral. tapering to an acute apex, in ventral view oblong posteriorly with apical portion rectangular; lateral sclerites about half length of terminal sclerite, in lateral view narrow and rounded posteriorly, anteriorly constricted at attachment to rounded anterior sclerite; in ventral view, lateral sclerites narrowly separated posteriorly, triangular in shape with outer margins concave; anterior sclerite round in ventral and lateral views. Type material. Holotype: Male, Alabama, Lawrence County, tributary to Bee Branch, below falls, T8S, R9W, S26, Bankhead National Forest, 21 May 1988, C. M. and O. S. Flint, Jr. (NMNH). Paratype: Alabama, same as above, 12 (USNM). Etymology. This species is named in honor of Mrs. Carol Flint, who along with her husband collected the type series and has contributed much through her collecting efforts to our knowledge of Trichoptera. Protoptila cahabensis, new species Figs. 3, 4 Diagnosis. In overall appearance, this species resembles both P. palina Ross and P. lega Ross. Males of the new species are separated from both P. palina and P. lega 312 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) by the ventrally curving rods of the preanal appendages and the truncated apex of the phallus. Females of P. cahabensis are separated from P. palina and P. lega by the elongate lateral lobes of sternite VIII and the lack of internal sclerotized plates. In addition to this new species, which is restricted in its distribution to a small section of the Cahaba River, only three other Protoptila have been collected in Alabama: P. maculata (Hagen) in north Alabama, P. georgiana Denning in a portion of the Alabama Piedmont and the widespread P. palina. Description. Male: Length 3.0-4. 5 mm. In alcohol, head and thorax reddish brown dorsally. Abdomen, legs, palpi and antennae yellow brown. Front wings brown with thin white transverse band at midlength, hind wings uniformly light brown. Abdom- inal sternum VIII long, reaching to tip of phallus; in ventral view with deep mesal incision apically. Sternite IX narrow, extending about half length of preceding seg- ment. Tenth tergite divided into lateral arms, each narrowed and beaklike apically. Preanal appendages with long, ventrally curving rods, extending nearly length of segment X, extent of bending variable from slight to abrupt. Phallus with apex nearly truncate, ventral lip rounded, basally bearing a setose lobe and sinuate ventral pro- jection. Female: Length 3. 5-4. 5 mm. Overall appearance and coloration similar to male. Abdominal sternite VIII deeply and broadly incised mesally, lateral lobes elongate. Vaginal apparatus rectangular, sclerotized laterally, lacking mesal sclerotization. Type material. Holotype: Male, Alabama, St. Clair County, Cahaba River at Coun- ty Highway 10, near Whites Chapel, T16S, RIE, S28, 25 August 1981, S.C. Harris (NMNH). Paratypes: Alabama, same as above, but 24 May 1981, 3(56 (SCH); same as above, but 13 August 1981, 1166 (NMNH, INHS, FSCA, UA); same as above, but 9 October 1982, 4666 999 (NMNH, INHS, FSCA, UA, SCH). Etymology. Named for the Cahaba River. Hydroptila micropotamis, new species Fig. 5 Diagnosis. A member of the H. waubesiana group of Marshall (1979), this species resembles H. gunda Milne in the lateral elongation of segment VIII, and both H. Ouachita Holzenthal and Kelley and H. oakmulgeensis Harris in the general config- uration of segment X and the inferior appendages. These characters taken in com- bination render the species distinct. The genus Hydroptila is represented by 47 species in Alabama (Harris, 1986). Description. Male: Length 2. 0-2. 6 mm. Antennae 27-segmented. Brown in alcohol. Abdominal sternum VII with short, apicomesal process. Segment VIII with rounded mesal excisions both dorsally and ventrally; in lateral view narrowing distally to rounded apex. Segment IX short and completely retracted within VIII; dorsally reduced to a narrow bridge. Segment X in dorsal view narrowed basally, forked posteriorly, each arm with rounded apex; in lateral view narrow over entire length, upturned apically and acute. Subgenital plate short; in ventral view generally trian- gular, narrowing posteriorly, bearing two setae at apex. Inferior appendages rectanglar in dorsal and ventral views with mesal incision apically; in lateral view narrow over length, with 3 -pronged apex. Phallus tubular, widest basally, gradually tapering to narrow apex; thin titillator at midlength encircling shaft. Type material. Holotype: Male, Alabama, De Kalb County, Little River at Canyon 1989 NEW TRICHOPTERA FROM ALABAMA 313 Fig. 5. Hydroptila micropotamis, male genitalia. A, lateral; B, dorsal; C, ventral; D, phallus. Park, 4 miles E Dog Town, T8S, R9E, SIO, 22 June 1987, S.C. Harris (NMNH). Paratypes: Alabama, same as above, 1705(3 (NMNH, INHS, FSCA, UA, SCH), De Kalb County, Little River at mouth of Bear Creek, 4 miles E Dog Town, 22 June 1987, 3855, S.C. Harris (SCH). Etymology. From the Greek, little river. Ochrotrichia weoka, new species Fig. 6 Diagnosis. In overall appearance, this species resembles O. xena Ross. It differs primarily in the trapezoidal shaped inferior appendages and the dorsolateral bulbous appearance of segment X. This species brings to 1 1 the total number of Ochrotrichia collected in Alabama (Harris, 1986). Description. Male: Length 2.5 mm. Antennae 27-segmented. Brown in alcohol. Abdominal segment VIII square. Segment IX generally square laterally; dorsum deeply incised; sternum rounded anteriorly, slightly incised posterolaterally. Segment X narrow in lateral view, distally produced into rounded lobe, heavily sclerotized dorsally; in dorsal view parallel-sided basally, narrowing to rounded apex, more 314 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Fig. 6. Ochrotrichia weoka, male genitalia. A, lateral; B, dorsal; C, ventral; D, phallus. heavily sclerotized distally, with heavy peg-like seta subapically. Inferior appendages in lateral view trapezoidal in shape, with peg-like setae dorsally and ventrally at apex; in ventral view peg-like setae distally at mesal margin. Phallus simple, tubular, triangular at apex. Type material. Holotype: Male, Alabama, Elmore County, Fisher Creek on un- marked county road, 3.5 miles southwest Weoka, T20N, RISE, S36, 29 April 1987, S. C. Harris and P. E. O’Neil (NMNH). Etymology. Named for the Alabama town of Weoka, located near the type locality. Ceraclea alabamae, new species Fig. 7 Diagnosis. This species is in the C. fulva group of Morse (1975) with similarities to both C. transversa (Hagen) and C. latahensis (Smith). In common with these species are the long phallic parameres and long mesal spine on the inner surface of the inferior appendage. Ceraclea alabamae is easily distinguished by the acute, up- turned apex of the tenth tergum. Thirteen species of Ceraclea have been identified from Alabama. Description. Male: Length 6. 0-9.0 mm. In alcohol, head, thorax and palps reddish brown. Legs and antennae pale brown. Wings uniformly light brown, with no dis- 1989 NEW TRICHOPTERA FROM ALABAMA 315 Fig. 7. Ceraclea alabamae, male genitalia. A, lateral; B, segments IX and X, dorsal; C, inferior appendage, caudal; D, apex segment X, caudal; E, phallus intact, lateral; F, phallus distended, lateral; G, phallus intact, ventral. cernable pattern. Ninth abdominal segment generally triangular, extending anteriorly to middle of segment VIII. Segment X in dorsal view divided into three lobes distally, lateral lobes with acute, sclerotized dorsal projections, mesal lobe about half length of lateral lobes and rounded apically; in caudal view lateral lobes crescent shaped, with apices bent outward; in lateral view apex of segment X sharply upturned to acute point. Inferior appendages in lateral view with basal projection tipped with long, thick spines, dorsal lobe curving ventrad; harpago extending posteriorly beyond dorsal lobe; in caudal view mesal ridge with long spine, originating from short, truncate, apical projection. Phallus with lateral parameres nearly as long as entire phallic apparatus, extending slightly beyond apex; in ventral view parameres curving mesad at apex. Type material. Holotype: Male, Alabama, De Kalb County, West Fork of Little River at DeSoto State Park, T6S, RlOE, S20, 22 June 1981, S. C. Harris (NMNH). Paratypes: Alabama, same as above, 19$$ (NMNH, INHS, FSCA, UA, SCH), De Kalb County, Little River at Canyon Park, 4 miles E Dog Town, 22 June 1987, 8 156, S. C. Harris (NMNH, INHS, UA), De Kalb County, Little River at mouth of Bear Creek, 4 miles E Dog Town, 22 June 1987, 8755, S. C. Harris (FSCA, UA, SCH). Etymology. Named for its occurrence in Alabama. 316 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) ACKNOWLEDGMENTS The Geological Survey of Alabama provided equipment and facilities during the course of the study and is gratefully acknowledged. The late Dr. Don Denning kindly verified the unique- ness of Pwtoptila cahabensis and helped in the description. Dr. O. S. Flint, Jr. kindly made available for description the new species of Rhyacophila and offered comments on diagnostic characters. K. C. McGilfen of the Illinois Natural History Survey loaned type material of both Pwtoptila lega and P. palina. Dr. J. C. Morse reviewed a preliminary draft of the manuscript and offered useful comments. Also due thanks are P. E. O’Neil for assisting in field collections, Peggy Marsh for typing the manuscript, and Ruth Turner for photographing the plates. This paper represents contribution number 1 1 7 from the Aquatic Biology Program, University of Alabama. LITERATURE CITED Harris, S. C. 1986. Hydroptilidae (Trichoptera) of Alabama with descriptions of three new species. J. Kansas Entomol. Soc. 59:609-619. Marshall, J. E. 1979. A review of the genera of the Hydroptilidae (Trichoptera). Bull. British Mus. (Natur. Hist.) Entomol. 39:135-239. Morse, J. C. 1975. A phylogeny and revision of the caddisfly genus Ceraclea (Trichoptera, Leptoceridae). Contrib. Am. Entomol. Inst. 11:1-97. Schmid, F. 1 970. Le genre Rhyacophila et la famille des Rhyacophilidae (Trichoptera). Mem. Entomol. Soc. Can. 60:1-230. Schmid, F. 1980. Genera des Trichopteres du Canada et des Etats adjacents, pt. 7. In: Les insectes et arachnides du Canada. Agric. Canada, Ottawa, 296 pp. Received January 12, 1989; accepted March 28, 1989. J. New York Entomol. Soc. 97(3):3 17-331, 1989 AN UNUSUAL BLACK FLY (DIPTERA: SIMULIIDAE), REPRESENTING A NEW GENUS AND NEW SPECIES B. V. Peterson Systematic Entomology Laboratory, PSI, Agricultural Research Service, USD A, % National Museum of Natural History, NHB-168, Washington, D.C. 20560 Abstract.— male, female, and pupa of Piezosimulium jeanninae are described and illus- trated. These specimens (consisting of various portions of 2 males, 1 female, and associated pupal parts) represent a new genus and new species in the family Simuliidae. The new genus is readily distinguished from all other genera of the family by the presence of a conspicuous sperm pump, and a distinct, sclerotized, setose plate situated ventrally between the bases of the male gonocoxites. This is the first record of such structures in the family Simuliidae. The sclerotized plate is of unknown homology, but it might be a remnant of stemite 9 (hypandrium). A combination of several other characters, especially the slender katepisternum, the erect to semi-erect pile on the scutum, the large calypter, and the small eyes of both sexes are also characteristic. Under the present family classification this genus is a member of the tribe Prosimuliini, and is possibly the sister group to the other taxa in this assemblage, and possibly the sister group to all known black flies. Specimens of the following taxon have been known since early 1983, when they were received for identification along with a series of other species from Mr. Bruce Wahle, at the time a graduate student at the University of Colorado. The male was immediately recognized as undescribed but was put aside for later description with a number of other new species. At that time, I had not removed the existing portions of the fully developed male from the pupal skin, and it was only some time later, as I was looking more closely at the species, that I saw the very distinct sperm pump within the male abdomen. Subsequent attempts by the author, and others from the Institute of Arctic and Alpine Research at the University of Colorado, failed to collect additional specimens of this unique species. In view of the fact that neither a sperm pump nor a prominent sclerotized plate in the terminalia, as described below, have been reported for the family Simuliidae, and because additional specimens of this new genus and species might not be collected again for many years, as happened with Parasimulium Malloch, it is deemed worthwhile to describe this new taxon on the basis of the material at hand. Moreover, 2 major works have recently been completed dealing with various aspects of the phylogeny of the family Simuliidae (Currie, 1988, Ph.D. thesis; Wood and Borkent, in press), and it seems expedient to publish this information as it should shed new light on the phylogeny of the family and its relationships with other members of the Culicomorpha, especially the Chi- ronomoidea. It must be recognized that if, or when, additional specimens are col- lected, these descriptions will require modification and additions. Piezosimulium, new genus Type species: Piezosimulium jeanninae Peterson, by present designation. Characters of the genus. Antenna with 9 flagellomeres. Occipital foramen larger 318 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) than normal with combined hypostomal bridge and postgenal areas distinctly longer than dorsomedial occipital sclerite. Wing veins with hne setae only; Rs with a long, somewhat obscure fork; base of R setose; calypter subquadrate, somewhat larger than in other black flies. Calcipala and pedisulcus absent. Scutum with moderately long, erect to semi-erect pile, no recumbent setae. Scutellum depressed and flattened, nearly rectangular, slightly pointed posteromedially. Katepisternum moderately slender and narrowly rounded ventrally (but not as slender or narrowly rounded as in Parasi- mulium), with a slight depression but without a distinct sulcus or groove separating upper and lower portions. Male: Eyes small with resulting broadened orbits and vertex; facets of eye gradually increasing in size dorsally without a sharp line of demarcation between smaller lower facets and larger upper facets; number of larger, upper facets reduced as compared to male eyes of other black flies. Eyes narrowly but distinctly separated along midline, at medioventral angle by at least width of an eye facet, and at mediodorsal angle by width of about 3 or 4 large, dorsal facets; frontal area between eyes with sparse, short setae; inner margin of eye at level of antenna with a narrow, facetless, crescent-shaped, thickened margin (=ocular triangle or notch) (Figs. 1 , 2). Body of ventral plate of aedeagus, in terminal view, somewhat W-shaped while, in ventral view, it is more M-shaped, with 2 sublateral, ventrally directed liplike lobes separated by a deep median groove. Median sclerite of aedeagus broadly trough-like, distal arms barely indicated, gonopore large. Gonocoxites each with a prominent ventromedial lobe that articulates with a prominent, sclerotized, setose plate that is situated anteriorly between these lobes (Figs. 3-5). A moderately large, well-developed sperm pump present with internal spicules, a short but distinct ejaculatory apodeme, and a long sperm duct (Figs. 6-8). Female: Eyes small, widely separated, accompanied by broader than usual orbits, frons, and vertex; vertex slightly raised ridgelike, and, in front view, appearing broadly peaked dorsomedially; in dorsal and lateral views, eyes bulging out from head more than in most other species; ocular triangle present but small, a second small, facetless area present on inner dorsal angle of eye. Each claw with a tiny, erect, subbasal tooth. Pupa (only pieces available): Respiratory organ (gill) with 14 filaments. Larva unknown. Etymology. The generic name is derived from piezo, Gr., meaning to squeeze, in combination with Simulium, and refers to the presence of a ‘squeezing’ organ or sperm pump in the male. Gender, neuter. Piezosimulium jeanninae, new species Figs. 1-15 Description. MALE (preserved in alcohol): General body color blackish brown. Head (Figs. 1 , 2) relatively small, with a slightly darkened spot along posterolateral margin of eye which might be a vestige of a stemmatic bulla. Clypeus small but strongly convex and protrudent well beyond front margin of eye, greatest width and length subequal. Occipital condyle small but conspicuous, strongly sclerotized. Oc- ciput with dense, long, erect, black setae. Antenna yellowish; pedicel and first flagello- mere subequal in length but pedicel slightly wider; fine pubescence pale yellowish, first flagellomere with a series of short, black, stout setae, these present also on ventral surface of other flagellomeres. Palpus yellowish, with black setae; palpomere 5 about % longer than palpomere 3. Sensory vesicle small, with a small, round mouth. Antepronotum well developed, with moderately large lateral lobes connected by 1989 NEW GENUS OF SIMULIIDAE 319 Figs. 1, 2. Piezosimulium jeanninae. Male. 1. Head, lateral view. 2. Head, front view. 320 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) a broader than usual dorsomedial straplike portion; postpronotum somewhat re- duced, subtriangular in shape; anteromedial portion of scutum broadly rounded and strongly arched; all preceding thoracic sclerites densely covered with short to mod- erately long, erect black setae, apparently without recumbent setae. Scutellum yel- lowish, brownish along margins, densely covered with long, whitish setae, most of which have dark bases. Postnotum somewhat narrowed and strongly convex. Anepi- sternal membrane broader dorsally, slightly tapered ventrally so it is roughly wedge- shaped; mesepimeral tuft dark, extending ventrally more than Vi height of posterior portion of mesepimeron. Wing membrane hyaline but with a faint grayish tinge; veins pale grayish; base of costa with predominantly pale whitish setae but some setae with black bases and some entirely black; Sc with long setae ventrally; stem vein with whitish setae having dark bases; R, and R4+5 setose both dorsally and ventrally, R2+3 bare dorsally, setose ventrally near tip; MA darkly pigmented; setae on other veins dark; fringe of calypter and anal lobe whitish. Halter shorter than in most other species; knob white, stem brownish yellow, its outer edge with a basal tuft of short, pale setae which is contin- uous with a row of similar setae extending dorsally to base of knob; at both ends of this series, setae are in double rows while medially they are in a single row; inner edge of stem with 2 setae at base, and with a single row of setae beginning at base of knob and extending nearly to top of knob, inner surface of knob with a patchlike series of minute setae. Legs in poor condition, but, in teneral state, appearing to be more yellow than body, rather slender and delicate; hind basitarsus about 3 times as long as broad. Claw long, slender, and slightly curved near tip, with a tiny basal tooth; dorsobasal toothlike process crescent shaped, about as broad as claw and about Vi as long as claw; empodium relatively long, slender, mostly yellowish, tip darkened and plumose. Abdomen brownish black; basal scale (tergite 1) narrow, not as well developed as in other black flies, reduced dorsomedially to a narrow, straplike strip connecting somewhat broader lateral portions, at narrowest with only 1 or 2 rows of moderately long, dark setae, lateral portions more setose; remaining tergites all broad, sclerotized, with pale hind margins, and rather densely covered with moderately long, dark setae; tergite 10 small, subquadrate, smaller than cercus. Stemite 1 small, crescent shaped, lightly sclerotized; sternite 2 rectangular, weakly sclerotized, and bearing a patch of moderately long, dark setae (in other species sternite 2 is usually membranous and often bare); sternites 3-7 moderately sclerotized, densely covered with long, dark setae; sternite 8 more densely sclerotized, nearly bare except for about 1 8 short, dark setae posteriorly. Pleural membrane of segments 4-8 with a faint, but distinct, sha- greened area consisting of numerous, closely placed, minute granules; spiracles of segments 4-7 placed in or next to these areas; pleural membrane of each segment sparsely setose. Terminalia as in Figures 3-9. Stemite 9 a slender, heavily sclerotized rodlike structure that is continuous dorsolaterally with tergite 9; slightly broader midventrally and with a short but distinct, irregularly pointed process on each side ventrolaterally (Fig. 5). Gonocoxite (Figs. 5, 6) stout, slightly wider than greatest length, with a distinct constriction, or depression, at about apical % and sparsely covered with pile distal to this constriction; inner proximal corner produced as a short but prominent, stout, sclerotized process that apparently articulates medially with a large, somewhat 1989 NEW GENUS OF SIMULIIDAE 321 Figs. 3-5. Piezosimulium jeanninae. Male. 3. Sclerotized, setose plate, ventrolateral view. 4. Terminalia, lateral view. 5. Terminalia, ventral view. Abbreviations: aed memb, aedeagal membrane; cere, cercus; goncx, gonocoxite; gonpr, gonopore; gonst, gonostylus; m scl, median sclerite of aedeagus; pm, paramere; st 9, stemite 9 (hypandrium); tg 10, tergite 10; vmlb, ventromedial lobe of gonocoxite; v pit, ventral plate of aedeagus. 322 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Figs. 6-9. Piezosimulium jeanninae. Male. 6. Abdomen, ventral view, showing position of sperm pump in relation to abdominal segments and terminalia. 7. Sperm pump, dorsal view (?). 8. Sperm pump, ventral view (?) (it is uncertain which is true dorsal and true ventral because 1989 NEW GENUS OF SIMULIIDAE 323 ovate, cuplike sternite (possibly a remnant of stemite 9 (hypandrium)); this stemite (Figs. 3-5) with anterior margin heavily sclerotized and bearing a short, submedian apodemelike process, ventral surface bearing a patch of long setae, posterior margin narrowly but heavily sclerotized and situated close to anteroventral surface of ventral plate of aedeagus (Figs. 4, 5). Gonostylus (Figs. 5, 9) short, stout, a curved triangular shape, slightly but distinctly longer than greatest width at base; pointed apically with 1 small, apical, and 1 subapical spine; moderately setose. Ventral plate of aedeagus (see generic description) (Figs. 4, 5) broad; in ventral view, M-shaped, widest at junction with basal arms, tapering distally, distal margin bilobed with a deep con- cavity between lobes, these lobes densely setose, proximal margin of ventral plate convex; basal arms short, slender and straight; in lateral view, with a deep ventral lip that is about % as deep as ventral plate is long, arm wide (Fig. 4). Stem of median sclerite trough-like, broader than medial groove of ventral plate, rather heavily scler- otized, distal arms essentially undeveloped, gonopore broad (Figs. 4, 5). Plate of endoparameral organ subtriangular, moderately sclerotized, connecting to tip of arm of ventral plate by a long, slender, heavily sclerotized rod, and to base of gonocoxite by a short, broad arm (Figs. 4, 5). Sperm pump (Figs. 6-8) lightly sclerotized but conspicuous, moderately large, about Vi as wide as abdominal segment, floating free in abdominal cavity in, or somewhere near, segment 6; bilobed with each lobe again curved, twisted or folded so it in turn is somewhat bilobed; 1 main lobe with a short but well-developed and heavily sclerotized ejaculatory apodeme on anterior border, area surrounding ejaculatory apodeme distinctly brownish; other lobe with a some- what T-shaped sclerotized thickening near where sperm duct emerges; a sclerotized, tubelike process, with internal circular thickenings, also present which seems to join 2 main lobes together internally (this not clearly discernible in specimen at hand); inner surface of sperm pump with numerous moderately long spicules (or setae), and with 2 relatively long (but shorter than spicules), stout, dark spines on inner anterior surface of lobe not bearing ejaculatory apodeme, this portion of lobe distinctly brown- ish. FEMALE (preserved in alcohol): General body color blackish brown; wing crum- pled; most of abdomen missing. Head (Figs. 10, 13) relatively small; frons at vertex about V4 wider than at narrowest point, and nearly Vi width of head; about twice as wide as long; ventral area of frons, just dorsal to antennae, produced as 2 small but distinct, subshining humps, or calli, separated by a depressed area along midline; this midline bare, remainder of frons densely covered with moderately long, erect to semi-erect, black pile. Clypeus small, strongly convex; slightly wider than long; mod- erately covered with moderately long, black setae; in lateral view, both lower margin of frons (humps or calli) and clypeus projecting well beyond anterior margin of eye. Postcranium with combined ventral hypostomal bridge and postgenal areas about % longer than dorsomedial occipital sclerite. Occiput densely covered with long, black pile; usual long, stiff, postocular setae absent or undifferentiated. Antenna with scape the abdominal muscles were almost completely deteriorated and the sperm pump was free to move around inside the abdomen). 9. Right goxocoxite and gonostylus, dorsal (inside, upper) view. 324 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Figs. 10-14. Piezosimulium jeanninae Peterson. Female. 10. Head, lateral view (antennae missing). 1 1. Tarsal segments of hind leg. 12. Claw, lateral view. 13. Head, front view (antennae missing). 14. Distal 3 palpomeres. Abbreviations; clw, claw; sb tth, subbasal tooth; sen ves, sensory vesicle of third palpomere. 1989 NEW GENUS OF SIMULIIDAE 325 and pedicel brown, slightly darker than flagellum which is more yellowish brown; pedicel and first flagellomere subequal in length and width; fine pubescence pale yellow, longer setae dark. Mandible and blade of maxilla present but weakly scler- otized; mandible with, at most, faint marginal undulations and a few very minute, irregular, serrations apically; blade of maxilla without retrorse teeth but may have 2 or 3 minute, setalike projections apically. Palpus short, pale yellowish, distal 2 pal- pomeres slightly lighter than palpomere 3; with pale and black setae; palpomere 5 slightly longer than 3, and about A longer than palpomere 4. Sensory vesicle small, slightly less than V3 as long as its segment, rather centrally situated, its neck short, arising anterodorsally and extending vertically, with an enlarged round mouth. Ci- barium weakly developed, median proximal space deep, broadly U-shaped, dorso- lateral arms long, slender, only moderately sclerotized, nearly as long as remainder of cibarium. Antepronotal lobes basically as in male. Postpronotum small, distinctly paler than scutum. Anteromedial portion of scutum broadly rounded, slightly elevated ridgelike. All 3 of these thoracic sclerites densely covered with short to moderately long, erect to semi-erect, black setae, apparently without recumbent setae. Scutellum and post- notum essentially as in male. Anepisternal membrane rather broadly rectangular, pale whitish; mesepimeral tuft dark. Precoxal bridge present but weakened at points of attachment with proepistemum. Katepisternum, in lateral view, with anterior margin nearly straight, lower portion without the usual lobelike appearance; in ventral view, ventral margin only weakly bilobed. Wing in poor condition, presumably as in male. Halter short, knob white, stem pale brownish yellow, with short, pale setae. Legs in poor condition but all appearing to have a similar color pattern as follows: coxae and trochanters brown, those of forelegs each with a yellowish patch (this might darken with time); femora yellowish, narrowly brown distally; tibiae yellowish but with both bases and apices brown; basal tarsal segment of each leg yellowish on basal half and brown distally, remaining tarsal segments yellowish brown; all leg segments with dark setae. Claw short, weak, evenly curving from base, with a variably devel- oped but small, subbasal tooth that is difficult to see except under high magnification; empodium as in male. Basal scale (tergite 1) of abdomen narrow, yellowish, with a fringe of long pale yellow pile; remainder of abdomen missing. PUPA: Respiratory organ (Fig. 1 5) relatively short, consisting of a short, slender base densely covered with minute granules, and with 14 filaments arranged in 3 groups all on short petioles: a dorsal group of 6 filaments in 3 pairs, a mediolateral group of 4 filaments in 2 pairs, and a ventral group either in 2 pairs (2 + 2) or branching 3 + 1 (d-v); filaments brownish gray basally and covered with minute granules, these granules rapidly decreasing in number and filaments becoming paler whitish distally. Head smooth, without pattern or granules; antenna of male reaching nearly to hind margin of head; antenna of female reaching hind margin of head; 2 sublateral setae present on each side near ventral margin of clypeus. Dorsum of thorax smooth, with only a faint, superficial, reticulate pattern; 2 long, simple, dorsal trichomes present on each side of thorax. Abdominal tergites 1 and 2 dorsally on each half with 5-7 short, fine setae, these usually bent or sinuous. Cocoon a thick, loosely woven, shapeless sleeve that covers most of pupa. 326 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Fig. 15. Piezosimulium jeanninae Peterson. Pupa. 15. Respiratory organ (gill), right lateral view. Holotype. Male (in fragmentary condition but abdomen well preserved), with por- tions of associated pupal pelt. North Boulder Creek, Boulder County, Colorado, elevation 3,600 m ( 1 1 ,8 1 1 ft), 11 September, 1981, John Bushnell (code 7). Deposited in the National Museum of Natural History, Washington, D.C. Paratypes. Head and thorax of 16, 12, both removed from pupal skins (both in poor condition), same data and depository as for holotype. Etymology. This species is named for, and dedicated to, my wife, Jeannine, who has been of inestimable help to me for nearly forty years. Biological notes. The type locality is a small stream originating from the glacier on Niwot Ridge, near the Continental Divide, at an elevation of about 3,600 m (1 1,81 1 feet), in the controlled access area of the watershed from which the city of Boulder gets its drinking water. This stream continues in a variable path descending from its origin through a series of alpine lakes, named Green Lakes 2 through 5, and Albion Lake. The water temperature of the stream remains fairly constant in any given locality varying only from about l°C near its origin to about 10°C near Albion Lake. It passes over a rocky bed with some moss and some trailing grasses, and through shifting muddy areas on its way down the slopes. Elgmork and Saether (1970) give an excellent description of North Boulder Creek and its environs. Other species 1989 NEW GENUS OF SIMULIIDAE 327 found in this stream include an undescribed species of Prosimulium Roubaud (near P. neomacropyga Peterson; reported as P. ursinum Edwards by Elgmork and Saether (1970), and Saether (1970)), P. travisi Stone (almost certainly the species reported as P. esselbaughi Sommerman by Elgmork and Saether (1970) and Saether (1970)), and a new species of Metacnephia Crosskey. The specimens of Piezosimulium jeanninae were taken in association with speci- mens of the undescribed species of Prosimulium mentioned above and were not separated at first because both have 14 filaments in the respiratory organ (gill). However, the respiratory organs of the two species can be distinguished by the different textures of their component filaments and their overall shapes (see fig. 75, in Peterson, 1970). Specimens of the new species of Metacnephia, also taken in the same collection, were fewer in number than the new Prosimulium species but were more numerous than the specimens of P. travisi collected. However, subsequent collections resulted in large numbers of all stages of the new Metacnephia species. Remarks. These descriptions are based, as noted previously, on fragmentary adult material salvaged from within pupal skins. Some of the fragments were in good condition, especially the abdomen and terminalia of the holotype male, and could be seen clearly within the pupal skins. At the same time, most portions of the specimens were quite fragile and in poor condition, especially since the specimens were partially decayed at the time they were collected. Handling and preparing this material for description and illustration also caused some damage to the specimens. Piezosimulium belongs to the Prosimuliini as defined by Crosskey (1969), and as outlined in his recent (1987) checklist of world black flies. It would also be included in this taxon within the more restricted definition of the tribe given by Currie (1988). The features that are most characteristic and immediately distinguish males of this monotypic genus from all other known black flies are the conspicuous baglike struc- ture that is here considered to be a sperm pump, and the distinct sclerotized, setose plate situated ventrally between the bases of the gonocoxites. This is the first record of such structures in the family Simuliidae, and I am not aware of any such sclerotized plate in any other dipteran. Piezosimulium also is readily distinguished from the other genera of Prosimuliini (and from all other simuliid genera) by a combination of several characters, but especially by the slender katepistemum (which is somewhat intermediate in development between that of the very reduced state found in Parasi- mulium (Peterson 1981), and the more usual development found in other genera such as Prosimulium, and the more apomorphic Simulium Latreille); the erect to semi-erect pile on the scutum (apparently without short recumbent setae) like in Gymnopais Stone, Crozetia Davies, and a few others; and the small and bulging eyes of both sexes (this character is also found in some other genera but the pattern of the large and small facets of the male eye is quite different from other known groups, and none seem to have eyes that so noticeably bulge from the head yet with the clypeus and lower frontal calli that project well beyond the anterior margins of the eyes). Other features such as the larger quadrate calypter; the enlarged occipital foramen with the accompanying reduction of the dorsomedial occipital sclerite, and the con- comitant longer hypostomal bridge and postgenal areas in both sexes; and the narrow, strongly arched condition of both the clypeus and postnotum in both sexes, probably are diagnostic in combination. However, it is impossible to evaluate the range of 328 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Table 1. Distribution of some Character States in Simuliidae. Character state Piezosimulium Prosimuliini Parasimulium Other Simuliidae 1 + - ± — (head distinctive) 2 + + - + 3 + - - - 4 + - ± - 5 + ± - - 6 + - + - 7 + - + - 8 + - - - 9 + + 7 + 10 + - - - 11 + - - - Character states in Character states in other black Piezosimulium jeanninae ( + ) fly genera (-) Head 1 . $ eyes small, narrowly separated dorsal- ly along medial margins; upper omma- tidia gradually enlarging dorsally with- out a sharp line of demarcation separating them from smaller, lower ommatidia. 2. 9 eyes small and bulging. 3. Clypeus small, strongly convex. 4. Occipital foramen enlarged with accom- panying reduction of median occipital sclerite, and longer hypostomal bridge and postgenal areas. Thorax 5. Scutum dorsally with only erect to semi-erect setae (apparently without short, recumbent setae). 6. Postnotum narrow, strongly convex. 7. Katepisternum moderately slender, nar- rowly rounded ventrally, with a slight depression but without a distinct sulcus or groove separating upper and lower portions. 8. Calypter enlarged, more quadrate. Abdomen 9. $ abdominal sternite 2 sclerotized and setose. 10. Sperm pump present. 1 1 . Setose, sclerotized plate present antero- ventrally between bases of gonocoxites. 6 eyes large, contiguous dorsally along medi- al margins; upper ommatidia abruptly en- larging dorsally and separated from small- er lower ommatidia by a distinct line of demarcation. 9 eyes usually large and not bulging. Clypeus larger, more flattened (less convex). Occipital foramen not enlarged, median oc- cipital sclerite not reduced, and hyposto- mal bridge and postgenal areas not elongate. Scutum dorsally without erect setae, but with abundant short, recumbent setae. Postnotum broader, more flattened (less strongly convex). Katepisternum broader and broadly rounded ventrally, with a distinct sulcus or groove separating upper and lower portions. Calypter smaller, more rectangular. (5 abdominal sternite 2 usually membranous, and bare or setose. Sperm pump absent. Without a setose, sclerotized plate antero- ventrally between bases of gonocoxites. 1989 NEW GENUS OF SIMULIIDAE 329 variability, and the integrity of such morphological characters on the basis of the material at hand. Depending on how the above mentioned characters are regarded, one could suggest that Piezosimulium really is the sister group of all other Simuliidae and should be placed in a separate subfamily; or that it is an aberrant, highly derived entity belonging to the Prosimuliini but on an evolutionary dead end. Another possibility is that it represents an oddity (of a type probably not to be encountered again) due to some miscued developmental factors. Neither of the hrst two possibilities can be unequiv- ocally proclaimed until the character states of the adults and pupa are more fully known, and the larva is discovered and studied. The remarkably well-developed features of the male terminalia strongly argues against these features being artifacts or malformations resulting from the presence of mermithid nematode parasites or due to some other unknown factors that might cause unnatural development. It is difficult to conceive of a well-developed ejaculatory apodeme, albeit small, arising as the result of the presence of some parasite or miscued developmental factors especially since the presence of a sperm pump and ejaculatory apodeme has not been reported before in the Simuliidae. The same can be said about the presence of the setose, sclerotized plate situated between the bases of the gonocoxites, with its short but well-developed apodeme. None of the specimens of the associated species in this collection were parasitized or showed any evidence of parasites of any kind whose presence might have initiated the formation of such structures. It is equally difficult to conceive of the possibility that the baglike sperm pump is really some expression of 1 or more malformed spermathecae even though it possesses internal spicules (and 2 minute spinules) similar to those found in the spermathecae of some species. Again, it is difficult to conceive of the present specimens representing some type of sexual mosaic especially when such features as the sperm pump, ejaculatory apodeme, and the unique sclerotized plate have been unknown in the family. Some form of sperm pump is found in most Mecoptera, all Siphonaptera, and is present in many of the families of lower Diptera that are widely recognized by dipterists as being among the most primitive of the living Diptera (see Downes, 1968). Downes (1968) also mentioned that the process of sperm transfer by sperm pump occurs in a rather precise, closed system as a fairly widespread primitive feature, and that the process usually cannot be completed very quickly. This is consistent with what can be surmised of P. jeanninae. In both sexes the eyes are greatly reduced which suggests that mating takes place on the ground over a period of time rather than in flights of short duration as occurs in the majority of other black flies. The high altitude habitat, with its relatively short summer season, cooler temperatures and more inclement weather patterns, is less conducive to mating during flight than it is in the more favorable conditions found in less severe habitats. The small eyes and the narrow postnotum suggest this species is not a strong flyer. Also, as the female is nonblood-feeding, I suspect that if this species flies at all it is only for short distances at any one time. The majority of other black flies are strong flyers and transfer sperm by means of a double-chambered spermatophore, usually, but not always, during a short mating flight. For such species at least initial coupling takes place in flight and mating may or may not be completed while still in the air. Downes (1968) called this process an open system that generally is faster and less precise than in the closed system by sperm pump. So far as is known, at least some representatives 330 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) of the other three families now included in the Chironomoidea, i.e., the Thaumaleidae (Downes, 1968:613; McAlpine, 1981), Ceratopogonidae, and Chironomidae, effect sperm transfer by a gelatinous, double-chambered spermatophore, but not by a sperm pump. The unique sclerotized plate situated between the anteroventral (medial) comers of the gonocoxites, and anteroventral to the base of the ventral plate of the aedeagus, appears to be a relic (not just an artifact) possibly from stemite 9 (the hypandrium). This sclerite bears a distinct patch of moderately long setae ventrally and has a short, but distinct, submedian apodeme on its anterior margin which probably is a point of articulation with the main body of sternite 9. Additional specimens of P.jeanninae will be necessary for study before any homology of this structure can be resolved. As far as I am aware, none of the other three families presently constituting the superfamily Chironomoidea, has any structure that even vaguely resembles the scler- otized plate found in P. jeanninae, and, in fact, nothing similar is present in any other Diptera known to me. Table 1 summarizes the character states mentioned above and their distributions generally within the family. It is not intended for concluding phylogenetic relation- ships; such conclusions would only be speculative until the larva of this species is discovered and studied and the adults and pupa are more completely known. ACKNOWLEDGMENTS I am grateful to B. Wahle, Longmont, Colorado, and J. Bushnell, Department of E.P.O. Biology, University of Colorado, Boulder, Colorado for providing the specimens on which this paper is based. I am also grateful to the following individuals from the Institute of Arctic and Alpine Research (INSTAAR), University of Colorado, Boulder, Colorado: J. Halfpenny who arranged my initial transportation into the area, and T. Mihuc who accompanied me into the watershed on my first trip; M. Losleban who provided the necessary permits to enter and collect in the restricted area of the Boulder City Watershed and provided transportation for subsequent trips. I am also grateful to S. K. Wu, Zoology Section, University of Colorado Museum, Boulder, Colorado, who accompanied me on my second trip into the area. I am most grateful to Linda H. Lawrence, staff artist. Systematic Entomology Laboratory, who painstakingly prepared the accompanying illustrations. I thank W. N. Mathis, Department of Entomology, Smithsonian Institution, Washington, D.C.; T. Pape, Zoological Museum, Copenhagen, Denmark; M. F. Mickevich, Maryland Center for Systematic Entomology, University of Maryland, College Park, Maryland; N. E. Woodley and M. E. Schauff, Systematic Entomology Laboratory, Washington, D.C., and two unknown reviewers who read and commented on the manuscript. LITERATURE CITED Crosskey, R. W. 1 969. A re-classification of the Simuliidae (Diptera) of Africa and its islands. Bull. Brit. Mus. (Nat. Hist.), Entomol. Suppl. 14:1-195; 1 pi. Crosskey, R. W. 1 987. 32 An annotated checklist of the world black flies (Diptera: Simuliidae). Pages 425-520 In: K. C. Kim and R. W. Merritt (eds.). Black Flies. Ecology, Population Management, and Annotated World List. Pennsylvania State Univ., University Park and London. Currie, D. C. 1988. Morphology and systematics of primitive Simuliidae (Diptera: Culico- morpha). Unpublished Ph.D. Thesis, University of Alberta. Downes, J. A. 1 968. Notes on the organs and processes of sperm-transfer in the lower Diptera. Can. Entomol. 100:608-617. 1989 NEW GENUS OF SIMULIIDAE 331 Elgmork, K. and O. A. Saether. 1 970. Distribution of invertebrates in a high mountain brook in the Colorado Rocky Mountains. Univ. Colo. Studies, Ser. Biol. 31:1-55. McAlpine, J. F. 1981. Chapter 2, Morphology and terminology-adults. In: J. F. McAlpine, B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, and D. M. Wood (eds.). Manual of Nearctic Diptera, Vol. 1. Res. Br., Agr. Can. Monogr. 27:9-63. Peterson, B. V. 1970. The Pwsimulium of Canada and Alaska (Diptera: Simuliidae). Mem. Entomol. Soc. Can. 69:1-216. Peterson, B. V. 1981. Chapter 27, Simuliidae. In: J. F. McAlpine, B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, and D. M. Wood (eds.). Manual of Nearctic Diptera, Vol. 1. Res. Br., Agr. Can. Monogr. 27:355-391. Saether, O. A. 1970. Chironomids and other invertebrates from North Boulder Creek, Col- orado. Univ. Colo. Studies, Ser. Biol. 31:57-114. Wood, D. M. and A. Borkent. Chapter 1 14, Phylogeny and classification of the Nematocera. In: J. F. McAlpine, B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, and D. M. Wood (eds.). Manual of Nearctic Diptera, Vol. 3. Res. Br., Agr. Can. Monogr. (in press). Received November 23, 1988; accepted March 14, 1989. J. New YorkEntomol. Soc. 97(3):332-340, 1989 BUTTERFLY EXPLOITATION OF AN ANT-PLANT MUTUALISM: ADDING INSULT TO HERBIVORY P. J. DeVries’ and I. Baker^ ’Department of Zoology, University of Texas, Austin, Texas 78712, and Smithsonian Tropical Research Institute, Box 2072, Balboa, Panama ^Department of Botany, University of California, Berkeley, California 94720 Abstract. — The myrmecophilous butterfly caterpillar of Thisbe irenea is shown to gain growth benefits from not only feeding on leaf tissue, but by also drinking the extrafloral nectar of its hostplant. Since both the plant and caterpillar use ants as defenses, it is suggested that a conflict is generated between plant and herbivore for the attentions of ants, and that such conflicts may be widespread in ant-plant, and ant-herbivore systems. It is further suggested that this study points to the possibility that in such systems, 2-species mutualisms may be susceptible to invasion and exploitation by a third species. It is well documented that both plants and insect herbivores may form mutualisms with ants. Ants provide plants with protection against herbivores (see reviews in Buckley, 1983a; Beattie, 1985; Koptur, 1984), and ants provide insects with benefits that include protection against predators and parasitoids, faster growth rates and higher reproductive success (Banks and Nixon, 1958; Bartlett, 1961; Bristow, 1984; Cottrell, 1984; DeVries, 1987; Pierce et ah, 1987). In these mutualisms insects provide ants with secretions directly through specialized organs (Mittler, 1958; Way, 1963; Cottrell, 1984; Fiedler and Maschwitz, 1988; DeVries, 1988; Wilson, 1971), whereas plants may either provide secretions directly through extrafloral nectaries (Beattie, 1985), or may attract ants indirectly via honey-dew secreting Homoptera (Messina, 1981). Among butterflies the habit of associating with ants, or myrmecophily, is best known from the Lycaenidae whose larvae have specialized ant-organs for associating with ants (Cottrell, 1984). Larval ant-organs are so widespread within the Lycaenidae that myrmecophily is thought to have played an important role in lycaenid evolution (Hinton, 1951; Vane-Wright, 1978; Pierce, 1984). Some species in the Riodinidae also have larvae that associate with ants and possess ant-organs that are analogous, but not homologous, to those found on lycaenids (Cottrell, 1984; DeVries, 1988). Because they are considered to share a close relationship to the lycaenids (Ehrlich, 1958; Kristensen, 1976; Harvey, 1987; but see Robbins, 1988), assumptions about the evolution of myrmecophily in riodinids are based primarily on studies of lycaenids (Pierce, 1987). However, the biology of ant association in most riodinid species is unknown (see Ross, 1966; Callaghan, 1986; DeVries, 1987). This paper presents observations and experiments done that probe hostplant use and ant association in the myrmecophilous riodinid butterfly Thisbe irenea (Stoll), and extends these observations to other ant-insect systems. The purpose of this paper is to show that some ant-associated caterpillars not only feed on plant tissues but also feed from extrafloral nectaries on the hostplant, thus exploiting the basis of the mutualism between plants and ants. We suggest that when plants and caterpillars 1989 BUTTERFLIES AND ANT-PLANT MUTUALISM 333 both have mutualistic associations with the same species of ants, a conflict is generated between plant and herbivore for the attentions of ant mutualists, and that such conflicts may be widespread in many ant-plant, and ant-herbivore systems. Although traditionally ant-plant mutualisms and ant-insect mutualisms have been considered separately, this study points to the possibility that 2-species mutualisms may be susceptible to invasion and exploitation by a third species. MATERIALS AND METHODS From September 1985 to September 1986, and intermittently from August 1987- October 1988, one of us (PJD) studied the wide ranging riodinid butterfly T. irenea on Barro Colorado Island, Panama, and on surrounding mainland habitats. Here T. irenea caterpillars feed only on saplings and seedlings of the euphorbiaceous pioneer tree Croton billbergianus (Robbins and Aiello, 1982; DeVries, 1987). The vegetative surfaces of the hostplant are patrolled by ants attracted to EFN’s located at the base of each leaf, and these ants tend T. irenea caterpillars. All five larval stages feed on developing leaves, but only fourth and fifth instar caterpillars can eat all types of leaves; earlier instars must feed on new developing leaves. Infestations of T. irenea caterpillars commonly remove from 18-38% of the total leaf area of small C. bill- bergianus plants, occasionally killing them (DeVries, unpublished). Upon reaching third instar, specialized ant-organs become functional that allow caterpillars to attract and maintain the presence of ants (DeVries, 1988). The major ant species tending both T. irenea and the EFN’s of C. billbergianus at the study site was Ectatomma ruidum (Formicidae: Ponerinae), and these ants protect larvae from predators in exchange for secretions provided by ant-organs (DeVries, 1987). From weekly and bi-monthly censuses of marked C. billbergianus plants, and observations on potted plants in an ambient temperature laboratory, it was estab- lished that T. irenea caterpillars of all instars rest with their heads on or immediately adjacent to EFN’s of the hostplant (Fig. 1). First through third instar caterpillars spend most of their time, day and night, on or near EFN’s, whereas fourth and fifth instars hide on the stem during the day and crawl up on the leaves to feed at night. The habit was observed so regularly as to suggest that caterpillars were drinking extrafloral nectar. The following experiments were conducted to determine whether the presence of EFN’s and ants affected larval growth. Twelve potted hostplants (paired by size, leaf number and leaf maturity) were ringed near their bases with Tanglefoot (Trade mark) to eliminate access to the foliage by crawling insects. One half of the plants had the EFN’s excised, the other half did not. Pairs of plants were placed on either side of six captive colonies of E. ruidum ants maintained in plastic tubs. Ants from the colonies were allowed access to the plants by placing wooden bridges above the Tanglefoot into the plastic tub. Each plant received one larva of T. irenea; all larvae were the same instar and weight. Every 48 hours during the following 12 days caterpillars were weighed to the nearest milligram. A simultaneous experiment was set up in the same manner, except that larvae were grown without allowing ants to tend them. All experiments were done in an ambient temperature laboratory. To examine if larvae drank nectar, and if nectar quality had an effect on growth, larvae were grown using experimental nectars without ants. Experimental nectaries were made from capillary tubes sealed into the sides of Petri dishes (DeVries, 1987). 334 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) First Second Third Fourth Fifth LARVAL INSTARS Fig. 1. Summary of 1,378 diurnal observations of Thisbe irenea caterpillars taken from diurnal censuses. NEC = caterpillars found with head on or immediately next to an EFN; LEAF = caterpillars found on leaf tissue; STEM = caterpillars found resting on woody plant parts away from leaves. Late fourth and all fifth instar caterpillars rest on the stem during the day, but are active at night when they are frequently found with their heads on or near EFN’s. Nectars were made by adding 5 drops of red liquid food coloring to 400 ml of distilled water. They were then mixed well, and fractioned into two containers. To one con- tainer, 20% sucrose (by volume) was added to mimic extrafloral nectar. This was designated the experimental nectar. The other container, the control nectar, did not have added sucrose. Twenty four larvae, paired by weight and instar, were given standardized leaf sections and placed separately in Petri dishes fitted with experi- mental nectaries. Twelve larvae were placed in dishes fitted for experimental nectar, twelve were placed in dishes fitted for control nectar. All were kept in a constant humidity chamber. Every 24 hours the nectars were withdrawn and replenished, and every 48 hours larvae were weighed. Consumption of fluid was measured as milli- meters traveled down the capillary tubes every 24 hours. The effects of EFN’s, ants, and nectar quality on larval growth through time were analyzed using a 3 way or 2 way Repeated Measures Design ANOVA (Winer, 1971). The factors in the analyses were: time, EFN’s, and presence of ants for the 3 way analysis, and time and type of nectar for the 2 way analysis. Volumes of experimental and control nectar consumed were compared using a two-tailed, paired /-test (Sokal and Rohlf, 1981). Ants often tended T. irenea larvae with greater frequency and fidelity than they did the EFN’s of the hostplant (DeVries, 1988), suggesting that larvae are more attractive to ants than EFN’s. To compare contents of caterpillar secretion and plant 1989 BUTTERFLIES AND ANT-PLANT MUTUALISM 335 extrafloral nectar, sample secretions were taken with micropipettes and spotted on chromatography paper. Amino acid and sugar concentrations were then analyzed using flourometric methods (Baker and Baker, 1976). RESULTS Two results suggested that T. irenea caterpillars drank extrafloral nectar and gained growth benefits from it (Fig. 2). First, caterpillars raised on plants with natural EFN’s gained weight significantly faster (F[5,100] = 6.107, P < .0005) than those raised on plants with EFN’s removed (Fig. 2a). The presence of ants also contributed to weight gain (F[5,100] = 2.821, P < .05), an effect also documented for other ant associated insects (Way, 1963). However, the presence of both ants and EFN’s produced no significant increase in growth rate above the level achieved in the presence of EFN’s alone (F[5,100] = 1.114, P = .358). Second, caterpillars raised with experimental nectars rested with their heads on or near the end of the artificial nectary, and recovery of food coloring in frass and epidermis demonstrated that caterpillars imbibed the nectar. Caterpillars raised with artificial nectar containing sugar imbibed significantly more fluid (paired-/ [10] = 6.897, P < .005), and increased weight significantly faster (F[6,132j = 8.241, P < .0005) than those raised with the control solution containing no sugar (Fig. 2b). The mean concentrations in 15 of 1 7 amino acids examined, as well as the total amino acid concentration (DeVries, 1988) were significantly higher in T. irenea caterpillar secretions than in extrafloral nectar of the hostplant (Fig. 3). Glutamic acid and methionine were found as non-measurable traces in caterpillar secretion, and in extrafloral nectar occurred at concentrations of .0 1 1 and .004 micrograms/ microlitre respectively. T. irenea secretions contained almost no sugars, in contrast to the high concen- trations found in extrafloral nectar of C. billbergianus (DeVries, 1988). Since these caterpillars feed at EFN’s but do not secrete sugars to ants, this suggests that cater- pillars metabolize the sugars taken in as extrafloral nectar, but that sugar plays a minor role in caterpillar secretion as an ant attractant. DISCUSSION We have shown that, in addition to feeding on leaf tissue, myrmecophilous larvae of T. irenea gain growth benefits from drinking the extrafloral nectar of their hostplant (Fig. 2). Drinking extrafloral nectar explains why these larvae typically rest with their heads on or near EFN’s (Fig. 1). Since the ant-organs, which become functional in the third instar, attract and maintain protective ants (DeVries, 1988), the contribution of extrafloral nectar to accelerated growth should greatly benefit young caterpillars by permitting them to reach the third instar quickly. Our results show that the larval section of T. irenea contains significantly higher amino acid concentrations than the extrafloral nectar of C. billbergianus (Fig. 3), but that larval secretion contains almost no sugars. Thus, the high amino acid content of larval secretion is likely to be a factor influencing the preference of E. ruidum ants for tending T. irenea caterpillars over the EFN’s of C. billbergianus (DeVries, 1988). In contrast to T. irenea, some lycaenid caterpillar secretions may have amino acid and sugar concentrations similar to extrafloral nectar (Maschwitz et al, 1975; Pierce, 336 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Effects of nectar and ants on Thisbe irenea larvae 0 2 4 6 8 10 12 14 INTERVALS (days) Fig. 2. Summary of the growth responses of Thisbe irenea caterpillars to EFN’s, ants, and experimental nectars. A. Caterpillar grown on potted plants with captive ant colonies. B. Cat- erpillars grown without ants on standardized leaf tissue and experimental nectars. 1983). Different ant taxa may show demonstrable preferences for different amino acid and sugar concentrations (Lanza and Krauss, 1984; Lanza, 1988), and recently a broad taxonomic pattern of caterpillar-ant associations was shown to be explained, in part, by the feeding ecology of ants (DeVries, 1 987). We suggest that further insights 1989 BUTTERFLIES AND ANT-PLANT MUTUALISM 337 Amino acid Figure 3. Concentrations in micrograms per microlitre of 15 amino acids in caterpillar secretion (N = 5) and extrafloral nectar (N = 4). Concentrations of amino acids varied among individual caterpillars, but not among individual plants (DeVries, 1988). into the taxonomic patterns of caterpillar-ant associations may be gained by analyzing secretions from many species of caterpillars. For example, butterfly caterpillars like T. irenea that associate with ants in a subfamily comprised of predaceous species may produce secretions with a nutrient content that approximates arthropod prey items— high amino acid concentrations and low sugar content. In contrast, caterpillar species associating with ants that typically harvest secretions (e.g., Azteca spp, Iri- domyrmex spp. [Dolichoderinae], Carnponotus spp. Oecophylla spp. [Formicinae]) may produce secretions with amino acid and sugar concentrations similar to extra- floral nectar. A theoretical understanding of both ant-plant and ant-insect mutualism results from considering how two-species mutualisms evolve and are maintained (May, 1976; Goh, 1979; Addicott, 1981; Pierce and Young, 1986). However, the butterfly caterpillars described here use the mutualism between plants and ants, and the basis of this mutualism for their own benefit: both ants and extrafloral nectar benefit the 338 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) growth and survival of herbivorous caterpillars. While we cannot reject the possibility that this system is a 3-way mutualism (i.e., that plant, ant, and caterpillar all benefit from association), it is unlikely that substantial loss of leaf area to caterpillars benefits the plant (DeVries, unpublished). Hence, this study suggests that a third species has invaded a 2-species mutualism. Similar invasions are likely to occur between ant- plant mutualisms and herbivores because many ant-attracting insect herbivores feed on plants with EFN’s. Our current understanding of many ant-plant mutualisms suggests that ants protect plants from herbivores (Beattie, 1985; Atsatt and O’Dowd, 1976). Mutualism can occur between ant-attracting Homoptera and plants without EFN’s because Ho- moptera may act as surrogate EFN’s (Messina, 1981). However, mutualism is unlikely to occur between plants with EFN’s and insects that attract ants (Buckley, 1983b); especially those insects with chewing mouthparts. When lycaenid caterpillars not only attract ants as defenses for themselves, but also specialize on new shoots or young leaves of plants with EFN’s (Pierce, 1985), we suggest that a potential conflict is generated between plant and herbivore for the attention of ants; the plant stands to lose meristems and future photosynthetic potential to an herbivore invading a two-species mutualism. In ant associated riodinid butterflies the conflict between plant and herbivore may be stronger because their larvae commonly feed on extra- floral nectar in addition to leaf tissue (DeVries, 1987, and unpublished). These cat- erpillars not only feed on young meristematic tissues and benefit by using the plant’s ant-guards for protection, but they also exploit the currency of the plant-ant mu- tualism (extrafloral nectar), thereby adding insult to herbivory. Thus, although it is typical to consider the evolutionary stability of two-species mutualism only in the context of both species, our findings suggest that two-species mutualisms may be vulnerable to invasion and exploitation by a third species. ACKNOWLEDGMENTS We thank M. Aide, J. Bull, D. Feener, L. Gilbert, N. Greig, B. Hawkins, P. Harvey, S. Hubbell, E. Leigh, J. Longino, M. Singer, W. Sousa, and P. Ward for comments on the manu- script. N. Greig and T. Gutierrez assisted DeVries with field work. This study was funded in part by a Smithsonian predoctoral fellowship to DeVries and is part of his doctoral dissertation. University of Texas, Austin. This paper is dedicated to the memory of L. Breau and J. Pastorius. LITERATURE CITED Addicott, J. F. 1981. Stability properties of 2-species models of mutualism: simulation studies. Oecologia 49:42-49. Atsatt, P. R. and D. J. O’Dowd. 1976. Plant defense guilds. Science 193:24-29. Baker, H. K. and I. Baker. 1976. Analysis of amino acids in nectar. Phytochemical Bull. 9: 4-7. Banks, C. J. and H. L. Nixon. 1958. Effects of the ant Lasius niger (L.) on the feeding and excretion of the bean aphid Aphis fabae Scop. J. Exper. Biol. 35:703-71 1. Bartlett, B. R. 1961. The influence of ants upon parasites, predators, and scale insects. Ann. Ent. Soc. Amer. 54:543-551. Beattie, A. J. 1985. The Evolutionary Ecology of Ant-Plant Mutualisms. Cambridge Univ. Press, Cambridge. Bristow, C. M. 1984. Differential benefits from ant attendance to two species of Homoptera on New York ironweed. J. Animal. Ecol. 53:715-726. Buckley, R. 1983a. Ant-plant interactions in Australia. W. Junk Publishers, The Hague. 1989 BUTTERFLIES AND ANT-PLANT MUTUALISM 339 Buckley, R. 1983b. Interaction between ants and membracid bugs decreases growth and seed set of host plant bearing extrafloral nectaries. Oecologia 58: 132-136. Callaghan, C. J. 1986. Notes of the biology of Stalachtis susanna (Lycaenidae: Riodininae) with a discussion of riodinine larval strategies. J. Res. Lep. 24:258-263. Cottrell, C. B. 1984. Aphytophagy in butterflies: its relationship to myrmecophily. Zool. J. Linn. Soc. 79:1-57. DeVries, P. J. 1987. Ecological aspects of ant association and hostplant use in a riodinid butterfly. Ph.D. thesis, Univer. of Texas, Austin. DeVries, P. J. 1988. The ant associated larval organs of Thisbe irenea (Riodinidae) and their effects on attending ants. Zool. J. Linn. Soc. 94:379-393. Ehrlich, P. R. 1958. The comparative morphology, phylogeny, and higher classification of the butterflies (Lepidoptera: Papilionoidea). Univ. Kansas Sci. Bull. 39:305-370. Fiedler, K. and U. Maschwitz 1988. Functional analysis of the myrmecophilous relationships between ants (Hymenoptera: Formicidae) and lycaenids (Lepidoptera: Lycaenidae). II. Lycaenid larvae as trophobiotic partners of ants— a quantatative approach. Oecologia 75:204-206. Goh, B. S. 1979. Stability in models of mutualism. Am. Nat. 1 13:261-275. Harvey, D. J. 1987. The higher classification of the Riodinidae (Lepidoptera). Ph.D. thesis. University of Texas, Austin. Hinton, H. E. 1951. Myrmecophilous Lycaenidae and other Lepidoptera- a summary. Proc. Trans. S. Lond. Entom. and Nat. Hist. Soc. 1949-1950:1 1 1-175. Koptur, S. 1984. Experimental evidence for defense of Inga (Mimosoideae) saplings by ants. Ecology 65:1787-1793. Kristensen, N. P. 1976. Remarks on the family-level phylogeny of butterflies. Z. Zool. Syst. Evolforsch. 14:25-33. Lanza, J. 1988. Ant preferences for Passiflora nectar mimics that contain amino acids. Bio- tropica 20:341-344. Lanza, J. and B. R. Krauss 1984. Detection of amino acids in artificial nectars by two tropical ants, Leptothorax and Monomorium. Oecologia 63:423-425. Maschwitz, U., M. Wurst and K. Schurian. 1975. Blaulingraupen als Zuckerlieferanten fur Ameisen. Oecologia 18:17-21. May, R. M. 1976. Models for two interacting populations. Pages 49-70 in: R. M. May (ed.). Theoretical Ecology. Saunders, Philadelphia. Messina, F. L. 1981. Plant protection as a consequence of an ant-membracid mutualism: interactions on goldenrod {Solidago sp.). Ecology 62:1433-1440. Mittler, T. E. 1958. Studies on the feeding and nutrition of Tuberolachnus salignus (Gmelin) (Homoptera, Aphidae) II. The nitrogen and sugar composition of ingested phloem sap and excreted honeydew. J. Exp. Biol. 35:74-84. Pierce, N. E. 1983. The ecology and evolution of symbioses between lycaenid butterflies and ants. Ph.D. thesis. Harvard Univ. Pierce, N. E. 1984. Amplified species diversity: a case study of an Australian lycaenid butterfly and its attendant ants. Symp. Roy. Ent. Soc. Lond. 1 1:196-200. Pierce, N. E. 1985. Lycaenid butterflies and ants: selection for nitrogen fixing and other protein rich food plants. Am. Nat. 125:888-895. Pierce, N. E. 1987. The evolution and biogeography of associations between lycaenid but- terflies and ants. Oxford Surveys in Evol. Biol. 4:89-1 16. Pierce, N. E. and W. R. Young. 1986. Lycaenid butterflies and ants: two-species stable equilibria in mutualistic, commensal, and parasitic interactions. Am. Nat. 128:216-227. Pierce, N. E., R. L. Kitching, R. C. Buckley, M. F. J. Taylor, and K. F. Benbow. 1987. The costs and benefits of cooperation between the Australian lycaenid butterfly, Jalmenus evagoras, and its attendant ants. Behav. Ecol. & Sociobiol. 21:237-248. Robbins, R. K. 1988. Comparative morphology of the butterfly foreleg coxa and trochanter (Lepidoptera) and its systematic implications. Proc. Entomol. Soc. Wash. 90:133-154. 340 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Robbins, R. K. and A. Aiello. 1982. Foodplant and oviposition records for Panamanian Lycaenidae and Riodinidae. J. Lep. Soc. 36:65-75. Ross, G. N. 1966. Life history studies of a Mexican butterfly. IV. The ecology and ethology of Anatole rossi, a myrmecophilous metalmark. Ann. Ent. Soc. Amer. 59:985-1004. Sokal, R. R. and F. J. Rohlf 1981. Biometry. W. H. Freeman, San Francisco, California Vane-Wright, R. I. 1978 Ecological and behavioural origins of diversity in butterflies. Symp. Roy. Ent. Soc. Lond. 9:56-70. Way, M. J. 1963. Mutualism between ants and honeydew producing Homoptera. Ann. Rev. Entom. 8:307-344. Wilson, E. O. 1971. Insect Societies. Harvard University Press, Cambridge. Winer, B. J. 1971. Statistical Principles in Experimental Design. McGraw-Hill, New York. Received March 25, 1989; accepted April 19, 1989. J. New York Entomol. Soc. 97(3):341-346, 1989 SENSORY STRUCTURES ON THE OVIPOSITOR OF THE BALL GALL FLY EURO ST A SOLIDAGINIS (FITCH) (DIPTERA: TEPHRITIDAE) Edward Ritter and Carey E. Vasey Biology Department, SUNY College of Arts and Science, Geneseo, New York 14454 Abstract.— Yht structural features of the ovipositor, its associated sensilla, and its inversion membrane were investigated by scanning electron microscopy. Anteriorly directed, scale-like outgrowths of the integument arranged in triangular-shaped quadrants cover the inversion membrane. Campaniform and trichoid mechanosensilla are found on the inner valves of the ovipositor while a series of shallow pits containing ampullae-like sensory structures occur on both the dorsal and ventral surfaces of the distal portion of the organ. The ventro-lateral margins near the ovipositor tip have distinct depressions on either side which contain two types of chemosensilla. The number of these chemosensilla is not consistent between specimens. In most cases, there are two sensilla on each side but in some specimens there are two sensilla in the right pit and three in the left. The possible functional role of all these sensilla during oviposition is postulated. The ball gall fly, Eurosta solidaginis (Fitch) is known to oviposit in several species of goldenrod (Felt, 1940). In western New York State, Uhler (1951) demonstrated a distinct preference by the fly for Solidago canadensis L. over several species of goldenrod. However, in central Pennsylvania, Greenwald, McCrea and Abrahamson (1984) and Abrahamson, McCrea and Anderson (1989) have shown that Solidago altissima L. is the preferred host plant for this fly. Studies conducted by Miller (1959) in the South revealed that when S. altissima occurs with S. gigantea Ait. and S. ulmifolia Muhl., the fly will oviposit on all three. No hypothesis has been put forth of how E. solidaginis can differentiate between goldenrod species for oviposition. Schoonhaven (1983) noted that a number of dip- teran species use chemoreceptors on tarsi, labella, and ovipositors in finding suitable host plants for oviposition. The structure and possible functions of chemosensilla, together with mechanosensilla, have been recently reported on the ovipositors of two species ofTephritidae: Urophora affinis Frauenfeld (Zacharuk et al., 1986) and Rhag- oletis pomonella (Walsh) (Stoffolano and Yin, 1987). In this paper, we describe the sensilla present on the ovipositor of another trephitid, Eurosta solidaginis and pos- tulate the manner in which they function during egg laying. MATERIALS AND METHODS Twelve specimens of Eurosta solidaginis initially used in this study were obtained from the insect collections of the State University of New York, College of Arts and Science at Geneseo. These flies had been reared from galls collected locally on Sol- idago canadensis. An additional 1 2 specimens were obtained from galls on S. altis- 342 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Figs. 1-9. 1. Ventral view of ovipositor and inversion membrane x 50. Bar = 67 ixm. O = ovipositor, formed from fusion of outer valves. Arrow indicates inner valves. S = inversion membrane with scale-like outgrowths of the integument arranged in triangular-shaped quad- rants. 2. Enlarged view of scale-like outgrowths of integument on inversion membrane x200. Bar = 18.7 nm. 3. Fusion of valves to form ovipositor x400. Bar = 8.65 ixm. Arrows indicate 4 outer valves (ov) of ovipositor iv = inner valve. Note trichoid sensilla on right inner valve. 4. Sensilla on ventral surface of inner valves x400. Bar = 8.6 Aim. D = dome-shaped sensilla. T = trichoid sensilla of inner valves. Also note small trichoid sensilla on lateral margin of 1989 OVIPOSITOR STRUCTURE IN EURO ST A 343 sima in February 1988 by Dr. Warren G. Abrahamson of Bucknell University. When received, the galls were placed in rearing cages at room temperature for 10-14 days, at which time the flies emerged. All specimens were air dried for 7-10 days and attached to stubs, gold-coated on a Polaron diode sputterer and examined with an ISI-Alpha-9 Scanning Electron Microscope. The dried specimens gave a clearer pic- ture of components than did fresh material and, thus, critical point drying was not used. RESULTS The ovipositor and inversion membrane (Fig. 1) corroborate the description cited by Uhler (1951:9) for the external genital system. He noted the “sharp, pointed, chitinized ovipositor” and that the “ovipositor sheath (i.e., inversion membrane) . . . bears chitinized projections on its surface.” These are scale-like outgrowths of the integument and are directed anteriorly. They are arranged in triangular shaped quad- rants (Fig. 2). The significance of this scale pattern is not known. A ventral view of the ovipositor is clearly seen in Figures 1 and 3. Although the valves appear fused, they are not. Specimens softened by boiling in potassium hy- droxide reveal a membrane between them and the dorsal part of the ovipositor. Anterior to the point of contact of the inner valves, three small sensilla may be seen on each valve (Fig. 4). The sensilla are evenly spaced and are different with regard to type and arrangement on each valve. The right inner valve contains two cam- paniform sensilla and one trichoid sensillum, while the left inner valve contains three trichoid sensilla. All the sensilla are directed medially (Fig. 4). The fusion of the outer valves and cerci forms the pointed ovipositor which Uhler (1951) described. On the outer, lateral margin of the expanded ovipositor near the base, is a small trichoid sensillum (Figs. 6, 7). Three more sensilla are located along the outer lateral margin between the base of the ovipositor and the point of contact of the ventral valves. Comparable structures were not discemable on the dorsal or ventral portions of the valves. However, posteriorly to the point of fusion near the tip of the ovipositor, a number of small ampulloid sensilla may be seen on both the ventral and dorsal surface (Figs. 8, 9). On the ventro-lateral margins near the posterior region, where the ovipositor narrows to form the sharp tip, there are two pits con- taining chemosensilla. The number of these sensilla was not consistent for all spec- imens. In most cases, each pit contained two sensilla. However, in three of the 24 specimens we examined, there were two sensilla in the right pit and three in the left (Fig. 9). outer valves, just below figure number. 5. Ventral genital opening x400. Bar =11.2 nm. 6. Trichoid sensillum on outer valve near base, x 1 ,000. Bar = 4.5 Atm. 7. Enlarged view of trichoid sensillum on outer valve, x 3,000. Bar = 1.04 A^m. 8. Dorsal view of posterior portion of ovipositor x700. Bar =5.12 /xm. Note ampulloid sensilla arranged in pairs. 9. Ventral view of apical portion of ovipositor, x 1,000. Bar = 3.46 ixm. Chemosensilla in pits occupy lateral margins. The usual number of sensilla is two in each pit. However, in three of the 24 specimens examined, there were 3 in one pit and two in the other. Notice difference in shape of sensilla. Ampulloid sensilla are also visible on ventral surface. A = ampulloid sensillum. B = chemo- sensilla in pits. 344 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) DISCUSSION The structural relationships seen in the ovipositor and inversion membrane of Eurost a are quite different from that described from such diverse forms as Biosteres longicaudatus (Braconidae), a parasite of the Caribbean fruit fiy Anastrephia (Greany et aL, 1977), the carrot fly Psila rosae (Psilidae) and the cabbage root fly Delia brassicae (Anthomyiidae) (Behan and Ryan, 1977), the blow fly Phormia regina (Calliphoridae) (Wallis, 1962), the sorgum shootfly Antherigona soccata (Anthomyiidae) (Ogwaro and Kokwaro, 1981), and the face fly Musca autumnalis (Muscidae) (Hooper et aL, 1972). The ovipositors of these insects are all segmented structures and contain setal and sensillar patterns that appear to be much more similar to each other than to Eurost a. This is because the ovipositor of tephritids is homologous with only segments eight and beyond in the flies mentioned above (Dr. Allen Norrbom, 1986, pers. comm.). While the ovipositor structure is quite different, many of the sensory struc- tures appear similar and are probably functionally similar as well, even though verification must await further experimentation. Trichoid sensilla have been repeatedly shown to function as mechanoreceptors (Wallis, 1962; Behan and Ryan, 1977; Ogwaro and Kokwaro, 1981; Hooper et al., 1972; Zacharuk et al., 1986; Stoffolano and Yin, 1987). Thus, one can readily pos- tulate that the function of the small trichoid sensilla on the lateral margin of the ventral valve of the ovipositor is to transmit positional information to the fly through deformation of the sensilla as the ovipositor is inserted into the goldenrod tissue. Similarly, the trichoid sensilla on the ventral surface of the two inner valves would be presumed to be mechanoreceptors in the more immediate vicinity of egg release. This would be in agreement with what Stoffolano and Yin (1987) postulated for the trichoid mechanosensilla found on the ovipositor of Rhagoletis pomonella. The two campaniform sensilla on the right inner valve are remarkedly similar in appearance to the “dome-shaped” sensilla shown by Greany et al. (1977) at the tip of the ovipositor of Biosteres longicaudatus. Similar campaniform sensilla were re- ported on the ovipositor of Urophora affinis by Zacharuk, et al. (1986) and on the oviposition apparatus of R. pomonella by Stoffolano and Yin (1987). Greany et al. (1977) also showed TEM micrographs with a definite pore at the tip of the sensillum and concluded it had a chemosensory function— monitoring the chemical environ- ment in the vicinity of oviposition. Schoonhaven (1983) noted that Anastrepha suspensa (Loew) (Tephritidae) can measure acidity and the presence of oviposition pheromone in fruit with its ovipositor. Similar assumption can be made for Eurosta, with the “dome-shaped” sensilla monitoring the chemical environment in the im- mediate vicinity where the egg would be released. However, confirmation of this would necessitate TEM studies. Chemical monitoring of oviposition sites and/or host plant tissues undoubtedly also occurs in the chemosensilla near the tip of the ovipositor. The sensilla in each pit are not all the same. In each pit, the “terminal-most” sensillum is broadly rounded at its tip while the one(s) anterior to it have a slender curved tip with a suggestion of a pore at the bend. Although no specific pore has yet been clearly demonstrated in these particular sensilla, similar structures in other species have been shown to have pores and to have a chemosensory or gustatory function. The differences some- times in number and shape of these sensilla in their respective pits would indicate, however, that they probably are sensing different chemicals, or the pH, etc. in the 1989 OVIPOSITOR STRUCTURE IN EUROSTA 345 environment of the host plant tissue. Interestingly, the large chemosensilla which Wallis (1962) illustrated on the anal plate of the blow fly Phormia are similar to the anteriormost chemosensilla occurring in the pits near the tip of the ovipositor of Eurost a. In Phormia, these sensilla have a gustatory function, enabling the fly to taste the ovipositional substrate. The probable function of the small ampullae-like structures which occur on the dorsal and ventral surfaces of the posterior portion of the ovipositor are more difficult to ascertain. There does not appear to be the pore relationship one would expect if they were chemosensory, nor can one detect micro-hair-like structures that might suggest a mechanosensory function. It is possible that they are part of a water-sensing system, but there is no proof for it at this time. What is clear is that there are a sufficient number of different types of sensilla to account for the fly’s ability to control oviposition and egg release. It also can account for the times in which Eurosta has been known to pierce goldenrod tissue but not release eggs (Uhler, 1951; Abrahamson, pers. comm., 1985). ACKNOWLEDGMENTS We thank Dr. Allen Norrbom, Systematics Entomology Laboratory, USDA, and Dr. John Stoffblano, Department of Entomology, University of Massachusetts, and Dr. Robert Reason, Biology Department, SUNY College of Arts and Science, Geneseo, New York, for their advice and suggestions after reading the manuscript. We are also indebted to Dr. Warren C. Abra- hamson, Biology Department, Bucknell University, for providing some of the specimens used in this study. The technical assistance of Ms. Elizabeth Netzband is also greatly appreciated. LITERATURE CITED Abrahamson, W. G., K. D. McCrea and S. S. Anderson. 1989. Host preference and recognition by the goldenrod ball gallmaker Eurosta solidaginis (Diptera: Tephritidae). Amer. Midi. Nat. 121:322-330. Behan, M. and M. F. Ryan. 1977. Sensory receptors on the ovipositor of the carrot fly Psila raffle (Diptera: Psilidae) and the cabbage root fly Delia brassicae (Di^XtrSi: Anthomyiidae). Bull. Entomol. Res. 67:383-389. Felt, E. P. 1940. Plant Galls and Gall Makers. Haefner, New York. Greany, P. D., S. D. Hawke, T. C. Carlysle and D. W. Anthony. 1977. Sense organs in the ovipositor of Biosteres (Opius) longicaudatus, a parasite of the Caribbean fruit fly Anas- trepha suspensa. Ann. Entomol. Soc. Amer. 70:319-321. Greenwald, L. L., K. D. McCrea and W. G. Abrahamson. 1984. Meetings Ecol. Soc. Amer., Ft. Collins, Colorado, Abstracts p. 102. Hooper, R. L., C. W. Pitts and J. A. Westfall. 1972. Sense organs on the ovipositor of the face fly, Musca autumnalis. Ann. Entomol. Soc. Amer. 65:577-586. Miller, W. E. 1959. Natural history notes on the goldenrod ball gall fly, Eurosta solidaginis (Fitch) and on its parasites Eurytoma obtusiventris Gahan and E. gigantea Walsh. J. Tenn. Acad. Sci. 34:246-251. Ogwaro, K. and E. D. Kokwaro. 1981. Morphological observations on sensory structures on the ovipositor and tarsi of the female and on the head capsule of the larva of the sorghum shootfly, Atherigona soccata Rondani. Insect Sci. Applications 2:25-32. Schoonhaven, L. M. 1983. The role of chemoreception in host plant finding and oviposition in phytophagous Diptera. Proc. CEC/IOBC Symposium. Athens, Nov. 1982. Stoffblano, J. G., Jr. and L. R. S. Yin. 1987. Structure and function of the ovipositor and 346 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) associated sensilla of the apple maggot, Rhagoletis pomonella (Walsh) (Diptera: Te- phritidae). Int. J. Insect Morphol. Embryol. 16:41-70. Uhler, L. D. 1951. Biology and ecology of the goldenrod gall fly, Eurosta solidaginis (Fitch). Cornell Univ. Agri. Exp. Sta. Memoirs. 300:1-51. Wallis, D. I. 1 962. The sense organs on the ovipositor of the blow fly, Phormia regina Meigen. J. Ins. Physiol. 8:453-467. Zacharuk, R. Y., R. W. K. W. Lee, and D. E. Berube. 1986. Fine structure of sensilla on the ovipositor of the tephritid fly Uwphora affinis. Can. J. Zool. 64:973-984, Received July 23, 1986; accepted January 25, 1989. J. New York Entomol. Soc. 97(3):347-357, 1989 REVIEW OF NEARCTIC RHICNOCOELIA AND CALLIMERISMUS WITH A DISCUSSION OF THEIR PHYLOGENETIC RELATIONSHIPS (HYMENOPTERA: PTEROMALIDAE) Steven L. Heydon Department of Entomology, NHB; Mail Stop 165, Smithsonian Institution, Washington, D.C. 20560 Abstract.— The genera Rhicnocoelia Graham and Callimerismus Graham are delimited and their relationships with other genera of the Pteromalidae are discussed. Both genera are reported from the Nearctic region for the first time. Rhicnocoelia has two North American species— i?. punctifrons n. sp., and the Holarctic species R. constans (Walker). A key to separate these species is given. The first host record for Rhicnocoelia, from a puparium of the chloropid Meromyza americana Fitch, is presented. Callimerismus has one Nearctic species, C. inusitatus n. sp. Callimerismus latipennis (Ashmead) is removed from the genus, but its correct generic place- ment is uncertain. If one compares the numbers of species and genera of the pteromalid subfamily Miscogasterinae in the Palearctic region (summarized in Graham, 1969) with the number of genera and species in the Nearctic region (summarized in Burks, 1979), it might be surmised that the Nearctic miscogastrine fauna is depauperate relative to the Palearctic fauna. My continuing study of the Nearctic Pteromalidae has shown that this is not true, and that nearly all the Palearctic miscogasterine genera are present in the Nearctic region as well. This paper documents the first Nearctic records of two genera, Rhicnocoelia Graham and Callimerismus Graham. Rhicnocoelia is represented by the Holarctic species R. constants (Walker) and the new Nearctic species R. punctifrons. Callimerismus is represented by the new Nearctic species C. inusitatus. Callimerismus latipennis (Ashmead) previously transferred to Callimer- ismus by Hedqvist (1969) is removed from that genus. The generic placement of new Nearctic miscogasterine species is often difficult because they show morphological variation not found in their Palearctic counterparts. As these new species are described, generic concepts need to be carefully reviewed. The following reanalysis of the characters used to separate Rhicnocoelia from Cal- limerismus is necessary since many of the characters given by Graham (1956b) to differentiate these two genera are either matters of degree rather than discrete dif- ferences, or now need to be supplemented based on the characters of the new species described in this paper. Several characters Graham used to separate Rhicnocoelia from Callimerismus are not as distinctive now that more material of these genera is available. 1 . Graham stated that the pronotal collar of Rhicnocoelia lacks the distinct transverse anterior Carina found in Callimerismus. I find the strength of development of this carina varies greatly between specimens of C. fronto (Walker). Furthermore, C. inusitatus and an undescribed species of Callimerismus from Sweden in my personal collection have 348 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) the neck and collar meeting at a right angle, but clearly not carinate. Thus, the presence or absenee of a pronotal carina is of little value for separating these two genera. 2. Graham reported that the spiracles of Rhicnocoelia are smaller and more circular than those of Callimerismus. However, these differences in spiracular shape can be readily seen only when a series of specimens of both genera are available for com- parison because of the variability in spiracular shape even in different specimens of the same species. 3. Graham also reported that Rhicnocoelia has a rugose propodeum, while the propodeum of Callimerismus lacks rugae. However, the holotype of C. inusitatus has distinct rugae on the propodeum, and I find there is nearly as great a difference in the degree of rugosity of the propodeum between different specimens of Rhicnocoelia constans and R. punctifrons as between those species and specimens of Callimerismus species. 4. Graham reported that the postmarginal vein is as long as the marginal vein in Callimerismus, but shorter than this in Rhicnocoelia. I find little difference in the relative lengths of the marginal and postmarginal veins in these two genera; the postmarginal vein is sometimes shorter than the marginal vein in both genera. Some of the differences between these genera recognized by Graham are still valid. 1. Rhicnocoelia is somewhat unusual because the costal cell of the fore wing lacks any ventral setae in its basal half (Figs. 1, 3). Callimerismus has the more common condition with a ventral row of setae extending the length of the costal cell (Fig. 7). To my knowledge, among the Miscogasterinae, an incomplete ventral row of setae in the costal cell occurs only in Rhicnocoelia and Bubekia Dalla Torre. 2. The hind margin of the first gastral tergum in Rhicnocoelia is emarginate medially (Fig. 5) (except in R. grahami); in Callimerismus, it is always straight (Fig. 6). 3. Graham also notes that the petiole of Rhicnocoelia differs from that of Callimerismus because the petiole of Rhicnocoelia is shorter and not distinctly sculptured (Fig. 9). But the differences between the petioles are more fundamental than this. The short, smooth petiole of Rhicnocoelia resembles those of the plesiomorphic miscogasterine genera such as Seladerma Walker and Lamprotatus Westwood (compare Figs. 8, 9). It is not sclerotized ventrally (Fig. 1 0) and its basal bracing consists of lateral reflexed lobes (Fig. 9). The structure of the petiole of Callimerismus is synapomorphic with that of the more advanced miscogasterine genera such as Sphegigaster Spinola and others in the Sphegigasterini {sensu Graham 1969). The petiole of Rhicnocoelia is reticulate dorsally, elongate, tubular but with a median ventral sulcus, and is braced basally by an anteriorly directed flange which runs continuously laterally and ventrally except where it is interrupted medially by the sulcus (Fig. 1 1). Despite the plesiomorphic petiole, Rhicnocoelia is morphologically, and probably phylogenetically, transitional between the more primitive miscogasterine genera and the more apomorphic genera grouped in the Sphegigasterini with whom Rhicnocoelia shares several apomorphic character states. These character states include the de- velopment of a horizontal posterior strip on the pronotum (called the collar), notauli that are relatively shallow and groovelike rather than deep and furrowlike, a scutellum that is only about as long as wide rather than elongate, a postmarginal vein at most about as long as the marginal vein rather than distinctly longer, a basal vein without obvious pigmentation or tracheolation, and a reduction in the distribution of setae in the basal cell. 1989 REVIEW OF RHICNOCOELIA AND CALLIMERISMUS 349 For a detailed but preliminary evaluation of the exact phylogenetic relationships of Rhicnocoelia and Callimerismus, see Heydon (1988). MATERIALS AND METHODS Terminology in this paper generally follows that of Graham (1969), except that genal concavity is used instead of genal hollow, the lower ocular line (abbreviated LOcL) is an imaginary line across the face between the most ventral point of the orbits, club is used instead of clava, the elongate raised sensilla on the antennal flagellar segments are called multiporous plate (abbreviated MPP) sensilla, and the middle body tagma including the thorax and propodeum is called the mesosoma. In addition, the gastral terga are numbered 1-7 beginning with the first tergite after the petiole. The following abbreviations are used: median ocellar diameter is MOD, ocel- ocular distance is OOL, posterior ocellar distance is POL, lateral ocellar distance is LOL, antennal funicular segments are FI through F6, and the gastral terga are T1 through T7. Sculpturing is defined according to Harris (1979). The units of mea- surement given in the descriptions can be converted to millimeters by multiplying by 0.02. The acronyms used for the various museums can be found in the Acknowl- edgments; the acronym for my personal collection is SLH. My taxonomic arrange- ment of genera discussed generally follows Graham (1969) except Merismus and Callimerismus are transferred from the Miscogasterini to the Sphegigasterini because the petiole of these genera is sclerotized ventrally and has a basal ventral flange. GENERIC REVISIONS Rhicnocoelia Graham Megorismus TYvomson, 1876:220, 240 [«cc Walker, 1846]. Rhicnocoelia Graham, 1956b:262-263. Type species: Pteromalus constans Walker, 1836 (original designation). Peck, Boucek, and Hoffer, 1964:35, 38 (key). Graham, 1969:151, 168-171 (key, synonymy). Boucek, 1988:242, 469-470, 498. Dogmiella Delucchi, 1962:7. Type species: Dogmiella viridis Delucchi, 1962 (not seen, accepted on the authority of Graham, 1969). Graham (1956b) described Rhicnocoelia with the type species Pteromalus constans Walker, 1836. He also transferred Larnprotatus coretas Walker, 1848 and six other Walker species into Rhicnocoelia in that paper. Graham (1969) retained R. coretas as a valid species, but the other seven Rhicnocoelia species were collapsed into the two species R. constans and R. impar (Walker), 1836 and Graham further expressed some doubt whether even these two species were really separable. Graham (1969) also placed the monotypic genus Dogmiella Delucchi, with its type species, D. viridis Delucchi, 1962 in synonymy with Rhicnocoelia. Rhicnocoelia constans, R. coretas, R. grahami Boucek, 1970 and R. impar all occur in Europe, and R. viridis occurs in Morocco. Rhicnocoelia incisa Boucek, 1988 was recently described from Australia. I extend the range of this genus to the New World with the description of a new Nearctic species, R. punctifrons n. sp., and the first Nearctic record of R. constans. Description. Species metallic blue or green to dark green. Head with clypeus co- riaeeous, apical margin with three distinct and asymmetrically arranged denticles 350 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Figs. 1-7. Rhicnocoelia constans (Walker). 1, Female fore wing. Rhicnocoelia punctifwns n. sp. 2, Female head (anterior view). 3, Female fore wing. Rhicnocoelia constans (Walker). 4, Female head (anterior view). 5, Female gaster (dorsal view). Callimerismus inusitatus n. sp. 6, Female whole body. 7, Female fore wing. 1989 REVIEW OF RHICNOCOELIA AND CALLIMERISMUS 351 Figs. 8-11. Petioles. 8, Seladerma sp., female (dorsolateral view). 9, 10, Rhicnocoelia con- stans (Walker), male. 9, (dorsolateral view). 10, (anterior view). 1 1, Callimerismus sp., female (ventrolateral view). (Figs. 2, 4); gena with weak concavities or lacking them entirely. Antenna inserted one torular diameter above LOcL; antennal formula 1:1:2:6:3; scape cylindrical, length approximately 5-6 x width; flagellum usually clavate in females, parallel-sided in males; MPP sensilla prominent; female club with small patch of micropilosity ventrally on apical segment, no terminal spine. Male maxilla without lobes off stipites, palps slender. Mandibles with three teeth on left, four on right. Mesosoma elongate, length 1.6 or more times width (but usually 2 x as long as wide or more); pronotum long (length about Vi width), with short horizontal collar lacking anterior transverse Carina; mesoscutum alveolate, notauli traceable to hind margin but shallow poste- riorly; prepectus flat, without distinct carina setting off posterior comer; scutellum just longer than wide, no anterior median sulcus present, frenal sulcus distinct, frenum reticulate (reticulations either impressed or raised); propodeum reticulate-mgose, with median portion projecting between hind coxae, nucha set off anteriorly by sharp Carina from which a series of short carinae extend anteriorly, median carina rugiform when traceable, plicae fading out in anterior Vs, spiracles circular or shortly ovate. Legs stout, fore femur four or less times as long as wide. Fore wing (Figs. 1 , 3) with marginal vein approximately as long as postmarginal vein; stigma hardly wider than 352 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) stigmal vein; costal cell with broad basal interruption in ventral setal row; basal cell usually setose distally; speculum open posteriorly. Petiole very short, conical, smooth, unsclerotized ventrally, with a pair of lateral reflexed lobes (Figs. 9, 10). Gaster tapering in females, ovate in males; hind margin of T 1 usually shallowly emarginate (Fig. 5), straight in R. grahami. A dorsal view of R. incisa is given in Boucek (1988: 498), and of R. grahami in Boucek (1970:51). Discussion. Rhicnocoelia and Callimerismus are phenetically similar genera. These two genera have nearly identical head structure; a similar elongate mesosoma which is alveolately sculptured dorsally; a long pronotum which is half as long as wide; shallow notauli; similar propodeal structure with the median portion reticulate, pro- truding posteriorly far between the hind coxae, and with the region between the spiracular sulci raised above the level of the lateral regions; and an elongate and rather acuminate gaster. Characters to separate these genera are discussed in the introduction of this paper. Biology. The hosts of the described species in this genus are unknown; however, a male of an undescribed Nearctic species in the USNM collection from Danville, Pennsylvania, was reared from a puparium of the wheat stem maggot, Meromyza americana Fitch (Diptera: Chloropidae). KEY TO NEARCTIC SPECIES OF RHICNOCOELIA GRAHAM 1 . Scutellum and frenum both coriaceous; eye height less than 2.7 times the malar distance (Fig. 2); frons and vertex with piliferous punctures across the entire width; basal cell usually bare (Fig. 3) punctifrons - Scutellum alveolate, frenum coriaceous; eye height 2.8 or more times malar distance (Fig. 4); piliferous punctures restricted to inner orbits; basal cell setate at least over apical one-fourth (Fig. 1) constans Rhicnocoelia punctifrons, new species Figs. 2-3 Description. Holotype, female. Color: Head, scape, pedicel, mesosoma, gaster me- tallic green; head and pedicel with coppery, gena with red reflections; flagellum dark brown; mandible amber, teeth dark reddish brown. Legs with femora, tibiae orange- yellow, middle femur with metallic green strip mid-dorsally; pretarsi dark brown, fore tarsi brown, middle and hind basitarsi cream-colored, remainder orange-yellow. Wing veins yellow-brown, parastigma and stigma reddish brown. Sculpture: Clypeus finely coriaceous, remainder of head coriaceous, frons and vertex with scattered punctures along orbits; pronotum, anterior part of mesoscutum imbricate; disc of middle lobe of mesoscutum alveolate; side lobes, axilla, scutellum, frenum coriaceous; dorsellum alveolate; propodeum alveolate rugose; gastral Tl-3 smooth, T4-7 coriaceous; prominent MPP sensilla give the antennal flagellum coarse texture. Structure: Body length 3.1 mm. Head (Fig. 2) width 1.2 x height (43.5:35.0), 1.8 x length (43.5:24.0); eye height 1.3 x length (21:16), 2.5 x malar distance (21.0:8.5), length 2.7 X temple length (16:6); ratio of MOD, OOL, POL, LOL as 2.5:7.5:11.0: 5.5; torulus located a little over one diameter above LOcL. Antenna with length of 1989 REVIEW OF RHICNOCOELIA AND CALLIMERISMUS 353 pedicel plus flagellum 1.3 x head width (55.0:43.5); ratio of lengths of scape, pedicel, anelli, Fl-6, club as 16.0:6.0:2.0:7.5:6.5:6.0:5.5:5.0:5.0:8.0; widths of FI, F6, club as 4. 0:5. 0:5. 5; MPP sensilla long, in single irregular row on FI, two partially over- lapping rows on F2-6; apical club segment with ventral patch of micropilosity. Mesosoma 2.2 x as long as wide (75:34); dorsellum nearly ‘/z length of frenum; propodeum with series of short longitudinal carinae anteriorly, rugae diverging from sharp median carina; spiracles ovate, 1 x own diameter from anterior margin of propodeum, callus with very few setae. Legs stout, fore femur length 3.4 x width (3 1 : 9). Fore wing (Fig. 3) length 2.8 x width (130:47); ratio of submarginal, marginal, postmarginal, stigmal veins as 55:22:24:12; basal cell and basal vein bare. Gaster broadly fusiform in dorsal view, length 1.8 x width (55:31); length 1.1 x height (55: 50); hind margin of T1 with broad median emargination. Allotype, male. Color: Similar to holotype female except legs brownish yellow, middle femur with dorsal brown patch, not metallic. Structure: Body length 2.9 mm. Head width 1.8 x length (37.0:20.5). Antenna with lengths of pedicel plus flagellum 1.9 X head width (72:37); ratio of lengths of scape, pedicel, anelli, Fl-6, club as 8.5: 5.5:1.5:9.0:9.0:9.0:8.0:8.0:7.0:15.0; widths of FI, F6, club as 3. 5:4. 0:4.0; funicular segments cylindrical, thickly covered with fine light colored hairs which are curved toward the apex so that their tips lie parallel to long axis of the segment, MPP sensilla in two irregular rows on each segment. Fore wing with basal vein with one seta on left wing and three on right wing. Gaster ovate, length 1.9 x width (55:29). Variation. Specimens of both sexes may have the pronotum, side lobes of the mesoscutum, and axilla bluish green to brilliant blue. The male from Colorado has extensive red areas on the face. The body length of females varies from 2.7 to 3.5 mm; males from 2.4 to 3.2 mm. The temple length is up to 0.55 times the length of the eye in females; in males, the ratio of temple length to eye length varies between 0.53 and 0.69. Females usually have a more or less well developed patch of metallic coloration on the posterior surface of the middle femur, but this patch is nearly lacking in 2 of the 7 type females. The allotype is the only specimen in the paratype series with setae on the basal vein. Diagnosis. This species keys to R. constans in Graham (1969) but is readily dis- tinguishable from that species by the large head (head width being only about two times length), the lack of raised sculpturing on the head and scutellum (including frenum), the lack of setae on the basal cell, and by having the parastigma and stigma distinctly darker than the marginal and stigmal veins. Males of both R. punctifrons and R. constans have cylindrical antennal funicular segments thickly covered with fine light colored hairs which are curved toward the apex so that their tips lie parallel to the long axis of the segment. Etymology. The species name is a conjunction of the Latin words punctum, meaning small hole, and frons, meaning brow or forehead, and refers to the prominent pilif- erous punctures on the head of this species. Type Material. The holotype female (USNM), allotype male (USNM), and five female and seven male paratypes (USNM) were collected at Springer, New Mexico in 1909 by C. N. Ainselie. Two additional paratypes were collected as follows (USNM): United States. COLORADO: Fort Collins, 10- VIM896, M; Moffat Co., 12. Host. Unknown. 354 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Rhicnocoelia constans (Walker) Figs. 1, 4-5, 9-10 Pteromalus constants Walker, 1836:468-469. Lectotype, $; BMNH, Hym. Type No. 5.1696 [examined]. Graham, 1956b:263 (synonymy). Pteromalus cliens Walker, 1836:469 [type not seen]. Graham, 1956b:263 (synonymy). Megorismus chloris Thomson, 1876:241 [type not seen]. Graham, 1956b:263 (syn- onymy). Rhicnocoelia constans (Walker): Graham, 1956b:263; 1969:169-170. Boucek, 1970: 48-49; 1977:51. Graham (1969) also cites the possible synonymy of Pteromalus vindalius Walker, 1839; Pteromalus archidemus Walker, 1839; Pteromalus orsippus Walker, 1839; Lamprotatus labaris Walker, 1848; and Pteromalus phalasarna Walker, 1848 with Rhicnocoelia constans. The type material of the first four of these species consists of unassociated males which he says are difficult to positively determine to species. Diagnosis. Characters to distinguish R. constans from R. punctifrons are given in the discussion section for that species. Material Examined: Palearctic Region (BMNH, CNC, INKS, SLH, USNM): En- gland, Sweden, France, Germany, Hungary, Cyprus. Nearctic Region (CNC, USNM): United States. ALASKA: Anchorage, 3 1 VIM 95 1, 19. MICHIGAN: Delta Co., 25- VIIM952, 19. Canada. ALBERTA: McMurray, 7- VIII- 1953, 1<5; 10- VIIT 1953, 19. The following unassociated males probably belong to this species (CNC, USNM): United States. COLORADO: IS; Doolittle Ranch, Mt. Evans, 10 VIIT1961, 16. Canada. YUKON TERRITORY: Mile 87, Dempster Highway, 4-8 VIIM973, 16. Callimerismus Graham Callimerismus Grakava, 1956a:78. Type species Merismus fronto Walker , 1833 (orig- inal designation). Graham, 1969:151, 176-179. Callimerismus Graham was created for a single British species, Merismus fronto Walker, 1833 (Graham, 1956a). Graham (1969) later described a second species, Callimerismus suecicus, from Sweden. I am extending the geographic range of this genus to the Nearctic region with the description of a new Nearctic species, Calli- merismus inusitatus n. sp. Dipara latipennis Ashmead, 1890 was transferred to Callimerismus by Hedqvist (1969). This species is known only from the male holotype and its generic placement is problematic. In most respects, it resembles the other Callimerismus species, but it differs from them in three critical ways. The petiole of C. latipennis is structured much as in MiscogasterWalker— slender, elongate, sculptured, but slightly convergent anteriorly and with no basal bracing of any kind. The other species of Callimerismus have a basal flange bracing the petiole basally (Fig. 1 1), a characteristic of genera in the Sphegigasterini. Callimerismus latipennis also has a shiny swollen apical ventral shiny strip on the scape (called a boss) which is characteristic of many genera of the Miscogasterini such as Seladerma, Sphaeripalpus Forster, Stictomischus Thomson, Miscogaster, Lamprotatus, and Skelocerus Delucchi. A boss is very rare in any species of the Sphegigasterini. Finally, the hind margin of the first tergum is sinuous laterally, again similar to the first tergum in Miscogaster; the hind margin of the first tergum 1989 REVIEW OF RHICNOCOELIA AND CALLIMERISMUS 355 is straight in other Callimerismus species. Callimerismus latipennis will not be treated as a Callimerismus species further in this paper. Its eventual generic placement will have to await a critical study of the genera similar to Miscogaster. Description. Body color metallic blue or green. Head with anterior margin of clypeus with three distinct asymmetrically arranged apical denticles; mouth margin arched laterally; gena with short concavity. Antenna inserted well above LOcL; antennal formula 1:1:2:6:3; scape cylindrical, length about 5x width; MPP sensilla in single row; female club lacking any micropilosity or terminal spine. Mandibles with three teeth on left, four on right. Mesosoma elongate; pronotum elongate (length about Vi width), collar short and weakly carinate or crested anteriorly; mesoscutum alveolate, notauli complete but sometimes shallow posteriorly; preprectus nearly flat, reticulate, lacking carina delimiting hind comer; scutellum just longer than wide, without an- terior median sulcus, with distinct frenal sulcus, frenum reticulate; propodeum re- ticulate, protruding between hind coxae, portion between spiracular sulci distinctly raised above lateral regions, median carina complete or not, plicae incomplete, spi- racles ovate. Fore wing (Fig. 7) with marginal vein longer than postmarginal vein; stigma twice as wide as stigmal vein; costal cell with complete ventral row of setae; basal cell bare; basal vein setate; speculum open posteriorly. Petiole cylindrical, longer than wide, reticulate; sclerotized ventrally; lateral setae present; with basal flange. Gaster with hind margin of T 1 straight. Diagnosis. Callimerismus is probably most closely related to a group of genera including Merismus Walker, Toxeuma Walker, and Cryptoprymna Forster. All these genera have an elongate propodeum that is distinctly raised between the spiracular sulci and extends posteriorly between the hind coxae. Callimerismus differs from Merismus by having the anterior transverse carina on the pronotal collar developed to a much lesser degree (if at all) and by having the prepectus uniformly reticulate; Merismus has the prepectus smooth and with a carina setting off its hind comer. Callimerismus differs from Toxeuma by having the anterior margin of the clypeus with three asymmetically arranged denticles, and the anterior transverse carina on the pronotal collar developed to a much lesser degree; Toxeuma has no anterior denticles on the clypeus, and a strongly developed anterior transverse carina on the collar. Callimerismus differs from Cryptoprymna by having a metallic green or blue body, distinct denticles on the anterior margin of the clypeus, no large patch of micropilosity on the female club, and the first gastral tergite not covering the entire gaster; Cryptoprymna species are black, have a weakly truncate clypeus, a large patch of micropilosity on the female club, and T1 virtually covers the gaster. The differences between Callimerismus and the phenetically similar genus Rhicnocoelia are given in the discussion section for that genus. Callimerismus inusitatus, new species Figs. 6-7 Description. Holotype, female. Color: Clypeus blue in lower half; remainder of head yellow-green, vertex with coppery reflections; pronotum, pleural region yellow- green; dorsum of mesosoma green; propodeum, petiole blue-green; gaster dark brown, basal half of T1 with blue-green reflections, T4-7 with yellow-green reflections. An- tenna with scape yellow, remainder brown. Legs yellow-brown, femora and middle 356 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) and hind tibiae with weak yellow-green reflections, pretarsi dark brown. Wing veins yellow-brown. Sculpture: Clypeus smooth; remainder of head, pronotum, mesoscutum, scutellum, axilla, dorsellum alveolate; propodeum alveolate with a few fine but sharp rugae; petiole finely alveolate; Tl-5 smooth, T6-7 coriaceous. Structure: Body length 2.2 mm. Head width 1.3 x height (32:35), 1.7 x length (32: 19); eye height 1.3x length (18:14), 5.1 x malar distance( 18.0:3. 5), length 4. Ox temple length (14.0:3.5); genal concavity extending Vi malar distance; ratio of MOD, OOL, POL, LOL as 3:4:7:4; distance from median ocellus to front of head equal to distance from lateral ocellus to occiput; torulus inserted 2 x inside diameter above LOcL. Antenna with length of pedicel plus flagellum 0.9 1 x head width (29:32); scape cylindrical, length 5.0 x width (10:2); ratio of lengths of scape, pedicel, anelli, Fl-6, club as 10.0:4.0:1.5:2.5:3.0:3.0:3.0:3.0:3.0:8.0; widths of FI, F6, club as 3. 0:3. 5:4.0; funicular segments cylindrical, MPP sensilla sparse, length about equal to length of funicular segments. Mesosoma length 2.0 x width (29:24); collar with anterior edge crested; propodeum with nuchal region set off anteriorly by fine carina, spiracles 1 x own diameter from anterior margin of propodeum. Fore wing (Fig. 7) length 2.4 x width (86:36); ratio of lengths of submarginal, marginal, postmarginal, stigmal veins as 33:19:16:8; basal vein with one complete and a partial second row of setae. Petiole length 1 .2 x width (8. 5:7.0); median carina weak; pair of fine setae extending laterally. Gaster fusiform; length 2.1 x width (41.0:19.5). Variation. The body length of the paratype female is 2.4 mm. Its body color is generally similar to the holotype, but the scape is brown apically, the legs are darker with brown femora and tibiae, and the lateral regions of the scutellum are blue. The combined length of the scape and pedicel is 0.91 times the head width. The head width is 1.8 times its length. The dorsellum of the holotype is obscured, but in the paratype, the dorsellum is bandlike, carinate anteriorly, and short [its width 3.6 times its length (9.0:2. 5)]. The petiole length is 1.3 times its width in the paratype. Discussion. This species will key out with C. fronto in the key to European species given by Graham (1969). Callimerismus inusitatus differs from C. fronto in the following: 1 . The scutellum in C. inusitatus is moderately arched; it is strongly arched in C fronto. 2. The scape of C. inusitatus is non-metallic; it is metallic in C. fronto. 3. The eye height is about five times the malar distance in C. inusitatus', it is 3.5 times the malar distance in C. fronto. 4. The petiole has a single or no pairs of lateral setae in C inusitatus', there are several pairs present in C. fronto. Type material. The holotype (CNC) is a female, from Ancaster, Ontario, Canada, and was collected 24 June 1955 by O. Peck. The paratype female (CMNH) is from Keron Hill, Pittsburgh, Pennsylvania, and was collected on 16 June 1940 by G. E. Wallace. Etymology. The species name comes from the Latin word inusitatus, meaning rare or unusual, and refers to the rareness of this species. Host. Unknown. ACKNOWLEDGMENTS I thank the following persons and institutions for loan of materials: Dr. J. S. Noyes, British Museum (Natural History) (BMNH), London; Dr. G. Wallace, Carnegie Mu- 1989 REVIEW OF RHICNOCOELIA AND CALLIMERISMUS 357 seum of Natural History (CMNH), Pittsburgh; Dr. G. P. G. Gibson, Canadian Na- tional Collection (CNC), Ottawa; Dr. W. E. LaBerge, Illinois Natural History Survey (INHS), Champaign; and Dr. E. E. Grissell, United States National Museum (USNM), Washington, D.C. I thank Dr. Christer Hansson for his gift of Palearctic material; the Illinois Natural History Survey, the Canacoll Foundation, the H. H. Ross Me- morial Foundation, the Smithsonian Institution, and Sigma Xi for supporting this research; Dr. J. S. Noyes for use of his card hie; Drs. Z. Boucek, E. E. Grissell and W. E. LaBerge for insightful comments; and the staff at the Center for Electron Microscopy of the University of Illinois for use of the SEM. LITERATURE CITED Boucek, Z. 1970. Contribution to the knowledge of Italian Chalcidoidea, based mainly on a study at the Institute of Entomology in Turin, with descriptions of some new European species (Hymenoptera). Mem. Soc. Entomol. Ital. 49:35-102. Boucek, Z. 1 977. A faunistic review of the Yugoslavian Chalcidoidea (Parasitic Hymenoptera). Acta Entomol. Jugoslav. 13(Suppl.):l-145. Boucek, Z. 1988. Australasian Chalcidoidea (Hymenoptera). A biosystematic revision of genera of fourteen families, with a reclassihcation of species. CAB International Institute of Entomology, Wallingford, England. Burks, B. D. 1979. Family Pteromalidae. Pages 768-835 in: K. V. Krombein, P. D. Hurd, Jr., D. R. Smith, and B. D. Burks (eds.). Catalog of Hymenoptera in America North of Mexico. Volume 1, Symphyta and Apocrita (Parasitica). Smithsonian Institution Press, Washington, D.C. Delucchi, V. 1962. Hymenopteres Chalcidiens du Maroc II. Pteromalidae (suite). A1 Awamia 4:7-25. Graham, M. W. R. de V. 1956a. A revision of the Walker types of Pteromalidae (Hym., Chalcidoidea). Part I (including descriptions of new genera and species). Entomol. Mon. Mag. 92:76-98. Graham, M. W. R. de V. 1956b. A revision of the Walker types of Pteromalidae (Hym., Chalcidoidea). Part 2 (including descriptions of new genera and species). Entomol. Mon. Mag. 92:246-263. Graham, M. W. R. de V. 1969. The Pteromalidae of Northwestern Europe (Hymenoptera: Chalcidoidea). Bull. Br. Mus. (Nat. Hist.) Entomol. Suppl. 16:1-908. Harris, R. A. 1979. A glossary of surface sculpturing. Occ. Pap. Entomol. St. Calif Dept. Fd. Agric. 28:1-31. Hedqvist, K.-J. 1969. New genera and species of Diparini with notes on the tribe (Hym., Chalcidoidea). Entomol. Tidskr. 90:174-202. Hey don, S. L. 1988. The Sphegigasterini: A cladistic analysis and generic classification with reviews of selected genera (Hymenoptera: Pteromalidae). Ph.D. Thesis, Univ. Illinois at Urbana-Champaign, Urbana. Peck, O., Z. Boucek, and A. Hoffer. 1 964. Keys to the Chalcidoidea of Czechoslovakia (Insecta: Hymenoptera). Mem. Entomol. Soc. Canada. 34:1-170. Thomson, C. G. 1876. Hymenoptera Scandinaviae. 4. Pteromalus (Svederus). pp. 193-259. Lund. Walker, F. 1836. Monographia Chalcidum. Entomol. Mag. 3:465-496. Received June 3, 1988; accepted March 13, 1989. J. New York Entomol. Soc. 97(3):358-362, 1989 SEED PREDATION BY A BRACONID WASP, ALLORHOGAS SP. (HYMENOPTERA) Margarete V. DE Macedo and Ricardo F. Monteiro Universidade Federal do Rio de Janeiro, I.B., Departamento de Ecologia, CP 68020, Ilha do Fundao, Rio de Janeiro, RJ, CEP 21941, Brasil Abstract. — record the first case of phytophagy among the Braconidae. The larvae of an unidentified species of Allorhogas have been observed feeding on Pithecellobium tortum (Le- guminosae) seeds in restinga of Barra de Marica, Rio de Janeiro, Brazil. Ecological and biological aspects of the species are described. In a study of seed predation of a leguminous plant, Pithecellobium tortum, Martins, we observed a braconid wasp, Allorhogas sp., feeding on immature seeds. This genus belongs in the Doryctinae, an ectoparasitoid subfamily in the Braconidae, which usually attack wood-boring and bark-mining Coleoptera larvae (Capek, 1970). The phytophagous habit was not recognized at first because the literature considers all the Braconidae to be parasites or hyperparasites upon other insects (Matthews, 1974). Despite the fact that the Ichneumonoidea have been studied for a long time only 10 to 20% of the described species have their hosts recorded (Iwata, 1976). Thus it is reasonable to expect that the number of known cases of phytophagy in the Braconidae may increase with more studies on the biology of this group. METHODS Samples of fifteen fruits were collected biweekly from February to June 1986 from each of seven P. tortum plants, a spreading shrub up to three meters in height. The study was conducted in the restinga (coastal scrub) of Barra de Marica in the city of Marica, Rio de Janeiro, Brazil (22°57'S, 42°50'W). About 200 fruits at varying stages of maturation were collected from various individuals and dissected under magnifying lenses to observe larval behavior. RESULTS The following insects emerged from fruits of P. tortum: Allorhogas sp. (Braconidae), Eurytoma sp. (Eurytomidae), Merobruchus boucheri Kingsolver (Bruchidae) and Eupelmus amicus Girault (Eupelmidae). Emergence occured in two different intervals. When fruits were immature, only braconids and eurytomids emerged. When fruits were ripe, bruchids and eupelmids emerged. The sequential emergence of these two groups had a slight overlap due to a time lag in the fruiting period of plants and in the development of fruits from the same plant (Fig. 1). It is noteworthy that bruchid eggs, which are laid on the pod surface, were only found on well developed fruits, after the emergence of Allorhogas and Eurytoma. As Allorhogas sp. and Eurytoma sp. are found simultaneously in the fruits of P. tortum, it was necessary to distinguish between their immature forms. The description 1989 BRACONID SEED PREDATION 359 Days after the beginning of the fruiting period Fig. 1 . Time variation of insect species obtained from P. tortum fruits each biweekly sample consisted of 105 fruits from seven trees. of the two different larvae and the obtention of adults from these described forms showed that the braconid larvae have sclerotized cephalic structures about the mouth- parts while the eurytomid larvae lack these structures. It was thus possible to identify the larvae easily, in the lab, even when they were newly hatched. In the laboratory we verified that all attacked immature seeds had the same kind of damage, which suggested that there was only one phytophagous species. This was confirmed by direct observations of Eurytoma larvae externally parasitizing braconid pupae (Fig. 2). The larva consumes its host within three days and pupates in the following day. This observation, however, served only to prove that Eurytoma could parasitize the braconid pupae, but did not prove that Allorhogas was phytophagous. In July 1988 we observed that almost all of fifty sampled P. tortum fruits from the restinga of Barra de Marica, wQiXhQv Allorhogas nor Eurytoma were found. Thirty young fruits were collected and exposed to Allorhogas adult females reared from P. 360 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Fig. 2. Sequence of development of the parasite larva, Eurytoma sp. (EU), on its host, Allorhogas sp. pupa (AL). Note the division on the seed— C. tortum seeds from another region (restinga of Arraial do Cabo, RJ, about 70 km from Marica). Two weeks later we dissected the fruits and found only Allorhogas larvae in the great majority of the seeds. BIOLOGY OF ALLORHOGAS SP. We observed larvae of allorhogas at different stages of development. Oviposition occurs on young, full-length fruits which thicken considerably before maturation. At this stage the seed is still small; it has a soft coat and a large amount of endosperm, but its embryo is as yet poorly developed. At the beginning of its development the braconid larva induces a division in the seed in which it is located and stays in one of the two parts (Fig. 2C). We noticed that even when a seed is attacked by two or more larvae, the divisions occur in such a way that there always is an undamaged region. In 98% of cases (N = 150) the embryo is found in this undamaged region. Initially, the larva feeds on endosperm, and later on the seed coat. The larva never attacks the embryo or the funicle. 1989 BRACONID SEED PREDATION 361 Close to pupation the larva spins a silk cocoon around itself. The newly formed pupa is yellow; later on, the eyes, antennae, and some regions of the thorax and abdomen become darker. Development is completed in 20 days. The adult chews a hole in the fruit with its mandibles and emerges. DISCUSSION According to Marsh (1979) host records indicate that some Allorhogas species are associated with gall making insects, especially cynipids. However, Marsh states that there is no confirmation that these species are parasites of the gall makers and that they are probably parasites on other insects. Guimaraes (1957, unpublished thesis) obtained a different undescribed species of Allorhogas (cited as A. muesebecki Guimaraes, 1957) from galls in Anemopaegma mirandum Alph. DC. (Bignoniaceae). Since this species was the only one to emerge from the galls studied, he concluded that the galls were induced by the braconid. Whitehead (1975) obtained a species of Allorhogas from fruits of Lysiloma and Albizzia (Leguminosae: Mimosoideae) and stated that the species could be a specialist parasite on Merobruchus, a bruchid obtained from the same fruits. By comparing the systems observed by Whitehead in Lysiloma and Albizzia with the one in P. tortum we find a clear similarity. It may thus be possible that the Allorhogas species found by Whitehead is also phytophagous. Extensive ecological and and taxonomic studies on Allorhogas are of high interest. These will improve our understanding of the biology and ecology of the Braconidae. As to the Eurytoma species found, the fact that its larvae were observed feeding on small larvae of Allorhogas sp., which are certainly not sufficient for their complete larval development, leaves open the possibility that the larvae complement their diet with plant tissue. Such an observation was made by Phillips (1927) for Eurytoma parva (Girault). ACKNOWLEDGMENTS This study received financial support from FINEP and CNPq scholarships to M. V. Macedo. V. Graf, P. M. Marsh, C. R. Brandao, T. M. Lewinsohn, G. W. Fernandes, and an anonymous reviewer offered valuable criticism of the manuscript. P. M. Marsh, R. W. Clarson, and J. Kingsolver provided the insect identifications. LITERATURE CITED Capek, M. 1970. A new classification of the Braconidae (Hymenoptera) based on the cephalic structures of the final instar larva and biological evidence. Can. Entomol. 102:846-875. Guimaraes, J. A. 1957. Contribui^ao ao estudo da Cecidologia Brasiliana. Thesis, UFRRJ, Brasil. Iwata, K. 1976. Evolution of Instinct— Comparative Ethology of Hymenoptera. Amerind Publishing Co., New Delhi. Marsh, P. M. 1979. Family Braconidae. In: K. V. Krombein, P. D. Hurd Jr., D. R. Smith, and B. D. Burks (eds.). Catalog of Hymenoptera in America North of Mexico, Vol. 1, Smithsonian Institution Press, Washington, D.C., 1,198 pp. Matthews, R. W. 1974. Biology of Braconidae. Ann. Rev. Ent. 19:15-32. 362 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Phillips, W. J. 1927. Eurytoma parva (Girault) Phillips and its biology as a parasite of the wheat jointworm, HarmoUta tritici (Fitch). Jour. Agr. Res. 34:743-758. Whitehead, D. R. 1975. Parasitic Hymenoptera associated with bruchid infested fruits in Costa Rica. J. Wash. Acad. Sci. 65:108-1 16. Received October 6, 1988; accepted January 25, 1989. BOOK REVIEWS J. New York Entomol. Soc. 97(3):363-364, 1989 Asa Fitch and the Emergence of American Entomology: With an Entomological Bibliography and a Catalog of Taxonomic Names and Type Specimens.— Jeffrey K. Barnes. The University of the State of New York, Albany, New York State Museum Bulletin No. 461, 1988. This is an admirable biography of “the first salaried professional entomologist in the United States [whose] career established the model for professional entomologists in the civil service.” (hi) Commencing in 1854, Fitch’s reports as New York State Entomologist established the problem centered approach which was followed by subsequent government entomologists. By the close of his career in the 1870’s, Fitch had amassed the largest collection relating to agricultural entomology in the country. Jeffrey Barnes, an employee of the New York State Museum, wrote this biography on the Sesquicentennial of the New York Geological and Natural History Survey (1836-1986). He utilized his intimate knowledge of the political and institutional developments of the New York survey, the state museum, the state cabinet, and other New York institutions to explain Fitch’s achievements within the scientific, educational, and agricultural context of the state and the nation. Drawing upon primary sources such as the Fitch correspondence and diaries housed in the archives of the Sterling Memorial Library of Yale University, the Museum of Comparative Zoology, the New York State Museum, plus extensive reading of the agricultural press, Barnes strikes a judicious balance in his discussion of Fitch the individual and Fitch the participant in American agricultural change and scientific advancement. The biography adds significant new information to what was known of Fitch from standard biographical sources like Arnold Mallis, American Entomologists. For ex- ample, Fitch’s appointment in 1854 as entomologist of the New York State Agri- cultural Society has often been cited as the first instance of a salaried “professional” entomologist in the United States. Barnes’ explanation of the appointment and the developments preceding it make it clear that Fitch’s activities in agricultural reform— specifically his agricultural survey of Washington County— must be seen as the pro- totype for the investigation of insect pests in the United States. The most puzzling aspect of Fitch’s life is his relative lack of contact with other American entomologists who organized their discipline. Fitch had little contact with the Entomological Society of Pennsylvania in the 1 840’s, the American Entomological Society in the 1860’s, and the Entomological Club of the American Association for the Advancement of Science in the 1870’s. Among American entomologists, who were typically prolific letter writers, Fitch was notorious for leaving letters unan- swered. Toward the end of his career, Fitch received visits from C. V. Riley, J. A. Lintner, and P. R. Uhler (1870) and John H. Comstock (1873) who wished to learn more about him and his collection, yet even these visits led to no lasting contact. Fitch, the pioneer, remained the loner among his colleagues. Barnes explains that Fitch’s neighbors’ perceived him as the eccentric “bug catcher of Salem,” but he could perhaps have also explained Fitch’s lack of involvement with other entomol- ogists. 364 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Of special interest to entomologists are the two appendices which list Fitch’s entomological publications and his contributions to entomological taxonomy. Barnes also discusses the unfortunate dismemberment and loss of much of Fitch’s extensive collection. We need more biographies like this one. Charles V. Riley, Benjamin D. Walsh, Asa Spring Packard, and John L. LeConte come to mind as major figures in nineteenth century American entomology for whom we need scholarly biographies. — Sorensen, University of Alaska Southeast, 11120 Glacier Highway, Juneau, Alaska. J. New York Entomol. Soc. 97(3):364-365, 1989 Insect Flight: Dispersal and Migration. — W. Danthanarayana, ed. 1986. Springer- Verlag, Berlin. This collection of papers covers a wide variety of topics on insect migration by flight, ranging from biochemical and physiological to ecological and evolutionary. By presenting viewpoints from very different perspectives, this volume succeeds in providing a more holistic view of the field. My only criticism is that care was not taken to make all papers accessible to as wide an audience as possible. There is a tendency to forget that technical terms, such as “hypertrehalosemia,” may not be familiar to non-physiologists, or that many of us do not know how to interpret a radar photo. On the positive side, the volume is permeated with awareness that field studies are essential for understanding insect migration. As Taylor notes (Chap. 20), “migration is not easy to create indoors.” A particularly good example of how lab studies, field behavior observations, and radar tracking can complement each other in presenting a unified picture of migration is found in Gatehouse’s chapter on the African ar- myworm (Chap. 9). Development of radar technology is responsible for many recent advances in our understanding of insect migration under natural conditions. This is attested by the fact that results from radar tracking are used in most chapters to illustrate one point or another, while three chapters (6, 13, and 16) are exclusively dedicated to it. Another recurring theme throughout the book is that insect migration is an ad- aptation for dealing with environments that vary in time and space. Gatehouse (Chap. 9) argues that rigid genetic determination of propensity to migrate evolves not when reliable environmental cues are absent, but when they are irrelevant. This situation may arise in a species with ubiquitous host plants, since the balance between the costs of staying and leaving is in favor of the latter, especially if the species is subject to heavy mortality from natural enemies. In a similar vein, Dixon and Howard (Chap. 10) review polymorphism in migratory propensity exhibited by many aphids. They show that this polymorphism among the members of a clone is programmed. While subscribing to the view that insect migration represents an adaptive syn- drome, Dingle (Chap. 2) nevertheless cautions that the knowledge of evolutionary and genetic aspects of insect migration is still in embryonic form. For example, we do not know why migrants are not selected out of many “pied paper” Lepidoptera (these insects migrate north in the spring, but are caught by winter before they can migrate south). Gibo (Chap. 1 2) treats the famous exception to this rule, the monarch 1989 BOOK REVIEWS 365 butterfly which possesses a remarkable directional control enabling it to return to its Mexican overwintering sites in the fall. Risks of dispersal, however, are very real. Many migrating insects are carried to hostile environments where they perish, such as snow-covered peaks (Edwards, Chap. 14). Curiously, the fallout of these “derelicts of migration” becomes the basis of a scavenger and predator community, and may hasten the recolonization of huge disturbed areas such as the blast zone of Mt. St. Helens. In a very interesting chapter, Farrow (Chap. 13) explains the concepts of the boundary-layer and synoptic scale meteorology, and how atmospheric processes help us understand migration of micro-insects (insects that are less than 1.5 cm in length). Maybe someone familiar with meteorology would not be impressed, but I found this chapter very instructive. By flying at night insects appear to utilize the nocturnal temperature inversion in the planetary boundary level. Several chapters deal with night-flying insects. Danthanarayana (Chap. 7) discusses the lunar periodicity in insect flight. Riley and Reynolds (Chap. 6) give an overview of various cues that may be used by night-flying insects for orientation, while Danthanarayana and Dashper (Chap. 8) focus on one of these cues: polarized light from natural sources. Mikkola (Chap. 1 1) describes the effect of wind on insect migrations into Finland. Endocrine stimulation of migratory behavior, and endocrine control of flight me- tabolism are discussed in Chap. 3 and Chap. 4, respectively. Four chapters deal with applied topics: Chap. 1 5 and 1 6 with migration in Heliothis zea and relatives. Chap. 17 with dispersal models of agricultural pests, and Chap. 18 with insects of public health importance. Several chapters are concerned with developing techniques for tracking migrating insects. Sounds produced by insects are important because wing- beat frequencies of insects can be detected by radar (Belton, Chap. 5). Lingren et al. (Chap. 19) describe the uses of night- vision equipment for observation of insect behavior at night. In summary, this volume will be of interest to many workers in the field of insect flight. While not attempting to give a comprehensive overview of all aspects of insect flight, this book (especially if read in combination with a more physiologically ori- ented volume of the same title {Insect Flight. Goldsworthy, G. J., and Wheeler, C. H. (eds.). 1989. CRC Press, Boca Raton) serves a valuable function by bringing together different perspectives on insect migration, ranging from biochemical to evolutionary. In this age of increasing specialization, such attempts should be en- couraged.—Turchin, So. Forest Exp. Station, Forest Service- USDA, Pineville, Louisiana 71360. J. New YorkEntomol. Soc. 97(3):365-366, 1989 Biogeography and Taxonomy of Honeybees. — Friedrich Ruttner. 1988. Springer- Verlag, New York, xii + 284 pp. $87.50 (hardcover). The biology and morphometries of honeybees have received increasing attention over the past decade because of the problems encountered in the Western Hemisphere with the Africanized honeybee. For such a small genus. Apis is one of the most widespread and, in human eyes, dominant groups of insects. However, up until 366 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Ruttner’s work, there has been no thorough, comparative source which summarizes information on the species and races of honeybees. The book is basically organized in 3 parts. First, a general overview of the genus Apis is given, which includes discussions of biology, phylogeny, and geographic vari- ation. Second, are detailed discussions of the distribution, morphology, biology, and to some extent physiology of each species. Finally, the last chapters, comprising nearly half the book, are devoted to discussions of each geographic/morphometric race of Apis mellifera L. recognized by the author. The text is clearly written but terse. In fact some discussions become almost telegraphic. I find this a pleasant change from similar types of studies which tend to expand a small amount of information into a large amount of verbiage. However, some chapters, particularly the introductory ones, would have benefitted from more extensive discussions. Somehow one paragraph on the ecology of honeybees seems inadequate. It is easy to see where the author’s interests lie, as comparatively speaking, he waxes poetic in his discussions of the races of Apis mellifera. Overall the book is well organized, beautifully illustrated and informative. The chapters on species and races of Apis provide information on each of these groups in a clearly organized, comparative manner. For each species there is a dis- cussion of morphology, particularly that of the male terminalia; distribution; behav- ior, including comparisons of dance language, foraging activities, swarming and col- ony defenses; morphometries; physiology; parasites, and relationships with man. Discussions of mellifera races are arranged by general geographic region, and include taxonomic problems, precise distributions, morphometric characteristics and be- havioral peculiarities. I have several complaints with this work. The first is a matter of nomenclature. Throughout the text Ruttner refers to “races of honeybees” and yet the names he uses are clearly subspecies. He even proposes the name Apis mellifera macedonica ssp. nova as a new subspecies, but in the diagnosis he calls it a race. I find this confusing, as strictly speaking, races are not given official names according to the Zoological Code of Nomenclature. There is also the phrase: “two new Braula species of the genus Megabraula . . .” which is unclear and may just be a problem of trans- lation. Second, the study of Winston and Michener (1977) is cited repeatedly as the final word on the phylogenetic relationships of the apid subfamilies. The later study by Kimsey ( 1 984) made some corrections of the earlier paper and provided additional characteristics. Ruttner actually does cite this 1 984 paper in his references but I could find no reference to it in the text. Finally, to my knowledge, no one has been able to demonstrate resource competition between honeybees and native bees. The state- ment: “Therefore it seems that the bigger [Apis] species avoid disastrous competition by shifting to other more distant [food] sources” has to be my favorite. I do have one final criticism, which is addressed to the publisher: the price of this volume is outrageous, and will discourage many interested individuals from buying a copy .—Lynn S. Kimsey, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138. LITERATURE CITED Kimsey, L. S. 1984. A re-evaluation of the phylogenetic relationships in the Apidae. Syst. Ent. 9:435-441. 1989 BOOK REVIEWS 367 Winston, M. L. and C. H. Michener. 1977. Dual origin of highly social behavior among bees. Proc. Natl. Acad. Sci. USA 74:1 135-1 137. J. New YorkEntomol. Soc. 97(3):367-370, 1989 Tree and Shrub Insects of the Prairie Provinces. — W. G. H. Ives and H. R. Wong. 1988. Information Report NOR-X-292, Northern Forestry Centre, Canadian For- estry Service, Edmonton, Alberta, xi + 327 pp. This pictorial guide to identification of arthropods that feed on trees and shrubs in the Prairie Provinces culminates the authors’ work in that region of Canada; each has devoted about 40 years to the study of forest entomology. They have produced an outstanding manual, one useful to researchers, extension entomologists, horti- culturists, and plant inspectors working in regions well beyond the area of coverage. Species affecting native trees and shrubs of the forest vegetation are emphasized (some common exotic plants of field and farm shelterbelts are included), but this volume will interest those who work with ornamentals. Tree and Shrub Insects of the Prairie Provinces is aesthetically pleasing through- out—from its striking and unique cover illustration (larva of forest tent caterpillar on a black background), well-conceived, consistent layout of species write-ups, to the superb color plates. Included among the 1 17 plates featuring about 1,100 pho- tographs are some of the best images of insects and their injury that we have seen. Depth of field and lighting are remarkably consistent, and the reproduction is ex- cellent. Even their poorest figures, perhaps 45D (carpenter ant nest) and 61G (gypsy moth larva), are better than the majority of those in some publications. Despite numerous color plates, the book is functional, not extravagant. It was laid out with the reader in mind; as an example, the 8" x 11" pages have margins sufficient for making notes. Paper stock is coated and heavy enough to minimize “see-through”; the binding (perfect, double score hinged) should hold up under extensive use. A one-page introduction covers objectives and organization of the manual. Intro- ductory material also includes line drawings of adult and immature insects showing various structures (setae of the geometrid larva in Figure IH cannot be seen, and ocelli might have been indicated for some larvae) and a short glossary. Cremaster is defined as the “terminal abdominal segment” of a pupa but actually is the apex or hooked process of the last segment. Insects that feed on conifers and on hardwoods are segregated in the table of contents and arranged in each category by plant part attacked (e.g., seeds and cones, foliage, buds or shoots). Mites, caterpillars, sawffies, beetles, and other groups are broken out under plant parts, and each arthropod group is further subdivided to facilitate identification. For instance, lepidopterans attacking conifer foliage are sep- arated into budworms, caterpillars, loopers, miscellaneous larvae, and tube makers. Caterpillars, for example, are further subdivided into the spruce harlequin and its allies, hairy or spiny larvae, and colorful or bizarre larvae. 368 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Some 600 species are covered in the book’s main portion. A dark green page precedes the section on arthropods of conifers; a paler green page introduces species of hardwoods so that a reader can locate these major sections in thumbing through the book. One page (always the recto) is devoted to each group of species associated with various parts of conifers or hardwoods. Headings are the same throughout: Distribution, Hosts, and Importance; Life Cycle and Appearance; Damage; and Bib- liography (numbers only, which refer to the 657 references cited in the terminal bibliography). Each page of text is accompanied by a color plate on the facing page (verso), making it convenient for the reader. An increase in type size from that used (between 8 and 9 point) would have made the text easier to read but, owing to the length of some write-ups, was not feasible with their one-page format (a decrease in leading might have allowed 10-point typeface to be used). Information given for each species is necessarily brief but useful and generally accurate. The writing style is good, although on several pages an irritatingly large number of sentences begin with “The,” sometimes five or six in succession; in many cases its use is unnecessary. There are fewer typographical errors and misspellings than in most books of comparable size; we found “damge” (p. 7), “Osborne” (rather than Osborn) on p. 117, and “sp.” (p. 97) when the plural spp. was needed. Latin names of most taxa, apparently those under discussion, are in bold italics, whereas those noted in passing or not being emphasized are merely italicized. Usage occa- sionally seems inconsistent and might have been explained in the introduction. Com- mon names for a few species differ from those used in other Canadian publications, for instance, “eastern pine shootborer” and “pine rootcollar weevil” (rather than shoot borer and root collar as in Rose and Lindquist, 1973). A few other minor problems we noted were redundancies like “oval in shape,” “reddish in color,” and similar constructions. For Figure 23C, the legend notes “larval case at arrow,” but the arrow is missing. An inconsistency occurs in reference to length of the pentatomid Banasa dimidiata: 7-10 mm on p. 65 but 8.5-1 1 mm on p. 119. An agromyzid fly is referred to inappropriately as a “midge” (p. 213), and it is inaccurate to characterize the life history of the tarnished plant bug, Lygus lineolaris, as similar to that of the boxelder bug, Leptocoris (or Boisea) trivittatus (p. 1 19). The statement that each female aphid lays a single egg on host tissue (p. 107) is misleading when applied to all species under discussion (mainly aphidines); it is accurate in reference to the Pemphiginae. Coverage of the book, as expected, reflects the authors’ research interests and is most complete for defoliators, notably lepidopteran and sawfly larvae. Beetles and most other groups are covered adequately, but treatment of the Homoptera and Heteroptera seems spotty. In the former group only one leafhopper associated with hardwoods is mentioned; among Heteroptera, only three mirids are listed, but nu- merous species are associated with shrubs and trees in the Prairie Provinces. Several other heteropterans could have been included, e.g., the coreid Leptoglossus occiden- talis Heidemann, a pest of conifer seed, and the lygaeid Kleidocerys resedae (Panzer), which develops extensively in birch catkins. In addition, only one eriophyid species associated with conifers is included. The bibliography is not meant to be exhaustive, but several key references on particular groups are omitted, e.g., Bailey (1951) for Tingidae and Wood (1982) for Scolytidae. Also useful might have been the guide to plant abnormalities caused by 1989 BOOK REVIEWS 369 eriophyid mites (Keifer et al., 1 982) and booklet on insect and mite galls of the Pacific Northwest (Larew and Capizzi, 1983). Particularly surprising is the omission of several relevant Canadian works, including Rockburne and Lafontaine (1976), Kelton (1980), McAlpine et al. (1981), and Hamilton (1983, 1985). Many of these omissions are from the ’80’s and, although a few works from 1 985- 1987 are cited, we suspect the authors decided not to incorporate all pertinent lit- erature published during the latter stages of manuscript preparation. A cutoff date for literature perhaps could have been mentioned in the introduction. Three useful, easy-to-read indexes conclude the book: taxonomic, diagnostic, and insect. Author names might have been spelled out in the taxonomic index instead of using cryptic abbreviations like “K. & Y.” or “Bsk.”; complete names, however, are provided in the text. The list of arthropod names is uncommonly free of mis- spellings; it appears that the authors tried to obtain current names. In checking the first group appearing in the taxonomic index — Eriophyidae, for which nomenclatural confusion abounds— we found only a few problems. Aceria fraxinijlora (Felt) is a junior synonym of A. fraxinivorus (Nalepa), calaceris Keifer should be placed in Aceria rather than Eriophyes, parentheses should be added to the author’s name in A. parapopuli (Keifer), and parentheses should be deleted from authors’ names in E. emarginatae Keifer and E. padi Nalepa (J. W. Amrine, Jr., personal communication). The problems and perceived deficiencies just noted are mostly trivial— they do not seriously detract from what is an outstanding volume. But with books of fewer pages and no color sometimes selling for more than $100, what does this quality reference with extraordinary color plates cost? It is available at no charge— and we would recommend it enthusiastically even if it sold at a typical market price.— /I. G. Wheeler, Jr., Bureau of Plant Industry, Pennsylvania Department of Agriculture, Harrisburg, Pennsylvania 17110, and Gregory A. Hoover, Department of Entomology, Pennsylvania State University, University Park, Pennsylvania 16802. LITERATURE CITED Bailey, N. S. 1951. The Tingoidea of New England and their biology. Entomol. Amer. 31:1- 140. Hamilton, K. G. A. 1983. Introduced and native leafhoppers common to the Old and New worlds (Rhynchota: Homoptera: Cicadellidae). Can. Entomol. 1 15:473-51 1. Hamilton, K. G. A. 1985. Leafhoppers of ornamental and fruit trees in Canada. Agric. Can. Publ. 1779/E. 71 pp. Keifer, H. H., E. W. Baker, T. Kono, M. Delfinado and W. E. Styer. 1982. An illustrated guide to plant abnormalities caused by eriophyid mites in North America. U.S. Dep. Agric., Agric. Res. Serv. Agric. Handb. 573, 178 pp. Kelton, L. A. 1980. The Insects and Arachnids of Canada, Part 8. The Plant Bugs of the Prairie Provinces. Res. Branch Agric. Can. Publ. 1703. 408 pp. Larew, H. and J. Capizzi. 1983. Common Insect and Mite Galls of the Pacific Northwest. Oregon State Univ. Press, Corvallis, 80 pp. McAlpine, J. F., B. V. Peterson, G. E. Shewell, J. R. Vockeroth and D. M. Wood, eds. 1981. Manual of Nearctic Diptera. Vol. 1. Res. Branch Agric. Can. Monogr. 27, 674 pp. Rockburne, E. W. and J. D. Lafontaine. 1976. The Cutworm Moths of Ontario and Quebec. Res. Branch Agric. Can. Pub. 1593, 164 pp. 370 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(3) Rose, A. H. and O. H. Lindquist. 1973. Insects of Eastern Pines. Dep. Environ. Can. For. Serv. Publ. 1313, 126 pp. Wood, S. L. 1 982. The bark and ambrosia beetles of North and Central America (Coleoptera: Scolytidae), a taxonomic monograph. Great Basin Nat. Mem. 6, 1359 pp. INSTRUCTIONS TO AUTHORS The Journal of the New York Entomological Society is devoted to the advancement and dissemination of knowledge of insects and related taxa. The costs of publishing the Journal are paid by subscriptions, membership dues, page charges, and the proceeds from an endowment established with bequests from the late C. P. Alexander and Patricia Vaurie. The Journal will consider for publication manuscripts of any length dealing with original research in entomology. Longer papers will be printed as articles, shorter ones as “scientific notes.” Book reviews will be solicited by the Book Review Editor. Manuscripts should be submitted in duplicate to: Dr, Randall T. Schuh, Editor, Journal of the New York Entomological Society, c/o Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024. All material must be double-spaced (including references, tables, captions, etc.) and prepared in the format of a recent issue of the Journal. Longer manuscripts intended for submission as articles should be accompanied by a brief abstract. Footnotes should be avoided. Tables should be prepared as separate pages; they should be kept to a minimum because of the high cost of typesetting, but may be submitted as photographically reproducible material (see below). The list of references is headed “Literature Cited” and should follow the format indicated in the CBE Style Manual or as found in a recent issue of the Journal. Illustrations (originals whenever possible) should be submitted flat, never rolled or folded, with the manuscript. 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Society members will be charged a fee of $20.00 per printed page and $5.00 per plate of figures. Non-members will be charged $55.00 per printed page and $5.00 per plate of figures. Member authors who do not have institutional funds may petition to the Society for waiver of page charges for no more than eight pages on a once a year basis. Because of limited funds, all such requests will be handled on a first-come first-serve basis. Authors will receive a reprint order blank with the proofs. Reprints are ordered directly from the printer with no benefit accruing to the Society. Journal of the New York Entomological Society VOLUME 97 JULY 1989 NO. 3 CONTENTS Annotated checklist of the Thysanoptera of Bermuda Sueo Nakahara and Daniel J. Hilburn 25 1-260 Annotated checklist of the whiteflies (Homoptera: Aleyrodidae) of Bermuda Sueo Nakahara and Daniel J. Hilburn 261-264 Blissus breviusculus: New distribution records of a little-known chinch bug (Het- eroptera: Lygaeidae) A. G. Wheeler, Jr. and Jonathan E. Fetter 265-270 Three new species of Lincus (Hemiptera: Pentatomidae) from palms L. H. Rolston 271-276 Reconstitution of Coleometopini, Tenebrionini and related tribes in America north of Colombia (Coleoptera: Tenebrionidae) John T. Doyen 277-304 Pityogenes bidentatus (Herbst), a European bark beetle new to North America (Coleoptera: Scolytidae) E. Richard Hoebeke 305-308 New Trichoptera from Alabama S. C. Harris 309-316 An unusual black fly (Diptera: Simuliidae), representing a new genus and new species B. V. Peterson 317-331 Butterfly exploitation of an ant-plant Mutualism: Adding insult to herbivory P. J. DeVries and I. Baker 332-340 Sensory structures on the ovipositor of the ball gall fly Eurost a solidaginis (Fitch) (Diptera: Tephritidae) Edward Ritter and Carey E. Vasey 341-346 Review of Nearctic Rhicnocoelia and Callimerismus with a discussion of their phylogenetic relationships (Hymenoptera: Pteromalidae) Steven L. Heydon 347-357 Seed predation by a braconid wasp, Allorhogas sp. (Hymenoptera) Margarete V. de Macedo and Ricardo F. Monteiro 358-362 Book Reviews Asa Fitch and the Emergence of American Entomology: With an Entomological Bibliography and a Catalog of Taxonomic Names and Type Specimens Conner Sorensen 363-364 Insect Flight: Dispersal and Migration Peter Turchin 364-365 Biogeography and Taxonomy of Honeybees Lynn S. Kimsey 365-366 Tree and Shrub Insects of the Prairie Provinces A. G. Wheeler, Jr. and Gregory A. Hoover 367-370 Vol. 97 OCTOBER 1989 No. 4 Journal of the New York Entomological Society (ISSN 0028-7199) Devoted to Entomology in General JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Editor: Randall T. Schuh, Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024 Book Review Editor: David A. Grimaldi, Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024 Publications Committee: Louis Trombetta, St. Johns University, Queens, New York, Chairman; Alfred G. Wheeler, Jr., Pennsylvania State Department of Agriculture, Harrisburg; Joseph M. Cerreta, St. Johns University, Queens, New York. The New York Entomological Society Incorporating The Brooklyn Entomological Society President: Dennis J. Joslyn, Department of Biology, Rutgers University, Camden, New Jersey 08102 Vice President: Durland Fish, Medical Entomology Laboratory, New York Medical College, Armonk, New York 10504 Secretary: Richard Falco, Westchester County Health Department, White Plains, New York 10601 Treasurer: Louis Sorkin, Department of Entomology, American Museum of Natural History, New York, New York 10024 Trustees: Class of 7955— Henry M. Knizeski, Jr., Mercy College, Dobbs Ferry, New York; Michael D. Schwartz, American Museum of Natural History, New York, New York; Class of 7959— Christine Falco, West- chester County Health Department, White Plains, New York; James S. Miller, Department of Entomology, American Museum of Natural History, New York, New York. Annual dues are $23.00 for established professionals with journal, $10.00 without journal, $15.00 for students with journal, $5.00 without journal. Sustaining memberships are $53.00 per year, institutional memberships are $125.00 per year, and life memberships are $300.00. Subscriptions are $40.00 per year domestic and $45.00 foreign. All payments should be made to the Treasurer. Back issues of the Journal of the New York Entomological Society, the Bulletin of the Brooklyn Entomological Society, Entornologica Americana, the Torre-Bueno Glossary of Entomology and other Society publications can be purchased from Lubrecht and Cramer, RD 1, Box 244, Forestburgh, New York 12777. Meetings of the Society are held on the third Tuesday of each month (except June through September) at 7 p.m. in the American Museum of Natural History, Central Park West at 79th Street, New York, New York. Mailed March 20, 1990 The Journal of the New York Entomological Society (ISSN 0028-7 199) is published 4 times per year (January, April, July, October) for the Society by Allen Press, Inc., 1041 New Hampshire, Lawrence, Kansas 66044. Second class postage paid at New York. New York and at additional mailing office. Postmaster: Send address changes to the New York Entomological Society. % American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024-5192. Known office of publication: American Museum of Natural History, New York, New York 10024. Journal of the New York Entomological Society, total copies printed 700, paid circulation 602, mail subscription 602, free distribution by mail 19, total distribution 621, 79 copies left over each quarter. THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER. NOW AVAILABLE ! Harper Lyme Disease Poster Charley Harper is a nationally known wildlife artist who has done commissioned work for the Audubon Society, National Park Service, Cincinnati Zoo, and many other conservation organizations. His limited edition prints, published by Framehouse Gallery of Houston, Texas, have frequently featured insects. Notable among these are "Water Striders" (1969), "Beetle Battle" (1971), and "The Last Aphid" (1981) which have become collector's favorites. The New York Entomological Society is proud to present this Harper poster on Lyme disease, which combines ad and science to deliver an important public health message. The printing is high quality and is appropriate for framing. Send $12.00 to Harper Poster, New York Entomological Society, c/o American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024. A Revised and Expanded Edition o THE TORRE-BUENO GLOSS ARi OF ENTOMOLOGY THE NEW YORK ENTOMOLOGICAL SOCIETY announces AMERICAN MUSEUM OF NATURAL HISTORY in cooperation with the Compiled by Stephen A. Nichols Managing Editor Randall T. Schuh First published in 1937, the Glossary of Entomology by J. R. de la Torre-Bueno, with some 8500 entries, has long been the standard for usage of entomological terms in the English language. Now available, after 50 years, is a revised and expanded edition, containing approximately 16,000 terms in 840 pages. The new Glossary serves as a basic reference in general entomology, listing thousands of new terms and definitions in the fields of insect morphology, behavior, classification, physiology, medical entomology, pathology, and applied entomology. In conjunction with the Glossary the Society has prepared an entomological spell- checker file containing over 9000 terms. This ASCII file can be added to most word processors and serve as part of your personal dictionary of technical terms. Please send payment for The Glossary made out to the New York Entomological Society at the address indicated below. Prices are as follows: TO ORDER: Glossary (member price) Glossary (non-members and libraries) Entomological Spell-Checker IBM Diskette 3-1/2 inch or 5-1/4 inch Mac Diskette 40.00 45.00 15.00 Send payment to: New York Entomological Society c/o Department of Entomology American Museum of Natural History Central Park West at 79th Street New York, New York 10024 J. New York Entomol. Soc. 97(4);37 1-393, 1989 ATLAS OF ANTENNAL TRICHOBOTHRIA IN THE PACHYNOMIDAE AND REDUVIIDAE (HETEROPTERA) Pedro W. Wygodzinsky and Sarfraz Lodhi Department of Entomology, American Museum of Natural History, New York, New York 10024 Abstract.— KniQnndiQ of Reduviidae comprising adults of 99 species in 19 subfamilies and nymphs of 25 species in 7 subfamilies, and one species of Pachynomidae were examined using compound light and scanning electron microscopy. Figures are presented indicating the numbers and distribution of trichobothria, the range being from 1-20 or more. Trichobothria are well known in the Heteroptera, occurring ventrally on the ab- domen in the Trichophora (Schaefer, 1975), on the femora in the Miridae (Schuh, 1975), on the scutellum in the Prostemmatinae (Nabidae) (Carayon, 1970), and elsewhere in other groups of true bugs, as well as in other groups of insects and arthropods (see Schuh, 1975, for a brief survey). Those occurring on the second antennal article of the Reduviidae were first described in detail by Lent and Wy- godzinsky (1979) for the Triatominae. Their first record of occurrence in the Pachy- nomidae was that of Carayon and Villiers (1968), who referred to the structures on the third antennal article as trichobothrioform sensilla, pending histological inves- tigation; they also documented the occurrence of trichobothria on the abdomen in this group. In the present paper we document the occurence of antennal trichobothria in adults of one species of Pachynomidae and 99 species of Reduviidae and nymphs of 25 species of Reduviidae. Because of the limited variation in numbers and distribution of the trichobothria and the limited understanding of phylogenetic relationships among the currently recognized subfamilies of Reduviidae, we have not attempted a detailed analysis of the taxonomic value of these structures.* Nonetheless some very general conclusions can be drawn. The presence of antennal trichobothria in the Pachynomidae and Reduviidae sug- gests a possible sister group relationship between the two taxa, as proposed by Schuh (1979). Within the Pachynomidae, which have five apparent antennal articles, the position of the trichobothrium suggests that the second article— the pedicel— is the one that has undergone subdivision. The occurrence of a single trichobothrium in the Pachynomidae and some first instar Reduviidae (no information is available for nymphal Pachynomidae) suggests that the primitive number is probably one. Con- ‘ Furthermore, because of the failing health of the senior author during the data-gathering phase of the project, it was not possible to prepare a more complete analysis of the available information. The results presented here are offered in the anticipation that future workers will use them as a base for additional work elucidating relationships between the Pachynomidae and Reduviidae, and among the higher groups of Reduviidae, on the basis of antennal trich- obothria as well as other structures. 372 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Fig. 1 . SEM micrographs of Pachynomidae. A-D. Aphelonotus sp. A. 1 40 x . B. 2,500 x . C. 750x. D. 650x. trary to Lent and Wygodzinsky (1979) it is evident that some first instar Reduviidae have more than one trichobothrium (e.g., Amphibolus Venator, Figs. IL-O), and it appears that all Reduviidae have antennal trichobothria as adults. The figures presented below are organized by subfamily, those groups with the lowest numbers first, those with higher numbers toward the end. To facilitate use of this work, we present the following list, organized alphabetically by subfamily and species, with the figure numbers. LIST OF TAXA STUDIED Family Pachynomidae Aphelonotus sp. (Figs. 1 A-D; 60-Q). Family Reduviidae Subfamily Apiomerinae Agriocoris sp. (Fig. 1 7H, I). Amauroclopius sp. (Fig. 17L). Apiomerus sp. (Fig. 1 7J, K). Calliclopius sp. (Fig. 1 7M). 1989 ANTENNAL TRICHOBOTHRIA 373 Fig. 2. SEM micrographs of Reduviidae. A, B. Eupheno pallens. A. 2,200 x. B. 440 x. C. Leogorus litura, 2,000 x . D. Phymata georgiensis. Heniartes erythromerus Spinola (Fig. 1 7F). Heniartes flavicans (Fabr.) (Fig. 1 7D, E). Heniartes maestralis Fracker & Bruner (Fig. 1 7B, C). Heniartes sp. (Fig. 17G). Manicocoris sp. (Fig. 17A). Micrauchenus sp. (Fig. 1 7N). Subfamily Centrocneminae Centrocnemis sp. (Fig. 6L). Subfamily Cetherinae Cethera sp. (Fig. lOA-C). Eupheno pallens (Laporte) (Figs. 2 A, B; lOH). Eupheno sp. (Fig. 2M). Subfamily Ectrichodinae Bayerus sp. (Fig. 1 3K, L). Cricetopareis tucumana (Berg) (Figs. 3C, D; 13B, C). 374 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) C D Fig. 3. SEM micrographs of Reduviidae. A, B. Barce sp. C-D. Cricetopareis tucumana. C. 400 X. D. 1,600 X. Daraxa sp. (Fig. 1 3G, H). Ectrichodia crux (Thunberg) (Fig. 1 3D, E). Katanga etiennei Schouteden (Fig. 13M). Pothea lugens (Fabr.) (Fig. 1 3 A). Racelda sp. (Fig. 1 31, J). Vilius sp. (Fig. 13F). Zirta limbata Breddin (Fig. 18C). Subfamily Emesinae Barce fraternus (Say) (Fig. 9D, E). Barce sp. (Fig. 2A, B). Bergemesa brachmanni (Berg) (Fig. 9C). Bettyella marita Wygodzinsky (Fig. 9B). Empicoris rubromaculatus (Blackburn) (Fig. 18 A, B). Ploiaria chilensis (Philippi) (Fig. 9A). Stenolemus pallidipennis McAtee & Malloch (Fig. 8L). 1989 ANTENNAL TRICHOBOTHRIA 375 E F Fig. 4. A. SEM micrographs of Reduviidae. Pygolampis sp., 1 00 x . B. Reduvius personatus, 1 ,900 X . C. Sphaeridops amoenus, 450 x . D. Sphaeridops amoenus, 1 , 1 00 x . E. Tribelocephalus sp., 400 X. F. Tribelocephalus sp., 2,000 x. Subfamily Harpactorinae Amphibolus Venator {¥\gs. 13H-I; 15I^P). Arilus sp. (Figs. 13L, M; 15U). Atrachelus cinereus (Fabr.) (Fig. 13C, D). 376 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Bactrodes femoratus (Fabr.) (Fig. 1 3N, O). Castolus ferox Banks (Fig. 1 5 J-K). Castolus lineatus Maldonado (Fig. 15R, S). Erbessus sp. (Fig. 16P). Havinthus rufovarius Bergroth (Fig. 1 6K). Notocyrtus (Figs. 15Q; 16E-G). IRepipta flavicens (Fig. 1 5T). Rhaphidosoma didiera Jeannel (Fig. 151). Ricolla quadrispinosa (Linne) (Fig. 15A-F). Sinea diadema (Fabr.) (Figs. 1 6 A, B; 1 8G). Yolinus sp. (Fig. 1 6J). Zelus exsanguis Stal (Fig. 15G-H). Subfamily Microtominae Homalocoris sp. (Fig. 8D). Microtomus gayi (Spinola) (Fig. 8C). Microtomus purcis (Drury) (Fig. 8A, B). Subfamily Phimophorinae Phimophorus spissicornis Bergroth (Fig. 18K). Subfamily Phymatinae Anthylla nervosopunctata Signoret (Fig. 6E). Macrocephalus tuberosus Westwood (Fig. 6G). Pamgreuocoris aethiopicus Carayon (Fig. 6F). Phymata georgensis Melin (Fig. 2D). Phymata pennsylvanica Handlirsch (Fig. 6A-D). Phymata stali Melin (Fig. 6H-I). Subfamily Physoderinae Physoderes sp. (Fig. 6J, K). Subfamily Peiratinae Brachysandalus ephippiger V\f\n\Q (Fig. 14L). Catamiarus brevipennis (Serville) (Fig. 14F). Chryxus sp. (Fig. 6M). Ectomocoris cordatus (Wolff) (Fig. 14H, I). Eusius rubricosus StM (Fig. 18E). Melanolestes abdominalis (Herrich-Schaeffer) (Fig. 14G). Nalata sp. (Fig. 14M). Peirates sanctus (F.) (Fig. 14E). Psophis sp. (Fig. 6N). Rasahus biguttatus (Say) (Fig. 14C). Rasahus hamatus (Fabr.) (Fig. 14J, K). Rasahus sulcicoUis (Serville) (Fig. 14A, B). 1989 ANTENNAL TRICHOBOTHRIA 377 Sirthenea flavipes (Stal) (Fig. 14D). Tydides rufus (Serville) (Fig. 14N). Subfamily Reduviinae Inara flavopicta Stal (Fig. lOD). Leogorrus formicarius (Fabr.) (Fig. 1 IF, G). Leogorrus litura (Fabr.) (Figs. 2C; 1 II^N). Microlestria sp. (Fig. 1 IR, S). Neivacoris sp. (Fig. 5N). Pasira perpusilla (Walker) (Fig. 1 IH, I). Reduvius personatus (Linne) (Figs. 4B; 1 1 J, K). Sphedanovarus camerunensis (Breddin) (Fig. 1 10~Q). Velitra philippina Stal (Fig. 1 1 A, B). Zelurus circumcinctus (Hahn) (Fig. 1 ID, E). Zelurus spinidorsis (Gray) (Fig. 1 8D). Zelurus weyrauchi Lent & Wygodzinsky (Fig. 1 1C). Subfamily Saicinae Gallobelgicus sp. (Fig. 9G, H). Oncer otrachelus acuminatus (Say) (Fig. 91, J). Polytoxus sp. (Fig. 18H). Saica apicalis Osborn & Drake (Fig. 9F). Subfamily Salyavatinae Salyavata nigrofasciata Costa Lima (Fig. 8K). Salyavatinae sp. (Fig. 8F-J). Subfamily Sphaeridopinae Sphaeridops amoenus (Lepeletier & Serville) (Figs. 4C, D; 8E). Subfamily Stenopodainae Apronius sp. (Fig. 1 8F). Ctenotrachelus sp. (Fig. 1 8 J). Kodormus bruneosus Barber (Fig. 7J, K). Oncocephalus nubilus Van Duzee (Fig. 7A-C). Pnirontis languida StM (Fig. 7F-I). Pygolampis sp. (Fig. 4A). Staccia sp. (Fig. 181). Stenopoda cinerea Laporte (Fig. 7D, E). Subfamily Triatominae Belminus peruvianus Herrer, Lent & Wygodzinsky (Fig. 1 21). Eratyrus mucronatus StM (Fig. 5L). Panstrongylus chinai (Del Ponte) (Fig. 1 2A, B). 378 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Panstrongylus megistus (Burm.) (Fig. 12C). Paratriatoma hirsuta Barber (Fig. 5P). Psammolestes arthuri (Pinto) (Fig. 50). Rhodnius neivai Lent (Fig. 5J). Rhodnius neglectus Lent (Fig. 5K). Triatoma barberi Usinger (Fig. 5H). Triatoma infestans (King) (Fig. 12D, E). Triatoma lecticularia (StM) (Fig. 5D). Triatoma lenti Sherlock & Serafim (Fig. 51). Triatoma longipennis Usinger (Fig. 5C). Triatoma maculata (Erichson) (Fig. 5B). Triatoma platensis Neiva (Figs. 5E; 12G, H). Triatoma protract a (Uhler) (Fig. 5G). Triatoma rubida (Uhler) (Fig. 5A). Triatoma vitticeps (Stal) (Figs. 5F; 12F). Subfamily Tribelocephalinae Tribelocephalus sp. (Fig. 4E, F). Tribelocephalinae sp. (Fig. 101). Subfamily Vescinae Pessoaia Ipiratoides Costa Lima (Fig. lOJ-M). Vescia sp. (Fig. ION, O). 1989 ANTENNAL TRICHOBOTHRIA 379 Fig. 5. Reduviidae— 5th instar nymphs. A. Triatoma rubida. B. Triatoma maculata. C. Triatoma longipennis. D. Triatoma lecticularia. E. Triatoma platensis. F. Triatoma vitticeps. G. Triatoma protracta. H. Triatoma barberi. I. Triatoma lenti. J. Rhodnius neivai. K. Rhodnius neglectus. L. Eratyrus mucronatus. M. Eupheno sp. N. Neivacoris sp. O. Psammolestes arthuri. P. Paratriatoma hirsuta. 380 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Fig. 6. Reduviidae—Phymatinae. A-D. Phymata pennsylvanica, S and 9. E. Anthylla ner- vosopunctata. F. Paragreutus aethiopicus. G. Macrocephalus tuberosus. H, I. Phymata stall. Reduviidae— Physoderinae. J, K. Physoderes sp. Reduviidae— Centrocneminae. L. Centroc- nemus sp. Reduviidae— Piratinae. M. Chryxus sp. N. Psophis sp. Pachynomidae. O-Q. Aphe- lonotus sp., article 3. 1989 ANTENNAL TRICHOBOTHRIA 381 Fig. 7. Reduviidae— Stenopodainae. A-C. Oncocephalus nubilis. D„E. Stenopoda cinerea. F-I. Pnirontis languida, S and 2. J, K. Kodormus bruneosus. 382 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Fig. 8. Reduviidae— Microtominae. A, B. Microtomus purcis. C. Microtomus gayi. D. Hom- alocorissp. Reduviidae— Sphaeridopinae. E. Sphaeridops amoenus. Reduviidae— Salyavatinae. F-J. Salyavatinae, lst-5th instars. K. Salyavata nigrofasciata. Reduviidae— Emesinae. L. Steno- lemus pallidipennis. 1989 ANTENNAL TRICHOBOTHRIA 383 Fig. 9. Reduviidae— Emesinae. A, Ploiaria chilensis. B, Bettyella marita. C. Bergemesa bachmanni. D, E. Barce fraternus. Reduviidae— Saicinae. F. Saica apicalis. G, H. Gallobelgicus sp. I, J. Oncer otrachelus acuminatus. 384 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) I Fig, 10. Reduviidae— Cetherinae. A-C. Cethera sp., 2 and $. Reduviidae— Reduviinae. D. Inara flavopicta. Reduviidae— Cetherinae. E-H. Eupheno pallens, $ and 2. 1. Tribelocephalinae sp. Reduviidae— Vesciinae. J-M. IPessoaia piratoides. N, O. Vescia sp., <3 and 2. 1989 ANTENNAL TRICHOBOTHRIA 385 Fig. 11. Reduviidae— Reduviinae. A, B. Velitra philippina. C. Zelurus weyrauchi. D, E. Zelurus circumcinctus. F. Leogorus formicarius. G. Leogorus formicarius, 5th instar. H, I. Pasira perpusilla. J, K. Reduvius personatus. l^N. Leogorus litura. O-Q. Sphedanovarus camerunensis. R, S. Microlestria sp. 386 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Fig. 12. Reduviidae— Triatominae. A, B. Panstwngylus chinai. C. Panstrongylus megistus. D, E. Triatoma infestans. F. Triatoma vitticeps. G, H. Triatoma platensis. I. Belminus peru- 1989 ANTENNAL TRICHOBOTHRIA 387 Fig. 1 3. Reduviidae— Ectrichodiinae. A. Pothea lugens. B, C. Cricetopareis tucumana, S and 2. D, E. Ectrichodia crux, S and 2. F. Vilius sp. G. H, Daraxa sp. I, J. Racelds sp. K, L. Bayerus sp. M. Katanga etiennei. 388 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Fig. 14. Reduviidae— Piratinae. A, B. Rasahus sulcicollis. C. Rasahus biguttatus. D. Sir- thenea flavipes. E. Peirates santus. F. Catamiarus brevipennis. G. Melanolestes abdominalis. H, I. Ectomocoris caudatus. J, K. Rasahus hamatus. L. Brachysandalus ephippiger. M. Nalata sp. N. Tydides rufus. 1989 ANTENNAL TRICHOBOTHRIA 389 Fig. 15. Reduviidae— Harpactorinae. A-F. Ricolla quadrispinosa. G, H. Zelus exsanguis. I. Rhaphidosoma didiera. J, K. Castolus ferox, $ and 9. 1^0. Amphibolus Venator, 1st instar. P. Amphibolus Venator, 5th instar. Q. Notocyrtus sp., 1st instar. R. Castolus lineatus, 1st instar. S. Castolus lineatus. T. IRepipta Jlavicans, 5th instar. U. Arilus sp., 5th instar. 390 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Fig. 16. Reduviidae— Harpactorinae. A, B. Sinea diadema, S and 9. C, D. Atrachelus ci- nereus, 9. E-G. Notocyrtus sp., $ and 9. H, I. Amphibolus Venator, $ and 9. J. Yolinus sp. K. Havinthus rufovarius. L, M. Arilus sp. N, O. Bactrodes femoratus. P. Erbessus sp. 1989 ANTENNAL TRICHOBOTHRIA 391 Fig. 17. Reduviidae— Apiomerinae. A. Manicocoris sp., S. B, C. Heniartes maestralis, 9 and 3. D, E. Heniartes flavicans, 3 and 9. F. Heniartes eythromerus, 9. G. Heniartes sp., 5th instar. H, I. Agriocoris sp., 9 and 3. J, K. Apiomerus sp., 9 and S. L. Amauroclopius sp., 9. M. Calliclopius sp., <5. N. Micrauchenus sp. 392 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Fig. 18. Reduviidae— various subfamilies. A, B. Empicoris rubromaculatus, 1st instar. C. Zirta limbata. D. Zelurus spinidorsis. E. Fusius rubricosus. F. Apronius sp. G. Sinea diadema, 9. H. Polytoxus sp. I. Staccia sp. J. Ctenotmchelus sp. K. Phimophorus spissicornis. 1989 ANTENNAL TRICHOBOTHRIA 393 LITERATURE CITED Carayon, J. 1970. Etude des Alloeorhynchus d’Afrique Centrale avec quelques remarques sur la classification des Nabidae (Hemiptera). Ann. Soc. Entomol. Fr. (N.S.) 6(4):899-931. Carayon, J. and A. Villiers. 1 968. Etude sur les Hemipteres Pachynomidae. Ann. Soc. Entomol. Fr. (N.S.) 4(3):703-739. Lent, H. and P. Wygodzinsky. 1979. Revision of the Triatominae (Hemiptera, Reduviidae) and their significance as vectors of Chagas’ disease. Bull. Amer. Mus. Nat. Hist. 163(3): 125-520. Schaefer, C. W. 1975. Heteropteran trichobothria (Hemiptera: Heteroptera). Int. J. Insect Morphol. Embryol. 4(3): 193-265. Schuh, R. T. 1975. The structure, distribution, and taxonomic importance of trichobothria in the Miridae (Hemiptera). Amer. Mus. Novitates 2585:1-26. Schuh, R. T. 1979. [Review of] Evolutionary Trends in Heteroptera. Part II. Mouthpart- structures and Feeding Strategies. Syst. Zool. 28:653-656. Received March 10, 1989; accepted September 21, 1989. New YorkEntomol Soc. 97(4):394-408, 1989 REVIEW OF THE NEW WORLD SPECIES OF THE GENUS NEOTTIGLOSSA KIRBY (HETEROPTERA: PENTATOMIDAE) D. A. Rider Department of Entomology, Louisiana Agricultural Experiment Station, Louisiana State University Agricultural Center, Baton Rouge, Louisiana 70803 Abstract.— five North American species of the genus Neottiglossa are reviewed. Diagnoses are provided for the genus Neottiglossa, for the two subgenera, and for the North American species. Neottiglossa trilineata (Kirby) and N. undata (Say) belong in the nominate subgenus, while N. cavifrons StSl, N. sulcifrons St^l, and N. tumidifrons Downes belong in the subgenus Texas Kirkaldy. Neottiglossa coronaciliata Ruckes, 1957 is placed as a junior synonym of Neotiglossa cavifrons StM, 1872; and N. californica Bliven, 1958 is placed as a junior synonym of N. undata (Say, 1832). A key is provided for the identification of species. The genus Neottiglossa Kirby belongs in the nominate tribe and subfamily of the Pentatomidae, and lacks a spine or tubercle at the base of abdominal segment 3 (2nd visible). Related Western Hemisphere genera occurring north of South America were keyed by Rolston and McDonald (1984). The North American species of Neottiglossa form two distinct species groups: N. trilineata (Kirby) and N. undata (Say) belong in the nominate subgenus, while N. cavifrons Stal, N. sulcifrons Stal, and N. tumidifrons Downes belong in the subgenus Texas Kirkaldy. Neottiglossa species are quite similar in appearance, which has resulted in confusion concerning the identity of several species. All New World species are herein reviewed, and a key is provided for the identification of subgenera and species. Neottiglossa Kirby, 1837 Pentatoma {Neottiglossa) Kirby, 1837:276. Aelioides Dohm, 1859:101; Mulsant and Rey, 1866:142. PlatysolenPiQhQv, 1860:82. Neottiglossa: Uhler, 1871:96-97, Stal, 1872a:36; Stal, 1872b: 18; Lethierry and Sev- erin, 1893:137; Summers, 1898:42; Jakovlev, 1903:325-327; Kirkaldy, 1904:280; Van Duzee, 1904:49-50; Kirkaldy, 1909:79; Zimmer, 1912:222, 230; Parshley, 1915:172; Van Duzee, 1917:47; Hart, 1919:179, 186-187; Stoner, 1920:62, 92; Parshley, 1923:758, 765; Blatchley, 1926:122, 148; Torre-Bueno, 1939:209, 226- 227; Froeschner, 1941:128, 130; Esselbaugh, 1946:690; Hoffman, 1971:29, 42; Furth, 1974:26, 37; McPherson, 1982:47, 71; Froeschner, 1988:586. Type species. Pentatoma {Neottiglossa) trilineata Kirby, 1837, by monotypy. Diagnosis. Small, less than 7 mm in length; elongate oval, strongly convex; usually fuscous to black with pale markings. Abdominal segment 3 (2nd visible) lacking basal spine or tubercle. Juga distinctly longer than tylus and contiguous in front of it. Propleura produced cephalad, antennifers exposed in lateral view. Ostiolar rugae small, each usually extending about three-fourths distance to lateral margin of meta- 1989 REVIEW OF NEOTTIGLOSSA 395 pleuron as small polished ridge. Scutellum broadly rounded apically, width at distal end of frena 0.5 or more width at base. Scutellum nearly as long as or longer than coria. Comments. Neottiglossa is closely related to the genus Aelia Fabricius, but can be separated from that genus by the smaller size and by differences in the structure of the propleura. Species of the genus Aelia are usually more than 8 mm in length and have the propleura covering the antennifers. Species of Neottiglossa are usually less than 7 mm in length, and the antennifers are exposed. Five species of Neottiglossa occur in the New World. These are placed into two subgenera. Neottiglossa cavifrons, N. sulcifrons, and N. tumidifrons belong in the subgenus Texas, while the nominate subgenus contains N. trilineata and N. undata. KEY TO THE NORTH AMERICAN SUBGENERA AND SPECIES OF Neottiglossa kirby 1 . Coxae pale yellow; evaporative surfaces pale yellow to brown-grey with contrasting black punctures subgenus Neottiglossa Kirby 2 Coxae fuscous to black; evaporative surfaces black with concolorous punctures . . subgenus Texas Kirkaldy 3 2(1). Dorsal surface of head and propleura mostly black with concolorous punctures . . trilineata (Kirby) - Dorsal surface of head and propleura with large areas of pale yellow to brown with black punctures undata (Say) 3(1). Apex of head broadly rounded, arcuate (Fig. 7); dorsal surface of head distinctly concave, thickly clothed with short inward-curving hairs cavifrons St^l Apex of head more tapering, narrowly rounded (Figs. 10, 14); dorsal surface of head not distinctly concave, or if somewhat concave then lacking short hairs 4 4(3). Boundary between dark surface of abdominal venter and pale lateral callus relatively straight, sharp (Fig. 1 3); trochanters dark fuscous; scutellum lacking medial pale line (Fig. 11) sulcifrons Sih\ Boundary between dark surface of abdominal venter and pale lateral callus diffuse, not well defined (Fig. 1 7); trochanters pale; scutellum usually with medial pale line (Fig. 15) tumidifrons Downes SnhgQnus Neottiglossa Kirby, 1837 Pentatoma {Neottiglossa) Kirby, 1837:276. Aelioides Dohm, 1859:101; Mulsant and Rey, 1866:142. PlatysolenPiebQY, 1860:82. Neottiglossa: Stal, 1872b: 18; Kirkaldy, 1904:280; Kirkaldy, 1909:79; Van Duzee, 1917:47; Stoner, 1920:92; Froeschner, 1988:586. Type species. Pentatoma {Neottiglossa) trilineata Kirby, 1837, by monotypy. Diagnosis. Coxae, trochanters, and base of each femur pale yellow, some fuscous markings medially on each femur. Evaporative surfaces fairly extensive, pale yellow to brown-grey with black punctures. Each ostiolar ruga auriculate with anterior por- tion produced laterad into small, obscure polished ridge. Juga only slightly elevated above surface of head; dorsal surface of head relatively flat, not concave. Segments 4-5 of antennae fuscous, distinctly darker than segments 1-3. Scutellum subequal in length or slightly shorter than coria. 396 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Comments. Previous workers have usually defined this subgenus by the slightly tumescent juga and by the auriculate nature of the ostiolar rugae. The jugal character can be hard to diagnose, and in most specimens each ostiolar ruga is produced laterad as a small, obscure, polished ridge. The two included species are, however, more closely related to each other than to the three remaining species. Specimens of this subgenus tend to be slightly larger and more broadly oval than specimens of the subgenus Texas. They can be easily recognized by the pale coxae and pale evaporative surfaces. Neottiglossa trilineata (Kirby, 1837) Figs. 1-3 Pentatoma {Neottiglossa) trilineata Kirby, 1837:276, pi. 6, fig. 6. Aelia trilineata: Dallas, 1851:224. Neottiglossa {Neottiglossa) undata (of authors, not Say): Stal, 1872b: 18 (part). Neottiglossa trilineata: Uhler, 1877:401; Uhler, 1886:5; Hussey, 1922:85-88, figs, la, Ic; Blatchley, 1926:148, 149-150; McPherson, 1970:52; McPherson, 1980:6; McPherson, 1982:72. Neottiglossa undata (of authors, not Say): Lethierry and Severin, 1893:138 (part). Neottiglossa {Neottiglossa) undata var. trilineata: Kirkaldy, 1909:80; Zimmer, 1912: 230. Neottiglossa {Neottiglossa) trilineata: Van Duzee, 1916:6; Van Duzee, 1917:48; Torre- Bueno, 1939:226; Froeschner, 1988:586. Diagnosis. Lateral margins of head tapering to narrowly rounded apex (Fig. 1); dorsal surface not concave, rather flat medially, lacking hairs, mostly fuscous or black with concolorous punctures, except sometimes with partial medial pale stripe basally. Scutellum with pale medial stripe from base to apical infuscated area (Fig. 2). Ventral surface of head black except bucculae pale; propleura black except area around coxal cavity, obscure area along posterior margin, and narrow lateral margin pale. Meso- pleura and metapleura black except area around coxal cavities and evaporative area pale. Boundary between dark abdominal surface and pale lateral margin abrupt, distinct, with only a few fuscous punctures in pale area. Femora with numerous fuscous spots medially; coxae, trochanters, and base of each femur pale. Medial excavation in posterior margin of pygophore relatively broad, shallow (Fig. 3). Types. Kirby (1837) described N. trilineata from a single specimen from Saskatch- ewan. The deposition of the type specimen is unknown. Specimens examined. 22 specimens collected from 18 May to 15 August. CAN- ADA: ALBERTA: Bilby; Whitford Lake. MANITOBA: Cedar Lake; Winnipeg. NORTHWEST TERRITORIES: Great Bear Lake. ONTARIO: Belmont. UNITED STATES: CALIFORNIA: Grass Lake. COLORADO: Boulder: Longs Peak Inn. Larimer: Fort Collins, Rist Cyn. IDAHO: Latah: 6 mi N Bovill. OREGON: Grant: Seneca. WYOMING. Sheridan: 42 mi W Sheridan, Prune Crk. Sublette: Green R. Lake. Teton: Yellowstone Natl. Park, Yellowstone Lake. Comments. This species is most closely related to N. undata from which it can be separated by the darker coloration of the propleura and the dorsal surface of the head. The relatively broad medial excavation in the posterior margin of the pygophore is also diagnostic. 1989 REVIEW OF NEOTTIGLOSSA 397 Figs. 1-6. Figs. 1-3. N. trilineata. 1. Head. 2. Habitus. 3. Pygophore, caudal view. Figs. 4- 6. N. undata. 4. Head. 5. Habitus. 6. Pygophore, caudal view. Neottiglossa undata (Say, 1832) Figs. 4-6 Pentatoma undata Say, 1832:8; Say, 1859:319-320. Neottiglossa undata:\J\i\QT, 1871:96-97; Uhler, 1872a:395; Uhler, 1872b:471; Uhler, 1876:284; Uhler, 1877:401; Uhler, 1878:376; Provancher, 1885:39; Uhler, 1886: 5; Van Duzee, 1889:2; Osborn, 1892:121; Lethierry and Severin, 1893:138 (part); 398 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Van Duzee, 1894:170; Wirtner, 1904:187; Van Duzee, 1904:49, 50; Van Duzee, 1905:548; Smith, 1910:133; Olsen, 1912:53; Zimmer, 1912:230; Van Duzee, 1912: 318; Torre-Bueno, 1913:58; Parshley, 1914:148; Parshley, 1915:175; Stoner, 1915a: 355; Stoner, 1915b:353-354; Parshley, 1917:22; Stoner, 1917:45; Torre-Bueno, 1918:25; Hart, 1919:186, 187; Hussey, 1922:85-88, figs, lb. Id; Parshley, 1923: 765; Stoner, 1925:57; Blatchley, 1926:148-149; Stoner, 1926:29; Brimley, 1938: 62; Furth, 1974:37-38; McPherson, 1970:52; McPherson, 1979:93; McPherson, 1980:6; McPherson, 1982:72. Aelia americana (of authors, not Dallas): Provancher, 1885:38. Mormidea undata: MacGillivray & Houghton, 1903:262-265. Neottiglossa {Neottiglossa) undata: Stal, 1872b: 18 (part); Kirkaldy, 1909:80; Van Duzee, 1916:6; Van Duzee, 1917:48; Stoner, 1920:93-94; Torre-Bueno, 1939:226; Froeschner, 1988:586. Neottiglossa calif or nica Bliven, 1958:12-13. NEW SYNONYMY. Neottiglossa {Texas) californica: Froeschner, 1988:587. NEW SYNONYMY. Diagnosis. Lateral margins of head tapering to narrowly rounded apex (Fig. 4); dorsal surface not concave, relatively flat medially, devoid of hairs; mostly pale yellow to brown with black punctures. Scutellum mostly pale except usually with small infuscated area distally and sometimes obscure infuscated areas basally; with distinct medial pale impunctate line (Fig. 5). Ventral surface of head fuscous except bucculae and surrounding area pale; thoracic pleura mostly pale with black punctures; evap- orative area rather extensive, pale. Boundary between dark abdominal surface and pale lateral margin usually indistinct, diffuse, sometimes a pale spot surrounding each spiracle and obscure pale areas in band just mediad of spiracles. Femora with a few fuscous spots medially, apical tarsal segment sometimes brown; coxae, tro- chanters, and base of each femur pale. Medial excavation in posterior margin of pygophore relatively narrow, deep (Fig. 6). Types. Say (1832) described this species from an unknown number of specimens from the Northwest Territory. The types are apparently no longer in existence, but because the identity of this species has never been in question, a neotype is not designated. Bliven (1958) described N. californica from 12 specimens from Humboldt County, California. The holotype and all 1 1 paratypes, which are housed in the California Academy of Sciences, San Francisco, were examined. There is no consistent difference between N. californica and N. undata. Specimens examined. 167 specimens collected from 7 April to 12 October. CAN- ADA: ALBERTA: Bilby. BRITISH COLUMBIA: Summerland; Thombull Mtn, Ter- race; Victoria; Wellington. MANITOBA: Cedar Lake. NEW BRUNSWICK: Monc- ton. NEWFOUNDLAND: Buchans Road; Kings Point Bog. NOVA SCOTIA: Halifax. ONTARIO: Crystal Bay; Dundas; 1 5 mi NW Ignace; Oakwood nr. Lindsay; Obatanga Provincial Park; Ottawa, Nepean; Ridgeway; Sault Ste. Marie; Swansea. QUEBEC: Luskville Falls. UNITED STATES: CALIFORNIA: Humboldt: Eureka; Fieldbrook; Shively. Mendocino: 15 mi E Fort Bragg. Shasta: 25 mi SE McCloud; Hwy 299, Hat Crk. COLORADO: Boulder: Gregory Cyn. Douglas: Prairie Park. CONNECTICUT: Luternk. New Haven: 2 mi E Seymour Marsh. IDAHO: Blaine: Bellevue. Bonner: Walsh Lake, Samuels. Elmore: Dixie. Idaho: 10.2 mi WSW Lolo Pass, Powell Pasture. Latah: Moscow. Lemhi: 5 mi S Salmon. INDIANA: Howard. Noble: Sylvan Lake. 1989 REVIEW OF NEOTTIGLOSSA 399 IOWA: Black Hawk: Cedar Falls. Boone: Pilot Mound. Cass: Atlantic. Iowa: Amana. Story: Ames. Webster: Fort Dodge. MAINE: Franklin: Farmington. MASSACHU- SETTS: Essex: Marblehead. Middlesex: Tyngsboro. Worcester: Worcester. MICHI- GAN: Clinton: Rose Lake Wildlife Expt Stn. Gogebic. Kalkaska. Oakland. MIN- NESOTA: Clearwater: Itasca Park. Itasca: Deer Lake. Polk: Crookston. MONTANA: Gallatin: nr. Bozeman; Gallatin R. Lincoln: Libby Crk, 5 mi S Libby. NEVADA: Washoe: Verdi. NEW HAMPSHIRE: Coos: Mt. Washington. Grafton: Franconia. NEW YORK: Pine Island; Yaphank. Columbia: Chatham. Erie: Lancaster. Monroe: Rochester. Orange: Greenwood Lk. Suffolk: Mastic Beach, Long Island. Tompkins: Ithaca. OHIO: Ashtabula: Crooked Crk Farm nr. Hartsgrove. OREGON: Browns- mead. Baker: Spring Crk. Benton: Corvallis. Columbia: Goble; Scappoose. Deschutes: Redmond. Douglas: 1 8 mi W Diamond; 20 mi E Reedsport, Spencer Crk; 7 mi NW Roseburg. Hood River: Parkdale. Jackson: Buckhom Mineral Sprg; 22 mi N Prospect. Klamath: 1 mi NW Bly, Meryl Crk; Crater Lake Natl Park; Crescent Lake; 12 mi SW Keno; Klamath Falls; Mares Egg Sprg; Sun Crk; Wood R. Sprg. Lake: Hart Lake. Multnomah: Gresham. Polk: Monmouth. PENNSYLVANIA: Wayne: Hawley; Honesdale. SOUTH DAKOTA: Lawrence: Savoy, Spearfish Cyn. UTAH: Cache: Logan; Providence. Emery: Green. VERMONT: Windsor: Woodstock. WASHING- TON: Wawai. Jefferson: Olympic Natl. Park. Okanagon: Okanagon nr. Buzzard Lake. WISCONSIN: Dane: Madison. American Falls. WYOMING: Albany: Snowy Range Mtns. Sheridan: Banner. Teton: Yellowstone Natl. Park. Whitman: 8 mi SW Pullman, Lyle Grove Biol. Area. Comments. This species is most closely related to N. trilineata from which it can be distinguished by the more extensive pale areas on the propleura and the dorsal surface of the head and by the relatively narrow medial excavation in the posterior margin of the pygophore. Subgenus Texas Kirkaldy, 1904 Neottiglossa (Melanostoma) Stal, 1872b: 18 (preoccupied). Neottiglossa {Texas) Kirkaldy, 1904:280 (replacement name); Kirkaldy, 1909:79, 80; Van Duzee, 1917:48; Stoner, 1920:92; Froeschner, 1988:586. Type species. Neottiglossa sulcifrons Stal, 1872, by subsequent designation (Kir- kaldy, 1909:XXX). Diagnosis. Coxae fuscous, trochanters and base of each femur often fuscous. Evap- orative surfaces reduced, black with concolorous punctures. Juga usually distinctly tumescent, in lateral view elevated above surface of head; dorsal surface of head sometimes concave. Each ostiolar ruga not auriculate, but produced laterad into relatively well-defined polished ridge. Segments 4-5 of antennae only slightly darker than segments 1-3. Scutellum longer than coria. Comments. The three species in this subgenus are easily recognized by the fuscous coxae and the black evaporative surfaces. Neottiglossa cavifrons S\k\, 1872 Figs. 7-9 Neottiglossa {Melanostoma) cavifrons Stal, 1872b: 18. Neottiglossa cavifrons: Uhler, 1886:5; Lethierry and Severin, 1893:137; Scott and 400 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Fiske, 1902:31; Hart, 1919:186, 187; Blatchley, 1926:148, 151; Downes, 1928:91, fig. 1; Brimley, 1938:62; Froeschner, 1941:130, 139; Esselbaugh, 1946:678, 680; Esselbaugli, 1948:28; Hoffman, 1971:42; Getting and Yonke, 1971:457-458;Furth, 1974:37, 38; McPherson and Mohlenbrock, 1976:153; McPherson, 1978:161; McPherson, 1979:93; McPherson, 1980:6; McPherson, 1982:71-72. Neottiglossa {Texas) cavifwns: Kirkaldy, 1904:280; Kirkaldy, 1909:80 (part); Van Duzee, 1917:48 (part); Torre-Bueno, 1939:227 (part); Froeschner, 1988:587. Neottiglossa coronaciliata Ruckes, 1957:41. NEW SYNONYMY. Neottiglossa {Texas) coronaciliata: Froeschner, 1988:587. NEW SYNONYMY. Diagnosis. Head broadly rounded apically, arcuate (Fig. 7); dorsal surface distinctly concave, clothed with numerous, silver, inward-curving hairs; juga completely black. Scutellum fuscous or black except two subbasal calloused spots and narrow lateral margins near apex pale, lacking pale medial line (Fig. 8). Ventral surface of head and thoracic pleura black except narrow lateral margin of propleura pale; ostiolar rugae and evaporative surfaces black with black punctures. Boundary between dark ab- dominal surface and pale lateral margin abrupt, distinct. Legs pale except coxae, trochanters, and base of each femur fuscous, sometimes a few small fuscous spots medially on each femur. Posterior margin of pygophore as in figure 9. Types. Stal (1872b) described N. cavifrons from at least one male and one female specimen from Texas. The types were not examined but the description is sufficient to identify this distinctive species. In the original description Stal states “Caput parte anteoculari concaviuscula, pilosa, ante medium utrimque rotundato-angustata.” There is only one Western Hemisphere species of Neottiglossa with the apex of the head rounded and the dorsal surface distinctly concave and pilose as in the above de- scription. The types are housed in the Swedish Museum of Natural History, Stock- holm. Ruckes (1957) described N. coronaciliata from two female specimens from Texas and Louisiana. Both specimens were examined and differ in no significant way from N. cavifrons. Both the holotype and paratype are conserved in the American Museum of Natural History, New York. Specimens examined. 244 specimens collected from 4 April to 29 June. UNITED STATES: ALABAMA: Butler: 1-65, 4.5 mi S Hwy 10. Lee: Auburn. Macon: 1-85, 5.1 mi E Shorter. ARKANSAS: Izard: Mt. View. Union: Smackover. GEORGIA: Jefferson: Louisville. KENTUCKY: Christian. LOUISIANA: Caddo: 2 mi S Shreve- port nr. Wallace Lake. Catahoula: Leland; Hwy 126, 6.8 mi W Hwy 124. Grant: Hwy 165, 6 mi N Pollock. Lincoln: 5 mi E Ruston. Morehouse: Bastrop; Chemin- A-Haut State Park. Natchitoches: Hwy 1 , 1 mi SE Galbraith; Kisatchie Natl. Forest, Red Bluff Campground. Ouachita: Bosco. St. Tammany. Webster: Shongaloo. MIS- SISSIPPI: Agricultural College. Hancock: I- 10 at Hwy 607. Lowndes: Columbus. MISSOURI: Monroe: Mark Twain State Park. New Madrid: 1-55, 5 mi S Hwy 62. Polk: 3 mi SE Flemington. Vernon: Osage Prairie. OKLAHOMA: Leflore: Page. TEXAS: Brazos: Minter Spgs. Shelby: Tenaha. Comments. This is the most distinctive species in the genus. It can be separated from all other New World species of Neottiglossa by the broadly rounded apex of the head and the distinctly concave surface of the head, which is clothed with nu- merous inward-curving hairs. 1989 REVIEW OF NEOTTIGLOSSA 401 Figs. 7-17. Figs. 7-9. N. cavifrons. 7. Head. 8. Habitus. 9. Pygophore, caudal view. Figs. 10-13. N. sulcifrons. 10. Head. 11. Habitus. 12. Pygophore, caudal view. 13. Lateral margin of abdominal venter. Figs. 14-17. N. tumidifrons. 14. Head. 15. Habitus. 16. Pygophore, caudal view. 17. Lateral margin of abdominal venter. Neottiglossa sulcifrons Stal, 1872 Figs. 10-13 Neottiglossa (Melanostoma) sulcifrons StM, 1872b: 18. Melanostoma sulcifrons: Uhler, 1876:284; Uhler, 1877:402. Neottiglossa sulcifrons: Uhler, 1886:5; Townsend, 1891:53; Lethieiry and Severin, 1893:138; Uhler, 1893:368; Uhler, 1894:228; Heidemann, 1899:217; Van Duzee, 1904:50; Crevecoeur, 1 905:232; Torre-Bueno and Brimley, 1907:442; Smith, 1910: 135; Zimmer, 1912:230; Torre-Bueno, 1913:58; Stoner, 1915a:355; Stoner, 1916: 402 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) 306; Stoner, 1917:45; Hart, 1919:186, 187; Hussey, 1921:8-15; Parshley, 1921: 13-24; Torre-Bueno, 1925:280; Whittaker, 1926:63; Blatchley, 1926:148, 150; Downes, 1927:3; Downes, 1928:91, fig. 2; Hendrickson, 1930:49-179; Harris, 1937: 169-176; Ruckes, 1937:32-36; Brimley, 1938:62; Froeschner, 1941:130, 138- 139; Esselbaugh, 1946:678; Esselbaugh, 1948:27-28; DeCoursey and Esselbaugh, 1962:331; DeCoursey and Allen, 1968:146; Hoffman, 1971:42; Furth, 1974:37, 38; McPherson and Mohlenbrock, 1976:153; McPherson, 1978:161; McPherson, 1979:93; McPherson, 1980:6; McPherson, 1982:71, 72. Neottiglossa {Texas) sulcifrons: Kirkaldy, 1904:280; Kirkaldy, 1909:80; Van Duzee, 1916:6; Van Duzee, 1917:48; Stoner, 1920:93, 94-95; Torre-Bueno, 1939:227; Froeschner, 1988:587. Diagnosis. Lateral margins of head tapering to narrowly rounded apex (Fig. 1 0); dorsal surface not distinctly concave, with at most a few outward-curving hairs apically; juga pale basally. Scutellum with medial Y-shaped fuscous band, lateral pale areas near apex more extensive, continuing cephalad to subbasal pale calloused spots, lacking pale medial line (Fig. 1 1). Ventral surface of head and thoracic pleura black except narrow lateral margin of propleura pale; ostiolar rugae and evaporative surfaces black with black punctures. Boundary between dark abdominal surface and pale lateral margin sharp, distinct (Fig. 1 3). Legs pale except coxae, trochanters, base of each femur, and several spots on superior surface of each femur fuscous. Posterior margin of pygophore as in Figure 12. Types. Stal (1872b) described N. sulcifrons from at least one male and one female specimen from Texas. The types were not examined, but the original description is adequate to fix this species. The types are conserved in the Swedish Museum of Natural History, Stockholm. Specimens examined. 1 09 specimens collected from 6 April to 10 October. UNITED STATES: ALABAMA: Macon: 1-85 at Hwy 81. Tallapoosa: Alexander City. ARI- ZONA: Cochise: Chiricahua Mtns, Coronado Natl. For., Sunny Flat. Maricopa: Scottsdale. Pima: Boboquivari Mtns, nr. Kitt Peak; 7 mi SE Continental; Lowell Ranger Stn; Santa Catalina Mtns, Bear Wallow; Santa Catalina Mtns, Sabino Cyn; Santa Catalina Mtns, Sycamore Cyn; Santa Rita Mtns, Parkers Ranch; Santa Rita Mtns, Sonoita; Tucson. ARKANSAS: Hempstead: Hope. Sebastian: Hacket. IOWA: Boone: Ledges State Park. Page: Shenandoah. Story: Ames. KANSAS: Riley: Man- hatten. LOUISIANA: E. Feliciana: Boy Scout Camp Avondale, Hwy 10 E of Clinton. Grant: Hwy 165, 6 mi N Pollock. Morehouse: Chemin-A-Haut State Park. Natchi- toches: Kisatchie Natl. Forest, Red Bluff Campground. Webster: Shongaloo. MIS- SISSIPPI: Madison: 1-55, 4 mi N Gluckstadt. Pike: Summit. Yazoo: 1-55, 2 mi S Hwy 432. MONTANA: Petroleum: 1.5 mi S, 5 mi W Winnett. NEBRASKA: Chase: 10 mi SW Imperial. Red Willow: Indianola. NEW JERSEY: Iona. Ocean: Lakehurst. NEW YORK: Pinelawn, Long Island. TEXAS: Bosque: 5 mi S Iredell. Brazos: College Station. VIRGINIA: Falls Church. WYOMING: Johnson: Bighorn Natl. Forest, Tie Hack Campground. Comments. This species is most closely related to N. cavifrons and N. tumidifrons. It can be separated from all other New World congeners by the combination of the following characters: dorsal surface of head not concave, lacking silvery hairs; evap- orative area black; coxae, trochanters, and base of each femur fuscous. 1989 REVIEW OF NEOTTIGLOSSA 403 Neottiglossa tumidifrons Downes, 1928 Figs. 14-17 Neottiglossa cavifrons (of authors, not StM): Van Duzee, 1904:49, 50; Stoner, 1926: 29; Blatchley, 1934:4. Neottiglossa {Texas) cavifrons (of authors, not StM): Kirkaldy, 1909:80 (part); Van Duzee, 1916:6; Van Duzee, 1917:48 (part); Torre-Bueno, 1939:227 (part). Neottiglossa tumidifrons Downes, 1928:90-92, fig. 3; Downes, 1935:46. Neottiglossa {Texas) tumidifrons: Froeschner, 1988:587. Diagnosis. Lateral margins of head tapering to narrowly rounded apex (Fig. 1 4); dorsal surface concave, lacking silver hairs; juga mostly pale yellow, distinctly tu- mescent. Scutellum mostly pale except one apical and two basal infuscated areas, with distinct medial pale line from base to apical dark area (Fig. 15). Ventral surface of head and thoracic pleura black except lateral margin of propleura and sometimes area around coxal cavities pale; ostiolar rugae sometimes pale; evaporative areas black with black punctures. Boundary between dark abdominal surface and pale lateral areas not distinct, diffuse (Fig. 1 7). Femora with a few obscure fuscous spots; coxae fuscous, trochanters and base of each femur pale. Posterior margin of pygophore as in Figure 16. Types. Downes (1928) described N. tumidifrons from 22 specimens with the ho- lotype from the Saanich District, British Columbia, Canada. The holotype is con- served in the Canadian National Collection, Ottawa. Specimens examined. 127 specimens collected from 5 March to 8 October. CAN- ADA: BRITISH COLUMBIA: Bear Hill; Summerland; Victoria. UNITED STATES: CALIFORNIA: Blue Cyn. Fresno: Friant. Humboldt: Eureka; Miranda mp 6.52, Avenue of the Giants. Lake: 0.2 mi W intersection of its. 20 & 53. Los Angeles: Griffith’s Park. Mariposa: Mormon Bar; Yosemite. Mendocino: 15 mi E Fort Bragg. Napa: N side Howell Mtn, 2 mi NNE Angwin. Riverside: Hwy 243, 2 mi N Poppet Flat. San Diego: Pauma. San Mateo. Santa Barbara: Santa Barbara. Shasta: Hat Crk; 25 mi SE McCloud. Siskiyou: Macdoel; 6 mi S Macdoel; Weed. Sonoma. Tulare: Visalia. IDAHO: Lenore. Cassia: 3.5 mi SE Basin. Latah: Kendrick; Moscow. Owy- hee: Hot Spring. Twin Falls: 9 mi SW Rogerson; Shoshone Falls. OREGON: Benton: Corvallis. Jackson: 2 mi S Ashland; Dead Indian Sprg; Griffin Crk; Prospect; 3 mi S Prospect; Shady Grove. Josephine: Cave Jnct; 7 mi W Grants Pass; Illinois R; 9 mi S Selma. Klamath: above Algoma; Buck Lake Road, 2.5 mi N Hwy 66; 12 mi SW Keno; Meryl Crk, 7 mi NW Bly; Modoc Point; 3 mi S Saddle Mtn. Lane: Eugene. Linn: Lacomb. Wallowa: Copper Crk at Snake R. Wasco: 1 2 mi S Dufur. Washington: Dilley. UTAH: Cache: Hyde Park; Providence. Salt Lake: Salt Lake City. Utah: Hobble Crk Cyn, 5 mi E Springfield. WASHINGTON: Pierce: Parkland. Thurston: Olympia. Whitman: Pullman. Yakima: Satus Crk; Tieton; Yakima. Comments. This species is most closely related to N. cavifrons and N. sulcifrons. It can be separated from all North American congeners by the following combination of characters: slightly concave dorsal surface of the head lacking short silvery hairs; evaporative area black; coxae black; and boundary between dark abdominal surface and pale lateral margin diffuse, indistinct. 404 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) ACKNOWLEDGMENTS I am indebted to the following individuals for the loan of specimens and other assistance relevant to this study: P. H. Amaud, Jr. (California Academy of Sciences, San Francisco); J. B. Chapin (Louisiana State University); R. Danielsson (Lund University, Lund, Sweden); J. E. Eger (personal collection, Tampa, FL); T. J. Henry (National Museum of Natural History, Washington, D.C.); M. A. Ivie (Montana State University, Bozeman); J. Loye (University of Utah, Salt Lake City); L. H. Rolston (personal collection. Baton Rouge, LA); and R. T. Schuh and M. D. Schwartz (American Museum of Natural History, NY). Special thanks go to P. H. Arnaud, Jr. and T. J. Henry for arranging the loans of the types of N. californica and N. coronaciliata, respectively. I would also like to thank J. B. Chapin, J. A. Moore, and L. H. Rolston (Louisiana State University, Baton Rouge) for their critical review of the manuscript. Approved for publication by the Director of the Louisiana Agricultural Experiment Station as manuscript number 89-17-3231. LITERATURE CITED Blatchley, W. S. 1926. Heteroptera or True Bugs of Eastern North America with Especial Reference to the Faunas of Indiana and Florida. Nature Pub. Co., Indianapolis, 1,116 pp. Blatchley, W. S. 1934. 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Soc. 97(4):409-416, 1989 ORIUS MINUTUS (LINNAEUS) IN NORTH AMERICA (HEMIPTERA: HETEROPTERA: ANTHOCORIDAE) John D. Lattin,* Adam Asquith, ^ and Steve Booth^ ‘Systematic Entomology Laboratory, Department of Entomology, Oregon State University, Corvallis, Oregon 97331 ^Department of Entomology, Oregon State University, Corvallis, Oregon 97331 Abstract. — The history and present distribution of Orius minutus (Linnaeus) in North America is summarized. The systematic position of O. minutus is reviewed, showing that this introduced species is the only member of the subgenus Heterorius Wagner in North America. The adult and nymph of O. minutus are illustrated and distinguished from the native sympatric species Orius tristicolor (B. -White). The ecology and potential benefits of O. minutus as a predator in agroecosystems are discussed. Orius minutus (Linnaeus) is a Palearctic species of Anthocoridae that has been introduced accidentally into western North America. According to Pericart (1972) its natural distribution is Europe and western Russia, North Africa, east to Turkestan, China and Siberia. Precise information is often lacking because of taxonomic con- fusion with other species of Orius in that vast area. Tonks (1953) first reported the species from North America, based on specimens collected on Lulu Island and Huntingdon, in the vicinity of Vancouver, British Columbia. The specimens were collected on raspberry and loganberry in 1951. An- derson (1962) added records from Victoria and the lower Fraser Valley from Van- couver to Hope, British Columbia and also reported the species from Seattle and Bothell, Washington and Albany, Oregon. He suggested that it had been recently introduced to the Pacific Coast region, stating that the earliest specimens he had examined had been collected in 1 939. Kelton (1963, 1978), Herring (1966) and Henry (1988) based their records on the publications of Tonks and Anderson. The earliest record we have seen is based upon a specimen in the OSU, Systematic Entomology Laboratory, collected in Seattle, Washington on 29 July 1930 by the late M. H. Hatch. Collection records are clustered in the following areas (earliest collection dates in parentheses): Seattle, Washington (1930); Vancouver, British Co- lumbia (1951); Portland, Oregon and the Willamette Valley, Oregon (1957) (Fig. 1). The earliest records for Hood River County, Oregon date from 1971. While the possibility of three separate introductions exists (Seattle, Vancouver and Portland), the species seems to occur chiefly in agroecosystems, particularly on fruit trees and shrubs, so that an alternative hypothesis would be a single introduction and estab- lishment followed by dispersal via movement of plant material through commerce and some normal dispersal. O. minutus seems well established from at least the Willamette Valley, Oregon north to the Vancouver, British Columbia region and east up the Columbia River to at least Hood River. 410 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Fig. 1. Distribution of Orius minutus in the Pacific Northwest with localities and dates of the earliest known specimens. Solid circles represent specimens examined. Solid triangles rep- resent literature records. SPECIMENS EXAMINED British Columbia. Milner, 1958 (G. J. Spencer) (UBC); Bowen Is., 24 April 1960 (G. J. Spencer) (UBC); Lions Bay, Vancouver, 22 May 1961 (G. J. Spencer) (UBC); Vancouver, 2 July 1961 (G. J. Spencer) (UBC); Vancouver, 2 July 1961 (G. J. Spencer) (UBC); Vancouver, 28 September 1976 (G. G. E. Scudder) (UBC); Van- couver, 23 October 1978 (J. Scudder) (UBC); Galiano Is., North end, 18 August 1982 (G. G. E. Scudder); Vancouver, 3 August 1983 (G. G. E. Scudder) (UBC); Spanish Hills, Galiano Is., 15 August 1983 (G. G. E. Scudder) (UBC); 10 mi E. Mission City, 31 July 1957, ex Hazel (N. Anderson) (OSU-SEL); OREGON: Benton Co.: 1 mi N Corvallis, Hwy 99W, ex. Corylus, 10 May 1989 (A. Asquith) (OSU-SEL); Clack- amas Co.: 8 mi E Oregon City, sweeping potatoes. 3 August 1962 (H. E. Morrison, R. F. Koontz) (OSU-SEL); Columbia Co.: Beaver, rose bloom, 6 July 1967 (K. Goeden) (OSDA); Hood River Co.: Hood River, 23 July 1971, ex. Aphid infested apple leaf (R. W. Zwick) (MCAREC); Hood River, 10 January 1972, ex. cardboard band of pear tree (G. J. Fields) (MCAREC); Hood Riv. Exp. Station, 30 April 1981, ex pear (MCAREC); Hood River, Power Block, 13 July to 18 September 1988, ex. pear (S. Booth) (MCAREC); Hood River, Power Block, MCAREC, 9 August 1988, 1989 ORIUS MINUTUS IN NORTH AMERICA 411 ex. oak (S. Booth) (OSU); Jefferson Co.: Culver, 20 July 1962, sweeping grass and potatoes (R. F. Koontz) (OSU-SEL); Linn Co.: 7 mi NE Albany, 5 June 1957, ex. raspberry (J. D. Lattin) (OSU-SEL); Yamhill Co.: McMinnville, Peavine Ridge, 29 May 1958, ex. Opulaster capitatus (K. Fender) (OSU-SEL); Washington Co.: 5 mi n N. Plains, 10 August 1960 (J. D. Lattin) (OSU-SEL); WASHINGTON: Seattle, 29 July 1930 (M. H. Hatch) (OSU-SEL); 7 May 1931; 2 July 1931 (M. H. Hatch) (OSU- SEL); 27 May 1933 (L. C. Snyder) (OSU-SEL); July 1961; Seattle (T. Kincaid) (OSU- SEL); U. W. Campus, 27 April 1937 (OSU-SEL); 8 April 1938 (E. Dailey); 7 April 1938 (Patterson) (OSU-SEL); 4 May 1948 (M. H. Hatch) (OSU-SEL); 12 April 1961; 1 May 1964. SYSTEMATIC POSITION The genus Orius was described by Wolff in 1811 and contains about 70 species (Pericart, 1972). Wagner (1952) proposed the subgenus Heterorius with Cimex mi- nutus Linneaus as the type. Kelton (1963) produced a synopsis of the species of Orius of Canada and the United States and included O. minutus. No mention of subgeneric placement was made for any of the species. Herring (1966) published a review of Orius for the Western Hemisphere and included a key to the subgenera following Wagner (1952). He stated that many of the New World species could not be placed in these four taxa, and remarked that all of the species treated but one fit the characters proposed for the subgenus Heterorius, but felt that the claspers of the males of these species did not conform to the European species included in Heterorius. Herring did not propose a revision of the Wagner classification to include the New World species. Henry (1988) followed Herring and cited the latter author’s remarks about the un- suitability of the Wagner classification for the North American fauna. Our investi- gations confirm Herring’s supposition; all palearctic species in the subgenus Heter- orius have a well defined spine at the base of the male clasper and all North American species lack this structure. Thus, as an introduction, O. minutus is the only repre- sentative of Heterorius, as presently defined, in North America. DIAGNOSTICS Adults. O. minutus adult females are large (2.05-2.60 mm total length) and broadly ovate (0.85-0.97 mm pronotal width) (Fig. 2). The head is dark brown to black with yellow antennae. The pronotum and scutellum are fuscous to piceous, with the hemelytra yellowish brown. The venter is fuscous to black, with the anterior and middle legs yellow and hind legs brown to fuscous. The dorsum is rather thickly clothed with long semi-erect golden setae. Adult males are more slender (0.7-0.82 mm pronotal width) with thicker antennae. The other common species of Orius in the Pacific Northwest, O. tristicolor (B.- White), is slightly smaller (1.82-2.29 mm total length) and more linear (0.66-0.83 mm pronotal width). It is dark brown to black on the head, pronotum, scutellum, clavus, the apical one third of the corium and the cuneus, with the antennae and basal half of the corium a contrasting pale white. The entire ventral surface is also solid black with only the front and middle tibia and tarsi white. The dorsal surface is covered with much shorter and more sparsely distributed decumbent pale setae. Nymphs. Fifth instar nymphs of O. minutus can be distinguished from O. tristicolor 412 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Fig. 2, Orius minutus. Dorsal habitus of adult female. 1989 ORIUS MINUTUS IN NORTH AMERICA 413 Fig. 3. Fifth instar nymphs of Orius spp. Dorsal habitus. A. O. tristicolor. B. O. minutus. by their broadly ovate form with the apical one third of the wing pads much darker than the rest of the dorsum (Fig. 3). O. tristicolor nymphs are slightly smaller, more linear and uniformly reddish brown in coloration. Badly bleached specimens can be distinguished by the maximum width of the pronotum, which in O. minutus is 0.70 mm or greater and in O. tristicolor 0.60 mm or less. Earlier instars are more difficult to distinguish, as both species are usually creamy white in color. O. minutus is always broader and more robust, the maximum width of the pronotum almost 1.5 times the width of the head. In minutus, the head is short, and the eyes almost touch the anterior margin of the pronotum. Early instar nymphs of tristicolor are more slender, the width of the head only slightly less than or equal to the width of the pronotum. The head is longer and the eyes are separated from the anterior margin of the pronotum by at least the width of the fourth antennal segment. FEEDING HABITS Orius minutus is chiefly predacious, as are most Anthocoridae (Pericart, 1972; Cobben, 1978; Kelton, 1978). Fulmek (1930) studied the development of this species 414 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) in Austria and reported several species of aphids as prey. Tonks (1953) reported . . mites, thrips, leafhoppers, and other small insects,” and Collyer (1953) reported the fruit tree red spider mite {Metatetranychus ulmi (Koch)) as prey in English orchards. Southwood and Leston (1959) also report mites and small insects as prey. Anderson (1962) observed O. minutus feeding on the two-spotted spider mite, Tetranychus telarius (L.) on raspberry in Orgeon. Pericart (1972) reported aphids, psyllids, jassids, thrips, mites and the eggs of pentatomids and Lepidoptera as prey items. Kelton (1978) listed a number of herbs and shrubs in British Columbia that provided a habitat for O. minutus while it fed on a variety of mites and insects. While the Anthocoridae are considered to be predators, considerable evidence indicates that some are partially phytophagous (see for example: Carayon and Steffan, 1959; Anderson, 1962; Chu, 1969; Fauvel, 1974; Pericart, 1972; Bachelor and Bar- anowski, 1975; Cobben, 1978). Several genera of the Oriini (Orius Wolff and Para- triphleps Champion) have been reported to feed on plant materials. Xambeu (1903) (cited in Butler, 1923 and Fulmek, 1930) reported Orius minutus feeding on plant fluids {Eryngium campestre) as well as aphids, as did Fulmek (1930) and Pericart (1972). Fauvel (1970) reported Orius vicinus (Ribaut) (also in the subgenus Heterorius) feeding on a number of plants besides feeding on mites and aphids, and provided interesting illustrations of pollen and beak tips, showing relative sizes. Carayon and Steffan (1959) found that Orius pallidicornis (Reuter) was primarily phytophagous, feeding on the pollen of Ecballium elaterium Rich. Dicke and Jarvus (1962) discov- ered the importance of corn pollen in building up populations of Orius insidiosus (Say) in Ohio. Bachelor and Baranowski (1975) were able to rear Paratriphleps lae- viusculus Champion successfully on the flowers and pollen of Manilkara zapotilla (Jacq.) Gilly in Florida. Any consideration of Anthocoridae as biological control agents should consider potential plant food sources as well as conventional prey items. LIFE HISTORY Orius minutus overwinters as a fertilized female, usually emerging from hibernation in March and April in Europe (Collyer, 1953). Some males may also hibernate. Egg laying occurs shortly after hibernation ends, the eggs being deposited in the midrib on the ventral side of the leaves of “host” plants (Collyer, 1953) or at the base of developing flower buds (Fulmek, 1930). Fulmek (1930) provides excellent figures of the eggs and nymphs of O. minutus, as does Pericart (1 972) who also includes drawings of the adult male and female. There are five nymphal instars. The developmental time from oviposition to adult varied from 24 to 30 days in England (Collyer, 1953). There appears to be continuous reproduction through the season until fall when the females that will overwinter are fertilized. At least two generations a year occur (Collyer, 1953), but possibly three or four (Pericart, 1972). Both phytophagous and predaceous feeding habits are recorded. The seasonal phenology of O. minutus in North America has not yet been detailed. Adults have been collected from early April to mid-September however, suggesting that the species is bivoltine. Orius minutus is a potentially beneficial introduction, particularly in cane and orchard crops. If it continues to expand its range and numbers in the Pacific Northwest it may prove to be an important addition to the predator complex of fruit crops. For 1989 ORIUS MINUTUS IN NORTH AMERICA 415 example, its present distribution in the fruit-producing Hood River Valley of Oregon appears to be limited. In a survey of the arthropod generalist predators inhabiting unsprayed pear (Booth, unpubl. data), this species was found almost exclusively at elevations less than 1,000 ft. At the Mid-Columbia Agricultural Research and Ex- tension Center, weekly observations were made in a small block of unsprayed D’ An- jou pear trees adjacent to a diverse vegetative community dominated by oak and pine. Although O. minutus nymphs were not distinguished from O. tristicolor, first generation adults appeared in mid- July, following O. tristicolor. Numbers of O. minutus exceeded those of O. tristicolor by early August, by which time O. minutus comprised about 10% of the total predators observed in that block. ACKNOWLEDGMENTS The following institutions kindly loaned us specimens for study: University of British Co- lumbia (UBC), G. G. E. Scudder and S. J. Cannings; California Academy of Sciences (CAS), P. H. Amaud Jr.; Oregon State Department of Agriculture (OSDA), R. L. Westcott. We also thank H. Riedl, Mid-Columbia Agricultural Research and Extension Center (MCAREC) for support and the loan of specimens. N. H. Anderson provided specimens of Orius nymphs. We thank B. B. Hall for the fine habitus drawing of adult O. minutus and A. C. Asquith for the illustrations of the nymphs. LITERATURE CITED Anderson, N. H. 1962. Anthocoridae of the Pacific Northwest with notes on distributions, life histories, and habits (Heteroptera). Can. Entomol. 94(1 2): 1325-1 334. Bachelor, J. S. and R. M. Baranowski. 1975. Paratriphleps laeviusculus, a phytophagous anthocorid new to the United States (Hemiptera: Anthocoridae). Florida Entomol. 58: 157-163. Butler, E. A. 1923. A biology of the British Hemiptera Heteroptera. London, 682 pp. Carayon, J. and J. R. Steffan. 1959. Observations sur le regime alimentaire des Orius et particulierement d'Orius pallidicornis (Reuter) (Heteroptera, Anthocoridae). Cah. Nat., Bull. N.P., n.s., 15:53-63. Chu, Y. 1969. On the bionomics of Lyctocoris beneficus (Hiura) and Xylocoris galactinus (Fieber) (Anthocoridae, Heteroptera). J. Faculty of Agriculture, Kyushu University 1 5(1): 1-136. Cobben, R. H. 1978. Evolutionary trends in the Heteroptera. Part II. Mouthpart-structures and feeding strategies. Mededelingen Landbouwhogeschool Wageningen 78-5. J. Veen- man and Zonen B.V., Wageningen, Nederland, 407 pp. Collyer, E. 1953. Biology of some predatory insects and mites associated with the fruit tree red spider mite (Metatetranychus ulmi (Koch)) in south-eastern England. II. Some im- portant predators of the mite. J. Hort. Sci. 28:85-97. Dicke, F. F. and J. L. Jarvis. 1962. The habits and seasonal abundances of Orius insidiosus (Say) (Hemiptera-Heteroptera: Anthocoridae) on com. J. Kansas Entomol. Soc. 35(3): 339-344. Fauvel, G. 1974. Sur falimentation pollinique d’un anthocoride predateur, Orius (Heterorius) vicinus Rib (Hemiptere). Ann. Zool. Ecol. anim. 6:245-258. Fulmek, L. 1930. Zur Kenntnis der Entwicklungsstadien von Triphleps minuta L. (Antho- coridae, Hemiptera, Heteroptera). Zeit. wiss. Insektenbiol. 25:82-88. Henry, T. J. 1988. Family Anthocoridae. Pages 12-28 in T. J. Henry and R. C. Froeschner (eds.). Catalog of the Heteroptera, or True Bugs, of Canada and the Continental United States. E. J. Brill, Leiden, 958 pp. 416 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Herring, J. L. 1966. The genus Orius of the Western Hemisphere (Hemiptera: Anthocoridae), Ann. Entomol. Soc. Am. 59(6): 1093-1 109. Kelton, L. A. 1963. Synopsis of the genus Orius Wolff in America north of Mexico (Heter- optera: Anthocoridae). Can. Entomol. 95(7):63 1-636. Kelton, L. A. 1 978. The insects and arachnids of Canada. Part 4. The Anthocoridae of Canada and Alaska. Heteroptera: Anthocoridae. Canada Department of Agriculture. Publ. 1639. 1-101 pp. Pericart, J. 1972. Hemipteres Anthocoridae, Cimicidae et Microphysidae de I’ouest-Palearc- tique. Faune de I’Europe et du Bassin Mediterraneen No. 7. Masson et Cie, Paris, 402 pp. Southwood, T. R. E. and D. Leston. 1959. Land and Water Bugs of the British Isles. F. Wame & Co., London, 436 p. Tonks, N. V. 1953. Annotated list of insects and mites collected on brambles in the Lower Fraser Valley, British Columbia, 1951. Proc. Entomol. Soc. Brit. Col. 49:27-28. Wagner, E. 1952. Die europaischen Arten der Gattung Orius Wff. (Hem. Het. Anthocoridae). Not. Entomol. 32:22-59. Wolff, J. F. 1801-1811. leones cimicum descriptionibus illustratae. J. J. Palm, Erlangen ( 1 800, 1:1-40; 1801, 2:43-84; 1802, 3:85-126; 1804, 4:127-166; 1811, 5:167-208). Xambeu, V. 1903. Moeurs et metamorphoses des Insects. Ann. Soc. Linn, de Lyon 50:208. Received May 4, 1989; accepted September 13, 1989. J. New YorkEntomol. Soc. 97(4):4 17-429, 1989 BIOLOGY OF LOPIDEA NIGRIDEA UHLER, A POSSIBLE APOSEMATIC PLANT BUG (HETEROPTERA: MIRIDAE: ORTHOTYLINAE) James D. McIver and Adam Asquith Systematic Entomology Laboratory, Oregon State University, Corvallis, Oregon 97331 Abstract.— The basic biology of Lopidea nigridea Uhler is described, including details of its growth, morphology, behavior, and ecology. Distribution and abundance of this univoltine, brightly colored plant bug were studied from May through August 1985, 1986, 1987 and 1988, at 1 5 sites within the cirque and valley of Pike Creek, on the eastern scarp of Steens Mountain, southeastern Oregon. At Pike Creek, L. nigridea occurs only on the legume Lupinus caudatus Kellogg and feeds on the stems, undersides of leaves, flower parts and developing seeds of its lupine host plant. This paper describes the relation between L. nigridea and its host plant, and identifies the principal species of visually oriented arthropod predators that occur on or visit this lupine. These predators may function as operators selecting for the evolution of aposematism in L. nigridea, and the biology of this plant bug species is placed within the context of how protective resemblance functions in a natural community. Lopidea nigridea Uhler (Miridae: Orthotylinae) is a brightly colored plant bug belonging to the tribe Orthotylini (Carvalho, 1958; Henry and Wheeler, 1988). Ge- neric relationships within this large complex tribe have yet to be resolved, but Lopidea appears related to those genera with sericeous scalelike setae and a single tergal process on the male genital capsule (Stonedahl and Schwartz, 1986). Lopidea is a New World genus with species occurring from Alaska to Honduras. The genus contains 103 species north of Mexico (Henry and Wheeler, 1988) and is now under review by the second author. As currently defined, Lopidea comprises those orthotylines with an oblique transverse suture on the gena and a single tergal process on the right side of the anterodorsal margin of the male genital aperture. Species vary considerably in size (3. 5-7. 7 mm), but most display some form of red- black, yellow-black or white-black color pattern. Lopidea nigridea Uhler belongs to a group of western species that are very similar in external morphology and general coloration. This species can be confidently dis- tinguished from related species only by examination of the male parameres— its highly variable coloration has contributed to the creation of at least 20 synonyms (Asquith, in press). Species determination was based on examination of type material of L. nigridea and its synonyms at the National Museum of Natural History, Washington, D.C., and the California Academy of Sciences, San Francisco, California, with special attention paid to morphological and color variation. From our study of museum specimens and literature records (Kelton, 1980), it appears that L. nigridea is widely distributed in western North America (Fig. 1), extending eastward through the north- ern plains. L. nigridea is the most common species of the genus west of the Rocky Mountains and has been collected from sea level to nearly 4,000 m in elevation. Although adults occasionally are found on a variety of different plants, in the western 418 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Fig. 1 . Distribution of Lopidea nigridea Uhler; Pike Creek study area, Steens Mountain, southeastern Oregon, •. part of its range it is typically associated with plants of the genus Lupinus L., whereas in the Rocky Mountains and northern plains it more commonly occurs on species of Astragalus L. At our study area in southeastern Oregon, adults of L. nigridea are large and robust with black calli, dark fuscous scutellum, clavus and medial aspect of the corium, with strongly contrasting red lateral margin of corium and pale red to white on the 1989 BIOLOGY OF LOPIDEA NIGRIDEA 419 Fig. 2. Adult male of Lopidea nigridea, dorsal habitus. 420 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Fig. 3. Fifth instar nymph of Lopidea nigridea, dorsal habitus. embolium and cuneus (Fig. 2). In other areas, this species may be almost solid bright red to dark fuscous with a bold white embolium and cuneus. The nymphs of L. nigridea possess similar contrasting colors, with late instars typically having a bright red abdomen with dark fuscous or black wing pads (Fig. 3). Although many species of Lopidea are bright red and black, information on their basic biology, especially as it relates to aposematism, is currently unavailable. Lopidea nigridea is ideal for the study of aposematism because it is easy to sample, easy to maintain in the laboratory, and is specific to a single species of lupine in southeastern Oregon, allowing accurate identification of the arthropod community with which it interacts. This paper describes the biology of L. nigridea and identifies the common arthropod species associated with it, including potential competitors and visually oriented predators. The study will serve as a base upon which more detailed work on aposematism in L. nigridea will depend. 1989 BIOLOGY OF LOPIDEA NIGRIDEA 421 Fig. 4. Maximum average per-plant abundance of Lopidea nigridea in samples of 50 Lupinus caudatus plants at 1 5 sites in the Pike Creek Drainage, Steens Mountain, southeastern Oregon, May-July 1987. Hatched areas denote primary sites. METHODS AND MATERIALS Study area. The field research was carried out from May through August of 1985, 1986, 1987 and 1988 on the east escarpment of Steens Mountain, southeastern Oregon (1 18®32'30"W; 42°32'30"N). Lopidea nigridea populations were studied at 1 5 sites along an altitudinal gradient from 1,353 m to 2,286 m (4,400-7,500 ft), within the Pike Creek drainage system (Fig. 4). From these 1 5 sites, we chose four primary sites for intensive study (JOE— 1,600 m, mPLAT— 1,661 m, CONF— 1,690 m, uHILL— 1,800 m), and from which to collect specimens for various aspects of the research. Plant communities of the primary sites at Pike Creek are dominated by sagebrush {Artemisia tridentata Nutt.) and rabbitbrush {Chrysothamnus nauseosus (Pall.) Brit- ton), with a variety of other shrubs, herbs and grasses intermixed (Great Basin Province; Franklin and Dyrness, 1973). On south-facing slopes and in flat open areas, the most common herbaceous plant is Lupinus caudatus Kellogg (Fabaceae), the only known host plant for Lopidea nigridea in the northern Great Basin Desert of south- eastern Oregon. 422 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Methods. Adults and 3rd, 4th, and 5th instars of L. nigridea were collected in the field for description and illustration. Some of these were reared on a lupine diet for estimates of instar duration at ambient field temperatures (range of highs: 23-32°C; range of lows: 5-1 3°C; 5-30 June 1987). Seventeen field-collected females were dis- sected to obtain estimates of fecundity. Behavioral observations were made of both nymphs and adult L. nigridea on their lupine host plant, and time budgets constructed for each set of observations. Casual observations were also made at irregular intervals to supplement the time budget data. Relative abundance, expressed as frequency of individuals per sample of 50 plants, was estimated for the four primary sites at regular intervals over the active portion of L. nigridea's life cycle (late April to early July 1986, 1987). To obtain a sample, bugs on each of the 50 plants were shaken onto a 0,75 x 0.75 m beating sheet, and then aged, recorded and returned to the plant. From these data, we also calculated 6 a “deme development index” (2^/^)’ where i = instar; ni = # individuals of age i=l i; N = total # individuals. This index provides an estimate of the effect of elevation on the speed and initiation of post-embryonic development in L. nigridea at Pike Creek. Distribution of L. nigridea was examined on a geographic scale (using information from the literature and from museum collections), over the fifteen sites at Pike Creek (using mean 50-plant sample densities), and among individual plants at each primary site (using mean and variance of 50-plant samples). The dispersion pattern of indi- viduals among plants within each site is described graphically, using a regression of Mean Sample Density {x = N/50) against Lloyd’s Mean Crowding Index (X* = x -\- {s^/x — 1); Lloyd, 1967. This regression is then compared to the co-occurring, ant- mimetic plant bug Coquillettia insignis Uhler (Phylinae). At five-day intervals from 29 May through 9 July 1985, 50-plant samples were taken from lupine at the CONF and mPLAT sites to identify the arthropods with which L. nigridea potentially interacts. Special attention was paid to other herbivores and visually oriented arthropod predators. Avian and lizard predators observed foraging on or around lupine are also reported. RESULTS AND DISCUSSION Life cycle. Overwintered eggs of L. nigridea begin to hatch during mid to late April at low elevations (<1,500 m), and early to mid May at high elevations (>1,750 m), and first instar nymphs can be found in the field until early to mid June. All instars of L. nigridea are active feeding stages, and development takes place during a time when host lupines are producing new growth, flowers, and seeds. Minimum time for development from early first instar to young adult is about 30 days. [Because we were unable to obtain reliable data on development in the lab, the estimate of 30 days is based on a comparison of field samples with samples taken for the sympatric plant bug Coquillettia insignis, for which we have good lab estimates of developmental time. The average C. insignis individual requires a minimum of 29.86 days to develop from early first instar nymph to adult in the lab (Mclver and Stonedahl, 1987a). Since the average speed of development for populations of C. insignis and L. nigridea 1989 BIOLOGY OF LOPIDEA NIGRIDEA 423 Fig. 5. Immature stages of Lopidea nigridea. A. Third instar nymph. B. Fourth instar nymph. C. Fifth instar nymph. D. Terminal abdominal stemites of 5th instar male. E. Terminal ab- dominal stemites of 5th instar female. in the field is the same at about 1 1 days per nymphal stage (see Fig. 7), we can infer that the minimum time required for a L. nigridea individual to develop is about 30 days.] By late June, at elevations less than 2,000 m, most individuals have reached the adult stage. Depending on plant condition, adults can be found on their host plants for nearly four weeks after the final molt. Females lay eggs from late June through most of July, usually on the same host plant upon which they developed. By early August, except in the most protected sites, lupine host plants have senesced, and the active portion of the life cycle of L. nigridea has ended, with the eggs overwintering to complete the life cycle. Description of immature stages. NYMPHS (Fig. 5a-c). Nymphal descriptions and illustrations are based on alcohol-preserved specimens displaying normal growth and orientation of the various body regions and appendages. Only 3rd-5th instars are treated, as these three stages adequately represent the variation normally observed in the juvenile form. Third through fifth instar nymphs are most easily differentiated by overall size and length of the wing pads. Sex can be determined by differences in 424 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) the position and shape of the sclerotized plates on the 8th and 9th abdominal seg- ments, and the larger size of females. All measurements are in millimeters. THIRD INSTAR (Fig. 5a). Length 1 .92-2.23; bright red; head and thoracic tergites darker red with scattered fuscous areas; dorsum with sparsely distributed black setae of various lengths; legs and antennal segments I-III also with black setae; antennal segment IV with more densely distributed, shorter pale setae. Head: subtriangular, vertical; dorsum dark red, ventral surfaces lighter; eyes large, bright red; width across eyes 0.73-0.74; only slightly broader than pronotum; vertex straight, with row of 6- 8 dark setae at posterior margin; frons broad, slightly convex; antennal fossa large, situated midway between eye and base of rostrum; tylus short, darker red; jugum large, well dehned, sub-rectangular, oriented vertically; lorum small, well defined; buccula short; gena well developed with oblique suture running from anteroventral base of antennal fossa to below posterior margin of eye, unpigmented. Rostrum: length 0.83-0.85, tip black, reaching past mesocoxae and almost to metacoxae. An- tenna: linear, segment I slightly broader than II-IV, narrowed basally; dark red, paler at joints; length of segment I, 0.25; II, 0.49-0.53; III, 0.47-0.52; IV, 0.40-0.41. Thorax: tergites dark red with fuscous areas concentrated at lateral aspects, stemites lighter red; pronotal disk broader than long (length 0.23-0.29; posterior width 0.65- 0.71), broadest posteriorly, anterior angles broadly rounded, posterior angles more acute, anterior and posterior margins nearly straight, calli indistinct; wing pads short, mesothoracic pair only slightly produced posteriorly, length 0.03-0.07. Abdomen: oblong-oval, deep red, midventral surface of stemites II-VII pale, weakly sclerotized; scent gland opening located between tergites III and IV, outlined with fuscous; ab- dominal sternite IX of female with dark brown sclerotized plates on either side of midline, male also with a pair of sclerotized plates on sternite IX but broader and slightly separated medially. Legs: coxae dark red with base and apex pale; trochanters pale; femora dark red with base, apex and ventral surface pale; tibia and tarsi uni- formly dark red; tarsi 2 segmented, segment I much shorter than segment II. FOURTH INSTAR (Fig. 5b). Similar to third instar except larger, and wing pads disproportionately longer. Head behind eye pale white, tylus and frons darker red, antennae closer to eye, antennal socket black; calli evident, dotted with fuscous; two plates on ventral surface of 9th abdominal segment more heavily sclerotized; coxae banded red and white, trochanters pale, femora and tibia darker, deep red to fuscous. Length 2.47-2.97. Head width across eyes 0.89-0.97. Rostrum: length 1.01-1.18. Antenna: I, length 0.30-0.33; II, 0.78-0.83; III, 0.68-0.75; IV, 0.42-0.48. Thorax: length of pronotal disk 0.39-0.40, posterior width 0.88-0.92, length of mesothoracic wing pad 0.31-0.38, reaching to anterior margin of second abdominal tergite. FIFTH INSTAR (Fig. 5c). Similar to fourth instar except body size larger and with antennal segment II longer and wing pads disproportionately much longer. Overall darker, antennal segment IV thinner, posterior angles of pronotum more rounded, posterior margin black, wing pads divergent and darker, medial areas black. Length 3.2-4.29. Head width 1.08-1.13. Rostrum: length 1.32-1.41. Antenna: I, 0.38-0.50; II, 1.14-1.28; III, 0.99-1.05; IV, 0.50-0.56. Thorax: length of pronotal disk 0.61- 0.64, posterior width 1 .09-1 .28; length of mesothoracic wing pad 0.85-1 .02, reaching to anterior margin of fifth abdominal tergite. Abdomen: sclerotized plates on ab- dominal sternite IX much larger than in earlier instars, female also with well-de- veloped plates on posterior half of sternite VIII (Fig. 5d, e). 1989 BIOLOGY OF LOPIDEA NIGRIDEA 425 INDIVIDUALS per PLANT Fig. 6. Dispersion of Lopidea nigridea individuals among plants, at low density sites (<1.5 individuals per plant) and high density sites (>1.5 individuals per plant). Each point represents a sample characterized by a mean density (x-axis) and a mean crowding index (y-axis; y = X* = Jc + (sVx - 1); dashed lines represent lines of slope 1. Fecundity estimate. A total of 17 adult females were collected at the HILL site on 1 8 and 26 July 1988, and dissected to determine egg load. Average egg load was 1 2.4 ±5.5 SD, with a range of 3 to 26 eggs per female. Distribution and abundance. Populations of L. nigridea can be found on Lupinus caudatus nearly everywhere the lupine grows in southeastern Oregon. At Pike Creek, L. nigridea is widely distributed on lupine from 1,300 to 2,250 m elevation. Among sites, however, L. nigridea is patchy in distribution, being very common at some localities and absent or rare at others (Fig. 4). Population density per site tends to remain constant over the years, so that a similar among-site distribution at Pike Creek was observed for all four field seasons (1985, 1986, 1987, 1988). Site constancy was extreme in some cases: at four sites within a 9 hectare area at Pike Creek, L. nigridea was never observed at uuHILL, HILL, and Fork, but achieved its greatest observed abundance for all four field seasons at the intermediate elevation site uHl (Fig. 4). Extreme site fidelity is probably due to a combination of factors, including the tendency for females to oviposit on the host plant upon which they develop, and the tendency for adults of both sexes to fly only short distances. Within a typical site, L. nigridea individuals are also patchily distributed, especially compared to the ant-mimetic plant bug Coquillettia insignis Uhler (Mclver and PER- PLANT ABUNDANCE 426 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) MAY JUNE JULY MAY JUNE Fig. 7. a. Number of Lopidea nigridea individuals per lupine plant from late April to late June 1987, at the four primary sites in the Pike Creek drainage system, b. Regressions of deme 6 development index n/N) versus time for the four primary sites in the Pike Creek drainage, i=l southeastern Oregon. Lattin, in press— Fig. 3). Regressions of Lloyd’s Mean Crowding Index on Density show that L. nigridea exhibits a wide range of densities among sites, a high average density per plant when it occurs, and a slope of density versus crowding index much greater than 1.0, all features of animals with aggregated distributions (Fig. 6). At some sites (e.g., uHILL), an average per-plant density of almost 10.0 individuals was consistently observed, nearly 1 0 x greater than any density observed for Coquillettia. Such extreme aggregation of individuals may be an adaptation related to aposematism (Edmunds, 1974): interestingly, L. nigridea is not only brightly colored and aggregated, but is known to be distasteful to some arthropod predators (Mclver, 1989; Mclver and Lattin, in press). The pattern of abundance of L. nigridea over time was strongly influenced by elevation (Fig. 7a). Elevation can be expected to influence temperature regimes, and so both the host plant and L. nigridea populations will tend to develop later at higher, cooler sites. When deme development index is regressed over time for the four primary sites, populations at higher sites clearly start later and develop faster than populations at lower sites (Fig. 7b). Presumably, this is because by the time eggs hatch at higher sites (early June at uHILL), mean ambient temperatures are higher, and so postembryonic development proceeds more rapidly compared to lower sites, where development starts much earlier (early May at JOE). 1989 BIOLOGY OF LOPIDEA NIGRIDEA 427 Behavior. Lopidea nigridea is an alert, visually oriented diurnal plant bug, the adults and nymphs of which feed on the higher stems, undersides of leaves, flower parts and developing seeds of its lupine host plant. Like many other foliage-feeding plant bugs, the feeding activity of L. nigridea often causes stippling and chlorosis on the upper surfaces of plant tissues (Knight, 1941). Six individuals were observed on lupine for a total of 279 minutes and five distinct behavioral activities were identified: rest = no observable movement, plant bug very responsive, body horizontal, pro- boscis tucked in; feed = no observable movement, bug much less responsive, body horizontal, proboscis inserted in plant tissue; groom = stereotypic grooming move- ments, use of mouthparts and tarsi of anterior legs to groom antennae, eyes, head, proboscis and legs with posture dependent on which body part is being groomed; run = rapid movement usually up or around stem or leaf, usually followed by rest; probe = periodic insertion of proboscis into plant tissue, usually followed by feeding. The six observed individuals spent most of their time resting (59%), followed by feeding (33%), running (6%), grooming (2%), and probing (0.4%). Compared to the sympatric ant-mimetic plant bug Coquillettia insignis, L. nigridea is much less active (Mclver and Stonedahl, 1987a), and individuals tend to use a single individual host plant for much of their active lives. Although on the scale of an individual lupine plant, populations of both adult and nymphal L. nigridea are commonly aggregated, no social behavior (other than mating) was ever observed. After mating, females insert their eggs into stems of their host plant, in a manner similar to other plant bug species (Mclver and Stonedahl, 1987b). Predators. Three species of lizards occur at Pike Creek (collared, western-fence, side-blotched), although none was observed foraging in or around lupine. Five species of insectivorous birds often foraged on the ground in the vicinity of lupine plants (lazuli bunting, rock wren, canyon wren, sage sparrow, green-tailed towhee); the towhee commonly perch on the lower stems of lupine. As yet, there is no evidence that vertebrate predators feed on plant bugs or have a significant impact on plant bug populations at Pike Creek. Nearly 20 species of arthropod predators forage on L. caudatus during the time when L. nigridea is active (Table 1; Mclver, 1987; Mclver and Stonedahl, 1987a). Most of these are visually oriented predators, some of which feed on L. nigridea in the field and in the laboratory. Probably the most important predators of L. nigridea are the flower spider Misumenops celer (Hentz) and the assassin bug Sinea diadema (Fabricius). The crab spider Xysticus montanensis Keyserling and most jumping spiders {Phidippus spp., Metaphidippus spp., Sassacus papenhoei G. & E. Peckham) typically reject L. nigridea after attack, presumably because of some unpleasant prop- erty (Mclver and Lattin, in press). The observed distastefulness, the red and black coloration, and the tendency for populations to be highly aggregated in dispersion, all strongly indicate that L. nigridea is an aposematic species. This study dealt with a population of L. nigridea near the middle of its geographic range and intermediate in color pattern. Comparative data are now needed on dif- ferent color forms of L. nigridea and on populations in different areas using alternate host plants. Our future research will focus on how aposematism influences survi- vorship in natural populations of L. nigridea, with an emphasis on how host plant chemistry contributes to variation in distastefulness and coloration observed over L. nigridea's geographic range. 428 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Table 1 . List of arthropod species with which Lopidea nigridea potentially interacts, observed on Lupinus caudatus from 29 May to 9 July 1985; relative abundance on 3,625 plants sampled by beat sheet and sweep net; and temporal range at Pike Creek, 1,500-2,200 meters in elevation. Taxa Abun- dance on 3,625 plants Temporal range Herbivores Lopidea nigridea Uhler 328 early May-mid July Coquillettia insignis Uhler 641 early May-mid July Other herbivores 3,899 Visual arthropod predators Misumenops celer (Hentz) & M. asperatus (Hentz) 126 mid May-late July Thanatus spp. immatures 8 late May Tibellus chamberlini Gertsch 7 mid May-late July Xysticus montanensis Keyserling 2 early May-late July Misumena vatia (Clerck) 2 Sassacus papenhoei G. & E. Peckham 33 late May-late July Phidippus texanus-group 2 Synageles occidentalis Cutler 2 mid May-late July Oxyopes scalaris Hentz 17 mid May-late July Sinea diadema (Fabricius) 32 early June-early August Reduviidae immatures 30 mid May-mid June Nabicula vanduzeei (Kirkaldy) & Nabis alternatus uniformis Harris 47 early June-early August Phytocoris sp. immature 2 Geocoris sp. immature 1 Total visual predators (16 spp.) 311 Total non- visual predators (6 spp.) Other visual predators observed since 1985: Phidippus johnsoni Peckham & Peckham, Metaphidippus hele- nae (Banks), Metaphidippus insignis-group, Tutelina similis (Banks) (only in riparian area). 47 ACKNOWLEDGMENTS We thank Bonnie B. Hall for the habitus illustrations of the adult and nymph, and Anna Asquith for the illustrations of the immature stages of L. nigridea. We also thank John D. Lattin, Oregon State University, for providing work space in the Systematic Entomology Lab- oratory and for providing funds for parts of this study. Other funds were provided by grants from the National Geographic Society (3358-86) and the National Science Foundation (BSR- 8700 1 79) to the first author. Specimens were received for study from the Heteroptera collections of the following institutions: American Museum of Natural History, New York (Randall T. Schuh); California Academy of Sciences, San Francisco (Paul H. Amaud, Jr.); Canadian National Collection, Ottawa (Robert Foottit); Colorado State University, Fort Collins (Boris C. Kon- dratieff); Los Angeles County Museum, Los Angeles (Roy Snelling); National Museum of Natural History; Washington (Thomas J. Henry); Oregon State University, Corvallis (John D. Lattin); University of California, Berkeley (Jerry A. Powell); University of California, Davis (Robert 1989 BIOLOGY OF LOPIDEA NIGRIDEA 429 O. Schuster); University of California, Riverside (Saul Frommer); University of Kansas, Law- rence (Robert W. Brooks); Washington State University, Pullman (Richard S. Zack). LITERATURE CITED Asquith, A. In press. Taxonomy and variation of Lopidea nigridea Uhler. Great Basin Nat- uralist. Carvalho, J. C. M. 1958. Catalogue of the Miridae of the World. Part III. Subfamily Ortho- tylinae. Arq. Mus. Nac., Rio de Janeiro 47:1-161. Edmunds, M. 1974. Defence in Animals. A Survey of Antipredator Defences. Longman, Harlow, Essex. Franklin, J. F. and C. T. Dymess. 1973. Natural vegetation of Oregon and Washington. USDA For. Serv. Gen. Tech. Report PNW-8. Henry, T. J. and A. G. Wheeler, Jr. 1988. Family Miridae. Pages 251-507 in: T. J. Henry and R. C. Froeschner (eds.). Catalog of the Heteroptera, or True Bugs, of Canada and the Continental United States. E. J. Brill, Leiden and New York, 958 pp. Kelton, L. A. 1980. The Plant Bugs of the Prairie Provinces of Canada. Heteroptera: Miridae. The Insects and Arachnids of Canada. Part 8. Agriculture Canada Research Publication No. 1703. Ottawa, 408 pp. Knight, H. H. 1941. The plant bugs, or Miridae, of Illinois. 111. Nat. Hist. Surv. Bull. 22:1- 234. Lloyd, M. 1967. Mean crowding. J. Anim. Ecol. 36:1-30. Mclver, J. D. 1987. On the myrmecomorph Coquillettia insignis Uhler: arthropod predators as operators in an ant-mimetic system. Zool. J. Linn. Soc. 90:133-144. Mclver, J. D. 1989. Protective resemblance in a community of lupine arthropods. Nat. Geog. Res. 5:191-204. Mclver, J. D. and J. D. Lattin. 1990. Evidence for aposematism in the plant bug Lopidea nigridea Uhler. Biol. J. Linn. Soc., in press. Mclver, J. D. and G. M. Stonedahl. 1987a. Biology of the myrmecomorphic plant bug Co- quillettia insignis Uhler (Heteroptera: Miridae: Phylinae). J. N.Y. Entomol. Soc. 95:258- 277. Mclver, J. D. and G. M. Stonedahl. 1987b. Biology of the myrmecomorphic plant bug Orectoderus obliquus Uhler (Heteroptera: Miridae: Phylinae). J. N.Y. Entomol. Soc. 95: 278-289. Stonedahl, G. M. and M. D. Schwartz. 1986. Revision of the plant bug genus Pseudopsallus Van Duzee (Heteroptera: Miridae). Am. Mus. Novit. 2842:1-58. Received February 6, 1989; accepted June 13, 1989. J. New YorkEntomol. Soc. 97(4):430-437, 1989 REDESCRIPTION OF PLATYNUS PROGNATHUS VAN DYKE (COLEOPTERA: CARABIDAE: PLATYNINI) AND CIRCUMSCRIPTION OF LINDROTH’S DECENTIS AND HYPOLITHOS GROUPS James K. Liebherr Dept, of Entomology, Comstock Hall, Cornell University, Ithaca, New York 14853-0999 Abstract. — Platynus prognathus Van Dyke is redescribed and reinstated in Platynus Bonelli. Shared-derived characters indicate that it is most closely related to other North American species of a redefined hypolithos group; P. hypolithos (Say), P. angustatus Dejean, P. cincticollis (Say), and P. mannerheimii (Dejean). The American hypolithos group can be diagnosed by spermathecal configuration, and the species are consubgeneric with the European Batenus Mot- schulsky {=Platynidius Casey NEW SYNONYMY). The American decentis group members share a fundamentally different spermathecal configuration that is also present in Platynus assimilis (Paykull), the type species of Platynus sensu stricto (=Limodromus Motschulsky). This spermathecal configuration is also present in other Neotropical lineages of the genus. In 1 926, Van Dyke described Platynus prognathus from a single specimen collected at St. Simon’s Island, Georgia. Csiki (1931) transferred P. prognathus to Agonum Bonelli sensu stricto, while considering Platynus Bonelli another subgenus of Agonum. Erwin et al. ( 1 977) retained the species in Agonum while elevating Platynus to generic rank. In this paper I redescribe the species, including external adult characters cur- rently used to distinguish among genera of platynine Carabidae. Based on shared- derived characters, the species is reinstated in Platynus, and is considered a member of a redefined hypolithos group. Lindroth’s (1966) key to North American species of the hypolithos and decentis groups is modified to facilitate identification of P. prog- nathus. The hypolithos and decentis groups of Lindroth (1966) are realigned based on their different spermathecal configurations. Based on spermathecal configuration, Euro- pean species in the subgenus Batenus Motschulsky (type species Harpalus livens Gyllenhal) are consubgeneric with species of the American hypolithos group. Several other Old World Platynus are classified as Batenus based on possession of this shared- derived, and highly unique spermathecal configuration. Based on spermathecal con- figuration, Platynidius Casey is considered a junior synonym of Batenus. The other large piceous Platynus species in America north of Mexico belong to the decentis group. These species exhibit a fundamentally different configuration of female re- productive tract also present in Platynus assimilis (Paykull) {=Carabus angusticollis F.), the type species of Platynus sensu stricto (=Limodromus Motschulsky). Other Mexican and Neotropical lineages of Platynus studied to date also possess the sper- mathecal configuration present in Platynus s.s. 1989 PLATYNUS PROGNATHUS 431 Figs. 1-2. Platynus prognathus Van Dyke. 1 . Head and pronotum, dorsal view. Left antenna omitted beyond scape. Apical setae and fine setae of antennomeres omitted. 2. Fourth meta- tarsomere of right hindleg, dorsal view, as = apical seta. Platynus prognathus Van Dyke Diagnosis. Distinguishable from all other North American Platynus by the elongate sickle-shaped mandibles (Fig. 1), narrow cordate pronotum with rounded basal an- gles, and narrow, elongate, parallel-sided and weakly convex elytra. Description. HEAD. Eyes convex, protruding beyond outlines of postocular area; frons convex, ocular grooves broad, slightly wrinkled, extending broadly to posterior clypeal margin; labrum weakly bisinuate; mandibles elongate, with narrowly acu- minate apices and strong retinacular teeth, tooth on the right mandible anterad that on left (Fig. 1); antennal segments filiform, basal 3 antennomeres glabrous except for apical setae, apical 8 antennomeres with apical ring of setae and covered with fine setae; constriction of neck shallow but visible in side view; mentum with well- developed acuminate median tooth, depressions on mentum broad and deep, lacking a central pit. PROTHORAX. Pronotum with rounded basal angles, a minute jag in lateral margin at posterior seta (Fig. 1); laterobasal depressions impunctate, margin laterad depressions narrowly reflexed at hind angles; basal marginal bead weak but evident across pronotal base; disc relatively convex, median longitudinal impression fine, traversed by irregular wrinkles; anterior transverse depression deep medially, delimiting a shiny median anterior callosity; anterior marginal bead and front angles lacking; lateral margin very narrow in front of anterior setae; prostemal projection 432 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) weakly depressed medially, unmargined. ELYTRA. Humeri narrow, sides sub-par- allel for much of length, each elytron about 3.9 x as long as wide; basal groove weak, slightly angulate between sixth and seventh striae; lateral margin narrow in front, slightly wider from basal ‘A to subapical sinuation; apex beyond sinuation narrowly rounded; elytral striae complete, impunctate though slightly wavering; elytral inter- vals weakly convex; basal seta set between base of scutellar and first striae; 3 or 4 dorsal elytral setae, the anterior in stria 3 at basal 1/6, the others in stria 2; a single seta in seventh stria mesad subapical sinuation; lateral series of 13 setae (5 or 6 anterior, 1 or 2 medial, 6 posterior). PTEROTHORAX. Metepistemum elongate, length along lateral edge 2 x maximum perpendicular width; flight wing brachypter- ous, 0.4 X length of elytron, with reduced venation and no reflexed apex. LEGS moderately elongate, tarsi stout, lacking internal or external dorsal sulci; fourth tar- somere with inner and outer apical setae, but no subapical setae (Fig. 2); apical tarsomere with very short ventral setae, not visible except at high magnification ( 1 25 X ); fourth tarsomere on middle and hind legs slightly asymmetrical, outer apical lobe longer and broader than inner lobe (Fig. 2); foretibia with weak anterior and posterior longitudinal sulci; mesocoxa bisetose, 1 ventral seta and 1 seta on mesocoxal ridge; mesofemur with 3 anteroventral setae; metacoxa bisetose; metafemur with 2 anteroventral setae. COLOR. Head capsule brunneous; labrum and mandibles rufous; palps and basal V/i antennomeres testaceous, apex of fourth antennomere and apical 7 antennomeres darker due to rufopiceous cast; elytral disc brunneous; lateral margin testaceous; venter rufous; coxae rufotestaceous; femora, tibiae and tarsi testaceous. MICROSCULPTURE. Frons and vertex with isodiametric microsculpture, slightly stretched transversely on neck; pronotal disc with transverse mesh microsculpture, mesh more isodiametric in anterior transverse depression; elytral intervals with well- developed isodiametric mesh, the cells regularly aligned transversely. LENGTH. 10.9 mm (male). Holotype. Male (C.A.S. No. 1861); GA: St. Simon Island, 22 Apr- 12 May 191 1, J. C. Bradley; Van Dyke Collection. Notes. Van Dyke incorrectly sexed the holotype as a female. The front tarsi are dirty, but the squamose male adhesive setae can be seen on several tarsomeres. Moreover, the apical abdominal stemite bears 2 apical setae indicating that the specimen is a male. The holotype is somewhat teneral and was not dissected. Fattig (1949) reported collecting P. prognathus from Dalton, Georgia, in July. This material was apparently destroyed before the Fattig collection was deposited at the University of Georgia (C. Smith, pers. comm.) PHYLOGENETIC PLACEMENT AND CLASSIFICATION OF PLATYNUS Within Platynini, P. prognathus can be characterized as a Platynus based on the constricted neck, median mentum tooth, glabrous basal 3 antennomeres, unmargined pronotal projection, externally suleate foretibia, and smooth claws. If the genus Agon- urn is considered the sister to Platynus (Liebherr, 1986), and the subtribe Sphodri is considered the sister group to the subtribe Platyni (Liebherr, 1986; Casale, 1988), a eonstricted neck can be considered a shared-derived character of Platynus species. Various clades within Platynus have lost the constricted neck (e.g., the Platynus ovatulus group [Liebherr, 1988]), but this appears to be correlated with smaller body size. 1989 PLATYNUS PROGNATHUS 433 Figs. 3-4. Female reproductive track of Platynus species, ventral view. 3. P. hypolithos (Say). 4. P. decentis (Say), be = bursa copulatrix; gel = basal gonocoxite; gc2 = apical gonocoxite; mo = median oviduct; sg = spermathecal gland; sp = spermatheca; vl = ventral lobe of bursa. Within North American Platynus, P. prognathus shares rounded pronotal basal angles, a derived state, with P. mannerheimii, P. angustatus, and P. hypolithos. The latter 3 species plus P. cincticollis (Say) share a very distinctive spermathecal con- figuration, in which the spermatheca is sclerotized and appressed to the surface of the bursa copulatrix (Fig. 3). The spermathecal gland enters the dorsal surface of the sclerotized spermatheca. The bursa has a greatly expanded ventral lobe, several times as long as the spermatheca. P. prognathus shares flattened elytra with P. angustatus and P. hypolithos. The brachypterous wings of P. prognathus are more reduced than the full wings of P. cincticollis or the slightly reduced wings of P. mannerheimii, but longer than the vestigial wings of P. hypolithos and P. angustatus. Metepistemal development and wing configuration are correlated, with elongate metepistema present in P. cincticollis, P. mannerheimii, and P. prognathus, and quadrate metepistema in the other two 434 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Figs. 5-6. Female reproductive tract of Platynus species, ventral view. 5. P. livens (Gyl- lenhal). 6. P. assimilis (Paykull). Scale bar applicable to both figures. species. Thus, based on external characters, P. prognathus is the sister species to the sister pair P. angustatus and P. hypolithos. The spermathecal configuration observed in P. hypolithos and allies also occurs in the European P. livens Gyllenhal (Fig. 5) (Schuler, 1963). P. livens is the type species of the Platynus subgeneus Batenus Motschulsky 1864. P. hypolithos was assigned the type species of the genus Platynidius Casey 1920 by Lindroth (1966). Based on spermathecal configuration, Batenus and Platynidius represent the same phyletic line, and Platynidius should be considered a junior synonym of Batenus. The other large piceous Platynus from America north of Mexico are members of the decentis group, females of which exhibit a tubular spermatheca with a basally entering duct (Fig. 4). Other New World Platynus investigated to date uniformly exhibit similar tubular spermathecae (e.g., Liebherr, 1987, 1989). The spermathecal configuration of the decentis group species is shared with Platynus assimilis (Paykull) (Fig. 6), the type species of Platynus Bonelli 1810 (=Limodromus Motschulsky 1864). Based on spermathecal configuration, the decentis group species are members of the same phyletic line comprising European Platynus s.s. Jeannel (1942) used Platynidius for generic placement of American species allied to P. hypolithos, as well as any vestigially-winged European ""Platynus.'" Macropterous 1989 PLA TYNUS PROGNA THUS 435 Table 1. Classification of Platynus prognathus plus other species of Platynus treated by Lindroth (1966) along with flight-wing configuration; +/+, macropterous and most likely func- tional flight wings; +/-, flight wings present but reduced, flight not recorded; flight wings vestigial. decentis group hypolithos group trifoveolatus Beutenmuller (— /— ) hypolithos Say (-/-) ovipennis Mannerheim (—/—) angustatus Dejean (-/-) decentis Say (+/-) prognathus Van Dyke (+/-) brunneomarginatus Mannerheim (+/+) mannerheimii Dejean (+/— ) opaculus LeConte (+/+) cincticollis Say (+/+) parmarginatus Hamilton (+/+) tenuicollis LeConte (+/+) pecki group agilis LeConte (-/-) pecki Barr (-/— ) species were classified separately in the genus Agonum, subgenus Platynus. Wing reduction occurs in many lineages of Carabidae (den Boer et al., 1980), and the morphological changes correlated with wing loss cause convergence on particular syndromes of body shape in distantly related clades (Casale, 1988:106-107). In Platynus, wing loss in a number of taxa is correlated with what can be termed a rhadiniform body shape, i.e., rounded humeri, flattened elytra, widened apical half of the elytra, and lengthened head and legs. Such changes are predictable based on wing configuration. The differences in reproductive tract configuration outlined above, however, are more complex and have not been observed repeatedly in different groups. They involve the shape of the spermatheca, its level of sclerotization, the position of the spermathecal gland duct entrance to the spermatheca, and the presence or absence of a large ventral lobe of the bursa situated basal to the spermatheca. Moreover, the uniqueness within the Platynini and the high level of similarity for the spermathecal configuration observed in the American hypolithos group species and European species such as P. livens (Figs. 3, 5), supports its status as a synapo- morphy for these taxa. Based on a limited survey of Old World species, Platynus praedator Andrewes, P. scrobiculatus F., and P. willbergi Reitter share the appressed spermathecal configu- ration of P. livens. P. praedator possesses fully-developed flight wings, whereas the other two species are vestigially- winged. Clearly, other Old World species should be studied to determine the distribution of the two fundamentally different reproductive tract configurations. For the North American fauna, Lindroth’s (1966) hypolithos and decentis species groups can be realigned based on spermathecal condition (Table 1). Placement of P. prognathus in the hypolithos group is based on derived states of external characters shared with P. hypolithos and P. angustatus. These characters include pronotal shape, brachyptery, and flattened elytra. It is acknowledged that description of the female reproductive tract will be necessary to confirm this placement. When this classifi- cation is adopted, it is apparent that flight wing condition is not correlated with either species group. 436 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Lindroth (1966) includes the Californian P. agilis LeConte in the hypolithos group. This species is much smaller and more pallid than species in the hypolithos group, and has a tubular spermatheca not unlike that of decentis group species. It is cla- distically similar to P. {Microplatynus) pecki Barr (1982), plus other undescribed Mexican species. It is hereby removed from the decentis group and is considered a member of the pecki group (=s.g. Microplatynus). IDENTIFICATION OF P. PROGNATHUS Because Platynus is currently recognized as a genus distinct from Agonum (White- head, 1973; Liebherr, 1986), Lindroth’s (1966) key and classification must be mod- ified. Within Lindroth’s monograph, species 64-70 of his Agonum, plus other species on pp. 641, 645 and 646, should be considered Platynus. In order to distinguish P. prognathus from other Platynus treated by Lindroth (1966:559-560), the following modification can be inserted at couplet 32. Figure numbers with an asterisk refer to Lindroth (1966). 32. Prothorax with completely disappeared hind angles (Figs. 1, 321a*) 32a - Hind angles of prothorax at least suggested (Figs. 321 b-P) 33 32a. Mandibles elongate, sickle-shaped (Fig. 1 ). Elytra narrow, parallel-sided, apex narrow with well-developed subapical sinuation. Legs testaceous, body brunneous P. prognathus Mandibles broad to near apex, the acuminate tip short. Elytra broadest at apical % of length, apex broadly rounded, subapical sinuation weak. Legs and body concol- orous, rufopiceous P. mannerheimii ACKNOWLEDGMENTS I thank David H. Kavanaugh (California Academy of Sciences, San Francisco) for loaning specimens of P. prognathus and P. assimilis, and Cecil Smith (University of Georgia, Athens) for information about the Fattig specimen(s) of P. prognathus. P. Morvan (Clamart, France) was kind enough to exchange specimens of Old World Platynus. This research was supported by Hatch Project NY(C) 139406. LITERATURE CITED Barr, T. C., Jr. 1982. Microplatynus, a new subgenus from the mountains of New Mexico (Coleoptera: Carabidae). Coleopt. Bull. 36:98-101. Casale, A. 1988. Revisione degli Sphodrina (Coleoptera, Carabidae, Sphodrini). Mus. Reg. Sci. Nat. Torino Monogr. 5, 1,012 pp. Csiki, E. 1931. Carabidae: Harpalinae 5. Coleopt. Cat. 1 15:739-1022. den Boer, P. J., T. H. P. Van Huizen, W. den Boer-Daanje, and B. Aukema. 1980. Wing polymorphism and dimorphism in ground beetles as stages in an evolutionary process. Entomol. Gener. 6:107-134. Erwin, T. L., D. R. Whitehead, and G. E. Ball. 1977. Family 4. Carabidae, the ground beetles. Biol. Res. Inst. Am., Kinderhook, New York, 68 pp. Fattig, P. W. 1949. The Carabidae or ground beetles of Georgia. Emory Univ. Bull. No. 7, 62 pp. Jeannel, R. 1942. Coleopteres Carabiques II. Faune de France 40:573-1 173. Liebherr, J. K. 1986. Cladistic analysis of North American Platynini and revision of the 1989 PLATYNUS PROGNATHUS 437 Agonum extensicolle species group (Coleoptera: Carabidae). Univ. Calif. Publ. Entomol. 106, 198 pp. Liebherr, J. K. 1987. A taxononomic revision of the West Indian Platynus beetles (Coleoptera: Carabidae). Trans. Am. Entomol. Soc. 1 12[1986]:189-268. Liebherr, J. K. 1988. Biogeographic patterns in West Indian Platynus carabid beetles (Co- leoptera). Pages 121-152 in: J. K. Liebherr (ed.). Zoogeography of Caribbean Insects. Cornell Univ. Press, Ithaca, New York. Liebherr, J. K. 1989. Redefinition of the Platynus jaegeri group and taxonomic revision of the Mexican and Central American species (Coleoptera: Carabidae: Platynini). Trans. Am. Entomol. Soc. 1 14[ 1988]: 167-2 14. Lindroth, C. H. 1966. The ground beetles of Canada and Alaska. Part 4. Opusc. Entomol. Suppl. 29:409-648. Schuler, L. 1963. Les organes genitaux femelles chez les Pterostichidae de France. Les tribus Anchomenini et Sphodrini (suite). Le cas des Patrobidae (Col. Carabiques). Bull. Soc. Entomol. France 68:13-26. Van Dyke, E. C. 1926. New species of Carabidae in the subfamily Harpalinae, chiefly from western North America. Pan-Pacif. Entomol. 2:1 13-126. Whitehead, D. R. 1973. Annotated key to Platynus, including Mexisphodrus and most “Co/- podes," so far described from North America including Mexico (Coleoptera: Carabidae: Agonini). Quaest. Entomol. 9:173-217. Received February 6, 1989; accepted March 14, 1989. J. New York Entomol. Soc. 97(4):438-447, 1989 AGGREGATION AND PREDATOR AVOIDANCE IN WHIRLIGIG BEETLES (COLEOPTERA: GYRINIDAE) K. VuLiNEC^ and M. C. Miller^ ^Cincinnati Museum of Natural History, and Department of Biological Sciences, University of Cincinnati, Cincinnati, Ohio 45221,^ and ^Department of Biological Sciences, University of Cincinnati, Cincinnati, Ohio 45221 Abstract. — Whirligig beetles (Coleoptera: Family Gyrinidae) aggregate on the surface of ponds, lakes, and streams. This study examines how these aggregations protect the beetles from pre- dation. The more beetles in an aggregation, the more quickly the group as a whole responds to the approach of stimuli. Experiments indicate that individual beetles either sight a stimulus themselves, or respond to waves generated by fleeing conspecifics. The distance between two beetles is important in determining how quickly a blinded beetle reacts to wave cues. Two hypotheses can explain the warning mechanism used by aggregations. (1) A high contact rate between aggregation members leads to increased physiological arousal which allows more rapid individual response, or (2) Environmental scanning is enhanced with the addition of more eyes to the group. Evidence from laboratory experiments supports the latter explanation. Whirligig beetles (Coleoptera: Family Gyrinidae) live in an exposed habitat: the water surface of ponds, lakes, and streams, where they aggregate in large rafts, some- times in multi-species groups. Rafts of 20,000 individuals have been reported (Hein- rich and Vogt, 1980). Beetles remain in these rafts all day, dispersing at dusk to forage singly. Most rafts appear to occur in the same location day after day, but individual beetles move around from one raft to another without apparent pattern (Heinrich and Vogt, 1980). Although highly conspicuous, gyrinids are not common prey of aquatic vertebrate predators (Benfield, 1972). In this paper, we examine the role of gyrinid aggregation in predator avoidance. Several hypotheses have been offered for this aggregating behavior. Brown and Hatch (1929) suggest it to be an orientation behavior due to habituation to certain visual patterns in the environment. On the other hand, Benfield (1972) and Heinrich and Vogt (1980) suggest a defensive function for gyrinid aggregations. Gyrinid beetles exude a strong-smelling secretion from the pygidial glands believed to be a defensive substance (Benfield, 1972; Meinwald et al., 1972; Miller et al., 1975; Newhart and Mumma, 1978; Heinrich and Vogt, 1980; Dettner, 1985). To demonstrate the nox- ious quality of the substance, Benfield (1972) fed gyrinids to fish and found (after a number of trials) that the fish rejected the beetles on sight. He hypothesized that the aggregations serve to advertise the gyrinids’ unpalatability. Heinrich and Vogt (1980) suggested that the groups occur in areas where there are no predators or where the predators have already learned to avoid the beetles. Defense appears to be the main function of these aggregations. Interactions among ^ Current address: Department of Ecology and Evolution, University of Chicago, Chicago, Illinois 60637. 1989 WHIRLIGIG BEETLES 439 individual beetles appear limited to maintaining interindividual distance, or sexual signaling (Kolmes, 1983a; see also Freilich, 1986), which excludes sociality as the function of the aggregations. The groups are not mating swarms, because aggregations occur throughout the months the adults are active, and not just during the mating season (Istock, 1967). In addition, pond-dwelling gyrinids disperse at night to forage singly (Heinrich and Vogt, 1 980; but see Kolmes, 1 983b; Vulinec and Kolmes, 1 987), suggesting that the rafts do not function in foraging. This study examines how gyrinid beetle aggregations function in predator avoid- ance by providing an early warning of predator approach. We first examined whether aggregation does allow an early warning of predator approach (group effect). We also determined the mechanism of information transfer among aggregation members. Finally, we tested two competing hypotheses that explain how early warning is ac- complished: (1) Beetles in large groups are more physiologically aroused, or (2) Beetles in large groups have more eyes available to scan the environment. MATERIALS AND METHODS Group effect. We performed a field experiment to test the effect of group size on group avoidance response. A human was used as a predator stimulus in this exper- iment to insure constant approach speed. One of us approached different-sized groups of gyrinids that were aggregated on ponds. Speed of approach was approximately constant at 1 50 cm/s. A point in the middle of the group was noted by an observer viewing through a Super-8 tripod-mounted camera. Measurement could then be made from this point to the location of the experimenter at the time that the entire group’s defensive movements began. Defensive movements can be easily distinguished from random swimming and the group reacts almost instantaneously. We obtained group sizes from the Super-8 film viewed with a stop-action projector. Thirty-five different groups were tested over a 4-day period. A regression was performed between reaction speed and group size and the best fit line was obtained by the least squares method. A t-iest was performed to determine the significance of the slope. Information transfer. We used temporarily blinded beetles to determine if beetles need to see the stimulus to react, and if the proportion of sighted beetles in a group is important to the speed of group reaction. Beetles were blinded by placing them in a foil-lined finger bowl and exposing them to a 150-watt photoflood lamp for 10 minutes (Kolmes, 1983b). We considered the beetle blinded if it did not react to a hand waved over it. Blinding was effective for 1 0 to 15 minutes after treatment and no experiments were run for longer than five minutes with the same beetles. In the first experiment, a varying proportion of 20 beetles was blinded. They were all placed in a large white porcelain testing arena (115 cm x 54 cm x 30 cm depth), which allowed good visibility; water level was maintained at 10 cm and temperature at 15°C. A predator stimulus (human hand) was shown from above and we recorded the time (up to a maximum of 60 seconds) at which all beetles in the group began moving defensively. Beetles were not reused, so the sample sizes of the groups were necessarily small. We conducted a laboratory experiment to examine the effect of a disturbed beetle’s proximity on the reaction speed of another. The initial distance between the two beetles was delimited by using three sizes of finger bowls (100, 200, or 300 mm in diameter) filled with 2 cm of 1 5°C water. The blinded beetle was placed in a bowl 440 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) and allowed approximately 3 minutes to acclimate, until it came to rest near the side of the bowl; the sighted beetle was then placed in the finger bowl at the farthest point from the blinded beetle. The sighted beetle immediately began defensive swimming: a rapid zig-zag movement very different from non-defensive swimming. This allowed us to record the time between the release of the sighted beetle and the initiation of defensive swimming by the blinded beetle. If the beetles made physical contact before the blinded beetle began defensive movements, or if the sighted beetle dove under the water, the trial was excluded. This experiment was conducted seven times for each size finger bowl, with different beetles each time. Data were analysed using a one-way analysis of variance. Chemical cues, in addition to tactile ones, might be used by gyrinid beetles to gain an early warning of danger. To determine if pygidial gland secretions are used in this context, we performed two tests: (1) A beetle’s secretion was milked from the pygidial glands onto a cotton swab. This secretion, mostly norsesquiterpene (Miller et al., 1975), is normally released whenever a beetle is held. The swab was carefully dipped into the water of an artificial pool near an aggregation of five beetles. We repeated this procedure with five different beetles and five different groups. (2) A beetle was held and squeezed to release its secretion, and the tip of its abdomen was placed in the water near an aggregation of five beetles. This trial was repeated five times with different beetles each time. We used only above water stimuli to invoke defensive swimming. We attempted to simulate underwater predation by moving a predator model (plastic fish) beneath a suspended glass bowl containing beetles. We got no response from any beetle even when the model was backlit. Hypothesis 1: increased physiological arousal. All beetles (Dineutes hornii Roberts 1895) used in the laboratory studies were collected with a dip net from five different sites in southwestern Ohio between August 1980 and June 1983. These beetles were placed into plastic quart containers half-filled with water, then kept in aquaria in the lab. Care was taken to disturb or handle them as little as possible. We performed a laboratory experiment to examine the hypothesis of physiological arousal. In the first experiment, we placed beetles of three different group sizes (1, 10, 20) in the water-filled testing arena and allowed them one hour to acclimate. A predator stimulus (in this case, the senior author) was shown from above. We recorded the time from the initiation of the stimulus to the defensive reaction of one beetle by timing the first beetle sighted on looking into the arena. Direction of sight was shifted to a different part of the arena for the beginning of every trial, which effectively randomized the trials. Because these beetles perform specific swimming movements in response to novel or sudden stimuli, we obtained accurate response times with a 0. 1 second stop watch. Each group size was tested 20 times with beetles randomly drawn from a common pool of 73 beetles. Data were analysed by a one-way analysis of variance. Hypothesis 2: environmental scanning. To determine if the beetles’ reaction speed is independent of the speed of a predator’s approach, we placed six beetles in a 20 cm diameter finger bowl that was half-filled with water. A predator stimulus (a black paper square 3 cm x 3 cm attached to a string) was lowered from a height of 90 cm toward the beetles at three different speeds, approximately 23 cm/s, 40 cm/s, and 77 cm/s. We then measured the distance of the black square from the beetles when all 1989 WHIRLIGIG BEETLES 441 Fig. 1 . Distance (m) a predator stimulus could approach a group of gyrinids before elicting a defensive response as a function of group size. Line is fitted by eye. six had begun defensive movements (which occurred within milliseconds of one another). All three approach speeds were tested 1 2 times with new beetles each time. Data were analysed by a one-way analysis of variance. RESULTS Group effect. When dilFerent-sized groups of gyrinids were approached by a human (with a constant approach speed), the reaction speed of the whole group (i.e., until the last beetle reacted) varied with group size (Fig. 1). Larger groups reacted signif- icantly faster to a predator stimulus than smaller groups {t = 7.5, df = 32, P < 0.00 1 , based on a linear regression [y = 0.03 lx + 5.48; r = 0.798]). Information transfer. Twenty beetles in a group with 0% or 10% blinded all reacted very quickly to a predator stimulus. When a greater percentage of beetles were blinded (25%-50%), reaction speed was much more variable; however, in all trials, every beetle in the group reacted with defensive swimming before 60 seconds had elapsed. When 75% or more were blinded, entire group reaction did not occur within the 60- second limit (Fig. 2). The size of the arena, and so presumably the distance between two beetles, is important in determining the speed of reaction (Fig. 3). The farther away a beetle is initially from its blinded neighbor, the longer it will take that neighbor to react (F = 24, df = 2 and 18, F < 0.01, ANOVA). 442 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) % BEETLES BUNDED Fig. 2. Response time (s) of twenty beetles as a function of the proportion of blinded beetles. Error bars include ±1 standard error. The N for each treatment is as follows: 0% = 13, 10% = 2, 25% = 5, 50% = 5, 75% = 2, 90% = 2, 100% = 2. Response time of all treatments with 75% or more beetles blinded was actually greater than 60 seconds; however, the observation period was terminated at that time. No beetle showed any reaction to the pygidial gland secretion either on a cotton swab or from the abdomen of a conspecific. Hypothesis 1: increased physiological arousal. Individual beetles from larger groups do not react faster than those in small groups (F = 0.9, df = 2 and 57, NS, ANOVA). Mean response times are 1.4 second, 1.5 second, and 0.9 second, for beetle group sizes of 1, 10, and 20. Hypothesis 2: environmental scanning. The beetles’ reactive distance varied in- versely with the approach speed of the predator stimulus (F = 21.43, df = 2 and 33, P < 0.001, ANOVA). Mean distances are 21.5 cm, 16.8 cm, and 8.2 cm respectively, for model speed of approach of 23 cm/s, 40 cm/s, and 77 cm/s. DISCUSSION Group effect and information transfer. Our field experiment demonstated that large groups of gyrinids are better able to avoid predators than small groups or single 1989 WHIRLIGIG BEETLES 443 DIAMETER OF ARENA (mm) Fig. 3. Response time (s) of a blinded beetle to defensive swimming of an intact beetle as a function of the diameter (mm) of the arena (initial distance between the two). Error bars include ± 1 standard error. individuals, as anyone who has tried to collect gyrinids with a dip net can verify. Experiments with the blinded beetles indicate that beetles react to the defensive movements of other beetles even when they cannot see the stimulus themselves. These beetles are reacting only to surface waves propagated by the defensive swim- ming reactions of the sighted beetles. In order to react with defensive swimming, a beetle must feel a neighbor’s waves from within a certain distance, an effect that may be due to wave attenuation, which occurs at distances of more than six body lengths (Tucker, 1969). When 0 to 10% of the group were blinded, the reaction speed of the entire group was less than 5 seconds, however, total group response was quite variable when 25% to 50% of the group were blinded. This result may indicate that there is 444 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) a threshold level for reaction to visually perceived stimuli that is lower than that for tactile stimuli. A response threshold in gyrinids may be a means of energy conser- vation. Alternatively, a large variance in interindividual distance within a partially blinded aggregation will result in a large variance in response time, due to the rela- tively slow speed of waves on the water surface. If response thresholds are to be demonstrated to mediate gyrinid escape behavior, this artifact must be experimentally or statistically factored out, a procedure that was beyond the scope of the present study. The distance between beetles is important in determining a beetle’s reaction speed, which suggests that a close aggregation may well be adaptive in transmitting predator defense information. Interindividual distance of aggregation members rarely exceeds 7 cm (Vulinec and Kolmes, 1987), the upper limit of wave propagation (Tucker, 1969). Because blinded beetles reacted to wave motion, we infer that tactile cues are very important in the defensive response of gyrinids, but that visual perception of a neighbor’s defensive swimming is not. Although gyrinids react quickly to perceived visual stimuli above the water surface, the function of the lower eyes is undetermined. Spectral sensitivity and electrophysiological studies of gyrinids reveal little difference between the two pairs of eyes (Carthy and Goodman, 1964; Bennett, 1967). Ana- tomical studies on the lower eye (Carthy and Goodman, 1964) do not indicate that it is adapted for sight under water, and our attempts to simulate the approach of a predator under the water surface dieted no reaction. A possibility is that the lower eyes are used during flight (D. Fong, pers. comm.). We also found that the pygidial gland secretion elicits no defensive response from a group of beetles. Defensive behavior occurred only in response to visual stimuli, or another beetle’s wave motion. Hypothesis 1: increased physiological arousal. In our experiments, individual bee- tles in large groups did not react any faster to stimuli than those in small groups or those that were solitary. This result indicates that beetles in larger groups are not more physiologically aroused than single beetles. Furthermore, pond gyrinids in aggregations contact each other at relatively low frequencies; less than 0.5 per minute (Vulinec and Kolmes, 1987). This low contact rate is unlikely to result in increased arousal. Contacts between individuals are also non-random (Freilich, 1986; see also Foster and Treheme, 1982), a behavior incompatible with a hypothesis of physio- logical arousal. Hypothesis 2: environmental scanning. Pulliam’s model (1973) explains why birds may aggregate in large numbers while foraging. According to this model, the reaction speed of all individuals to a predator is faster in large groups than in small because, in a large group, more eyes are available to scan the environment; therefore predators will be detected sooner. There are a number of empirical studies that support this hypothesis. Powell (1974) demonstrated that birds in flocks spend less time individ- ually in surveillance, but are able to detect a predator sooner than single birds. Similarly, Kenward (1978) found that a trained goshawk’s attacks on groups of pigeons became less successful the larger the group size. Sticklebacks actively pursue stray Daphnia in preference to a school (Milinski, 1977a, b), and predators such as squid, cuttlefish, pike, and perch experienced lowered success in capture the larger the group size of prey fish (Neill and Cullen, 1974). Our data demonstrate that large groups of gyrinids respond to a predator stimulus 1989 WHIRLIGIG BEETLES 445 more quickly than small groups. Because this finding also supports the physiological arousal hypothesis, we needed to show that the speed of the beetles’ response varied with the speed of predator approach. If a group’s response time to a stimulus is independent of the approach speed of that stimulus, and is based instead on intrinsic factors, there should be no difference in the time of response (reactive distance) of a group of five insects to a stimulus that moves toward them at different speeds (Tre- heme and Foster, 1980). Since we found that the reactive distance of the beetles varied inversely with the stimulus approach speed, and that individual beetles in large groups did not react more quickly than solitary beetles, we suggest that envi- ronmental scanning and not physiological arousal is responsible for the decreased response time in larger groups. Additionally, insects that rely on environmental scanning for early warning of danger will react quickly if they happen to see the danger themselves. Thus, there should be a great deal of variability in the response time of solitary insects, depending on whether they see the stimulus or not. However, in the fastest cases, a solitary insect should react as quickly as a group. In fact. Figure 1 shows that the fastest solitary insect responds as quickly to a stimulus as beetles in groups up to about 50 members. Predatory success on prey groups may be influenced by three factors: the dilution effect, increased detection capabilities of the prey, and the increased confusion of predators by many rapidly moving prey (Bertram, 1978). There is evidence that individuals in a group are protected just by being surrounded by conspecifics. For example, Foster and Treherne (1981) showed that fish attacks per individual Halob- ates robustus declined with increasing group size, an effect that is independent of any avoidance behaviors of the prey. The zig-zag swimming motion of whirligig beetles may serve to confuse predators. Additionally, these beetles swim extremely fast, with bursts up to 144 cm/s (Vulinec, 1987), a speed that approaches the burst swimming speed of possible fish predators (approximately 200 cm/s; Lagler et al., 1977). It seems likely that all three factors are important to whirligig beetle defense, with increased detection capabilities the front-line defense. The difference between ver- tebrate prey groups and these aquatic insects in their use of this defense is the use of substrate vibrational cues of danger, rather than visual or auditory ones. Tucker (1969) suggested that beetles use their own waves to echolocate, and Kolmes (1983b) found that beetles located prey by surface waves. Surface vibrational cues may also be used in precopulatory communication (Kolmes, 1985). Our data indicate that waves on the surface are also used as an early warning system. The transmission of the impulse would spread rapidly through the group, on an order similar to the “Trafalgar effect” observed in Halobates robustus by Treherne and Foster (1981), although without the necessity of actual contact between insects. The mean responsive distance of gyrinid groups in the field reaches an apparent asymptote above 60 to 80 individuals. This plateau indicates an upper limit beyond which adding more individuals does not contribute to increased detection capabilities for group members, and may explain why huge rafts of gyrinids are often divided into units of 50 to 100 individuals (Heinrich and Vogt, 1980). The pygidial secretion may play a significant role in whirligig beetle defense (Ben- field, 1972; Heinrich and Vogt, 1980). It is not known if the secretion is released into the water during escape and prior to capture. This possibility needs to be in- vestigated before pooling of defensive secretions can be proposed as an explanation 446 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) of gyrinid aggregations. Alternatively, early release of the secretion may assist the beetles’ movement across the water surface (Vulinec, 1987). Our data suggest that gyrinid beetles have an effective pre-attack defense. The pygidial substance may be used as a last resort defense, after the beetle has actually been captured. The hypothesis of environmental scanning is an extension of Hamilton’s selfish herd hypothesis (Hamilton, 1971). Grouping benefits the individual, both by de- creasing its chances of being singled out by a predator, and by increasing the number of eyes available to watch for a predator. Thus, a group can detect a predator sooner than a single individual. This tactic is especially effective for an animal in an exposed habitat such as the surface of a pond. The benefits of aggregation to gyrinid beetles are further enhanced by their ability to detect a neighbor’s defensive movements and react accordingly, whether or not they have sighted the stimulus themselves. Fossil gyrinid morphology (Hatch, 1927) suggests that gyrinid beetles were gen- eralized swimmers before they were surface swimmers. Because many other aquatic beetles and many terrestrial Adephaga possess chemical defenses and pygidial gland secretions (Blum, 1981; Dettner, 1985), we suggest that the chemical defense was present before water surface living evolved. Therefore, the following scenario is proposed for the evolution of gyrinid defenses: terrestrial existence ^ chemical de- fense ^ aquatic existence ^ exploitation of the water surface ^ aggregation as the primary defense in response to the exposed habitat. ACKNOWLEDGMENTS We wish to thank the Cincinnati Nature Center and Dravo Inc. for the use of their ponds. Jeff Burgess, Curt Meininger, Gail Stratton, and Patty Westlake assisted with the field work. Tom Kane, Monte Lloyd, and Jo Ann White provided valuable suggestions on earlier drafts of the manuscript, and Anita Buck assisted with editing. We would also like to thank T. Eisner and B. Heinrich for their comments. This research was partially supported by a grant from the Mining and Mineral Resources Research Institute. LITERATURE CITED Benfield, E. F. 1972. A defensive secretion of Dineutes discolor (Coleoptera: Gyrinidae). Ann. Entomol. Soc. Amer. 65:1324-1327. Bennett, R. R. 1967. Spectral sensitivity studies on the whirligig beetle, Dineutes ciliatus. J. Insect Physiol. 13:621-633. Bertram, B. C. R. 1978. Living in groups: predators and prey. Pages 65-96 in: J. R. Krebs and N. B. Davies (eds.), Behavioral Ecology: An Evolutionary Approach. Blackwell Scientific Publications, Oxford. Blum, M. S. 1981. Chemical Defenses of Arthropods. Academic Press, New York. Brown, C. R. and M. H. Hatch. 1929. Orientation and ‘fright’ reactions of whirligig beetles (Gyrinidae). J. Comp. Psychol. 9:159-189. Carthy, J. D. and L. J Goodman. 1964. An electrophysiological investigation of the divided eye of Gyrinus bicolor F. J. Insect Physiol. 10:431-436. Dettner, K. 1985. Ecological and phylogenetic significance of defensive compounds from pygidial glands of Hydradephaga (Coleoptera). Proc. Acad. Nat. Sci. Phil. 137:156-171. Foster, W. A. and J. E. Treheme. 1981. Evidence for the dilution effect in the selfish herd from fish predation on a marine insect. Nature 293:466-467. Foster, W. A. and J. E. Treherne. 1 982. Reproductive behaviour of the ocean skater Halobates robustus (Hemiptera: Gerridae) in the Galapagos Islands Oecologia 55:202-207. 1989 WHIRLIGIG BEETLES 447 Freilich, J. F. 1986. Contact behavior of the whirligig beetle Dineutus assimilis (Coleoptera: Gyrinidae). Entomol. News 97:215-221. Hamilton, W. D. 1971. Geometry for the selfish herd. J. Theor. Biol. 31:295-31 1. Hatch, M. H. 1927. A revision of fossil Gyrinidae. Bull. Brooklyn Entomol. Soc. 22:89-97. Heinrich, B. and F. D. Vogt. 1980. Aggregation and foraging behavior of whirligig beetles (Gyrinidae). Behav. Ecol. Sociobiol. 7:179-186. Istock, C. A. 1967. Transient competitive displacement in natural populations of whirligig beetles. Ecology 48:929-937. Ken ward, R. E. 1978. Hawks and doves: attack success and selection in goshawk flights at woodpigeons. J. Anim. Ecol. 47:449-460. Kolmes, S. A. 1983a. Precopulatory behavior of the whirligig beetle Dineutes discolor (Co- leoptera: Gyrinidae). J. N.Y. Entomol. Soc. 91:273-279. Kolmes, S. A. 1983b. Ecological and sensory aspects of prey capture by the whirligig beetle Dineutes discolor (Coleoptera: Gyrinidae). J. N.Y. Entomol. Soc. 91:405^12. Kolmes, S. A. 1985. Surface vibrational cues in the precopulatory behavior of whirligig beetles. J. N.Y. Entomol. Soc. 93:1 137-1 140. Lagler, K. F., J. E. Bardach, R. R. Miller, and D. R. M. Passino. 1977. 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N. 1974. Experimental analysis of the social value of flocking by starlings (Sturnus vulgaris) in relation to predation and foraging. Anim. Behav. 22:501-505. Pulliam, H. R. 1973. On the advantages of flocking. J. Theor. Biol. 38:419-422. Roberts, C. H. 1 895. Species of Dineutes of America north of Mexico. Trans. Amer. Entomol. Soc. 22:279-288. Treheme, J. E. and W. A. Foster. 1980. The effects of group size on predator avoidance in a marine insect. Anim. Behav. 28:1 1 19-1 122. Treheme, J. E. and W. A. Foster. 1981. Group transmission of predator avoidance behaviour in a marine insect: the Trafalgar effect. Anim. Behav. 29:91 1-917. Tucker, V. A. 1969. Wave-making by whirligig beetles (Gyrinidae). Science 166:897-899. V ulinec, K. 1987. Swimming in whirligig beetles: a possible role of the pygidial gland secretion. Coleopts. Bull. 41:151-153. Vulinec, K. and S. A. Kolmes. 1987. Temperature, contact rates, and interindividual distance in whirligig beetles (Gyrinidae). J. N.Y. Entomol. Soc. 95:481^86. Received May 24, 1989; accepted September 13, 1989. J. New York Entomol. Soc. 97(4):448-454, 1989 FIRST RECORD OF THE PALEARCTIC SPECIES OXYPODA OP AC A (GRAVENHORST) FROM NORTH AMERICA (COLEOPTERA: STAPHYLINIDAE: ALEOCHARINAE) E. Richard Hoebeke Department of Entomology, Comstock Hall, Cornell University, Ithaca, New York 14853-0999 Abstract. — Oxypoda opaca (Gravenhorst), a widespread Palearctic species, is reported for the first time from North America. The species is redescribed and diagnostic characters are provided to distinguish it from other Oxypoda species occurring in eastern North America. Distinguishing features are illustrated with line drawings and scanning electron photomicrographs. During the past several years, the unsorted Staphylinidae in the collections of North Carolina State University, Clemson University and the University of Vermont have been submitted to me for identification. Specimens of a large, but previously unrecognized, species of Oxypoda Mannerheim were found periodically in this ma- terial. These were subsequently identified by the author as Oxypoda opaca (Grav- enhorst), a species not previously reported from North America. Additional speci- mens have been found among the unidentified Aleocharinae in the collection of Cornell University, and only very recently have been collected by the author in New York State. The purpose of this paper is to record the presence of this Palearctic aleocharine in eastern North America, to redescribe and illustrate the species, and to present diagnostic characteristics to allow identification of the adults. Oxypoda, a cosmopolitan and predominantly temperate genus, is represented in the world by more than 350 recorded species. In North America, few, if any, of the nearly 1 00 recognized species (Moore & Legner, 1975; See vers, 1978) can be identified accurately since keys to species do not exist, and original descriptions and illustrations are inadequate to make precise determinations. North American Oxypoda have not been revised since the work of Casey (1893, 1906, 1911) who provided descriptions to the majority of the species (37 names in the East and 47 in the West). The remaining Nearctic species (12) have been named by Bernhauer (1905, 1907), Blatchley (1910), Erichson (1839), Maklin (1853), Notman (1920) and Sachse (1852). Members of Oxypoda can be recognized easily by the diagnostic combination of: strongly fusiform body shape; frontal suture present (though difficult to see); infraor- bital Carina sharply defined; hypomera not visible in lateral view; strong sinuation of outer, posterior margin of elytra; 5,5,5 tarsal formula; and first tarsomere of hind tarsus longer than tarsomeres II-IV combined. Oxypoda opaca (Gravenhorst) Aleochara opaca Gravenhorst, 1802:89. Oxypoda opaca: Mannerheim, 1831:483. Redescription. Dark ferrugineous-brown; head, basal areas of elytra adjacent to scutellum, and basal impressions of abdominal terga III-V dark brown to piceous. 1989 OXYPODA OPACA IN NORTH AMERICA 449 Figs, 1-4. Oxypoda opaca. 1. Labrum, dorsal aspect. 2. Right mandible, dorsal aspect. 3. Maxilla, dorsal aspect. 4. Labium, ventral aspect (setae of disc of mentum, except for 3 major setae near antero-lateral angles, not included in illustration). Paratergites, apical ‘/2-V3 of terga III-VII and apical Vs of sterna III-VII somewhat paler, rufobrunneous. Basal 3 antennal articles, mouthparts, and legs rufotestaceous. Length 3. 5-4. 5 mm. HEAD. Suborbicular, partially concealed by prothorax (Fig. 5). Eyes moderately large, slightly longer than length of temples; temples sharply margined below by infraorbital carina; dorsal surface densely and finely punctured and pubescent; pu- bescence (Fig. 5) subappressed, uniformly distributed, directed more or less anteriorly in most speeimens; surface between punctures with obsolete microsculpture of ir- 450 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Figs. 5-8. Oxypoda opaca. 5. Head, dorsal aspect. 6. Pronotum, dorsal aspect. 7. Left elytron, dorsal aspect. 8. Abdominal terga III-V. regular, transverse waves of microlines. Antenna 1 1 -segmented, relatively slender and long, reaching anterior V3 of elytra; articles I-III elongate. III slightly longer than II; articles IV-X becoming increasingly transverse; article XI conical, symmetrical, pointed apically, as long as or longer than IX + X combined. MOUTHPARTS. Labrum as in Figure 1 . Right mandible (Fig. 2) with small tooth at middle of inner margin, indistinctly crenulate before tooth; tooth absent from left mandible; ventral (condylar side) and dorsal (abcondylar side) molar regions smooth, without rows of denticles. Maxilla (Fig. 3) with galea slightly longer than lacinia; lacinia with compact row of large, somewhat recurved teeth on apical portion of inner surface, and patch of condensed spines and setae more basally; distal lobe of galea with compact patch of fine setae; maxillary palpus 4-segmented. Labium as in Figure 4; palpus 3-segmented, relatively slender, segment 1 longest, about equal in length to segments 2-3 combined; ligula broad, about as long as labial palpal segment 1 , shallowly bifid at apex, with 2 distinct pore-like structures at base; medial setae 2, bases moderately separated; longitudinal pseudopore field narrow, with many (approx. 15-20) large, closely spaced pseudopores forming a narrow longitudinal 1989 OXYPODA OPACA IN NORTH AMERICA 451 Figs. 9-12. Oxypoda opaca. 9. Mesostemal process. 10. Male tergite VIII. 11. Male stemite VIII. 12. Female stemite VIII. band; lateral fields with approximately 8-10 clumped pseudopores, 2 real pores and 1 setose pore distributed as in Figure 4. THORAX. Pronotum (Fig. 6) more or less transverse (L/W ratio = 0.71), about 1.4 times as wide as long; strongly convex in lateral view; anterior margin shallowly emarginate; antero-lateral angles sharply rounded and distinct; sides broadly rounded, broadest at base; posterior angles more broadly rounded; posterior margin broadly arcuate, with latero-posterior margin slightly sinuate in some specimens; hypomera not visible in lateral view; punctures and pubescence of dorsum dense, moderately coarse and uniformly distributed; pubescence (Fig. 6) along median line directed posteriorly in lower % and directed anteriorly in anterior V3, other pubsecence swirling laterally from median row [=Type III of Lohse, 1974 (Fig. 188, pg. 125)]; surface microsculpture similar to that of head. Elytra (Fig. 7) about 1.2 times as long as 452 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Figs. 13-17. Oxypoda opaca. 13. Male stemite IX. 14. Median lobe of aedeagus, lateral aspect. 15. Median lobe of aedeagus, ventral aspect. 16. Apical lobe of paramere. 17. Sper- matheca. pronotum and slightly wider at base than pronotum; each elytron uniformly convex; outer apical angles very strongly and broadly sinuate (Fig. 7); surface finely, but densely punctured and pubescent; pubescence uniformly directed posteriorly; integ- ument between punctures with minute microsculpture of wavy transverse micro- lines. Prostemum transverse, slightly cusped at middle. Mesocoxae narrowly sepa- rated. Mesosternal process (Fig. 9) slender, elongate, extending about 0.80 times length of mesocoxal cavity, apex acutely pointed. Metastemal process very short, extended about 0.14 times length of mesocoxal cavity, anterior margin broadly round- ed, margin defined by distinct bead. Mesocoxal acetabulae margined posteriorly by distinct carina. Ratio of mesosternal process : isthmus : metastemal process, 9. 1 : 1 : 1 .7. Tarsi 5,5,5 segmented, long and slender; first tarsomere long, slightly longer than articles II-IV combined. ABDOMEN. Abdomen (Fig. 8) tapered to apex, broad at base; first 3 visible tergites transversely impressed at base. Integument of abdomen with reticulate microsculp- ture. Apical margin of tergum VIII (males and females) shallowly emarginate at middle as in Figure 10. MALE. Sternum IX as in Figure 13. Median lobe of aedeagus as in Figures 14- 15. Apical lobe of paramerite as in Figure 16. FEMALE. Spermatheca as in Figure 17. Secondary sexual characteristics. Males: Apical margin of sternum VIII produced at middle (Fig. 1 1 ). Antennal article XI elongate, slightly longer than articles IX + X combined. Females: Apical margin of sternum VIII broadly rounded and with numerous close-set setulae (Fig. 1 2). Antennal article XI subequal in length to articles IX + X combined. 1989 OXYPODA OPACA IN NORTH AMERICA 453 Material examined. 2853, 1629. UNITED STATES: New York: Livingston Co., Letchworth St. Pk., 19-21 June 1981 (19)(CUIC). Tompkins Co., Ithaca, 1 November 1985 (355, 299) (CUIC); Ithaca, Forest Home, 10-17 May 1981, ex flight intercept trap (299) (CUIC); Ithaca, 10 March 1977, 1 7-24 & 24-31 May 1981 (255, 19) (CUIC); Town of Ulysses, N of Jacksonville, 6 June 1982, 15 June 1985, 22 June 1988, and 1 1 June 1989 (1355, 599) (CUIC); Danby, 2-3 April 1974 (15) (CUIC); 2 mi NW of West Danby, Vanbuskirk Gulf Rd., 29 June 1975 (19) (CUIC). North Carolina: Swaine Co., Clingmans Dome, 28 May 1936 (15) (NCSU). Wake Co., Clayton, 24 April 1964, ex chicken litter (355, 299) (NCSU); Clayton, 7 May 1964, under chicken feathers (555) (NCSU). South Carolina: Oconee Co., Mountain Rest, 24 April 1954 (19) (CUCC). Vermont: Caledonia Co., Saint Johnsbury, % mi E of Crow Hill, 30 May 1966, 1,140 ft (19) (UVCC). Voucher specimens of the North American populations are deposited in the col- lections of Cornell University, Ithaca, NY (CUIC); Clemson University, Clemson, SC (CUCC); North Carolina State University, Raleigh (NSUC); and the University of Vermont, Burlington (UVCC). The acronyms above follow Arnett & Samuelson (1986). Discussion. Adults of O. opaca can be readily distinguished from those of all other known North American species of Oxypoda by the following diagnostic combination: the larger body size (length 3. 5-4. 5 mm), the strongly fusiform body shape, the distinctive pubescence pattern of the pronotum, the long and slender antennae, the third antennal article slightly longer than the second article, and the distinctive characteristics of the male and female genitalia. This Palearctic (perhaps Holarctic) species is one of the most common and wide- spread of the genus and is found throughout Europe, Great Britain, North Africa, Asia Minor, Caucasus and Siberia (Horion, 1967). It is frequently found among haystack refuse, under vegetable detritus, in moss and fungi (Fowler, 1888), and in other decaying organic matter (Horion, 1967; Lohse, 1974). Specimens at hand from collections have been taken from a flight intercept trap and under an animal carcass in New York, and in chicken litter and under chicken feathers in North Carolina. In Tompkins Co., New York, in June 1989, numerous specimens were collected by the author beneath a very decayed pile of grass clippings. ACKNOWLEDGMENTS I thank the following individuals and institutions for the loan of specimens: M. W, Heyn, Clemson University, Clemson, South Carolina (CUCC); C. S. Parron, North Carolina State University, Raleigh (NCSU); and R. T. Bell, University of Vermont, Burlington (UVCC). I am also grateful to Drs. G. A. Lohse (Hamburg, W. Germany) and L. Zerche (Eberswalde, E. Germany) for confirming the identification of Oxypoda opaca. Drs. J. S. Ashe (University of Kansas, Lawrence) and J. H. Frank (University of Florida, Gainesville) kindly provided com- ments on a draft of this paper. LITERATURE CITED Arnett, R. H., Jr. and G. A. Samuelson. 1986. The Insect and Spider Collections of the World. E. J. Brill/Flora & Fauna Publications, Gainesville, Florida, 220 pp. Bemhauer, M. 1905. Neue Aleocharinen aus Nordamerika. Deutsche Entomol. Z. 1905:249- 256. 454 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Bemhauer, M. 1 907. Neue Aleocharini aus Nordamerika. (Col.) (3. Stuck.). Deutsche Entomol. Z. 1907:381-405. Blatchley, W. S. 1910. Aleocharinae. Pages 332-367 in: The Coleoptera or beetles (exclusive of the Rhynchophora) known to occur in Indiana with bibliography and description of new species. Indiana Dept. Geol. and Natur. Res. Bull. 1 . Casey, T. L. 1893. Coleopterological notices. V. Ann. N.Y. Acad. Sci. 7:281-606. Casey, T. L. 1906. Observations on the staphylinid groups Aleocharinae and Xantholinini, chiefly of America. Trans. Acad. Sci. St. Louis 16:125-434. Casey, T. L. 1911. New American species of Aleocharinae and Myllaeninae. Mem. Col. 2:1- 245. Erichson, W. F. 1839. Genera et species staphylinorum insectorum coleopterorum familiae, Pt. 1. Berlin, pp. 1-400. Fowler, W. W. 1888. The Coleoptera of the British Islands. A Descriptive Account of the Families, Genera and Species Indigenous to Great Britain and Ireland, with Notes as to Localities, Habitats, etc. Vol. 2. Staphylinidae. L. Reeve & Co., London, 444 pp. Gravenhorst, J. L. C. 1 802. Coleoptera microptera Brunsvicensia nec non exoticorum quotquot exstant in collectionibus entomologorum Brunsvicensium in genera familias et species distribut. Brunsvigae, 206 pp. Horion, A. 1967. Faunistik der Mitteleuropaischen Kafer. Bd. XI: Staphylinidae. 3. Teil: Habrocerinae bis Aleocharinae. 4 1 9 pp. Lohse, G. A. 1974. Aleocharinae, tribus 1-13 (Deinopsini-Falagriini). Pages 22-72 in: A. Freude, K. W. Harde and G. A. Lohse (eds.). Die Kafer Mitteleuropas, 5. Goeke & Evers, Krefeld. Maklin, F. G. 1853. [Description of new taxa]. Pages 95-273 in: C. G. Mannerheim, Dritter Nachtrag der Kaefer-Fauna der Nord-Amerikanischen Laender der Russichen Reiches. Bull. Soc. Imp. Nat. Moscou 26. Mannerheim, C. G. 1831. Precis d’un nouvel arrangement de la famille des brachelytres de I’ordre des insectes coleopteres. Mem. Acad. Imp. Sci. St.-Peterbourg. Vol. 1 (Mem. Sav. Etrangers), pp. 4 1 5-50 1 . Moore, I. and E. F. Legner. 1975. A Catalogue of the Staphylinidae of America North of Mexico (Coleoptera). Div. Agric. Sci., Univ. Calif, Special Publ. 3015. 514 pp. Notman, H. 1920. Coleoptera collected at Windsor, Broome Co., N.Y., 26 May to 5 June, 1918, with notes and descriptions. Jour. N.Y. Entomol. Soc. 28:178-194. Sachse, C. T. 1852. Neue Kafer. Stettiner Entomol. Z. 13:115-127, 142-149. Seevers, C. H. 1978. A generic and tribal revision of the North American Aleocharinae (Coleoptera, Staphylinidae). Fieldiana: Zool. 71:1-289. Received September 29, 1989; accepted October 30, 1989. J. New York Entomol. Soc. 97(4):455-470, 1989 TWO MOUTHPART MODIFICATIONS IN LARVAL NOTODONTIDAE (LEPIDOPTERA): THEIR TAXONOMIC DISTRIBUTIONS AND PUTATIVE FUNCTIONS G. L. Godfrey, ‘ J. S. Miller, ^ and D. J. Carter^ ‘Center for Biodiversity, Illinois Natural History Survey, Natural Resources Building, 607 E. Peabody Drive, Champaign, Illinois 61820; ^Department of Entomology, American Museum of Natural History, 79*^* Street at Central Park West, New York, New York 10024; and ^Department of Entomology, British Museum (Natural History), Cromwell Road, London SW7 5BD, England Abstract. — Two apomorphic features of the larval mouthparts in Notodontidae (Lepidoptera) are described and illustrated, and their taxonomic distribution within the family is documented. One hundred and fifty-four species of notodontid larvae were examined. These represent 90 genera and all currently-recognized subfamilies and tribes. The stipital lobe, a membranous projection on the dorsoposterior portion of the maxillary complex, varies in size and shape but occurs in 143 of the species studied. It does not occur in the Noctuidae, Lymantriidae, or Arctiidae. The majority of notodontid larvae exhibit a distinct developmental change in man- dibular morphology, from a serrate cutting edge in first instars to a smooth edge in later instars. A serrate mandible is typical of all instars in other noctuoids. Like the stipital lobe, presence of a smooth mandibular margin is broadly distributed among notodontids. Speculations con- cerning the functional significance of these mouthpart modifications are presented, and their potential use as synapomorphies for the family is discussed. The morphology of larval mouthparts has been quite thoroughly studied for the Noctuidae (e.g., Ripley, 1923), and mouthpart characters have been used in system- atic papers by numerous noctuid workers (e.g.. Crumb, 1929, 1956; Beck, 1960; Chu et al., 1963; Godfrey, 1972; Eichlin and Cunningham, 1978; McCabe, 1988; Mer- zheevskaya, 1988; Neil, 1988). Mouthparts of larval Notodontidae, in contrast, have rarely been investigated (Gardner, 1943, 1946; Godfrey, 1984; Weller, 1987). In this paper we discuss two modifications found in notodontid larvae. Grimes and Neunzig (1986) first described a structure, which they termed the stipital lobe, on the larval maxilla of two notodontid species, Macrurocampa mar- thesia (Cramer) and Schizura unicornis (J. E. Smith). The lobe is a projection on the stipes which obscures the dorsoposterior aspect of the galea. Grimes and Neunzig (1986) suggested that the stipital lobe may be uniquely derived for the Notodontidae because they did not find it in 23 other ditrysian families. Macrurocampa marthesia and Schizura unicornis are members of the tribe Het- erocampini (Forbes, 1948). To better understand the stipital lobe’s taxonomic dis- tribution and morphological variation, we examined additional species representing all five subfamilies and seven tribes currently recognized in the Notodontidae, in- cluding the Old World Thaumetopoeinae and Neotropical tribe Dioptini, the latter 456 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Figs. 1,2. 1. Heterocampa obliqua Packard, last instar (pinkish brown form) on Quercus macrocarpa Michx. (Fagaceae), central Illinois, USA. 2. Crinodes besckei (Hiibner), last instar (green form) on Gouania polygama (Jacq.) Urban (Rhamnaceae), Santa Rosa National Park, Guanacaste Province, Costa Rica. recently having been reduced from familial to tribal status (Minet, 1983; Weller, 1989). We also discuss a developmental change in mandibular morphology characteristic of notodontid caterpillars. The mandibles of almost all other Lepidoptera larvae, including noctuoids, bear a series of large teeth along each cutting edge (e.g., see figures in Peterson, 1962; Godfrey, 1972; McCabe, 1988; Neil, 1988). In most no- todontids, however, although the mandibles are toothed in first instars, the cutting edges are smooth in succeeding instars (Gardner 1943, 1946; Weller, 1987). After describing morphological variation in the stipital lobe and notodontid man- dible using Heterocampa obliqua Packard (Fig. 1) and Crinodes besckei (Hiibner) (Fig. 2) as examples, we document these character state distributions by examining larvae of 1 54 notodontid species. Finally, we offer speculations concerning the func- tional and phylogenetic significance of the two structures. MATERIALS AND METHODS Our study was based on alcohol-preserved specimens from the collections of J. G. Franclemont (Cornell University), Susan Weller (University of Texas, Austin), the American Museum of Natural History, the British Museum (Natural History), the Carnegie Museum of Natural History, the United States National Museum, and the Illinois Natural History Survey. Our species sample is broadly representative of the world fauna, and includes taxa from all biogeographic regions. We follow Forbes’ (1939, 1948) tribal and subfamilial arrangement of American notodontid species where possible, but the higher classification of the family is cur- rently rudimentary. Many of the genera we examined, especially from the Neotropics and Old World, have not been assigned to any tribe. The only available treatment for the world fauna (Gaede, 1934) recognized two subfamilies, the Notodontinae (360 genera) and the Pygaerinae (=Melalophinae) (6 genera), and it did not present a tribal classification. Because of these problems, we have simply arranged the genera examined in alphabetical order (Table 1), except that the Thaumetopoeinae and Dioptini are listed separately. We also made detailed examinations of larval mouth- 1989 MOUTHPARTS IN LARVAL NOTODONTIDAE 457 Table 1. Larval Notodontidae (final instar only) with stipital lobes present (L) or absent (N), and mandibular margins smooth (S) or toothed (T). Species Stipital lobe Mandib- ular margin® Collection” Afilia oslari Dyar L s JGF Antheua simplex Walker L s CMNH Cargida pyrrha (Druce) L T INHS, JGF Cerura vinula (Linnaeus) L S BMNH C sp. L S CMNH Cerurina marshalli (Hampson) L S BMNH Clostera albosigma (Fitch) L S INHS, JGF C. anachoreta (Denis & SchifFermiiller) L s BMNH C anastomosis (Linnaeus) L s BMNH C. brucei Hy. Edwards L s JGF C. curtula (Linnaeus) L s BMNH C. curtuloides (Erschoff) L s BMNH C. inclusa (Hiibner) L s INHS C. pigra (Hufnagel) L s BMNH Cnethodonta grisescens Staudinger L s BMNH Crinodes besckei (Hiibner) L T INHS Danima banksiae Lewin N S BMNH Dasylophia abbreviata Schaus L S BMNH D. anguina (J. E. Smith) L s INHS, JGF D. thyatiroides (Walker) L s JGF Datana contracta Walker L s USNM D. integerrima Grote & Robinson L s INHS, JGF D. major Grote & Robinson L s INHS, JGF D. ministra (Drury) L s BMNH, INHS D. perspicua Grote & Robinson L s USNM, INHS D. robusta Strecker L s USNM Desmeocraera ciprianii Berio L s BMNH D. latex Druce L s BMNH Didugua argent Hi nea Druce L s USNM Drymonia dodonaea (Denis & Schiffermiiller) L s BMNH D. dodonides Staudinger L s BMNH D. japonica (Wileman) L s BMNH D. ruficornis (Hufnagel) L s BMNH Dudusa synopla Swinhoe L s CMNH Eligmodonta ziczac (Linnaeus) L s BMNH Ellida caniplaga (Walker) L s INHS, JGF Epicerura ?tamsi Kiriakoff L s BMNH Epidonta brunneomixta (Mabille) N s BMNH Epodonta lineata (Oberthiir) L s BMNH Eufentonia nihonica (Wileman) L s BMNH Euhyparpax rosea Beutenmuller L s JGF Fentonia ocypete (Bremer) L s BMNH Furcula bicuspis (Borkausen) L s BMNH F. bifida (Brahm) L s BMNH F. borealis (Guerin-Meneville) L s INHS, JGF F. cinerea (Walker) L s INHS, JGF 458 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Table 1. Continued. Species Stipital lobe Mandib- ular margin® Collection*’ F. furcula (Clerck) L s BMNH F. scolopendrina (Boisduval) L BMNH Fusapteryx ladislai (Oberthiir) L s BMNH Gargetta ?divisa Gaede L s BMNH Gluphisia avimacula Hudson L s JGF G. lintneri (Grote) L s JGF G. septentrionis Walker L s INHS, JGF Goacampa variabilis Schaus L s INHS Harpyia microsticta (Hampson) L s CMNH H. umbrosa (Staudinger) L s BMNH Hemiceras nigrescens Schaus L s BMNH H. sp. L s INHS Heterocampa guttivitta (Walker) L s BMNH, INHS H. obliqua Packard L s INHS H. subrotata Harvey L s INHS H. umbrata Walker L s INHS Hippia packardii Morrison L s JGF Hupodonta pulcherrima (Moore) L s BMNH Hylaeora dilucida Felder L s BMNH Hyparpax aurora (J. E. Smith) L s JGF H. perophoroides Strecker L s USNM Hyperaeschra georgica (Herrich-SchafFer) L s BMNH, INHS, JGF Leucodonta bicoloria (Denis & Schiffermiiller) L s BMNH Leucophalera princei (Gundberg) L s BMNH Liparopsis postalba Hampson L s CMNH Lirimiris truncata (Herrich-Schalfer) L ? SJW Litodonta hydromeli Harvey L s USNM, SJW Lobeza suprema Schaus N s BMNH Lochmaeus bill neat a (Packard) L s INHS, JGF L. manteo Doubleday L s INHS, JGF Lophocosma atriplaga Staudinger L s BMNH Macrurocampa marthesia (Cramer) L s BMNH, INHS, JGF Microphalera grisea Butler L s BMNH Misogada unicolor (Packard) L s BMNH, INHS Nadata gibbosa (J. E. Smith) L s INHS, JGF Neostauropus basalis (Moore) L s BMNH Nerice bident ata Walker L s BMNH, INHS, JGF Notodonta dromedarius (Linnaeus) L s BMNH N. rothschildi Wileman L s BMNH N. scitipennis Walker L s JGF N. simplaria Graef L s JGF Nystalea nyseus (Cramer) L s BMNH N. sp. L s USNM Odontosia elegans (Strecker) L s JGF Oligocentria lignicolor (Walker) L s INHS, JGF O. semirufescens (Walker) L s USNM 1989 MOUTHPARTS IN LARVAL NOTODONTIDAE 459 Table 1. Continued. Species Stipital lobe Mandib- ular margin® Collection*’ Peridea anceps (Goeze) L s BMNH P. angulosa (J. E. Smith) L s INKS, JGF, USNM P. lativitta (Wileman) L s BMNH P. monetaria (Oberthiir) L s BMNH P. oberthueri (Staudinger) L s BMNH Phalera bucephala Linnaeus L s BMNH Pheosia fusiformis Matsumura L s BMNH P. gnoma (Fabricius) L s BMNH P. rimosa Packard L s BMNH, INHS, JGF P. tremula (Clerck) L s BMNH Psorocampa denticulata Schaus N s BMNH Pterostoma palpi na (Clerck) L s BMNH P. sinica Moore L s BMNH Ptilodon capucina (Linnaeus) L s BMNH P. hoegei (Graeser) L s BMNH P. jezoensis (Matsumura) L s BMNH Ptilophora plumigera (Denis & Schiffermuller) L s BMNH Pygaera timon Hiibner L s BMNH Quadricalcarifera punctatella (Motschulsky) L s BMNH Q. viridimaculata Matsumura L s CMNH Q. sp. L s CMNH Rosema sp. L s BMNH Schizura badia (Packard) L s USNM S. biedermani Barnes & McDunnough L s JGF S. concinna (J. E. Smith) L s BMNH, INHS S. errucata Dyar L s USNM S. ipomoeae Doubleday L s BMNH, INHS S. leptinoides (Grote) L s INHS, JGF S. unicornis (J. E. Smith) L s BMNH, INHS, JGF Scrancia stictica Hampson L s CMNH Seirodonta bilineata Packard L s BMNH Shachia circumscripta (Butler) L s BMNH Skewesia angustiora (Barnes & McDunnough) L s JGF Spatalia jezoensis Wileman L s BMNH Stauropus fagi (Linnaeus) L s BMNH Strophocerus pundulum (Schaus) L s BMNH Suzukiana cinerea (Butler) L s BMNH Symmerista albifrons (J. E. Smith) L s INHS, JGF S. canicosta Franclemont L s JGF S. leucitys Franclemont L s INHS, JGF Tarsolepis japonica Wileman & South L s CMNH Tecmessa elegans Schaus N s BMNH Theroa zethus Druce L T JGF Torigea straminea (Moore) L s BMNH Uropyia meticulodina (Oberthiir) L s BMNH Zaranga permagna (Butler) L s BMNH 460 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Table 1. Continued. Species Stipital lobe Mandib- ular margin® Collection*’ Thaumetopoeinae Anaphe panda (Boisduval) N ? BMNH E. melanosticta (Donovan) N ? BMNH Epicoma tristis Lewin N ? BMNH E. wilkinsoni Tams N ? BMNH Thaumetopoea pityocampa Denis & Schiffermiiller N 7 BMNH Discophlebia catocalina Felder N 7 BMNH Dioptini Cyanotricha necyria (Felder) L T AMNH Erbessa glaucaspis (Walker) L T BMNH Josia auriflua Walker L S BMNH J. cruciata Butler L S AMNH J. ligata Walker L S SJW J. flavissima (Walker) L S BMNH J. turgida Walker L S BMNH Phaeochlaena gyon (Fabricius) L T BMNH Phryganidia californica Packard L S AMNH, INHS, USNM Zunacetha annulata (Guerin-Meneville) L s AMNH Total number of genera examined = 90 Total number of species examined =154 “ In the thaumetopoeines and Lirimiris, scored as ‘ ‘?,” the mandibles have shallow dentations. These do not easily fit in either of the two categories. * Abbreviations; American Museum of Natural History, New York (AMNH); British Museum (Natural History), London (BMNH); Carnegie Museum of Natural History, Pittsburgh (CMNH); Illinois Natural History Survey, Champaign (INHS); John G. Franclemont Collection, Cornell University, Ithaca (JGF); Susan J. Weller Collection, University of Texas, Austin (SJW); United States National Museum, Washington, D.C. (USNM). parts in representatives of the other major noctuoid families, including the Lyman- triidae (2 species), Arctiidae (3 species), and Noctuidae (7 species). Mandibles were removed for study [see Godfrey (1972) for dissection procedure]. Stipital lobes were examined in two ways. All species were studied with dissecting stereomicroscopy while larvae were submerged in 70% ethanol. The lobes were fully exposed by removal of either the mandibles or the entire maxillary/hypopharyngeal complex. The stipital lobe in each of eight species was examined by scanning electron microscopy (SEM). For these, the entire maxillary/hypopharyngeal complex was removed, critical-point-dried, and mounted following the techniques outlined by Grimes and Neunzig (1986). The stipital lobes are membranous, and in some cases they collapsed during preparation; only those that retained their shape are figured with SEM in this paper. To observe the closing movement of the oral surfaces of the mandibles during adduction, we used 10% potassium hydroxide to clear the heads of last instar larvae of muscle tissue. We then cut away a small, posterior portion of the head capsule 1989 MOUTHPARTS IN LARVAL NOTODONTIDAE 461 Figs. 3-8. Scanning electron micrographs of last instar larval mouthparts. 3. Epicoma mela- nosticta (Donovan), left maxillary complex (dorsal; scale line = 100 /u). 4. Clostera albosigma Fitch, left maxillary complex (dorsal; scale line = 100 n). 5. Datana ministm (Drury), right maxillary complex (dorsal; scale line = 100 ix). 6. Heterocampa obliqua, right maxillary complex (dorsal; scale line = 50 ti). 7. H. obliqua, right maxillary complex (frontal; scale line = 40 m). 8. Phryganidia californica Packard, right maxillary complex (dorsal; scale line = 40 ti). Symbols: (H) hypopharynx; (M) mandible; (P) maxillary palpus; (SL) stipital lobe. 462 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) and inserted rnicroforceps through the enlarged area to grasp the mandibular adductor tendons and simulate the closing process. RESULTS Stipital lobes. Among the notodontid taxa examined, a stipital lobe occurs in 8 1 of 90 genera (90%) and in 143 of 154 species (92%) (Table 1). It is not found in the other noctuoid families we studied (see figures in Grimes and Neunzig, 1986). A small fold occurs on the dorsoposterior portion of the maxillary complex in Epicoma melanosticta (Donovan) (Fig. 3), a thaumetopoeine, but we do not consider this to constitute a lobe. There is no trace of a stipital lobe in the other thaumetopoeine genera we studied, Thaumetopoea, Discophlebia, and Anaphe. Five additional no- todontid genera lack the lobe. The size and shape of the stipital lobe varies significantly among Notodontidae. Relatively short, distally-rounded lobes characterize Clostera albosigma Fitch (Mela- lophinae) and Datana ministra Drury (Phalerinae) (Figs. 4, 5). The large lobe of Heterocampa obliqua (Fig. 6), which protrudes slightly from below the ventral margin of the mandible (Fig. 7), is similar to those of the two heterocampines studied by Grimes and Neunzig (1986). The type we observed most frequently has an acute tip, with the lobe often curving dorsally in front of the mandible (Figs. 15, 16). This type occurs in the Gluphisiini, Notodontini, Nystaleini, Hemiceratini, Dioptini (e.g., Zu- nacetha annulata Guerin-Meneville, Figs. 9, 10), and Cerurinae. A lanceolate lobe (Fig. 1 3) is found in Goacampa variabilis Schaus. In still other species, the stipital lobes are greatly swollen and curve upwards so far that they touch the distal margin of the labrum. Such lobes were found in the dioptines Erbessa glaucaspis (Walker) and Phryganidia californica Packard (Fig. 8), in the Old World genus Liparopsis, and in Ny staled nyseus (Cramer) (Nystaleini) (Figs. 17, 18). In all cases, the surface of the stipital lobe is spiculate (Fig. 1 1), and sometimes the spicules are quite prominent (e.g., Heterocampa obliqua. Figs. 6, 7). Stipital lobe shape does not appear to vary significantly within genera. Although stipital lobes can be most easily seen in last instar larvae, they occur in earlier instars as well. The lobes are present from the second to last instars of Het- erocampa obliqua and Crinodes besckei, but SEM revealed that they do not occur in the first instar. A second type of lobe was observed on the basolateral area of the stipes of Crinodes besckei (Fig. 1 2) and Nystalea nyseus. We call this the “basolateral lobe” to reduce potential confusion with the stipital lobe of Grimes and Neunzig ( 1 986). In C. besckei, it occurs on the maxilla of the first through fourth instars, but not the last. Early instars of N. nyseus were not available for study, but the lobe is prominent in the last instar. The internal morphology and function of the basolateral lobe are currently under study by G. Godfrey, J. B. Nardi, and D. H. Janzen. Mandibles. The first instar Heterocampa obliqua mandible bears five distal teeth and has a weakly concave oral surface with distally extending ridges (Fig. 22). How- ever, as is true for the majority of notodontid species, a dramatic change in man- dibular structure occurs during subsequent development. In the second through final instars, the teeth are lost and the distal edge is continuous, except for a slight notch near the dorsal comer. This notch demarks a low, dorsal tooth (Figs. 19-21, 23). 1989 MOUTHPARTS IN LARVAL NOTODONTIDAE 463 Figs. 9-14. Scanning electron micrographs of larval mouthparts. 9. Zunacetha annulata (Guerin-Meneville), labial and left maxillary complexes (frontal; scale line = 50 ii). 10. Zu- nacetha annulata, left maxillary complex (frontal; scale line = 25 n). 11. Zunacetha annulata, surface of stipital lobe, showing spicules (dorsal; scale line = 10 ij). 12. Crinodes besckei, first instar basolateral lobe (fronto-ventral; scale line = 100 ix). 13. Goacampa variabilis Schaus (dorsal; scale line = 50 fi). 14. C. besckei, last instar right mandible (oral view). Symbols: (BL) basolateral lobe; (H) hypopharynx; (M) mandible; (P) maxillary palpus; (SL) stipital lobe. Worn mandibles may lack the notch, making the distal edge appear entirely smooth. The series of oral ridges found in the first instar is also lost, being replaced by a single ridge that crosses the mandible between the ventral and distal comers. Associated but less pronounced changes include the following: (1) the adductor apodeme is positioned more distad of the rotational axis than in the first instar; and (2) the mandible protrudes and shows more medial curvature (Figs. 24, 25). 464 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) cephala Linnaeus (lateral). 17. Nystalea nyseus (Cramer) (lateral). 18. N. nyseus (frontal). Scale lines = 0.5 mm. Symbols: (L) labrum; (M) mandible; (SL) stipital lobe. A second type of mandibular morphology occurs among notodontids. First instar mandibles of Crinodes besckei (Fig. 27) are superficially similar to those of Hetero- campa obliqua, but in succeeding instars (Figs. 14, 26, 28, 29) they differ in significant ways. The most obvious differences are: (1) the distal, triangular teeth and oral ridges are retained; (2) platelike protuberances appear at the bases of the oral ridges; and (3) there is less mandibular protrusion and a more gradual medial curvature (Figs. 1989 MOUTHPARTS IN LARVAL NOTODONTIDAE 465 Figs. 19-23. Oral view of left mandibles of Heterocampa obliqua. 19. Second instar. 20. Third instar. 21. Fourth instar. 22. First instar. 23. Fifth instar. Scale line = 0.2 mm. Symbol: (D) dorsal tooth. 30, 31). In C. besckei the oral surface of the mandible becomes progressively more complex with each molt; “pockets” appear below the distal teeth (Fig. 14). These appear to receive the tips of the distal teeth on the opposing mandible (Fig. 32). DISCUSSION Mandibular modifications and functions. The teeth on the mandibular margin of first instar Heterocampa obliqua caterpillars are first used to chew an escape hole through the chorion, and then to slice and gouge leaf tissue between veinlets on the lower surfaces of fully expanded oak leaves (G.L.G., pers. obs.), leaving the top epidermal layer intact. The skeletonized leaf that results is typical of feeding damage caused by first instar notodontids (Riotte, 1 969; Godfrey and Appleby, 1 987; Weller, 1 987). Later instars clip through the leaf blade, beginning at the outer margin (Weller, 1987). This change in feeding behavior is correlated with loss of the mandibular Figs. 24, 25. Labrum and mandibles of Heterocampa obliqua (frontal view; labral and mandibular setae omitted). 24. First instar (scale line = 0.2 mm). 25. Second instar (scale line = 0.8 mm). Symbols: (L) labrum; (M) mandible. 466 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Figs. 26-29. Oral view of left mandibles of Crinodes besckei. 26. Second instar. 27. First instar. 28. Third instar. 29. Fourth instar. Scale line = 0.2 mm. Symbol: (D) dorsal tooth. teeth. Similar changes in mandibular structure were noted by Embree (1958) for larvae of Psilocorsis faginella (Chambers) (Lepidoptera: Oecophoridae). The feeding strategy of Crinodes besckei differs from Heterocampa obliqua. First instar caterpillars chew completely through the host leaf, Gouania polygama (Jacq.) Urban (Rhamnaceae), rather than skeletonizing it. The larva then enlarges the hole. Later instars clip through the leaf, either by chewing through the blade and then moving toward the margin, or by working toward the midrib from the margin. In both cases, larvae tend to avoid primary lateral veins (G.L.G., pers. obs.). First instar mandibles appear to be incapable of cutting leaf material after it has been removed from the blade. Heterocampa obliqua larvae feed on fully expanded oak leaves, which may present a greater ingestive and digestive problem than the tender, terminal Gouania polygama leaves on which first instar Crinodes besckei feed. By gouging out the softer tissue from oak leaves, first instar H. obliqua seem to avoid much of the indigestible material. Later instars of H. obliqua may partially masticate cut leaf tissue by pressing the distal edge of one mandible against the transverse oral ridge of the opposing one, a process observed by Bemays and Janzen (1988) for satumiids. In fifth instars, each mandible has an undulated area proximad of the transverse ridge (Fig. 23). These may have a mashing or crushing function; the surfaces appear to mesh as the man- dibles close. Figs. 30, 3 1 . Labrum and mandibles of Crinodes besckei (frontal view; labral and mandibular setae omitted). 30. First instar (scale line = 0.2 mm). 31. Last instar (scale line = 0.8 mm). Symbols: (L) labrum, (M) mandible. 1989 MOUTHPARTS IN LARVAL NOTODONTIDAE 467 Figs. 32, 33. 32. Mandibles of last instar Crinodes besckei in semiclosed position, viewed from the rear through head capsule. Scale line = 0.4 mm. Symbols: (P) pharynx, (AT) adductor tendon, (HC) head capsule. 33. "" Beborsteter Zapfen'" (bZ) on the maxilla of Talaeporia tubulosa (Retzius) (Psychidae), from Dampf (1910). The mandibular cutting edges of last instar H. obliqua and Crinodes besckei larvae slide past each other upon adduction. A similar mechanism was described for sa- tumiid and sphingid larvae by Bemays and Janzen (1988). Leaf cutting by satumiid mandibles is scissorlike (Makhotin and Davydova, 1961; Bemays and Janzen, 1 988). For C. besckei, the initial cut into the leaf blade appears to be facilitated by the serrate dorsal tooth. The dorsal comers of the mandibles are the first parts to meet during biting. The dorsal mandibular serrations illustrated by Bemays and Janzen (1988) may function in the same way. Leaf-edge clipping by the second to fifth instars of H. obliqua is theoretically enhanced by the increased distal, linear separation of the adductor apodeme from the rotational axis, and by the more medially-directed cutting edge on each mandible. In Crinodes besckei the plates and pockets proximad of the distal teeth appear to be modified for further mastication of food particles before they enter the gut. When the mandibles close, opposing sets of plates nearly inter-mesh, and opposing teeth insert into the pockets (Fig. 32). These actions parallel the process noted by Bemays and Janzen (1988) for sphingid mandibles. The oral plates may also help retain food particles in the oral cavity and thus assist the hypopharynx during ingestion. Our observations concerning developmental changes in feeding methods and man- dibular morphology of notodontid caterpillars fail to explain why later instars of other noctuoids, most of which clip through the leaf blade while feeding, retain the toothed mandible. Role of the stipital lobe. Grimes and Neunzig (1986) suggested that the stipital lobes aid the mandibles during mastication. However, because of their membranous nature, the lobes cannot be directly involved in chewing or crushing the leaf. The stipital lobes seem to form seals between the mandibles and maxillae, thereby helping hold leaf tissue within the oral cavity while the mandibles are acting. The large stipital lobes of Heterocampa obliqua (Fig. 6) and Nystalea nyseus (Fig. 1 7) may function not only as seals but, in conjunction with the deeply cleft labrum of these species 468 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) (Figs. 1 8, 25), as a channel for the leafs edge as it nears the mandibles. In hrst instar Heterocampa obliqua, which are leaf skeletonizers, the labrum is barely emarginate (Fig. 24) and the stipital lobe is absent. Phylogenetic implications. Although the Notodontidae is generally assumed to be a monophyletic group, few synapomorphies are known (Holloway et al., 1987). Per- haps the most frequently cited diagnostic character is the presence of two rather than one middorsal proprioceptor (MD) setae on abdominal segment 1 (Al) of the larva (Hinton, 1946; Common, 1979). However, dioptine larvae exhibit the plesiomorphic condition, a single MD seta on Al (J. Miller, unpubl. data; Weller, 1989). The only known adult synapomorphy is the presence of a ventrally-directed tympanal mem- brane (Common, 1979). The stipital lobe is a uniquely-derived character within the Noctuoidea. Dampf (1910) illustrated a similar structure, which he termed the "^beborsteter Zapfen'"^ (Fig. 33), on the larval maxilla of Talaeporia tubulosa (Retzius), a primitive psychid (Hinton, 1955), but Dampf s structure is almost certainly not homologous with the stipital lobe. Within the Notodontidae, a stipital lobe occurs in all but 9 of the 90 genera studied (Table 1). The Thaumetopoeinae, or Thaumetopoeidae of some authors (e.g., Kiriakoff, 1970), contains approximately 100 species, all restricted to the Old World. It is generally regarded as the most primitive notodontid group (Sick, 1940). Based on our exam- ination of six thaumetopoeine species in four genera, the stipital lobe is absent in that group (see Results and Table 1). These hndings suggest that the stipital lobe is a derived character supporting the monophyly of a clade which includes all noto- dontids exclusive of the Thaumetopoeinae. This further supports the hypothesis that thaumetopoeines represent the most basal notodontid lineage. Relationships among the remaining five genera that lack a stipital lobe, Danima, Epidonta, Lobeza, Psorocampa, and Tecmessa, are unknown. The first two are Old World taxa (Australia and Africa respectively), while the latter three are Neotropical (Gaede, 1 934). Our observations based on general adult appearance suggest that these five genera do not constitute a monophyletic group. However, a test of this hypothesis must await an improved classification for the Notodontidae. The presence of a smooth mandibular margin in final instar larvae is also taxo- nomically widespread among the Notodontidae (Table 1). A toothed mandible occurs only in some species of Dioptini, and in four additional genera, including Crinodes besckei, as well as two North American species of uncertain placement, Cargida pyrrha (Druce) (see also Godfrey, 1984) and Theroa zethus (Druce). Further research is required to determine whether these taxa form a monophyletic group. The thau- metopoeine mandible is problematical; it appears to be neither toothed nor smooth, but instead has three or four shallow dentations along its margin. Perhaps this rep- resents a transition state between the toothed mandible of noctuids, arctiids and lymantriids, and the smooth mandible found in higher notodontids. If true, this would imply that the mandibular serrations in species such as Crinodes are not homologous with those of other noctuoids. We hope that by describing the stipital lobe and notodontid mandible, we have helped provide two valuable diagnostic features for larvae of the Notodontidae. Furthermore, we hope our research will stimulate future study on the structure and function of noctuoid larval mouthparts. 1989 MOUTHPARTS IN LARVAL NOTODONTIDAE 469 ACKNOWLEDGMENTS Financial support for this study came from the Illinois Natural History Survey, the Kalbfleisch Fund (American Museum of Natural History), the Herbert Holdsworth Ross Memorial Fund (University of Illinois/Illinois Natural History Survey), the University of Illinois Research Board, Illinois Agricultural Experiment Station Project 12-361 (Biosystematics of Insects), and Insect and Disease Control (CFAA), Forest Insect and Disease Management Protection and Protection of Wood in Use (USDA Forest Service) (J. E. Appleby, principal investigator). The permission granted to G.L.G. by Servicio de Parques Nacionales de Costa Rica to work at Santa Rosa National Park is greatly appreciated, as is D. H. Janzen’s assistance with the work at Santa Rosa in 1986. Gratitude is owed to J. G. Franclemont for making his collection available, and to John Rawlins (Carnegie Museum), Alma Solis (USNM) and Susan Weller (University of Texas, Austin) for the loan of specimens. SEM’s were taken with the help of Andrew Simon (AMNH). Finally we thank J. E. Appleby, A. S. Hodgins, D. Janzen, S. Passoa, G. Robinson, W. G. Ruesnik, and M. J. Scoble for reviewing earlier drafts of this paper, and W. H. Allen, D. E. Dockter, D. H. Offner, L. M. Page, and K. R. Robinson for their comments and other forms of assistance. LITERATURE CITED Beck, H. 1960. Die Larvalsystematik der Eulen (Noctuidae). Abhandl. Larvalsyst. Ins. (Aka- demie-Verlag, Berlin) 4:1^06. Bemays, E. A. and D. H. Janzen. 1988. Satumiid and sphingid caterpillars: two ways to eat leaves. Ecology 69:1 153-1 160. Chu, H. F. C., C. L. Fang and L. Y. Wang. 1963. Fauna of Chinese Economic Insects. Vol. 7. Noctuidae (Immature Stages). Science Press, Peking, 120 pp. [In Chinese] Common, I. F. B. 1979. Lepidoptera. 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Expt. Sta. 274, 262 pp. Gaede, M. 1934. Notodontidae. In: E. Strand (ed.), Lepidopterorum Catalogus (59). Junk, Berlin, 351 pp. Gardner, J. C. M. 1943. Immature stages of Indian Lepidoptera (5). Indian J. Entomol. 5: 89-102. Gardner, J. C. M. 1946. Immature stages of Indian Lepidoptera (7). Indian J. Entomol. 7: 139-144. Godfrey, G. L. 1972. A review and reclassification of larvae of the subfamily Hadeninae (Lepidoptera, Noctuidae) of America north of Mexico. U.S. Dept. Agric. Tech. Bull. 1450, 265 pp. Godfrey, G. L. 1984. Notes on the larva of Cargida pyrrha (Notodontidae). J. Lepidop. Soc. 38:88-91. 470 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Godfrey, G. L. and J. E. Appleby. 1987. Notodontidae (Noctuoidea): The Notodontids and Prominents. Pages 524-533 in: F. W. Stehr (ed.), Immature Insects. Kendall/Hunt Publ. Co., Dubuque, Iowa, xiv + 754 pp. Grimes, L. R. and H. H. Neunzig. 1986. Morphological survey of the maxillae in last stage larvae of the suborder Ditrysia (Lepidoptera): mesal lobes (laciniogaleae). Ann. Entomol. Soc. Am. 79:510-526. Hinton, H. E. 1946. On the homology and nomenclature of the setae of lepidopterous larvae, with some notes on the phylogeny of the Lepidoptera. Trans. R. Entomol. Soc. London 97:1-37. Hinton, H. E. 1955. On the taxonomic position of the Acrolophinae, with a description of the larva of Acrolophus rupestris Walsingham (Lepidoptera: Tineidae). Trans. R. Ento- mol. Soc. London 107:227-231. Holloway, J. D., J. D. Bradley and D. J. Carter. 1987. C.I.E. Guides to Insects of Importance to Man, 1. Lepidoptera. C.A.B. International, Wallingford, U.K. Kiriakoff, S. G. 1970. Familia Thaumetopoeidae. In: P. Wytsman (ed.). Genera Insectorum, Lepidoptera. 219:54 pp. Makhotin, A. A. and E. D. Davydova. 1961. Morphology and functional importance of the elements of mouth apparatus in the caterpillars of some moths. Zool. Zhum. (Moscow) 40:1842-1857. McCabe, T. L. 1 988. Larval hosts of Anaplectoides and Aplectoides with notes on their biology (Lepidoptera: Noctuidae). J. N.Y. Entomol. Soc. 96:1-6. Merzheevskaya, O. I. 1988. Larvae of Owlet Moths (Noctuidae)— Biology, Morphology, and Classification (G. L. Godfrey, Scientific ed.). Translated from Russian by P. M. Rao. Smithsonian Institution Libraries and National Science Foundation, Washington, D.C. Minet, J. 1983. Elements sur la systematique des Notodontidae et nouvelles donnees con- cemant leru etude faunistique a Madagascar (Lepidoptera: Noctuoidea). Bull. Soc. Ento- mol. Fr. 87:354-370. Neil, K. A. 1988. Larvae of North American Leuconycta (Noctuidae). J. Lepidop. Soc. 42: 285-290. Peterson, A. 1962. Larvae of Insects. Part I, Lepidoptera and Plant Infesting Hymenoptera. Columbus, Ohio, 3 1 5 pp. Riotte, J. C. E. 1969. Rearing and descriptions of the early stages of the Nearctic species of Peridea, with special reference to P. basitriens (Lepidoptera: Notodontidae). Michigan Entomol. 1:351-356. Ripley, L. B. 1923. The external morphology and postembryology of noctuid larvae. Illinois Biol. Mon. 8:243-344. Sick, H. 1 940. Beitrag zur Kenntnis der Dioptidae, Notodontidae und Thaumetopoeidae und deren Verwandtschaftsbeziehungen zueinander. Zool. Jahrb. (Anat.) 66:263-290. Weller, S. J. 1987. Litodonta hydromeli Harvey (Notodontidae): description of life stages. J. Lepidop. Soc. 41:187-194. Weller, S. J. 1 989. The phylogeny of the Nystaleini (Lepidoptera: Noctuoidea: Notodontidae). Ph.D. dissertation. University of Texas, Austin, 396 pp. Received April 5, 1989; accepted September 13, 1989. J. New YorkEntomol. Soc. 97(4):47 1-474, 1989 ON VENEZUELAN LEPROLOCHUS (ARANEAE, ZODARIIDAE) Rudy Jocque* and Norman I. Platnick^ ^Musee Royal de I’Afrique Central, B-1980 Tervuren, Belgium, and ^Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024 Abstract. — A new species of the Neotropical spider genus Leprolochus, L. stratus, is described from inland Venezuela; records and illustrations are provided for related species. Members of the spider genus Leprolochus Simon are easily recognized by the transverse row of spines occupying the anterior margin of the carapace. Similar spines occur in the genus Cyrioctea Simon, but there they are situated between the two eye rows, rather than in front of the anterior row. Despite their similar appearance, the two genera do not appear to be sister groups (Jocque, 1987; Platnick, 1986; Platnick and Griffin, 1988). Cyrioctea is an austral genus, known from Queensland, Chile, Argentina, and Namibia; Leprolochus is Neotropical, extending from Panama and Trinidad to central Argentina (Jocque, 1988). In a recent revision, only three species of Leprolochus were reported (Jocque, 1988). Considering how little spider collecting has been done in much of South America, it is not surprising that a recent collection of material from inland areas of Venezuela included two species of Leprolochus, one of which is newly described below. All measurements are in mm; abbreviations used for eye and spination patterns are standard for the Araneae. We are grateful to Drs. J. and S. Peck for providing these, and many other fascinating, specimens to the American Museum of Natural History (AMNH); to Dr. A. Timotheo da Costa of the Museu Nacional, Rio de Janeiro, for loaning types; and to Dr. M. U. Shadab (AMNH) for supplying illustra- tions. Leprolochus stratus, new species Figs. 1-4 Types. Female holotype taken in a flight intercept trap in an evergreen forest 20 km N of Upata, Bolivar, Venezuela (21 June-12 July 1987; S. and J. Peck), and two male paratypes taken in a flight intercept trap in a forested woodland ravine at Puente Cocuizas, 70 km W of Ciudad Bolivar, Bolivar, Venezuela (19 June-August 1987; S. and J. Peck), deposited in AMNH. Etymology. The specific name is from the Latin stratus, meaning “with a saddle” and referring to the saddle-like depression at the base of the lateral apophysis of the male palpal tibia. Diagnosis. This species seems closely related to L. spinifrons Simon. Males can be distinguished by the long lateral tibial apophysis bearing a saddle-like dorsal depres- sion near its base (Fig. 2), females by the transversely oriented, corkscrew- shaped copulatory ducts (Fig. 4). Females of the other known species have longitudinally 472 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Figs. 1-6. 1^. Lepwlochus stratus, n. sp. 5, 6. L. parahybae Mello-Leitao. 1. Left male palp, ventral view. 2. Left male palp, retrolateral view. 3, 5. Epigynum, ventral view. 4, 6. Epigynum, dorsal view. oriented, less highly coiled copulatory ducts (for L. spinifrons and L. birabeni Mello- Leitao, see Jocque, 1988, figs. 8, 18; for L. parahybae Mello-Leitao, see Figs. 5, 6). FEMALE: Total length 5.42. Carapace 2.46 long, 1.83 wide, relatively broad in front, where narrowed to only 0.7 times maximum width, bearing row of 12 spines in front of eyes. Carapace and chelicerae medium brown; carapace darker in thoracic area, with pale brown median triangle in front of posterior margin. Endites and labium pale brown; sternum pale brown with darkened anterior margin. Femora medium brown suffused with dark brown; remainder of legs pale brown, tibiae with dark pro- and retrolateral blotches on both extremities of segment. Abdomen with pale dorsal pattern on dark sepia background, similar to that of L. spinifrons (Jocque, 1988, fig. 4); sides dark sepia with four broad, oblique, pale streaks on posterior half; venter pale, with row of three small dark spots between epigynum and tracheal spiracle, rims of which are slightly sclerotized and pale brown; spinnerets pale brown. Eye sizes and interdistances: AME 0. 1 2, ALE 0. 1 0, PME 0. 1 0, PLE 0. 1 2; AME-AME 0.07, AME-ALE 0.23, PME-PME 0.19, PME-PLE 0.32, ALE-PLE 0.05; MOQ an- terior width 0.76 times posterior width, 0.65 times length. Leg spination: femora: I, III dl-1-1-2, pi 1; II dl-1-1-2; IV dl-1-1-2, rl 1; patellae: I, II, IV dl-1, pi 1-1, rl 1; III dl-1, pi 1-1, rl 1-1; tibiae: I dl-1, pi 1-1-1, rl2; II dl-1-1, pi 1-1-1, v2, rl 1-1; III 1989 y^l^EZVELANLEPROLOCHUS 473 dl-1, pll-1-1, v2, rll-1; IV dl- 1-1, pll-1-1. vl-1, rll- 1; metatarsi: II dl; III dl. vl; IV 7 distal p. Measurements: I II III IV Palp Femur 1.42 1.46 1.54 1.67 0.66 Patella 0.58 0.62 0.63 0.67 0.42 Tibia 1.00 1.08 1.17 1.54 0.33 Metatarsus 1.50 1.50 1.96 2.42 — Tarsus 1.26 1.12 1.08 1.54 0.58 Total 5.76 5.78 6.38 7.84 1.99 Epigynum (Figs. 3, 4) ventrally with wide, featureless transverse slit near posterior margin; intromittent ducts coiled, with six transversely oriented loops, distal extrem- ities of ducts facing each other; spermathecae probably hidden within coils. MALE (values for second paratype in parentheses): Total length 2.50 (2.71). Car- apace 1.16 (1.25) long, 0.91 (0.87) wide, narrowed in front to 0.65 times maximum width, with row of nine spines in front of eyes. Coloration as in female except abdomen with pale brown dorsal scutum obscuring pattern. Leg spination: femora: Idl-2, pll;IIdl-l -2; IIIdl-1-2, plCIVdl- 1-1-2; patellae: I, III, IV dl-1, pi 1-1, rll; II dl-1-1, pll- .l, rll; tibiae: I dl-1, pll -1-1; II dl -1-11, pll-1 , v2; III dl-1. pll-1-1, rll;IVdl -1-1, pll-1-1 , vl, rl 1; metatarsi: II, IV 4 distal p, , III 5 distal p. Measurements: I II III IV Palp Femur 0.79 0.81 0.83 0.87 0.40 Patella 0.34 0.34 0.34 0.36 0.20 Tibia 0.63 0.59 0.61 0.79 0.16 Metatasus 0.71 0.77 0.87 1.21 — Tarsus 0.61 0.61 0.65 0.85 0.50 Total 3.08 3.12 3.30 4.08 1.26 Palpal tibia with four apophyses: one long terminal apophysis tapering to sharp point, reaching distal extremity of cymbium, excavated along its inferior margin, provided with saddle-like depression at dorsal base; two dorsal apophyses, mesal and dorsal, at base of terminal apophysis; fourth apophysis short, mesodorsal, ending in sinuous tip pointing backward (Fig. 2). Embolus originating on mesal side of bulb, long, whip- like; intertegular apophysis simple, straight, curved inward at tip; tegular apophysis membranous, poorly delimited (Fig. 1). Other material examined. None. Distribution. Known only from the type locality in inland Venezuela. Leprolochus spinifrons Simon New records: VENEZUELA: Bolivar: Rio Sipao, 110 km E Caicara, 17 June-4 Aug. 1987, flight intercept trap, gallery forest (S. and J. Peck, AMNH), 85, 22. Sucre: Carupano, 23 July 1987, elev. 80 m, thom-scrub litter (S. and J. Peck, AMNH), 12; 37 km W Carupano, 31 July 1987, elev. 50 m, thorn vine forest litter, humid ravine (S. and J. Peck, AMNH), 32. 474 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) LITERATURE CITED Jocque, R. 1987. Descriptions of new genera and species of African Zodariinae with a revision of the genus Hemdida (Araneae, Zodariidae). Rev. Zool. Afr. 101:143-163. Jocque, R. 1988. An updating of the genus Leprolochus (Araneae: Zodariidae). Stud. Neotrop. Fauna Environm. 23:77-87. Platnick, N. I. 1986. A review of the spider genus Cyrioctea (Araneae, Zodariidae). Amer. Mus. Novit. 2858:1-9. Platnick, N. I. and E. Griffin. 1988. On the first African and Australian spiders of the genus Cyrioctea (Araneae: Zodariidae). J. New York Entomol. Soc. 96:359-362. Received May 4, 1989; accepted June 30, 1989. J. New YorkEntomol. Soc. 97(4):475-478, 1989 KARYOTYPES OF THREE SPIDER SPECIES (ARANEAE: PHOLCIDAE: PHYSOCYCLUS) James C. Cokendolpher 2007 29th Street, Lubbock, Texas 7941 1 Abstract.— Three species of the spider genus Physocyclus {P. californicus, P. enaulus, P. sp.) were karyotyped using an air-drying, Giemsa staining method. All chromosomes were meta- centric, with males being 2n = 15 (N = 7 + XO) and females being 2n = 16. The karyotyping method resulted in slide preparations which were countable five years post-fixing. Members of the spider family Pholcidae are known cytologically by only four species. In each case, only males have been examined, and XO and XiX20 sex- determination mechanisms have been illustrated or reported (Sharma et al., 1959; Cokendolpher and Brown, 1985). Of these four species, reliable chromosome counts are available for only two species. Previous researchers have noted the difficulty in properly preparing pholcid chromosomes for accurate counting (Painter, 1914; Su- zuki, 1954). It is the purpose of this publication to record the karyotypes of two additional species including the first female karyotypes of two species. Additionally, data is provided on the stability of chromosome preparations made by an air-drying method. MATERIALS AND METHODS An adult male of Physocyclus californicus Chamberlin and Gertsch (9.5 km N Santa Isabella, San Diego Co., CA) and adult males and females of P. enaulus Crosby (Brackettville, Kinney Co., TX) and P. sp. (Lubbock, Lubbock Co., TX) were collected and returned to the laboratory. All specimens were dipped in a 0.005% colchicine/ Ringer’s solution and held for 24 hours to accumulate meiotic and mitotic meta- phases. Two males of P. sp. were karyotyped without colchicine treatment to ascertain effects of the drug on chromosomes (see Smith, 1965; Sharma and Sharma, 1972). Testis and ovary preparations were air-dried and stained with Giemsa following the procedure outlined by Cokendolpher and Brown (1 985). Preparations were not cover- slipped and were maintained at room temperatures. RESULTS AND DISCUSSION Chromosome preparations of a male P. californicus, two male P. enaulus, and four male P. sp. reveal 2n = 15, with n = 7 + XO (Figs. 1-4). These counts are based on 15, 14, and 31 nuclei, respectively, for the three species. Many meiotic nuclei were observed with 7:8 segregation with the X chromosome showing positive het- eropycnosis (Fig. 1). During diakinesis the X’s are isopycnotic (Fig. 2). Chromosome preparations of two P. enaulus (nine nuclei) and three female P. sp. (15 nuclei) reveal 2n = 16. The sex chromosomes are indistinguishable from auto- 476 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Figs. 1-3. Chromosomes of Physocyclus spp. males. 1. P. enaulus anaphase I. 2. P. sp. diakinesis. 3. P. sp. spermatogonial (?) metaphase. somes during mitotic metaphase, except for the size of the X’s. The three species of Physocyclus examined have only metacentric chromosomes. Bole-Gowda (1958) reported the haploid karyotype of Crossopriza lyoni (Blackwall) consisted of 1 3 metacentric autosomes and one metacentric X. In contrast, Sharma et al. (1959) recorded for the same species 11 metacentric autosomes and two ac- rocentric X chromosomes in the haploid set. Confirmation of this species’ haploid elements is needed. While Painter (1914) attempted to study Pholcus phalangioides (Fuessin) and Sper- mophora meridionalis Hentz, his admittedly unsatisfactory preparations revealed only the presence of two X chromosomes in the former species. The haploid set of Pholcus crypticolens Bdsenberg and Strand was reported by Suzuki (1954) to consist of 1 1 metacentric autosomes and two acrocentric X chro- mosomes. Cokendolpher and Brown (1985:figs. 1-3) illustrated the chromosomes of male P. sp. but did not comment on females or the sex-determining mechanism. The chro- mosomes illustrated in that publication were from the same population as samples reported upon herein. Male XO sex-determination mechanisms are uncommon in spiders (10% of 300 species) and are known only from ecribellate spiders: two species each from the Dysderidae and Segestriidae (Diaz and Saez, 1966; Benavente and Wettstein, 1980; Suzuki, 1954), two species of Lycosidae (Postiglioni and Brum-Zorrilla, 1981), eight species of Oxyopidae (Mittal, 1970), ten species of Thomisidae, one species each of Heteropodidae and Philodromidae (Bole-Gowda, 1952; Sokolow, 1962; Mittal, 1966), four species of Salticidae (Matsumoto, 1977), and possibly one Pholcidae (Sharma et al., 1959). Where the position of the centromere has been determined on spider species with XO males, all are acrocentric except for one Heteropodidae (Bole- Gowda, 1952; White, 1973) and possibly a Pholcidae (Sharma et al., 1959). In the case of Heteropoda sexpunctata (Simon) the male karyotype is 2n = 2 1 ( 1 9 metacentric and two acrocentric, with the X being metacentric). This rare condition is apparently 1989 KARYOTYPES IN PHOLCIDAE 477 =(( I) ('» II H i{ II ct \ • XX // f<« Figs. 4-6. Chromosomes of Physocyclus spp. 4. P. californicus male (2n = 1 5). 5. P. enaulus female (2n = 16). 6. P. sp. female (2n = 16). normal for the genus Physocyclus since all three species examined showed a male XO sex-determination system. While metacentric autosomes are rare in spiders (Sharma et al., 1959), the presence of metacentrics in both the autosomes and sex chromosomes has been recorded only in Heteropoda sexpunctata by Bole-Gowda (1952) and the questionable record by Bole-Gowda (1958) in Crossopriza lyoni. The air-drying, Giemsa staining method proved to be an effective method for preparing countable spreads. Previous researchers have noted difficulty in fixing and counting pholcid chromosomes. These problems are apparently overcome with the present method. Furthermore, preparations maintained for five years at room tem- peratures in dark slide boxes remained suitable for analysis. The chromosomes il- lustrated in Figure 4 were photographed three years post-fixing. The stability of the preparations may have been aided by the relatively arid environment of Lubbock. After examination the slides were tilted, allowing any excess immersion oil to run off the slides, but were otherwise not cleaned. Colchicine treated cells produced many more metaphase nuclei than untreated cells. Polyploids with countable chromosome numbers were present in low frequency (less than 15% of male nuclei counted). No other differences were noted between treated and non-treated chromosomes. ACKNOWLEDGMENTS I thank Nobuo Tsurusaki for his discussions on chromosomes, assistance with literature, and many helpful suggestions on the manuscript. Scott Stockwell kindly provided the living specimens of P. enaulus, and M. Kent Rylander (Texas Tech University) allowed the use of laboratory equipment and space in his care. Willis J. 478 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Gertsch is thanked for verifying the spider identifications. This study was supported in part by the Departments of Biological Sciences and Entomology, Texas Tech University. LITERATURE CITED Benavente, R. and R. Wettstein. 1980. Ultrastructural characterization of the sex chromo- somes during spermatogenesis of spiders having holocentric chromosomes and a long diffuse stage. Chromosoma 77:69-82. Bole-Gowda, B. N. 1952. Studies on the chromosomes and the sex-determining mechanism in four hunting spiders (Sparassidae). Proc. Zool. Soc. Bengal 5:51-70. Bole-Gowda, B. N. 1958. A study of the chromosomes during meiosis in twenty-two species of Indian spiders. Proc. Zool. Soc. Bengal 1 1:69-108. Cokendolpher, J. C. and J. D. Brown. 1985. Air-dry method for studying chromosomes of insects and arachnids. Entomol. News 96(3): 1 14-1 18. Diaz, M. O. and F. A. Saez. 1966. Karyotypes of South-American Araneida. Mem. Inst. Butantan Simp. Intemac. 33:153-154. Matsumoto, S. 1977. An observation of somatic chromosomes from spider embryo-cells. Acta Arachnol. 27:167-172. Mittal, O. P. 1966. Karyological studies on Indian spiders. IV. Chromosomes in relation to taxonomy in Eusparassidae, Selenopidae and Thomisidae. Genetica 37:205-234. Mittal, O. P. 1970. Karyological studies on the Indian spiders. VIII. Chromosomes in male germ cells of three species of the genus Oxyopes (Family Oxyopidae). Microscope 1970: 313-318. Painter, T. S. 1914. Spermatogenesis in spiders. J. Zool. Jahrb. Jena, Abt. Anat. 38:509-576. Postiglioni, A. and N. Brum-Zorrilla. 1981. Karyological studies on Uruguayan spiders. II. Sex chromosomes in spiders of the genus Lycosa (Araneae-Lycosidae). Genetica 56: 47-53. Sharma, G. P., B. L. Gupta and R. Parshad. 1959. Cytological studies on the Indian spiders. III. An analysis of the chromosomes in the male germ cells of the spider, Crossopriza lyoni (Blackwall), fam. Pholcidae. Res. Bull. Panjab Univ., N.S. Sci. 10:49-53, figs. 1-16. Sharma, A. K. and A. Sharma. 1972. Chromosome Techniques Theory and Practice. But- terworth and Co., London, 2nd ed., 575 pp. Smith, S. G. 1965. Heterochromatin, colchicine, and karyotype. Chromosoma 16:162-165. [Sokolow, I. L] 1962. [Studies on nuclear structures in Araneina. II. The sex chromosomes.] Tsitologia 4:617-625 (in Russian). Suzuki, S. 1954. Cytological studies in spiders. III. Studies on the chromosomes of fifty-seven species of spiders belonging to seventeen families, with general considerations on chro- mosomal evolution. J. Sci. Hiroshima Univ., ser. B, div. 1, 15:23-136, pis. 1-15. White, M. J. D. 1973. Animal Cytology and Evolution. University Press, Cambridge, 3rd ed., 923 pp. Received May 15, 1989; accepted June 6, 1989. NOTES AND COMMENTS J. New YorkEntomol. Soc. 97(4):479-482, 1989 A NEW STRUCTURE ON THE HIND LEGS OF MALE MONALOCORIS CARIOCA CARVALHO AND GOMES (HETEROPTERA: MIRIDAE) Monalocoris (Byrocorinae: Bryocorini) consists of eleven tropical and temperate species (Carvalho, 1957, 1981; Carvalho and Gomes, 1969). Only two {M. ameri- canus Wagner and Slater, 1952 and M. eminulus Distant, 1893) are known for North America; three {M. carioca Carvalho and Da Penha Gomes, 1971, A/, eminulus and M. pallidiceps Reuter, 1907) are neotropical. The Bryocorini are considered to be one of the most derivative tribes of the Miridae (Schuh, 1976). While examining samples of the South American species M. carioca (Fig. 1), author JMC noticed that adult males possess a depression on the prolateral surface of the slightly swollen hind femora. We decided to compare the femora of males and females of this species and of M. americanus, examine the fine details of the depression and suggest possible functions for it. Two adult pairs of both M. carioca and of M. americanus were studied with an IDS-DS 130 scanning electron microscope at the Electron Microscopy Laboratory, University of California (Berkeley). Since M. carioca is rare in collections, additional material of this species was examined only with light microscopy to prevent damage to the specimens. Confocal scanning microscopy and high voltage electron micros- copy were also employed to obtain correlative microscopical data for the innermost portions of the hairy depression in two hind femora of M. carioca. Confocal scanning microscopy, which allows optical sections of objects, is particularly useful when the structures of interest are relatively hidden (McCarthy and Walker, 1 988). High voltage electron microscopy, an instrument that can penetrate thick samples (Bastacky, 1986) and allows magnifications to about 1 0® x , was mainly employed to ascertain whether there were orifices on the leg depression. Collection data for the specimens examined were as follows: M. carioca, Colombia, Anolaima Cund., 10 Sept. 1965, J. A. Ramos (1<3, 39$) [JMC (=Jenaro Maldonado Capriles) collection]; Alban. Cund. {2$$) (JMC collection). M. americanus, USA, Pennsylvania: Centre Co., 2 mi N State College, ex Dryopteris sp., 10 June 1977 (Schuh, Henry, Wheeler) {16, 1$) (American Museum of Natural History = AMNH); New York: Albany Co., Rensselaerville, Huyck Preserve, ex Dryopteris sp., 29 June- 2 July 1977 (R. T. Schuh) (15, 19) (AMNH); Tennessee, Gatlinburg, Beach Gap, GSMNP, 5,500', S sweeps, 2 July 1947 (R. H. Whittacker) 2 1 h 5 53, 19 PARATYPE (AMNH, Donation from J. A. Slater Collection). These specimens are noted with a yellow label that reads VOUCHER SPECIMENS SEM studies Monalocoris (He- miptera: Miridae) hind femora. On male M. carioca, the hairy, oval, submedial depression is located on the pro- 480 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Figs. 1-6. 1-2. Monalocoris carioca. 1. Adult male (dorsal). 2. Hind femur of male (pro- lateral). 3-4. Depression on hind femur of male M. carioca. 3. Detail. 4. Setae on depression (note tubercles, seemingly flexible insertions and an orifice on one seta). 5. Hind femur of female M. carioca. 6. Slightly swollen hind femur of male M. americanus (prolateral, note absence of hairy depression). lateral face of the slightly swollen hind femora (Fig. 2). The depression measures about 280 x 80 x 50 ii. Confocal scanning microscopy and scanning electron mi- croscopy showed that the setae cover all of the depression but are most abundant on the lower two-thirds (Fig. 3). The setae are located on tubercles, are hollow and 1989 NOTES AND COMMENTS 481 their insertions seem flexible (Fig. 4). No evidence of orifices was found in the depres- sion. The hind femora of females are neither swollen nor do they have this depression (Fig. 5). Male Monalocoris americanus have a slightly and uniformly swollen hind femora (Fig. 6), but lack a depression, while female femora are neither swollen nor depressed. Apparently, this structure is unknown in other congeners. DISCUSSION In our literature search we were unable to locate references for a similar structure on the hind femora in any heteropterous family. Either this depression has been overlooked or it possesses no known homologues amongst the Heteroptera. Possible hypotheses concerning the functional significance of the depression are: 1 . olfactory or mechanoreceptive and 2. secretory. The setal morphology and their seemingly flexible insertion suggest an olfactory or mechanoreceptive function. However, the closely appressed setae covering the depression suggest a liquid retention function from a putative secretory organ, but the absence of orifices on the depression, through which a secretion could be oozed, questions (but does not necessarily reject, Schneider, 1966) this hypothesis. Also, it is strange to have a secretory opening or organ on a \Qg.—Jorge A. Santiago-Blay, Department of Entomological Sciences, University of California, Berkeley, California 94720 U.S.A., and Jenaro Maldonado Capriles, Ur- banizacion Aponte 61-1, Cayey, Puerto Rico 00633. ACKNOWLEDGMENTS Dr. R. T. Schuh (AMNH) lent the M. americanus specimens and granted permission to examine them on SEM. Dr. N. Fowler (R. J. Lee Group, Inc., Berkeley, CA) kindly made available a confocal scanning light microscope and operated a computer image enhancer that allowed us to obtain the first views of the femoral depression. Ms. A. Reynolds (Univ. California, Berkeley) trained author JASB in the use of the confocal scanning laser microscope. Dr. J. Bastacky (Donner Laboratory, Lawrence Berkeley Laboratory, Univ. California, Berkeley) op- erated the High Voltage Electron Microscope at the National Center for Electron Microscopy (Lawrence Berkeley Laboratory). Drs. R. F. Chapman (Division of Neurobiology, Univ. Ari- zona, Tucson) and R. L. Pipa (Univ. California, Berkeley), as well as an anonymous reviewer, read the typescript and suggested changes to it. To all of them our thanks. LITERATURE CITED Bastacky, J. 1986. High-voltage microscopy of unsectioned human lung alveolar walls. Pages 224-225 in: G. W. Bailey (ed.). Proceedings of the 44th Annual Meeting of the Electron Microscopy Society of America. Carvalho, J. C. M. 1957. Catalogo dos mirideos do mundo. Parte I. Subfamilias Cylapinae, Deraeocorinae, Bryocorinae. Arq. Mus. Nac. (Rio de Janeiro, Brasil) 44:1-158. Carvalho, J. C. M. 1981. The Bryocorinae of Papua New Guinea (Hemiptera, Miridae). Arq. Mus. Nac. (Rio de Janeiro, Brasil) 56:35-89. Carvalho, J. C. M. and I. da Penha Gomes. 1969. Mirideos neotropicais, CVII: descri^oes de cinco especies novas adicionais da Republica do Equador (Hemiptera). Rev. Brasil. Biol. 29:225-230. Carvalho, J. C. M. and I. da Penha Gomes. 1971. Mirideos neotropicais, CXXVIII: novo genero e novas especies de Bryocorini do Brasil (Hemiptera). Rev. Bras. Biol. 31:99- 102. 482 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Distant, W. L. 1880-1893. Insecta. Rhyncota. Hemiptera-Neuroptera. Biologia Centrali- Americana 1:1^62. McCarthy, J. J. and J. S. Walker. 1988. Scanning confocal optical microscopy. EMSA Bull. 18:75-79. Reuter, O. M. 1907. Capsidae in Brasilia collectae in Museo J. R. Vindobonensi aservatae. Ann. k. k. naturh. Hofmus. (Wien) 22:33-80. Schneider, D. 1966. Chemical sense communication in insects. Pages 273-297 in: Nervous and Hormonal Mechanisms of Integration. Symposia of the Society for Experimental Biology. Number XX. Academic Press Inc. New York. Schuh, R. T. 1976. Pretarsal structure in the Miridae (Hemiptera) with a cladistic analysis of the relationships within the family. Amer. Mus. Novit. 2601:1-39. Wagner, E. and J. A. Slater. 1952. Concerning some Holartic Miridae. Proc. Entomol. Soc. Wash. 54:273-281. J. New York Entomol. Soc. 97(4):482-483, 1989 RECORDS OF CHIMARRA SOCIA (TRICHOPTERA: PHILOPOTAMIDAE) FROM INTERIOR HIGHLAND STREAMS IN ARKANSAS AND MISSOURI The known distribution of Chimarra soda Hagen (Trichoptera: Philopotamidae) in North America includes southeastern Canada and the northeastern United States north of a line extending from the northwestern comer of Minnesota to central Pennsylvania, and southward in the Appalachian Mountains at least as far as Ten- nessee (see fig. 14, Lago and Harris, 1987). However, recent field work (and the discovery of some previously collected specimens) has revealed the presence of pop- ulations of C. soda in several streams in the Interior Highlands (Ozark and Ouachita Mountains) of Arkansas and Missouri. Faunistic investigations of intervening areas (Illinois, Ross, 1944, 1948; Kentucky, Resh, 1975; Arkansas, Unzicker et al., 1970) have failed to produce specimens of true soda and current work in Missouri and Arkansas has not revealed populations outside these mountain streams. Apparently these populations are relicts of what was once a much more widely distributed species, rather than representing a simple range extension. Among the specimens examined during this study was a series of C. soda from the Albert Pike Recreation Area in Montgomery County, Arkansas. The occurrence of soda at this locality prompted the reexamination of the paratype of C. parasoda recorded from the same vicinity by Lago and Harris (1987). This specimen proved to be C. soda, and its inclusion in the type series of parasoda was based on mis- interpretation of data recorded during initial examination. The dorsal aedeagal rods are slightly rotated so that the apices are nearly parallel (a condition seen in parasoda), but other characters are typical of soda (not parasoda as stated by Lago and Harris, 1987). Specimens on which the following records are based are housed in the insect 1989 NOTES AND COMMENTS 483 collections at the University of Arkansas (UA), Clemson University (CU) and Purdue University (PU). We wish to thank Dr. John C. Morse (Clemson) and Dr. Arwin V. Provonsha (Purdue) for allowing us to examine specimens in their care. Records. ARKANSAS: Clark Co., 3 mi. NE Amity, Caddo River, 9 July 1978, 35 19; 16 Aug 1978 (UA). Montgomery Co., Little Missouri River at Albert Pike Rec. Area, 30 May 1974, 15 (PU); 28 July 1980, 25; 19 Sep 1980, 725; 20 Sep 1980, 95; (CU). Pike Co., Glenwood, Hwy 270 at Caddo River, 27 Jul 1978, 225, 49 (UA). Saline Co., N. Fork Saline River, 17 Aug 1985, 45, 49 (UA). MISSOURI: Pulaski Co., Gasconade River, (T36N-R12W-Sec 5), 30 May 1986, 165; 26 Jun 1987, 25; 28 July 1985, 65; 28 Jul 1987, 85; 25 Sep 1987, 55 {\JA).—Paul K. Lago, Department of Biology, the University of Mississippi, University, Mississippi 38677, and Michael L. Mathis, Department of Zoology, and David E. Bowles, Department of Entomology, University of Arkansas, Eayetteville, Arkansas 72701. LITERATURE CITED Lago, P. K. and S. C. Harris. 1987. The Chimarra (Trichoptera: Philopotamidae) of eastern North America with descriptions of three new species. J. New York Entomol. Soc. 95: 225-251. Resh, V. H. 1975. A distributional study of the caddisflies of Kentucky. Trans. Kentucky Acad. Sci. 36:6-16. Ross, H. H. 1944. The caddisflies, or Trichoptera, of Illinois. Bull. Illinois Natur. Hist. Sur. 23, 326 pp. Ross, H. H. 1948. New Nearctic Rhyacophilidae and Philopotamidae (Trichoptera). Ann. Entomol. Soc. Amer. 41:17-26. Unzicker, J. D., L. Aggus and L. O. Warren. 1970. A preliminary list of the Arkansas Tri- choptera. J. Georgia Entomol. Soc. 5:167-174. J. New York Entomol. Soc. 97(4):483-485, 1989 POECILOCHIRUS MONOSPINOSUS (ACARINA: MESOSTIGMATA: PARASITIDAE), A PREDATOR OF HOUSE FLY IMMATURES: NEW LOCALITY RECORDS Poecilochirus monospinosus Wise, Hennessey, and Axtell is a recently described species of mite in the family Parasitidae (Wise et al., 1988). The mites live in chicken manure where they prey on saprophytic nematodes, other mites, and immature dipterans (Fig. 1). Both deutonymphs (Fig. 1) and females feed readily on eggs and first instars of the house fly, Musca domestica L. (Geden et al.; 1988, Wise et al., 1 988). Because of this, and because the mites occasionally are very abundant in newly accumulating poultry manure, P. monospinosus may play a role in the regulation of populations of flies associated with poultry production. Wise et al. (1988) described the species from mites collected from poultry manure in North Carolina, and sug- 484 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) Figs. 1-4. 1. Live Poecilochirus monospinosus deutonymph feeding on house fly eggs. 2. Intercoxal shield of P. monospinosus deutonymph. Specimen cleared in lactophenol and mount- ed in Hoyer’s medium. 3. Comparison of alcohol-preserved Macrocheles muscaedomesticae and P. monospinosus. Clockwise from upper left: M. muscaedomesticae female (ventral view), P. monospinosus female (ventral), P. monospinosus deutonymph (ventral), P. monospinosus deutonymph (dorsal), P. monospinosus female (dorsal), M. muscaedomesticae female (dorsal). Note darkened area on anterior region of intercoxal shield {is) of P. monospinosus deutonymph. 4. P. monospinosus deutonymph, ventral view. Note membranous projection {mp) on fixed digit of chelicerae. Specimen cleared in lactophenol and mounted in Hoyer’s medium. gested that the apparently limited distribution of the mites was due to a lack of sampling from other locations. We report here the occurrence of P. monospinosus from chicken manure in Massachusetts and New York. Geden and Stoffolano (1987) reported the occurrence of small numbers of un- identified Poecilochirus sp. from chicken manure in wide-span caged layer houses in Massachusetts. We have cleared and slide-mounted 10 deutonymphs, 5 females and 2 males from these collections and confirmed that they are P. monospinosus. These 1989 NOTES AND COMMENTS 485 mites were collected sometime between May and June 1980, from the Hill farm in Hubbardston, Massachusetts; we cannot determine the precise date of collection or the age of the accumulated manure owing to the manner in which the alcohol col- lections were pooled several years after the survey was completed. More recently, we collected ca. 1,000 cm^ of manure from a high-rise caged layer house in Wolcott, New York (Wegman’s farm) on 5 Jan. 1989 that contained large numbers of parasitid deutonymphs, as well as substantial numbers of other fly pred- ators (the mite Macrocheles muscaedomesticae [Scopoli] and the histerid Carcinops pumilio [Erichson]). The manure in the house had accumulated for approximately 10 months. After extracting the sample through Tullgren funnels into 70% ethanol, we cleared and identified deutonymphs, females, and males of P. monospinosus. A total of approximately 150 deutonymphs and 30 females were recovered from the sample. No attempt was made to quantify males or immatures other than the deu- tonymphs because of the presence of at least one other species of Parasitidae {Parasitus sp.) in the sample. P. monospinosus may be widely distributed throughout northeastern North Amer- ica, and a clearer picture of the mite’s distribution will emerge as other investigators examine the acarine fauna associated with poultry manure. Spot characters that can aid in locating and identifying the mites are: 1) observation of extremely fast-moving mites over and just under the manure surface (deutonymphs); 2) examination of alcohol-preserved deutonymphs for a characteristically darkened band on the inter- coxal shield (Figs. 2, 3) (diagnostic for all but one member of the genus); and 3) deutonymphs and females that are somewhat similar in size to female Macrocheles muscaedomesticae (Fig. 3). Proper identification requires clearing and examination of slide-mounted material for the characters described by Wise et al. (1988), especially the presence of a single, entire, membranous process on the apex of the fixed digit of the chelicerae of the deutonymphs (Fig. A). — Christopher J. Geden, Donald C. Steinkraus, and Donald A. Rutz, Department of Entomology, Comstock Hall, Cornell University, Ithaca, New York 14853-0999. LITERATURE CITED Geden, C. J., R. E. Stinner and R. C. Axtell. 1988. Predation by predators of the house fly in poultry manure: effects of predator density, feeding history, interspecific interference, and field conditions. Environ. Entomol. 17:320-329. Geden, C. J. and J. G. Stoflblano, Jr. 1987. Succession of manure arthropods at a poultry farm in Massachusetts, with notes on Carcinops pumilio sex ratios, ovarian condition and body size. J. Med. Entomol. 24:214-222. Wise, G. U., M. K. Hennessey and R. C. Axtell. 1988. A new species of manure-inhabiting mite in the genus Poecilochirus (Acari: Mesostigmata: Parasitidae) predacious on house fly eggs and larvae. Ann. Entomol. Soc. Am. 81:209-224. BOOK REVIEWS ARTHROPOD OVERVIEWS J. New York Entomol Soc. 97(4):486-487, 1989 Spiders. Webs, Behavior, and Evolution. — William A. Shear (ed.). 1986. Stanford University Press, Stanford, California. 492 pp. 1986. Hardbound: $55. Recently, a number of exciting new publications on arachnids have been published. This long-awaited volume is a welcome addition. It is the proceedings of a meeting of the American Arachnological Society at the University of Tennessee in 1981; the contributors were asked to expand the papers. The book includes 1 2 chapters written by 1 5 authors plus a summary chapter and glossary written by the editor. The chapters are listed below. Web-site selection: are we asking the right questions? Anthony C. Janetos. Habitat choice and utilisation in web-building spiders. Susan E. Reichert and Rosemary G. Gillespie. Transmission of vibration in a spider’s web. W. Minch Masters, Hubert S. Markl, and Anne J. M. Moffat. Effects of orb-web geometry on prey interception and retention. William G. Eber- hard. Prey specialisation in the Araneidae. Mark S. Stowe. Web building and prey capture in the Uloboridae. Yael S. Lubin. Social spider webs, with special reference to the web of Mallos gregalis. William James Tietjen. Web building and prey capture in communal orb weavers. George W. Uetz. Web building versatility and the evolution of the Salticidae. Robert R. Jackson. The role of silk in prey capture by nonaraneomorph spiders. Frederick A. Coyle. Web removal patterns in orb-weaving spiders. James Edwin Carico. The monophyletic origin of the orb web. Jonathan Coddington. The evolution of web-building behavior in spiders: a third generation of hypotheses. William A. Shear. The wealth of information presented in this volume defies a brief summary. Despite my taxonomic bent I was thoroughly impressed and delighted with most chapters. The book provides a vital coverage of many topics about the construction of webs and unlike some previous treatments deals with many groups, not just the orb- weavers— there are after all a few other spider groups that build webs, orbs and otherwise, even though they are not so elaborate or spectacular. This book marks the birth of the use of behavior as a character in the path to phylogenetic reconstruc- tion. The realisation that behavior is also a character suite that can contribute to unravelling complex phylogenetic knots finally dawns. Although Jonathan Codding- ton’s chapter causes the major upheaval in our concept of an “orb web” (they include also the deinopids), other authors have directed some energy to the integration of taxonomy/phylogeny/evolution and their specialisation of behavior. Robert Jackson takes the cue from Portia fimbriata, a bizarre web-building, spider-hunting salticid, and seeks to elaborate a hypothesis for the evolution first of the Salticidae and then 1989 BOOK REVIEWS 487 the entire Araneomorphae— “from what tiny seeds the mighty acorn grows.” Never- theless, many avenues remain to be explored. In the words of the editor, “This is a book of questions.” Many remain unanswered. I have no doubt that it will stimulate many new and exciting hypotheses for testing. The highlights for me were Fred Coyle’s chapter summarising data on the Mygalo- morphae and Mesothelae (Liphistiiidae) and presenting his own observations along- side them. Coyle’s work, as ever detailed and thorough, is the only such compilation on the much neglected Mygalomorphae. Finally, we see excellent photographs of the diplurid webs that trap a fascinating variety of prey and remain difficult to adequately describe. Equally, Jonathan Coddington’s photographs of the diverse webs of the many orb-weaving spider genera provide ample support for his complex and hard argued hypotheses. Only one thing detracted from the book. The taxonomic glossary provides much appreciated respite from the barrage of names. However, there are numerous errors in it. The Anyphaenidae and Amaurobioidae are listed separately and not crossrefer- enced. Cethegus, an Australian diplurid, steals from the Panamanian Diplura the title of being the most aerial of web-building mygalomorphs. The Liphistiidae are deemed to be “not clearly related to the Mygalomorphae or Araneomorphae,” the only other spider groups. However, Platnick and Gertsch’s (1976) hypothesis about the groups’ relationships remains uncontested. I guess others are also present but do not signif- icantly detract from the notion of a glossary or its function. Overall, I was thoroughly delighted with “Spiders. Webs, Behavior, and Evolu- tion.” The style and content lend themselves to reading by all arachnophiles, not just the academics and other professionals. Generally, the editing is very good, the book is a credit to Shear. I unreservedly recommend the volume.— J. Raven, Queensland Museum, PO Box 300, South Brisbane, 4101, Q. Australia. LITERATURE CITED Platnick, N. I. and W. J. Gertsch. 1976. The suborders of spiders: a cladistic analysis (Arach- nida, Araneae). Amer. Mus. Novitates 2807:1-15. /. New YorkEntomol Soc. 97(4):487-489, 1989 Evolution and Adaptation of Terrestrial Arthropods.— John L. Cloudsley-Thompson. 1988. Springer- Verlag, Berlin, Heidelberg, New York, x + 141 pp. $33.00 paper. This slim volume is designed to present “a concise synthesis of certain basic information required for BSc (Hons) and MSc (Entomology) examinations” (author’s preface), with a functional emphasis. The nine chapters cover (1) paleontology and phylogeny, (2) implications of life on land, (3) conquest of land by Crustacea, (4) insect phylogeny and origin of flight, (5) evolutionary trends in reproduction, (6) adaptations to extreme environments, (7) dispersal and migration, (8) defensive mechanisms, and (9) success of terrestrial arthropods. These are indeed important areas of functional and evolutionary entomology, ones with recent exciting discoveries 488 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) and conceptual advances. A concise and up-to-date synthesis of these topics would be welcome. Unfortunately, this book does not provide that synthesis. This book will be unintelligible to those who lack a thorough background in formal entomology. Anatomical terms and arthropod names are used without definition, description, or illustration, and concepts lack proper introduction as to their signif- icance for understanding arthropod evolution. The book is poorly edited, with ty- pographical errors on nearly every page and numerous misstatements of fact. The chapters are not well integrated, and there is insufficient recognition of important research since 1980. Cloudsley-Thompson’s decision to present little physiology and no biochemistry weakens his functional approach. The 86 line drawings are mostly modifications of previously published illustrations, typically of whole arthropods, that visually enhance the text but do not help in its clarification. The slender bibli- ography for each chapter (median of 14 references, 3 since 1980) provides only a cursory introduction to the literature. All but three of the chapters (1,4, and 9) are the topics of earlier books or reviews by this prolific author, which may explain the peripatetic nature of themes grouped under “evolution and adaptation.” Cloudsley-Thompson devotes much attention to Sidnie Manton’s theories of the evolution of Arthropoda, including diphyletic origin of mandibles, multiple origins of arthropodization, and myriapod-hexapod relationships. In contrast to Manton, he concludes that Arthropoda and Onychophora are separate phyla, the former con- taining the subphyla Uniramia, Crustacea, and Chelicerata, largely relying on the evidence provided in Gupta (1979). He concludes that the paranotal lobe theory of the origin of wings is the most widely held today but that Kukalova-Peck’s (1978) theory of wing origin from pleural gill plates is equally plausible. At one point, he considers the Paleodictyoptera to be the oldest and most primitive order of insects, having “generalized biting jaws” and comprising “an assemblage of primitive types” which gave rise to the Paraplecoptera and from them the Embioptera and Isoptera. However, in the next paragraph the Paleodictyoptera are correctly said to “have highly modified piercing and sucking mouthparts” and “were not ancestral to any modem insect orders.” Similar self- contradictions and non-sequiturs occur frequently in this book. Wing venation is analyzed according to Lameere’s and Tillyard’s modifications of the Comstock-Needham system, without recognition of Wootton’s (1979) important analysis or discussion of Kukalova-Peck’s (1978, 1985) hypotheses based on paleo- zoic fossils. The “Panorpoid complex” is said to include all endopterygote orders except the Hymenoptera and Coleoptera, following Tillyard, instead of the more widely accepted concept of Hinton which also excludes the neuropteroid orders. Throughout Cloudsley-Thompson’s discussion of arthropod evolution, there is no explicit use of a phylogenetic (Hennigian) approach and no new hypotheses as to systematic groupings or evolutionary lineages. Discussion of adaptations, especially those concerning reproduction, largely ignores the current revolution in sexual selection theory, neglecting even such important concepts as male-male competition and female choice. The primary functions of courtship are said to be appeasement and synchrony. The function of male swarming is implied to be unclear, and perhaps representing “a habit which has persisted long after its original function has disappeared” (p. 58). Courtship feeding by Panorpa scorpionflies is said to divert the female, with no recognition of Randy Thornhill’s 1989 BOOK REVIEWS 489 important studies on female choice in Mecoptera. Mating in Calopteryx damselflies is described without mention of Jonathan Waage’s studies on removal of previously deposited sperm by the penis. While R. Thornhill and J. Alcock’s The Evolution of Insect Mating Systems (Harvard University Press, 1 983) is cited in the bibliography, it appears to have been ignored in the preparation of the chapter on reproduction. Migration and dispersal are better presented, with documentation of inconspicuous as well as conspicuous migrations and acceptance of the adaptive value of leaving adverse conditions to the migrants themselves. However, monarch butterflies do not have some members of their population overwintering near the Canadian border; the presentation of the seasonal cycle of aphids is garbled; and there is no mention of phases of migratory locusts and little discussion of alary polymorphism in general and the environmental and physiological factors that control it. The short concluding chapter on success of terrestrial arthropods stresses the sig- nificance of the chitinous exoskeleton, small size, short life cycle, and “genetic adapt- ability,” which have enabled colonization of every conceivable terrestrial habitat. Morphological adaptation is illustrated by the evolution of sucking mouthparts in insects and vertebrate ectoparasitism in ticks and insects. This chapter, regrettably, mirrors the entire book— a tantalizing peek at an important subject, not so much erroneous as incomplete and out-of-touch with modem evolutionary biology. — George C. Eickwort, Department of Entomology, Comstock Hall, Cornell University, Ithaca, New York 14853. LITERATURE CITED Gupta, A. P., ed. 1979. Arthopod Phylogeny. Van Nostrand Reinhold Co., New York, 762 p. Kukalova-Peek, J. 1978. Origin and evolution of insect wings and their relation to meta- morphosis, as documented by the fossil record. J. Morph. 156:53-126. Kukalova-Peck, J. 1985. Ephemeroid wing venation based upon new gigantic Carboniferous mayflies and basic morphology, phylogeny, and metamorphosis of pterygote insects (Insecta, Ephemerida). Can. J. Zool. 63:933-955. Wootton, R. J. 1979. Function, homology and terminology in insect wings. Syst. Entomol. 4:81-93. J. New York Entomol. Soc. 97(4):489^91, 1989 Evolution and Adaptation of Terrestrial Arthropods.— John L. Cloudsley-Thompson. 1988. Springer- Verlag, New York, New York, 141 pp. $33 (paper). The phylum Arthropoda comprises, by far, the largest group of organisms on Earth. From their first appearance in the early Cambrian Period (570-480 million years before present), arthropods have radiated to fill ecological niches in virtually every comer of the globe, the Cmstacea reigning supreme in many marine habitats, while insects dominate the land. It is on land that the importance of the group is manifest. In terms of species diversity and numbers of individuals, the arthropods (the vast majority of which are insects) control the nature of life on the land surface; they are 490 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) the essential middle links, the great energy transducers, in the food webs of terrestrial ecosystems. Had the group never evolved, the world would be an unimaginably different place, and this reviewer would not now be here to expound on the merits of the following book. John L. Cloudsley-Thompson has produced a general synopsis touching on the major features of the evolution and ecology of terrestrial arthropods. According to the preface, the author’s intent was not to write a comprehensive treatise, but to bring together and summarize information on the origination of the terrestrial groups and the adaptations that have led to their pre-eminent success. The intended audience is undergraduate and beginning graduate students. Given the potential scope of the subject, the range of topics covered is necessarily broad. All of the classes, including those containing at least semiterrestrial species, are presented: Crustacea, Onychophora, Chilopoda, Diplopoda, Symphyla, Paurop- oda, Arachnida, Insecta, and the non-insectan hexapods, i.e., Collembola, Diplura, and Protura. The first part of the book deals generally with the transition of the arthropods from a marine or estuarine to a terrestrial existence and the particular enabling mechanisms evolved to permit this. Chapter 1 is an overview of the phy- togeny and fossil record of the land-dwelling forms, with brief mention of their aquatic progenitors. The earliest, unambiguous terrestrial animal records date from the Si- lurian Period, coincident with the spread of the early vascular land plants. A diverse fauna existed by Devonian times, including mites, spiders, scorpions, pseudoscor- pions, centipedes, various other myriopods, and possibly insects. Insects are un- equivocally represented in Carboniferous deposits. The second chapter discusses the crucial adaptive role played by the arthropodan exoskeleton and its relationships to the size and physiology of terrestrial forms. Considerations of allometric growth (scaling), water relations, respiration, nutrition, and excretion, as they relate to the arthropodan body plan, are noted. With this background covered. Chapter 3 outlines the mechanisms involved in the transition to land of the few crustacean groups that successfully completed the trip. The chapter title, “The Conquest of the Land by Crustacea,” however, is ill chosen; in their need for ready access to moist conditions for purposes of water balance, respiration, and reproduction, terrestrial crustaceans are, at best, imperfectly adapted to dry land, and in no way masters of their environments. In the second and main portion of the book are covered the major evolutionary innovations that have allowed arthropods to become the dominant life forms on the land. Surely, the most significant of these is the evolution of flight in the insects (some 99% of species belong to the winged groups). Phytogeny of the insects and the origin of flight, including brief exposition of the various theories erected to explain it, are covered in Chapter 4. In succeeding chapters are covered the problems of reproduction, extreme environments, dispersal and migration, and defense against enemies, respectively. In a concluding chapter, the author speculates briefly on the principal adaptations responsible for the success of terrestrial arthropods, focusing specifically on two insect innovations, chosen to exemplify the adaptability of ter- restrial arthropods: the evolution of sucking mouthparts, which have permitted access to a broadened range of life styles and diets (e.g., parasitism, pollination; blood, plant sap, nectar); and the adaptations inherent in an ectoparasitic way of life, illustrated by lice, fleas, and the bed bug. 1989 BOOK REVIEWS 491 Cloudsley-Thompson has produced a brief, mostly superficial discussion of the main problems involved in terrestrial arthropod existence. As such, the work will largely fulfill his prefaced objective, and serve primarily as a serviceable introduction to arthropod evolutionary ecology for college-aged zoology students and, perhaps, as a refresher for those a bit more knowledgeable in the field. For others, particularly practicing researchers, the book is clearly inadequate as a reference work. A number of important recent papers, which should have been included in a work of this nature, are not cited. For example, no reference is made in Chapter 1 to Retallack and Feake’s discovery of evidence for a possible Ordovician age for the first terrestrial arthropods (Science 235:61-63, 1987). Surprisingly, Kingsolver and Koehl’s brilliant hypothesis for the origin of the insect wing (Evolution 39:488-504, 1985) is absent from the discussion of insect flight in Chapter 4. In the section on cave life (Chapter 6), no mention is made of F. G. Howarth’s major contributions to the study of cavemicolous arthropods (e.g.. Annual Review of Entomology 28:365-389, 1983). In the discussion of diapause in the same chapter, there is no citation to Tauber, Tauber, and Masaki’s exhaustive review {Seasonal Adaptations of Insects, Oxford Univ. Press, 1 986), which covers the subject extensively. At the very least, mention of these significant sources could have been included as notes added in proof The literature included in bibliographies following each chapter is predominantly British (almost 60%), and comprises much somewhat dated, marginally relevant work, particularly that on flight. Also, apart from the many annoying typographical errors, which betray a lack of careful proofreading, errors of fact and interpretation appear. As examples, Cloudsley-Thompson reports (p. 124) that the sources of honey bee alarm pheromones are unknown. In fact, the glands producing these chemicals, associated both with the mandibles and the sting, have long been known and are well characterized, as summarized in M. L. Winston’s recent review {The Biology of the Honey Bee, Harvard Univ. Press, 1987). And the author’s hurried discussion of the attributes contributing to arthropod success on land (Chapter 9) was better and more elaborately framed a decade ago by Eisner and Wilson {The Insects, Freeman, 1977). Lastly, the price of the book, at $33.00 for the paperback version, may tend to place it somewhat out of reach for many potential readers, particularly students. In view of its shortcomings, one might well balk at paying such a price for such a cursory, incomplete treatment. Cloudsley-Thompson has a long, well deserved reputation for solid, scholarly work in arthropod ecology. One would expect rather more of him in producing this hook. — Thomas W. Culliney, 46 Vineyard Road, North Haven, Con- necticut 06473. ENTOMO-ECOLOGICAL ASSOCIATIONS J. New York Entomol. Soc. 97(4):492, 1989 Coevolution and Systematics. A. R. Stone and D. L. Hawksworth (eds.). 1986. Sys- tematics Association Special Volume No. 32. The re-assertion of systematics in its rightful place in evolutionary biology was recently engendered by advances in phylogeny reconstruction. Of particular import has been recent phylogenetic inquiry into the evolution of species’ interactions. Ehrlich and Raven’s model of coevolution was based on well-known parallels between lepidopteran taxonomy and that of their hostplants, though most of the subsequent research has been on microevolutionary aspects of the interactions. There is new interest in the macroevolutionary consequences of insect/plant interactions, however, and phytogenies can provide evidence on the sequence of reciprocal ad- aptations between species, their effect on diversification rates, and (the theme of this volume) the frequency of parallel phylogenesis between interacting lineages. Systematists have long suggested that the classifications of hosts and parasites should be reciprocally informative. The volume examines the “rules” for inferring parasite relationships from those of their hosts. The chapters cover a taxonomically diverse assemblage of both animal and plant parasites, and agree in finding little empirical support for parasitological “rules.” However, many more general phylo- genetic questions have emerged and evidence on these should illuminate the role of interactions in the diversification and macroevolution of characters affecting the interactions. The analyses and information in each chapter should give the volume lasting value to scientists interested in the evolution and ecology of antagonisms. Several chapters review the biologies of major higher taxa (e.g., Eastop on aphids, Beveridge on marsupial helminths). Some focus on rigorous analyses of “parallel phylogenesis” (e.g., Humphries et al. on Nothofagus and its herbivores; Lyal on bird/mammal lice; Thompson on Umbelliferae and herbivores). Most of the authors present the results of searches for vicariance patterns and little attention is paid to other possible sys- tematic regularities (i.e., directional change in parasite obligateness or specialization) in the phylogeny of particular antagonisms. Quantitative analyses of vicariance/ dispersal hypotheses are also absent, but the statistical methods for such tests are just now becoming available in the literature. Finally, many authors (e.g., Barrett, Parlevliet, on the genetics of host specificity) relate the persistence of particular interactions to their ecology or genetics. Thus, despite the volume’s emphasis on systematics (suggested by the title), the coverage of this new subdiscipline is synthetic, and future syntheses of these approaches in the study of particular systems should serve to invigorate our common field of evolutionary research. 1989 BOOK REVIEWS 493 J. New YorkEntomol Soc. 97(4):493-494, 1989 Biology of Mutualisms. D. H. Boucher (ed.). 1985. Oxford University Press (paper- back issued 1988). Since its publication, this excellent volume has been reviewed from an ecological perspective many times (e.g., Beattie, 1986; Goodman, 1986; Wilson, 1986; Thomp- son, 1987). I therefore depart from the traditional role of reviewers and take this opportunity to offer a phylogeneticist’s view of what might be included in the next volume on this fascinating and rich field of study. The included chapters in the Biology of Mutualisms, Ecology and Evolution volume are uniformly microevolutionary in treatment and thus parallel the state of the field— to date there have been no phylogenetic studies of mutualistic associations. At present, we have no information on, for example, which qualities of particular mutualisms reflect evolution in situ vs. phylogenetically ancient traits or how participants are gained or lost to such associations. While Ehrlich and Raven’s model of coevolution has served to organize research of antagonistic interactions at both micro- and macroevolutionary levels, no such organizing influence has been realized in the study of mutualisms. For whatever reason— if because mutualistic interactions are more often “diffuse” or involve dis- parate, comparatively little-known taxa— broad macroevolutionary patterns for mu- tualisms are less well known than for plants and herbivores. It is entirely possible that the associations of mutualistic species are not as evolutionarily persistent as for plant/herbivore interactions, perhaps for the reasons elaborated in chapters devoted to modelling the interactions by Dean, Keeler, Lane, Law, Post et al., Vandermeer et al., and Wolin. However, while phylogenetic evidence that would bear on the issue is virtually nonexistent, many of the papers in this volume make specific macroevo- lutionary predictions testable by phylogenies. If mutualisms evolve, then phylogenies should provide evidence on whether, for example, they become more efficient, or if there is directional evolutionary change in the obligate/facultative nature of inter- actions or in the degree of specialization (as postulated in the chapters by Cook, Law, Soberon and Martinez del Rio and by Templeton and Gilbert). More specifically, Janzen’s contribution offers innumerable hypotheses testable with phylogenies such as whether plants dispersed by animals sequentially evolve greater ornamentations, coupled with greater seed protection and whether lineages of seed-dispersing animals evolve greater “sloppiness” or efficiency (as predicted). For ant/plant mutualisms, one might ask whether plant-ants are derived only from arboreal ancestors and whether ant-plants are similarly derived from a few phylo- genetic sources “pre-adapted” to mutualisms. Does entry into such mutualisms in- fluence the subsequent diversification and distribution of either partner, or otherwise alter the scope of evolutionary opportunities? So far little is known of the marks of phylogenetic history on the structure, assembly and diversity of mutualistic com- munities. Thus, while the volume edited by Boucher should certainly serve to focus future research at microevolutionary levels (facilitated by the insightful overviews of the conceptual and historical bases of mutualism theory provided in chapters by Boucher 494 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 97(4) and Lewis), such inquiries should be enriched by systematic scrutiny of the macro- evolutionary questions suggested therein (and those common to antagonistic inter- actions as well). While a few of the major evolutionary ecological paradigms of the 60’s (e.g., plant/herbivore coevolution, insect sociality) are now receiving scrutiny by systematists, the theory of mutualisms nicely encapsulated in Boucher’s volume presents a rich source of untested evolutionary scenarios for the present generation of phylogeneticists. In sum, these volumes edited by Boucher and by Stone and Hawksworth should be on the bookshelf of any scientist interested in both the evolution and ecology of interspecific interactions. The future of evolutionary biology should thus be bright- ened by new syntheses of phylogenetic research with the population biologies of ecological associations.— D. Farrell, Department of Entomology, University of Maryland, College Park, Maryland 20742. REVIEWS CITED Beattie, A. J. 1986. Ecology 67:1435-1436. Goodman, M. J. 1986. Choice 23:1234. Thompson, J. N. 1987. Amer. Sci. 75:537-538. Wilson, D. S. 1986. Quart. Rev. Biol. 61:544. 1989 LIFE MEMBERS Dr. David G. Casdorph Dr. Howard E. Evans Mr. Irving Granek Dr. Donald F. J. Hilton Mr. Mark Indenbaum Dr. Gary G. Kennen Mr. Kikumaro Okano Dr. Nellie M. Payne Dr. L. L. Pechuman Dr. Kathryn M. Sommerman Mr. Hubert J. Thelen Dr. F. Christian Thompson Mr. George F. Townes Mr. Theodore H. Weisse REVIEWERS The Editorial Staff thanks the following individuals who reviewed manuscripts considered for publication in 1989: G. E. Ball, D. E. Bright, J. M. Carpenter, J. A. Coddington, R. W. Crosskey, D. C. Darling, O. S. Flint, R. C. Froeschner, E. Grissell, C. E. Griswold, S. W. Hamilton, T. J. Henry, L. H. Herman, R. W. Holzenthal, G. Imadate, S. A. Kolmes, J. K. Liebherr, J. Maldonado, J. D. Mclver, J. E. McPherson, J. S. Miller, M. Muma, A. Norrbom, J. Pericart, W. L. Peters, R. V. Peterson, N. I. Platnick, J. Rawlins, F. H. Rindge, W. H. Robinson, J. G. Rozen, Jr., M. D. Schwartz, J. A. Slater, R. R. Snelling, R. J. Snider, J. G. Stoffolano, G. M. Stonedahl, D. B. Thomas, Jr., H. Topoff, C. A. Triplehom, D. L. Wagner, D. Wahl, S. Weller, A. G. Wheeler, Jr., D. M. Wood, J. B. Woolley, J. Zrzavy. CHANGE OF EDITORSHIP As of January 1990 the editorship of the Journal of the New York Entomological Society will change. The new editor is: Dr. James K. Liebherr, Department of Ento- mology, Comstock Hall, Cornell University, Ithaca, New York 14853. Please direct all correspondence concerning the Journal to him. Dr. Liebherr will be assisted by E. Richard Hoebeke, also of the Department of Entomology, Cornell University. ERRATUM Volume 92(2): 131, line 3 of Acknowledgments should read Richard Rosenblatt, not Rosen Rosenblatt. HONORARY MEMBERS Dr. F. S. Chew Mrs. Wm. K. Firmage Dr. James Forbes Dr. John G. Franclemont Dr. James A. Slater Dr. R. Vishniac SUSTAINING MEMBERS Dr. R. C. Froeschner Dr. Ralph F. Kirchner NOTICE Copies of many reprints of the late Peter Ashlock and of E. P. Van Duzee are available for $5.00 within the U.S. and $10.00 outside the U.S. Send payment to Dr. Robert W. Brooks, Snow Entomological Museum, University of Kansas, Law- rence, Kansas 66045-2119. STATEMENT OF OWNERSHIP, MANAGEMENT, AND CIRCULATION 1. Title of Publication: Journal of the New York Entomological Society (ISSN 0028-7199). 2. Date of Filing: August 1989. 3. Frequency of Issue: Quarterly. 4. Complete Mailing Address of Known Office of Publication: 1 04 1 New Hamp- shire, Lawrence, Kansas 66044. 5. Complete Mailing Address of the Headquarters or General Business Offices of the Publishers: % Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024-5192. 6. Full Names and Complete Mailing Addresses of Publishers, Editors, and Man- aging Editor: Publisher: Allen Press, Inc. (for the New York Entomological Society), 1041 New Hampshire, Lawrence, Kansas 66044. Editor and Managing Editor: Randall T. Schuh, % Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024-5192. 7. Owner: New York Entomological Society (non-profit), % Department of Ento- mology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024-5192. 8. Known Bondholders, Mortgages, and Other Security Holders Owning or Hold- ing 1% or More of Total Amount of Bonds, Mortgages, or Other Securities: None. 9. Purpose: The purpose, function, and non-profit status of this organization and exempt status for federal income tax purposes have not changed during the preceding 12 months. 10. Extent and Nature of Circulation: Avg. no. copies Actual no. copies each issue of single issue during preceding published nearest to 12 mo. filing date A. Total no. copies (net press run) 700 700 B. Paid circulation 1. Sales through dealers and carriers. street vendors, and counter sales — — 2. Mail subscription 599 595 C. Total paid circulation 599 595 D. Free distribution by mail, carrier, or other means; samples, complimen- tary, and other free copies 18 18 E. Total distribution 608 613 F. Copies not distributed 1. Office use, left over, unaccounted. spoiled after printing 92 87 G. Total 700 700 {Continued from back cover) ENTOMO-ECOLOGICAL ASSOCIATIONS Coevolution and Systematics Biology of Mutualisms Honorary, Life, and Sustaining Members Reviewers for 1989 Statement of Ownership, Management, and Circulation Brian D. Farrell 492 Brian D. Farrell 493-494 495 495 496 INSTRUCTIONS TO AUTHORS The Journal of the New York Entomological Society is devoted to the advancement and dissemination of knowledge of insects and related taxa. The costs of publishing the Journal are paid by subscriptions, membership dues, page charges, and the proceeds from an endowment established with bequests from the late C. P. Alexander and Patricia Vaurie. 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Journal of the New York Entomological Society VOLUME 97 OCTOBER 1989 NO. 4 CONTENTS Atlas of antennal trichobothria in the Pachynomidae and Reduviidae (Heterop- tera) Pedro W. Wygodzinsky and Sarfraz Lodhi 371-393 Review of the New World species of the genus Neottiglossa Kirby (Heteroptera: Pentatomidae) D. A. Rider 394-408 Orius minutus (Linnaeus) in North America (Hemiptera: Heteroptera: Antho- coridae) John D. Lattin, Adam Asquith and Steve Booth 409-416 Biology of Lopidea nigridea Uhler, a possible aposematic plant bug (Heteroptera: Miridae: Orthotylinae) James D. Mclver and Adam Asquith 4 1 7-429 Redescription of Platynus prognathus Van Dyke (Coleoptera: Carabidae: Platyn- ini) and circumscription of Lindroth’s Decentis and Hypolithos groups James K. Liebherr 430-437 Aggregation and predator avoidance in whirligig beetles (Coleoptera: Gyrinidae) K. Vulinec and M. C. Miller 438-447 Fir st record of the Palearctic species Oxypoda opaca (Gravenhorst) from North America (Coleoptera: Staphylinidae: Aleocharinae) E. Richard Hoebeke 448-454 Two mouthpart modihcations in larval notodontidae (Lepidoptera): their taxo- nomic distributions and putative functions G. L. Godfrey, J. S. Miller and D. J. Carter 455-470 On Venezuelan Leprclochus (Araneae, Zodariidae) Rudy Jocque and Norman I. Platnick 47 1^74 Karyotypes of three spider species (Araneae: Pholcidae: Physocyclus) James C. Cokendolpher 475-478 Notes and Comments A new structure on the hind legs of male Monalocoris carioca Carvalho and Gomes (Heteroptera: Miridae) Jorge A. Santiago- Blay and Jenaro Maldonado Capriles 479-482 Records of Chimarra soda (Trichoptera: Philopotamidae) from interior highland streams in Arkansas and Missouri Paul K. Lago, Michael L. Mathis and David E. Bowles 482-483 Poecilochirus monospinosus (Acarina: Mesostigmata: Parasitidae), a predator of house fly immatures: new locality records Christopher J. Geden, Donald C. Steinkraus and Donald A. Rutz 483-485 Book Reviews ARTHROPOD OVERVIEWS Spiders. Webs, Behavior, and Evolution Robert J. Raven 486-487 Evolution and Adaptation of Terrestrial Arthropods George C. Eickwort 487-489 Thomas W. Culliney 489-491 {Continued on inside back cover) lES SMITHSONIAN INSTITUTION NOlifUliSNI NVINOSHilWS SBiavaaiT LIBF CO iSNI NVINOSHXmS S3 I 8 11 LIBRARIES SMITHSONIAN INSTITUTION NOIl _ m I ES SMlTHSONIAN^INSTITUTION^NOliniliSNI-NVmOSHillAJS S3IdVdan”j*‘BF c/> ^ ^ 2: r CO z: c/> z § %% ^ ° z ^ Z CO — Z 00 LSN! NVINOSHIIWS SHIdVdail LIBRARIES SMITHSONIAN INSTITUTION NOIl 5 CO ~ rn >Z^a VrT^ tn zT^!£?^oX ^ /<^ lES SMITHSONIAN INSTITUTION NOIlfUllSNI NVINOSHimS S3ldVdail LIB I m ^ ~ ^ CO X E CO ISN! 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