The PAN-PACIFIC ENTOMOLOGIST Volume 78 January 2002 Number 1 Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES (ISSN 0031-0603) The Pan-Pacific Entomologist EDITORIAL BOARD J. Y. Honda, Editor R. M. Bohart Z. Nisani, Assistant to the Editor J. T. Doyen R. E. Somerby, Book Review Editor J. E. Hafernik, Jr. Robert Zuparko, Treasurer Warren E. Savary Published quarterly in January, April, July, and October with Society Proceedings usually appearing in the following October issue. All communications regarding non-receipt of numbers should be addressed to: Vincent F. Lee, Managing Secretary; and financial communications should be addressed to: Robert L. Zuparko, Treasurer; at: Pacific Coast Entomological Society, Dept, of Entomology, California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599. 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POSTMASTER: Send address changes to the Pacific Coast Entomological Society, % California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599. This issue mailed 22 March 2002 The Pan-Pacific Entomologist (ISSN 0031-0603) PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, U.S.A. ® The paper used in this publication meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PAN-PACIFIC ENTOMOLOGIST 78(1): 1-6, (2002) SURVIVAL AND DEVELOPMENT OF LACAN OBI A SUBJUNCTA (GROTE & ROBINSON) (LEPIDOPTERA: NOCTUIDAE) LARVAE ON COMMON WEEDS AND CROP PLANTS OF EASTERN WASHINGTON STATE Peter J. Landolt USDA-ARS, Yakima Agricultural Research Laboratory, Wapato, WA, 98951 USA Abstract. —Ten common weed species, four tree fruit crops and four row crops were evaluated as hosts for larvae of Lacanobia subjuncta (Grote & Robinson), a noctuid moth pest of apple in eastern Washington. A separate comparative evaluation was made of the suitability of five varieties of apple as hosts for L. subjuncta larvae. Development was completed, from neonate larva to adult, on nine of ten weed species and seven of eight crops tested, indicating a broad potential host range for this insect. High rates of survival to adult, short developmental times, and large pupal weights were noteworthy on the weeds bindweed, dandelion, and mallow, and on potato. In the comparison of apple varieties, highest rate of survival to adult was with Red Delicious, greatest pupal weights were with Red Delicious, Gala, and Fuji, and shortest devel¬ opment times were with Gala and Golden Delicious. Strong seasonal variation (May versus July) was indicated in the quality of apple foliage as food for L. subjuncta larvae. Key Words: —Insecta, Lacanobia subjuncta, host plant, apple, potato. The noctuid moth Lacanobia subjuncta (Grote & Robinson) has recently be¬ come recognized as a significant pest of apple in eastern Washington and Oregon (Landolt 1998). The moth is widely distributed in North America (McCabe 1980), and has been present in irrigated areas of eastern Washington at least since the 1970s when it was collected in light traps in Yakima County by F. Howell (per¬ sonal communication). Following an apparent increase in damage to apple attri¬ buted to cutworms (Warner 1996), L. subjuncta was identified as the principal noctuid pest on apple in eastern Washington and adjacent areas of Oregon (Lan¬ dolt 1998). Lacanobia subjuncta is bivoltine (McCabe 1980), with a flight of moths from late May into mid June and again from late July into mid September in eastern Washington (Landolt 1998, Hitchcox 2000). Larvae can be found on apple trees from early June through July and again from late August until October (Hitchcox 2000). It is thought that the insect overwinters strictly as a pupa in soil. Most larvae held in the laboratory and fed on apple foliage went through 6 instars before pupating, although several larvae went through 7 (Hitchcox 2000). On apple, larvae of L. subjuncta are primarily foliage feeders and occasionally partially defoliate apple trees in commercial orchards. Damage to apple fruit oc¬ curs also, with larval feeding indicated by a hollowed out scoop in the surface of the apple that is somewhat characteristic of fruit feeding by other noctuids. The pest status for L. subjuncta on apple is due principally to their feeding on apple fruit and to problems in packing houses because of the presence of larvae on fruit (Warner 1998). The recently acquired pest status of this insect on apple in Washington and Oregon is not understood. Hypotheses to explain this apparent change in pest status include 1) resistance to commonly used pesticides together with escape 2 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1) from natural enemies, 2) shifts in larval host plants, 3) changes in geographic distribution, 4) past misidentification of cutworms and fruit worms damaging ap¬ ple, and 5) region-wide L. subjuncta moth population increases resulting in move¬ ment of moths into apple orchards. In order to consider the latter hypothesis, better information is needed on what plants may sustain L. subjuncta reproduction and may then be the principal host plants contributing to regional moth popula¬ tions. Larvae of L. subjuncta have been collected on a wide variety of plants, in¬ cluding trees, shrubs, and herbaceous plants (Godfrey 1972, Rings et al. 1992, Crumb 1956, Landolt 1998, and included references), indicating a potentially high degree of polyphagy. Recorded host plants include a number of agricultural crop plants, such as apple, cherry, peach, blueberry fruit and foliage, cabbage, aspar¬ agus, corn, and strawberry. However, the finding of larvae on a plant species is not necessarily a good indicator of suitable host plant status. The survival and development of larvae on plants along with the incidence of larvae on plants in the field would be better indicators of the importance of those plants as contrib¬ utors to populations of L. subjuncta. In a preliminary assessment of host suitability of common weeds, Landolt (1998) demonstrated that L. subjuncta larvae could complete development on the weeds Taraxacum officinale Weber (dandelion), Sonchus oleraceus L. (annual sowthistle), Convulvulus arvensis L. (field bind¬ weed), and Malva neglecta Walk (comon mallow), but with low (20 to 28) per¬ centages that survived to the adult stage. Reported here are the results of experiments evaluating the ability of larvae of L. subjuncta to develop on foliage of a larger number of plant species (18) that are commonly encountered in irrigated areas of eastern Washington. This evalu¬ ation included determination of larval survival to pupation and adult emergence, larval development time, and pupal weights for L. subjuncta fed on the foliage of each plant species or variety. The objective of the study was to determine if these plants sustain complete development of newly hatched larvae through to adult, and might then contribute in the field to populations of L. subjuncta. Ad¬ ditionally, this study sought to identify plant species that might be further eval¬ uated in the field as good hosts for L. subjuncta. Materials and Methods For each plant species and apple variety evaluated, data were obtained on sur¬ vival of larvae to pupation and adult emergence, on larval development time (egg hatch to entry into soil), and on pupal weight. Plants evaluated were weed species Sonchus asper (L.) Hill (spiny sowthistle), dandelion, common mallow, field bind¬ weed, Cirsium arvense (L.) Scop. (Canada thistle), Helianthus annuus L. (sun¬ flower), Chenopodium album L. (lambsquarters), Amaranthus retroflexus L. (red- root pigweed), Cardaria dr aba (L.) Desv. (hoary cress), and Kochia scoparia (L.) Schrad. (kochia), the crop plants Malus pumila Mill, (apple, var. Fuji), Pyrus communis L. (pear, var. Bartlett), Prunus armeniaca L. (apricot), Medicago sativa L. (alfalfa), Pisum sativum L. (dry peas, var. Columbian), Pisum sativum L. (suc¬ culent peas, var. Oregon Trail), and Solanum tuberosum L. (potato, var. Russet Burbank). In another experiment, the apple varieties Fuji, Gala, Braeburn, Red Delicious, and Golden Delicious were evaluated as hosts for L. subjuncta larvae. Adult L. subjuncta were obtained from a walk-in light trap at the Yakima 2002 LANDOLT: LACAN OBI A SUBJUNCTA BIONOMICS 3 Agricultural Research Laboratory southeast of Yakima, Washington, in an area of commercial irrigated apple and pear orchards. Female moths from the light trap were held for 24 h in 50 ml plastic snap-cap vials for oviposition. Females that did not oviposit within that time period were assumed to be unmated and were moved to 900 ml plastic tubs with ventilated lids. These tubs contained sugar water on cotton balls, water on cotton balls, and two males that had been collected in the light trap. Some of these females subsequently laid fertile eggs. Newly eclosed larvae from egg batches were used for the following experiments. The assay unit was a 300 ml waxed paper carton with a clear plastic lid, in which was placed plant foliage and one newly hatched larva. Cartons were held in a room on a 16:10 light:dark photoperiod, at 25° C and 60% RH. Plant foliage was added daily. Dried, brown, or moldy foliage was removed daily and new cartons were provided as needed when soiled by frass. When larvae reached about 3 cm in length, 2—3 cm of damp soil was placed in the carton and the plant foliage and larva were placed on top of this soil. Mature larvae entered the soil to pupate. Daily records were made of larval mortality, and larval movement into soil. Four to 6 days after larvae moved into soil, the soil was sifted to confirm the presence of a pupa, which was then weighed and transferred to a 30 ml paper cup held inside of the original waxed paper carton. Daily observations were made of pupae in order to record adult emergence. Data was not obtained on pupal duration because it was not possible to tell on what day a larva in the soil pupated. Dis¬ turbing the larva at that time might interfere with successful development. Plant foliage was obtained in the field as needed by cutting plants with scissors and transferring foliage in 3.6 liter Ziplock® plastic freezer bags held in a cooler until return to the laboratory. Bags of foliage were held in a refrigerator at 3° C for up to seven days and were accessed daily to obtain foliage that was provided to larvae. For weeds, plants were selected from areas not sprayed with insecti¬ cides. For crop plants, growers were consulted for information on the timing and application of insecticides to avoid the collection of foliage with pesticide resi¬ dues. For each plant species or apple variety evaluated, three sets of five larvae were tested, with each set of five larvae originating from the egg batch of a different female moth. These three sets of larvae were staggered in time (three weeks apart) so that the individual plants evaluated were different for each set of five larvae. Results Larvae of L. subjuncta developed to pupation and adult emergence on the following weeds: spiny sowthistle, dandelion, bindweed, lambsquarters, mallow, Canada thistle, hoary cress, pigweed, and kochia (Table 1). No larvae survived to pupation on sunflower. Highest percentages of larvae surviving to pupation were fed on field bindweed. Larvae also developed to pupation and adult emer¬ gence on the following crop plants: cherry, apple, pear, alfalfa, potato, dry peas, and succulent peas (Table 1). No larvae survived to pupation on apricot. Highest percentages of larvae that pupated were on cherry, pear, potato and alfalfa. Larvae developed to pupation and adult emergence on all 5 apple varieties tested, but with significant differences in percentage of pupation among these varieties (Table 2). Significantly more larvae on Gala apple foliage died than did on Golden Delicious foliage. 4 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1) Table 1. Mean (±SE) % pupation, adult emergence, pupal weight, and larval development times for Lacanobia subjuncta on 10 weedy plant and 8 crop species. Yakima, Washington, 2000. a Pupal weight Larval time Plant species % Pupated % Adult (mg) (days) Hoary Cress 66.7 + 13.3c 60.0 + 11.5 335 ± 8c 44.3 ± 1.6a Common Mallow 80.0 + 11.5bc 80.0 + 11.5 363 ± 15b 26.1 ± 0.7c Spiny Sowthistle 60.0 + 20.0c 60.0 + 20.0 436 ± 10a 26.9 ± 1.3bc Dandelion 86.7 + 6.7b 86.7 + 6.7 392 ± 5b 25.7 ± l.lbc Canada Thistle 46.7 + 6.7d 46.7 + 6.7 249 ± 16e 28.7 ± 1.5bc Common Sunflower 00.0 H- OO.Oe 00.0 + 00.0 _+. .+_ Lambsquarter 85.0 + 5.0b 73.4 + 6.7 342 ± lie 28.4 ± 1.6bc Redroot Pigweed 70.0 + lO.Od 70.0 + 10.0 280 ± lOde 35.1 ± 2.9ab Field Bindweed 100 + 00.0a 100 + 00.0 374 ± 8b 25.1 ± 0.6c Kochia 40.0 + 11.5d 40.0 + 11.5 270 ± 12e 35.8 ± 2.7ab Apple 66.7 + 6.7c 66.7 + 6.7 389 ± 7b 25.1 ± 0.7c Pear 100.0 H- 00.0a 100 + 00.0 333 ± 8c 26.5 ± 0.7bc Cherry 93.4 + 6.7ab 86.7 + 6.7 323 ± 12cd 30.4 ± 1.1b Apricot 00.0 H- OO.Oe 00.0 + 00.0 _+_ _+_ Potato 100 + 00.0a 100 00.0 377 ± 12b 24.7 ± 0.9c Alfalfa 100 H- 00.0a 100 + 00.0 297 ± 8d 29.9 ± 0.9b Dry Peas 86.7 + 13.3b 60.0 + 23.0 285 ± lOde 31.3 ± 1.0b Succulent Peas 73.4 + 17.6bc 66.7 -h 24.0 286 ± 8d 29.8 ± 0.9b a Means within a column followed by the same letter are not significantly different by Tukey’s test at P > 0.05. Mean development rates for larvae on weeds ranged from 25.1 days on bind¬ weed to 44.3 days on hoary cress (Table 1). Other weeds supporting rapid de¬ velopment of larvae were spiny sowthistle, dandelion, mallow, lambsquarter, and Canadian thistle. Mean development rates for larvae on crop plants ranged from 24.7 days for potato to 31.3 days on dry peas (Table 1). Other crops supporting rapid development were apple and pear. Among apple varieties tested, mean de¬ velopment times were similar for Gala, Red Delicious, Fuji and Golden Delicious, while development on Granny Smith foliage was significantly slower (Table 2). Also of interest, mean development time for larvae on Fuji apple foliage was 25.1 ± 0.7 days when evaluated in the first study and was then 38.2 ±1.0 days when evaluated in the second study, along with other apple varieties. The first evaluation used apple foliage collected during May, while the second evaluation used apple foliage collected during July. Table 2. Mean (± SE) % pupation, adult emergence, pupal weight, and larval development times for Lacanobia subjuncta on foliage of 5 apple varieties. Yakima, Washington, 2000. a Apple variety % Pupated % Adult Pupal weight (mg) Larval time (days) Gala 40.0 + 11.5b 33.3 + 6.7 336 ± 30ab 37.4 + 6.6bc Red Delicious 73.3 + 6.7ab 73.3 + 6.7 387 ± 9a 38.0 4- l.Obc Fuji 73.3 4- 13.3ab 60.0 + 11.5 357 ± 8ab 38.2 + 1.0b Golden Delicious 80.0 4- 20.0a 60.0 + 11.5 321 ± 12b 35.7 4- l.Obc Granny Smith 66.7 4- 13.3ab 50.0 -h 10.0 284 ± 22b 47.7 + 2.1a a Means within a column followed by the same letter are not significantly different by Tukey’s test at P > 0.05. 2002 LANDOLT: LACAN OBI A SUBJUNCTA BIONOMICS 5 Mean pupal weights for L. subjuncta reared on weeds ranged from 249 mg on Canada thistle to 436 mg on spiny sowthistle (Table 1). Other relatively heavy mean pupal weights were 392 mg for larvae reared on dandelion, 374 mg for larvae reared on bindweed, and 363 mg for larvae reared on mallow. In addition to Canada thistle, pigweed and kochia yielded low pupal weights (Table 1). Mean pupal weights for larvae fed crop plants ranged from 285 mg for dry peas to 377 mg for potato and 389 mg for Fuji apple (Table 1). When mean pupal weights for apple varieties were compared, they ranged from 284 mg for larvae fed Gran¬ ny Smith to 387 mg for larvae fed Red Delicious apple foliage (Table 2). Discussion These results clearly indicate a broad potential plant host range forL. subjuncta larvae and the potential for many plants in eastern Washington to contribute to regionally high population densities contributing to crop losses. Complete devel¬ opment from egg hatch to adult emergence was documented for 9 of the 10 weeds tested and 7 of the 8 crops tested, with sunflower and apricot not supporting larval development to pupation in this study. The plants selected for this study and supporting L. subjuncta development are taxonomically diverse and include the families Compositae, Cruciferae, Chenopodiaceae, Convulvulaceae, Malvaceae, Solanaceae, Rosaceae, and Leguminaceae. Undoubtedly, many other plants pre¬ sent in the region are probably equally suitable as hosts for L. subjuncta. Just as the collection of larvae on a plant species is not proof that the species is a good host plant, the demonstrations of survival and development to adult on these plant species does not demonstrate that L. subjuncta utilizes these plants. Additional information on patterns of adult female egg laying, of larval dispersal and movement under field conditions, and of larval numbers on these and other plant species would be more conclusive in assessing the potential of these plants as hosts. Such studies clearly should incorporate not only common weeds but additional crops that have not been reported to be infested with L. subjuncta. There were differences in the performance of larvae on Fuji apple foliage col¬ lected in May versus July that indicate possible seasonal variation in the suitability of foliage of apple as food for L. subjuncta larvae. There are two broods of L. subjuncta in Washington, with most larvae feeding in June/July and again in August/September (Hitchcox 2000). Larvae reared on Fuji apple foliage in May developed more rapidly and yielded larger pupae than larvae reared on Fuji apple in July. Despite these possible differences in apple suitability as a host for L. subjuncta, larvae are readily found on apple in the field during both time periods (Hitchcox 2000). Such differences in host suitability could be due to a variety of factors, such as accumulated leaf chemical defenses, increases in average leaf age, accumulated responses to disease and herbivore challenges, and nutritional chang¬ es in leaves. There is potential for much variation in foliage quality as food for larvae for each of the plant species tested and care must be exercised in using the data presented here for comparative purposes among plant species. Also of concern is the potential impact of the apparent variance of plant suit¬ ability as food forL. subjuncta larvae on phenology models used in IPMprograms for apple orchards. Such models may be used to predict L. subjuncta egg hatch for the purpose of accurately timing pesticide applications. If larval development rates are dependent in part on seasonal parameters affecting plant physiology, 6 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1) these must be incorporated into insect developmental models. Also, differential rates of development on different host plants will contribute to variance in adult emergence, oviposition, and egg hatch when multiple plants are used as hosts. Acknowledgment Technical assistance was provided by L. Biddick, J. Brumley, J. Dedlow, and D. Lovelace. This work was supported in part by funding from the Washington State Tree Fruit Research Commission. Literature Cited Crumb, S. E. 1956. The larvae of the Phalaenidae. USDA Tech. Bull. No. 1135. Washington D.C. Godfrey, G. L. 1972. A review and reclassification of the subfamily Hadeninae (Lepidoptera: Noc- tuidae) of America north of Mexico. USDA Tech. Bull. 1450. Hitchcox, M. E. 2000. Seasonal phenology and monitoring of Lacanobia subjuncta (Noctuidae: Lep¬ idoptera) in apple orchards of Washington State. M.Sc. Thesis, Washington State University, Pullman, WA. 75 pp. Landolt, P. J. 1998. Lacanobia subjuncta (Lepidoptera: Noctuidae) on tree fruits in the Pacific North¬ west. Pan-Pacific Entomol., 74: 32-38. McCabe, T. L. 1980. A reclassification of the Folia complex for North America (Lepidoptera: Noc¬ tuidae). N.Y. St. Mus. Bull. No. 432, Albany. Rings, R. W., E. H. Metzler, F. J. Arnold & D. H. Harris. 1992. The owlet moths of Ohio. Order Lepidoptera, family Noctuidae. Ohio Biol. Survey Bull. (NS), 9: 1-184. Warner, G. 1996. Goodbye codling moth, but hello cutworms. Good Fruit Grower, 47 (April 15): 17-18. Warner, G. 1998. New pest causes havoc in orchards and warehouses. Good Fruit Grower, 49 (May 15): 28-30. Received 29 June 2001; Accepted 16 Sept. 2001. PAN-PACIFIC ENTOMOLOGIST 78(1): 7-16, (2002) A NEW SPECIES OF HETEROSPILUS (HYMENOPTERA: BRACONIDAE) ASSOCIATED WITH THE DEATHWATCH BEETLE, HEMICOELUS GIBBICOLLIS (LECONTE) (COLEOPTERA: ANOBIIDAE) Brian J. Cabrera 1 , Paul M. Marsh 2 , Vernard R. Lewis 3 , & Steven J. Seybold 4 'Fort Lauderdale Research and Education Center, University of Florida, Fort Lauderdale, Florida 33314 2 P. O. Box 384, North Newton, Kansas 67117 department of Environmental Science, Policy and Management, University of California, Berkeley, California, 94720 departments of Entomology and Forest Resources, University of Minnesota, St. Paul, Minnesota 55108-6125 Abstract.—Heterospilus luridostigmus Marsh, a new braconid wasp species, is described. This wasp was found in abundance emerging from pieces of Douglas-fir, Pseudotsuga menziesii (Mir- bel) Franco, from an outdoor wooden deck in Daly City, California, that was infested with the deathwatch beetle, Hemicoelus gibbicollis (LeConte). Adult Heterospilus luridostigmus began emerging in May 1999 followed by emergence of Hemicoelus gibbicollis about 4 weeks later. Both species continued to emerge throughout the summer of 1999 and were the only species found in the boxes. During the summer of 2000, rearing boxes containing infested wood from the original source and from several homes in Alameda County, California yielded both Heter¬ ospilus luridostigmus and Hemicoelus gibbicollis as well as Odontocolon polymorphum Cushman (Hymenoptera: Ichneumonidae). Although we found no direct evidence of parasitism of Hemi¬ coelus gibbicollis, wasps in the genera Heterospilus and Odontocolon are known to parasitize anobiids, and Heterospilus flavicollis (Ashmead) is a parasitoid of an eastern deathwatch beetle species, Hemicoelus carinatus (Say). This suggests that Heterospilus luridostigmus is a parasitoid of Hemicoelus gibbicollis. The synchrony of emergence of these two species during both years also is indicative of a host/parasitoid relationship between the two species. The discovery of a new insect species in a heavily populated urban environment is noteworthy and serves as a vivid reminder of the untold number of insect species that have yet to be discovered. Key Words .—Insecta, Braconidae, Anobiidae, Ichneumonidae, deathwatch beetle, parasitoid, emergence hole. The genus Heterospilus Haliday is a member of the braconid subfamily Do- ryctinae. It can be identified by using the key to New World genera of Braconidae (Marsh 1997). In North America the genus is easily distinguished from the other doryctine genera by the reduction of fore wing vein 2RS (Fig. 1), which is always desclerotized and often completely absent, and the presence of a stigma in the hind wing of the male (not pictured). The small genus Pioscelus Muesebeck and Walkley, which also has fore wing vein 2RS absent, is separated by having no basal tubercle on the hind coxa. Heterospilus is the most species-rich genus in the Doryctinae, with an estimated 200 species in the Nearctic Region and 300 in the Neotropical Region. Most of these species are undescribed and the genus is badly in need of revision. All Heterospilus are idiobiont ectoparasitoids (Shaw & Huddleston 1991) and this genus also has the most diversified host range in the Doryctinae. Species of the genus parasitize a very wide range of endophytic, mostly stem-boring, hosts 8 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1) Figure 1. Wing venation of female Heterospilus luridostigmus, n. sp. (Marsh 1982, Shaw 1995) including those in the coleopteran families Anobiidae, Bostrichidae, Bruchidae, Buprestidae, Cerambycidae, Curculionidae, Languriidae, Mordellidae and Scolytidae, and those in the lepidopteran families Gelechiidae, Incurvariidae, Pyralidae and Tortricidae. Several other species have been reared from other hosts including stem-boring symphytan Hymenoptera. A few species are known to attack pemphredonine sphecid wasps (Marsh & Melo 1999). In North America, five species have been recorded from Anobiidae, including the new species described in this paper whose authorship is attributed solely to P. M. Marsh. V. R. Lewis obtained beetle-infested wood was obtained by V. R. Lewis. Wasps and beetles were reared and collected by V. R. Lewis and B. J. Cabrera. S. J. Seybold and V. R. Lewis are principal investigators on a deathwatch beetle, Hemicoelus gibbicollis (LeConte), pheromone project that includes the work presented in this paper. Heterospilus luridostigmus Marsh, NEW SPECIES Types. —Holotype, female. CALIFORNIA: SAN MATEO CO.: Daly City, 393 Oriente St., Oct. 1998, V. R. Lewis, ex Pseudotsuga menziesii (Mirbel) Franco boards from wooden deck. Deposited in NMNH, Washington, D.C. Paratypes. —CALIFORNIA: same data as holotype, 41 females, 7 males. De¬ posited in the California Academy of Sciences, San Francisco, NMNH, Washing¬ ton, D.C. and the University of Minnesota Insect Collection, University of Min¬ nesota, St. Paul. Description: — Female. Color: head, mesosoma and metasoma dark brown; scape, pedicel and basal flagellomeres light brown, apical flagellomeres dark brown; fore and middle legs yellow, femora marked with raised brown spot dorsomedially, tibiae marked with brown; hind leg with coxa and femur brown, trochanters yellow, tibia yellow marked with brown, tarsus yellow; wings slightly dusky, veins brown, stigma and vein C+SC+R light yellow, stigma sometimes nearly white. Body size: 2.5- 2002 CABRERA ET AL.: A NEW HETEROSPILUS SPECIES 9 4.0 mm. Head: face smooth and with dense long gold hair, frons and vertex transversely striate (Fig. 2), occasionally weakly so and nearly smooth; temple smooth; malar space two-thirds eye height; ocell-ocular distance about 3 times diameter of lateral ocellus; occipital carina not meeting hypostomal carina; 19-23 antennomeres. Mesosoma: pronotum rugose, with dense gold hair along weak pronotal groove; mesonotal lobes (Fig. 3) coriaceous, rugose along notauli, notauli scrobiculate, meeting before scutellum in rugose area with longitudinal short carinae, dense long gold hair along notauli; scutellum smooth; mesopleuron smooth, subalar area carinate, sternaulus deep and longitudinally striate, dense long gold hair on subalar area and along posterior edge; propodeum (Fig. 4) rugose, basal median areas smooth, median carina and areola distinct, dense gold hair laterally. Legs: hind coxa with small but distinct antero-ventral basal tubercle or tooth. Wings (Fig. 1): fore wing vein r as long as or slightly longer than 3RSa, vein 2RS indicated by weak infuscate line, vein r-m not sclerotized but distinct, vein m-cu arising distad from vein 2RS; hind wing vein M+CU longer than vein 1M, vein m-cu a distinct infuscated line. Metasoma (Fig. 5): first tergum longitudinally carinate, length slightly less than apical width, median raised area distinct, defined by carinae only on basal half; second tergum longitudinally carinate; third tergum smooth with carinate area across basal half; remainder of tergum smooth; ovipositor about two-thirds length of metasoma. Male .—Essentially as in female; body size 1.5-3.0 mm; 17-20 antennomeres; hind wing with elongate stigma. Biology .—Associated with adults of Hemicoelus gibbicollis (LeConte) (Cole- optera: Anobiidae) infesting Douglas-fir boards from a backyard deck. See details of biology below. Comments .—This species is distinctive by its light yellow to almost white stig¬ ma in the fore wing and by vein M+CU in the hind wing being longer than vein 1M. Although this hind wing venation is not typical for the genus, in all other characters the specimens are clearly congeneric. This species is easily distin¬ guished from H. baeticatus (Provancher), H. flavicollis (Ashmead) and H. lon- gicauda (Ashmead), which are also recorded as parasitoids of anobiids, by having the ovipositor shorter than the metasoma. This species has a similar ovipositor length to H. anobiidivorus Muesebeck but differs in the light stigma, completely carinate second metasomal tergum, shorter first metasomal tergum and longer antenna. Etymology. —The specific name is from the Latin luridus meaning pale yellow in reference to the pale yellow or almost white stigma in the fore wing. Biology and Observations Wood Collection. —Wood [Douglas-fir, Pseudotsuga menziesii (Mirbel) Franco] infested with the deathwatch beetle, Hemicoelus gibbicollis, was collected in Oc¬ tober 1998 from the deck of a home in Daly City, San Mateo County, California. The wood was cut into lengths of approximately 20-50 cm and stored at ambient temperature and relative humidity in a greenhouse at the University of California, Berkeley, in two wooden emergence boxes (122 X 122 X 122 cm) or in 58 liter plastic storage boxes with 5-cm diameter screen-covered holes in each end for ventilation. Wood moisture content was monitored with a moisture meter (Proti- meter Timbermaster, Protimeter Ltd., Marlow Bucks, England) and the wood was watered as needed to maintain approximately 14—17% moisture content (Suomi & Akre 1992a, b). Additional pieces of infested wood (predominantly P. menzie¬ sii) were collected in October, 1999 from several homes in Alameda County, California and kept outdoors on the premises of the University of California, Forest Products Laboratory, Richmond, California. In June 2000, this wood was also cut and stored in plastic storage boxes as previously described. 10 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1) Figures 2-5. Morphological characters of Heterospilus luridostigmus, n. sp. Figure 2. Vertex and frons. Figure 3. Mesonotum. Figure 4. Propodeum. Figure 5. Metasomal terga 1-4. 2002 CABRERA ET AL.: A NEW HETEROSPILUS SPECIES 11 Beetle and Wasp Emergence, 1999. —Boxes were checked occasionally for adult Hemicoelus gibbicollis (needed for extracting sex pheromone and for be¬ havioral assays) beginning in April 1999. Heterospilus luridostigmus appeared unexpectedly in the wooden emergence boxes on 3 May and continued to emerge through 2 Aug (Fig. 6A). The exact date of initial emergence is unknown because the boxes were not examined on a regular basis. The first Hemicoelus gibbicollis adults were found on 28 May with the exact date of emergence also unknown. The last adults were collected on 16 August. This emergence was in agreement with Suomi & Akre (1993a, b) who stated that normal emergence occurs during June, July, and August. We collected a total of 584 beetles (336 alive, 57.5% survival) and 179 wasps. Beetle and Wasp Emergence, 2000. —Beetles and wasps were first collected on 15 June. The number of emerged adults of both species was considerably lower than in 1999 (Figs. 6A and 6B), possibly because of the different collection source, differences in the host wood, or because of exposure to sub-optimal en¬ vironmental conditions prior to rearing. The wood had been kept outdoors for seven months before it was cut and placed in the rearing boxes. A total of 218 beetles (132 alive, 60.6% survival) and 32 wasps were collected. Several adult male and female Odontocolon polymorphum Cushman (Hymenoptera: Ichneu- monidae) were collected in addition to Heterospilus luridostigmus. Possible Parasitism of Hemicoelus gibbicollis. —The doryctine b raconids are generally considered to be ectoparasitoids of wood-boring beetle larvae (Marsh 1979). However, Suomi & Akre (1992a, b, c; 1993a, b) did not mention parasit- oids in their detailed descriptions of Hemicoelus gibbicollis biology and ecology. Furthermore, we did not directly observe wasps emerging from any life stage of Hemicoelus gibbicollis or from the wood and dissection of several small pieces of wood did not yield any parasitized larvae. The need for large numbers of adult H. gibbicollis made us reluctant to conduct a more thorough search that would have required destruction of more beetle-infested wood. However, we believe that Heterospilus luridostigmus is a parasitoid of Hemicoelus gibbicollis because: 1) adults of both species appeared in our rearing boxes in relative synchrony during both years; 2) a broad range of hole diameters was observed on the surface of the infested wood; and 3) other species of Heterospilus are idiobiont ectoparasi¬ toids of wood-destroying anobiids (e.g., Heterospilus flavicollis (Ashmead) on Hemicoelus carinatus (Say) [Drooz 1985], the most common wood-infesting deathwatch beetle in the northeastern United States [Simeone 1962], and Heter¬ ospilus longicauda (Ashmead) on Xyletinus peltatus (Harris) [Williams et al. 1979]). The only two live insect species to appear in 1999 in our rearing boxes were Heterospilus luridostigmus and Hemicoelus gibbicollis. Both species were col¬ lected again in 2000 from rearing boxes containing wood that was collected both years. In 1999, adult Heterospilus luridostigmus were first observed approxi¬ mately four weeks before the first emergence of Hemicoelus gibbicollis while the following year both species were first found on the same day. Peak emergence of both species was nearly synchronous in 1999, with the largest number of wasps emerging approximately two weeks before the largest number of beetles emerged (Fig. 6A). A similar trend was observed the following summer (Fig. 6B). The observation that Heterospilus luridostigmus preceded Hemicoelus gibbicollis in H. gibbicollis (Total number) tf. gibbicollis (Total number) Vol. 78(1) 12 THE PAN-PACIFIC ENTOMOLOGIST 80 70 60 50 40 30 20 10 0 1-May 16-May 31-May 15-Jun 30-Jun 15-Jul 30-Jul 14-Aug Date 50 40 30 20 10 0 B / . ii 4 I A iii ■A iii _ 1-May 16-May 31-May 15-Jun 30-Jun 15-Jul 30-Jul 14-Aug Date Figure 6. Emergence of beetles and wasps. A. Hemicoelus gibbicollis (LeConte) (bars), and Het- erospilus luridostigmus Marsh (line), summer 1999. □ Live beetles, ■ Dead beetles, A Live wasps. B. Hemicoelus gibbicollis (LeConte)(bars), and Heterospilus luridostigmus Marsh and Odontocolon polymorphum Cushman (line), summer 2000. □ Live beetles, ■ Dead beetles, A Live wasps. Wasps (Total number) Wasps (Total number) 2002 CABRERA ET AL.: A NEW HETEROSPILUS SPECIES 13 emergence in 1999 but not in 2000 might be attributed to laboratory worker inexperience, as both species are difficult to find amongst the pieces of wood in the rearing boxes. Hemicoelus gibbicollis is especially difficult to locate because emerged adults spend large periods of time seeking refugia in emergence holes. We also expected a much larger emergence of Hemicoelus gibbicollis during the summer of 2000. Dissection of several infested pieces of wood in mid-August of that year yielded beetle larvae of mixed size and, presumably, age, indicating that the infestations were still active. We speculate that the lower number of emerged beetles and the shorter emergence period during the second year were partly a result of parasitism. Williams et al. (1979) found that Heterospilus longicauda (Ashmead) accounted for 1.3—36.6% mortality of the potential population of their anobiid host, X. peltatus. We frequently observed small emergence holes in the infested wood, presum¬ ably made by Heterospilus luridostigmus, adjacent to much larger and more abun¬ dant emergence holes, presumably made by Hemicoelus gibbicollis (Fig. 7) ( Hem¬ icoelus gibbicollis is considerably larger than Heterospilus luridostigmus in cross- sectional area). Williams et al. (1979) reported that Heterospilus longicauda emer¬ gence holes were about one-eighth the size of X. peltatus emergence holes, that the wasp oviposited through the wood and onto the host larvae, and assumed there was one parasite larva per host. However, we measured holes (n = 434) from several pieces of wood collected in 1999 and instead of an expected bimodal distribution we obtained an approximately normal distribution ranging from 0.46 to 2.15 mm in diameter (Fig. 8). The lack of a distinct separation among emer¬ gence hole diameters prevents us from reporting species-specific emergence hole size ranges at this time. In contrast to Williams et al. (1979), our minimum emer¬ gence hole sizes all exceeded one-eighth of the maximum hole sizes. Our sample of emergence holes probably also contained holes made by O. polymorphum. The appearance of O. polymorphum from our rearings in 2000 reveals the possible existence of other parasitoids of Hemicoelus gibbicollis. The wood in the rearing boxes had been lying outdoors for seven months, thus making larvae and pupae of H. gibbicollis within the wood readily accessible to attack by Heteros¬ pilus luridostigmus and other opportunistic natural enemies. Alternatively, Het¬ erospilus luridostigmus and O. polymorphum may have located and colonized Hemicoelus gibbicollis while the wood was still in the structures from which it was removed. Odontocolon polymorphum has been collected in Oregon, Wash¬ ington, and British Columbia and has been associated with two wood-infesting anobiid species (one unidentified, the other Ptilinus basalis LeConte) (Carlson 1979). This paper represents the first report of O. polymorphum associated with Hem¬ icoelus gibbicollis. Ovipositor length and examples of other xoridine ichneumon- ids (Townes et al. 1960) suggest that O. polymorphum is ectoparasitic and ovi¬ posits through the wood surface onto larvae and pupae of H. gibbicollis. Whether H. gibbicollis is an obligate or facultative host for both wasp species remains to be determined. Although there is strong evidence for a host/parasitoid relationship between Hemicoelus gibbicollis and Heterospilus luridostigmus, di¬ rect observation of wasp oviposition behavior, emergence of adult wasps from the host or infested wood, detection of parasitized beetle larvae or larval beetle -F^ Figure 7. Adult emergence hole of Hemicoelus gibbicollis (LeConte) (A) and suspected emergence hole of Heterospilus luridostigmus Marsh (B). Bar = 1 mm. THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1) 2002 CABRERA ET AL.: A NEW HETEROSPILUS SPECIES 15 0.46 0.62 0.77 0.92 1.08 1.23 1.38 1.54 1.69 1.85 2.00 2.15 Emergence hole diameter (mm) Figure 8. Size distribution of emergence holes (n = 434) of Hemicoelus gibbicollis, Heterospilus luridostigmus, and Odontocolon polymorphum measured from several pieces of infested Pseudotsuga menziesii, collected in 1999 in Alameda Co., California. head capsules present in wasp cocoons is needed to confirm the existence of this proposed ecological relationship. Finally, it is noteworthy that a new insect species has been found in an urban environment. This discovery, in a heavily populated area and in close association with human dwellings, is a vivid reminder of the tremendous diversity of the Insecta and of the untold number of insect species that have yet to be discovered. Acknowledgment We thank G. Chow, L. Daniels, S. Garcia-Rubio & R. Raban for their assistance in the collection and processing of wood and the collection of wasps and beetles. We also thank D. Carver, Live Oak Structural Pest Control, Berkeley, California and S. Kala, Daly City, California for providing the infested wood. Dr. F. An¬ drews, California Department of Food & Agriculture, Sacramento, California, confirmed the identity of Hemicoelus gibbicollis. Dr. J. Luhman, Minnesota De¬ partment of Agriculture, St. Paul, Minnesota identified Odontocolon polymorphum and provided very useful information on the xoridine Ichneumonidae. Specimens of O. polymorphum and H. gibbicollis from this study were deposited in the University of Minnesota Insect Collection and specimens of H. gibbicollis were deposited in the California Academy of Sciences. The scanning electron micro¬ graphs were prepared by K Hampton, Department of Entomology, Kansas State University, Manhattan, Kansas. Figure 7 was taken by D. C. Blackford, University of Minnesota, Department of Entomology. We thank R. A. Wharton and an anon¬ ymous reviewer for their comments and suggestions on the manuscript. The work 16 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1) described in this paper was supported by California Department of Consumer Affairs, Structural Pest Control Board grant 849017-07 to VRL and SJS. Literature Cited Cadson, R. W. & B. D. Bunks. 1979. Family Ichneumonidae. pp. 315-740. In Krombein, K. V., P. D. Hurd Jr. & D. R. Smith (eds.). Catalog of Hymenoptera in America North of Mexico. Volume 1. Smithsonian Press, Washington D.C. Drooz, A. 1985. Insects of eastern forests. USDA Forest Service, Misc. Pub. No. 1426. Marsh, P. M. 1979. Family Braconidae. pp. 144-294. In Krombein, K. V., P. D. Hurd, Jr., D. R. Smith & B. D. Burks, (eds.). Catalog of Hymenoptera in America North of Mexico. Volume I. Smith¬ sonian Press, Washington, D.C. Marsh, P. M. 1982. Two new species of Heterospilus (Hymenoptera: Braconidae) from Mexico being introduced against the cotton boll weevil, Anthonomus grandis (Coleoptera: Curculionidae). Proc. Ent. Soc. Wash., 84: 849-854. Marsh, P. M. 1997. Subfamily Doryctinae. pp. 207-233. In Wharton, R. A., P. M. Marsh & M. J. Sharkey (eds.). Manual of the New World genera of the family Braconidae (Hymenoptera). Special Pub. Int. Soc. Hymenopterists No. 1. Marsh, P. M. & G. A. R. Melo. 1999. Biology and systematics of New World Heterospilus (Hyme¬ noptera: Braconidae) attacking Pemphredoninae (Hymenoptera: Sphecidae). J. Hym. Res., 8: 13-22. Shaw, S. R. 1995. Braconidae. pp. 431-463. In Hanson, P. & I. Gauld (eds.). The Hymenoptera of Costa Rica. Oxford University Press. Shaw, M. R. & T. Huddleston. 1991. Classification and biology of braconid wasps (Hymenoptera: Braconidae). Handbooks for the Identification of British Insects 7: 1-126. Simeone, J. B. 1962. Survey of wood-feeding Anobiidae in northeastern United States, including a study of temperature and humidity effects on egg development of Hadrobregmus carinatus (Say), pp. 326-335. 11th Int. Kongr. Entomol., Wien. Aug. 17-25 1960. Verh. Band II. Suomi, D. A. & R. A. Akre. 1992a. Characteristics of structures attacked by the wood-infesting beetle, Hemicoelus gibbicollis (Coleoptera: Anobiidae). J. Entomol. Soc. Brit. Columbia, 89: 63-70. Suomi, D. A. & R. A. Akre. 1992b. Control of the structure-infesting beetle Hemicoelus gibbicollis (Coleoptera: Anobiidae) with borates. J. Econ. Entomol., 85: 1188-1193. Suomi, D. A. & R. A. Akre. 1992c. Distribution of economically important, wood-infesting anobiid beetles in the Pacific Northwest. J. Entomol. Soc. Brit., Columbia, 89: 57-62. Suomi, D. A. & R. A. Akre. 1993a. Biological studies of Hemicoelus gibbicollis (LeConte)(Coleoptera: Anobiidae), a serious structural pest along the Pacific coast: adult and egg stages. Pan-Pacific Entomol., 69: 155-170. Suomi, D. A. & R. A. Akre. 1993b. Biological studies of Hemicoelus gibbicollis (LeConte)(Coleoptera: Anobiidae), a serious structural pest along the Pacific coast: larval and pupal stages. Pan-Pacific Entomol., 69: 221-235. Townes, H. M., S. Walley, L. Walkley, D. Habeck & G. Townes. 1960. Ichneumon flies of America north of Mexico. Vol. 2. Ephaltinae, Xoridinae, and Acaenitinae. USNM Bulletin No. 216, Part I, Washington, D.C. Williams, L. H., H. M. Barnes & H. O. Yates, III. 1979. Beetle ( Xyletinus peltatus) and parasite exit hole densities and beetle larval populations in southern pine floor joists. Environ. Entomol., 8: 300-303. Received 4 April 2001; Accepted 16 August 2001. PAN-PACIFIC ENTOMOLOGIST 78(1): 17-22, (2002) A NEW SPECIES OF YELICONES CAMERON (HYMENOPTERA: BRACONIDAE) FROM THAILAND Buntika Areekul 1 ’* & Donald L. J. Quicke 1 ’ 2 'Department of Biological Sciences, Imperial College at Silwood Park, Ascot, Berkshire, SL5 7PY, UK department of Entomology, The Natural History Museum, South Kensington, London, SW7 5BD, UK Abstract.—Yelicones siamensis Areekul & Quicke, NEW SPECIES is described and illustrated based on two adult females collected at light in Thailand. This wasp is the ninth species of Yelicones described from the East Palaearctic and Oriental regions. A modification to the key of Quicke et al. (1997: J. Nat. Hist. 31: 779-797) is included to differentiate Y. siamensis from similar species. Key Words.— Insecta, Hymenoptera, Braconidae, Yelicones, Thailand. Wasps of the genus Yelicones Cameron are solitary endoparasitoids of lepi- dopteran larvae, whose remains they mummify before pupating within the host (Quicke & Chishti 1997). For many years after its original description (Cameron 1887) the genus was known only from a handful of specimens from the New World (Shenefelt 1975, Quicke & Kruft 1995). However, over the last 20 years a number of new species have been described, extending the known range of Yelicones into the Indo-Australian, Afrotropical and Palaearctic regions. The ge¬ nus is now known to be widely distributed throughout the Old and New Worlds (Fischer 1961, 1962 [as Pectenopius Fischer]; Togashi 1980; Papp 1985, 1989, 1991, 1992; Belokobylskij 1993a, b; Quicke & Kruft 1995; Quicke et al. 1996, 1997, 1998; Quicke & Chishti 1997; Shaw 1998). In this paper a new species of Yelicones is described based on two female specimens from Thailand, the ninth for the East Palaearctic and Oriental regions (Quicke et al. 1997). It is being described because it has been used to generate DNA sequence data which will be published elsewhere as part of another study. Male morphology and biology are unknown. The genus Yelicones can be recog¬ nized using the keys of van Achterberg (1995), Chen and He (1997) or Shaw (1997). A brief diagnosis is provided below. Materials and Methods Two specimens were collected by light trapping in Chon Buri, Thailand and preserved in absolute alcohol. Three legs on one side of the body were taken from the paratype specimen for DNA sequencing and both specimens were then mounted for description and photography. Measurements were made with an eye¬ piece micrometer graticule. Terminology follows van Achterberg (1979, 1988) and Quicke et al. (1997). Genus Yelicones Cameron, 1887 Yelicones Cameron 1887: 387; van Achterberg, 1995: 147 (literature). Type spe¬ cies, Yelicones violaceipennis Cameron, designated by Viereck (1914). * Author for correspondence 18 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1) Figure 1. Yelicones siamensis NEW SPECIES, female holotype. Rhopalotoma Cameron 1911: 318. Type species, Rhopalotoma crassitarsis Cam¬ eron, monotypic. Pectenopius Fischer 1961: 156. Type species, Pectenopius paradoxus Fischer, original designation. Yelicones siamensis Areekul & Quicke, NEW SPECIES Types. —Holotype, female (Fig. 1); data: THAILAND. CHON BURI: Khao Kheow, 20—31 March 2001, D. L. J. Quicke and N. Laurenne, light trap; depos¬ ited: British Museum (Natural History). Paratype: same data as holotype, 1 fe¬ male; deposited Insect Collections of Chulalongkorn University, Bangkok, Thai¬ land. Description .—Female (holotype) Length. Body 4.5-5.0 mm, and fore wing 3.5 mm. (Fig. 1). Color. Yellow, antennae yellow basally, gradually brown on distal 0.4; wing veins dark brown, pterostigma basal 0.4 ivory, distal 0.6 dark brown (Fig. 1). Head. Antennae with 26 flagellomeres, terminal fla- gellomere pointed, approximately 2.3 X longer than wide; first flagellomere 1.1 X and 1.4X longer than the second and third respectively; first flagellomere 1.4X longer than wide; third flagellomere as long as wide; malar space unsculptured, length of malar space 0.04X height of eye; height of clypeus: inter-tentorial distance: tentorio-ocular distance = 1.0: 3.4: 0.8; clypeus slightly punctate, with long, dense setae; face with subtransverse carinae below the antennal sockets, punctate ventrally (Fig. 2), densely covered with long setae, with weak but distinct mid-longitudinal ridge (Fig. 3); height of eye: width of face: width of head = 1.0: 0.9: 1.7; length of face = 0.5X width of face; eyes glabrous; frons with sparse, long setae, impressed behind the antennal socket, mid-longitudinal ridge strongly developed, posteriorly with two curved transverse carinae; occiput and temples densely punctate; horizontal length of eye: horizontal length of head behind eye = 1.6: 1.0; post-ocellar length: trans- 2002 AREEKUL & QUICKE: A NEW YELICONES FROM THAILAND 19 Figures 2-6. Yelicones siamensis NEW SPECIES. Figure 2, front view of head. Figure 3, front and lateral aspect of head. Figure 4, mesoscutum. Figure 5, lateral view of prothorax. Figure 6, propodeum. 20 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1) Figures 7-8. Yelicones siamensis NEW SPECIES. Figure 7, fore wing. Figure 8, dorsal view of metasomal tergites 1-3. verse diameter of posterior ocellus: shortest distance between posterior ocellus and eye = 1.0: 2.0: 2.7; occipital carina nearly complete, absent for a small distance medially. Mesosoma. Shiny, densely punctured and setose, 1.8X longer than high; mesoscutum postero-medially with longitudinal groove¬ like impressions (Fig. 4); notauli weakly impressed throughout length of mesoscutum; scutellar sulcus with 7 carinae between the two outer ones; scutellum shiny and sparsely setose with small punctures; median area of metanotum medially without pit (Fig. 6); mesopleuron densely setose, with transverse carinae anteriorly, densely punctate posteriorly; precoxal suture weakly impressed, crenulate, upcurved posteriorly, impressed 0.8 length of mesopleuron (Fig. 5); propodeum strongly aerolate-rugose, antero- medially without a prominent U- or V-shaped carina (Fig. 6). Wings. Fore wing —length of veins SRI: 3SR: r = 3.3: 0.4: 1.0; vein 1-SR+M more or less straight; vein r arising 0.5 distance from base of pterostigma; length of veins 2-SR: 3-SR: r-m = 1.0: 1.0: 1.1; length of veins 2-SR+M: 2-M: m-cu = 1.0: 0.7: 0.7; length of veins 2-CU1: 3-CU1 = 2.4: 1.0; veins C+SC+R and 1-SR forming an angle of 60° (Fig. 7). Hind wing —length of veins lr-m: SC+R1 = 1.0: 1.8; vein 2-SC+R interstitial; vein SR posteriorly weak, more or less straight at apex; vein 2m-cu strongly postfur cal, length of vein 1M 4.4X vein 2M, vein 2m-cu more or less straight; marginal cell, basal cell and base of wing densely setose. Legs. Length of fore femur: tibia: tarsus = 1.0: 1.3: 1.0; fore femur 2.OX longer than maximum depth; fore tibia without mid-longitudinal ridge; hind femur 2.7X longer than maximum depth; length of hind femur: tibia: basitarsus = 1.8: 2.5: 1.0; hind basitarsus 3.4X longer than maximally depth. 2002 AREEKUL & QUICKE: A NEW YELICONES FROM THAILAND 21 Metasoma. Metasomal tergites shiny, first and second tergite with sparse setae, 3rd-8th tergites mod¬ erately setose; first and second tergite with punctate-rugulose sculpture; first metasomal tergite 1.3X wider than medially long, dorsal carina weakly impressed, uniting before the level of spiracles; second metasomal tergite 2.4X wider than medially long, without smooth triangular area anteriorly and with¬ out mid-longitudinal carina; second suture narrow, smooth; third metasomal tergite 2.4X wider than medially long; third tergite anteriorly finely punctate, posteriorly smooth (Fig. 8); 4th-6th metasomal tergites smooth. Tip of ovipositor pale. Diagnosis.—Yelicones siamensis Areekul & Quicke keys out to couplet 8 using the key to East Palaearctic and Oriental species of Yelicones (Quicke et al. 1997). It can be distinguished from Y. flavus Chen and Quicke by the following char¬ acters: face not transversely imbricate; mesoscutum postero-medially rugose; fore wing vein 1-SR+M more or less straight not sinuous; hind wing vein 2-SC+R interstitial not transverse; dorsal carinae of first metasomal tergite uniting anteri¬ orly not near mid-length, without median carina; second metasomal tergite ante- rior-medially without smooth triangular area and mid-longitudinal carina; color yellow, wings without distinct pattern of brownish blotches, pterostigma bicolored not unicolorous. Modification to the key to the species of Yelicones of the East Palaearctic and Oriental region (Quicke et al. 1997) to accommodate the new species. 8(7). Mesoscutum with line of notauli piceous on anterior two thirds and usually with a dark line connecting them in front of the posterior margin; propleuron and fore coxae whitish. Y. koreanus Papp Mesoscutum uniformly yellow-brown; propleuron and fore coxae brownish yellow . 8a 8a(8). Forewing with distinct pattern of brownish blotches; pterostigma uni¬ formly, pale brown; antennae with 33 flagellomeres; second meta¬ somal tergite antero-medially with smooth triangular area . . Y. flavus Chen and Quicke Forewing without distinct pattern of brownish blotches; pterostigma bicolored, whitish basally, brown distally; antennae with fewer than 33 flagellomeres; second metasomal tergite antero-medially without smooth triangular area . Y. siamensis NEW SPECIES Variation in Females .—Body length 4.5-6.0 mm; antennae with 26—28 flagel¬ lomeres; mesoscutum postero-medially with punctate-rugose sculpture or longi¬ tudinal groove-like impressions; scutellar sulcus with 6—7 carinae between the outer ones. Male .—Unknown. Distribution .—Thailand. Etymology .—The name is derived from the old name for Thailand. Material Examined .—See Types. Acknowledgment We thank Angoon Lewvanich and Sura Pimpasalee for help in collecting the specimens, and Apidet Singhaseni for permitting collecting at Khao Kheow Zoo. Robert Belshaw, Gavin Broad and David Orme reviewed the manuscript and provided helpful comments. Andrew Polaszek helped with Automontage imaging facilities. 22 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1) Literature Cited Achterberg, C. van. 1979. A revision of the subfamily Zelinae auct. (Hymenoptera, Braconidae). Tijdschr. Ent., 122: 241-479. Achterberg, C. van. 1988. Revision of the subfamily Blacinae Foerster (Hymenoptera, Braconidae). Zool. Verh., Leiden, 249: 1-324. Achterberg, C. van. 1995. Generic revision of the subfamily Betylobraconinae (Hymenoptera: Bra¬ conidae) and other groups with modified fore tarsus. Zool. Verh., Leiden, 298: 1-242. Belokobylskij, S.A. 1993a. New taxonomic data on the braconid fauna (Hymenoptera, Braconidae) of Vietnam. Russian Entomol. J., 2: 37-67. Belokobylskij, S.A. 1993b. Contribution to the taxonomy of Braconidae (Hymenoptera) of the Russian Far East. Russian Entomol. J., 2: 87-103. Cameron, P. 1887. Family Braconidae, Biologia Centrali-Americana. Hymenoptera, 1: 312-419. Chen, X. & J. He. 1997. Revision of the subfamily Rogadinae (Hymenoptera: Braconidae) from China. Zool. Verh., Leiden, 308: 1-187. Fischer, M. 1961. Zwei neue Opiinen Gattungen (Hym., Braconidae). Annin naturh. Mus. Wien, 64: 154-158. Fischer, M. 1962. Die Opiinae des Museo Civico di Storia Naturale in Genua (Hymenoptera, Bracon¬ idae). Annali. Mus. Civ. Stor. Nat. Giacomo Doria, 73: 71-97. Papp, J. 1985. Braconidae (Hymenoptera) from Korea, 7. Acta Zool. Hung., 31: 341-365. Papp, J. 1989. A contribution to the braconid fauna of Israel. Israel J. Ent., 22: 45-59. Papp, J. 1991. New braconid wasps (Hymenoptera: Braconidae) in the Hungarian Natural History Museum, 2. Annls hist.-nat. Mus. Natn. hung., 83: 145-167. Papp, J. 1992. New braconid wasps (Hymenoptera: Braconidae) in the Hungarian Natural History Museum, 3. Annls hist.-nat. Mus. Natn. hung., 84: 129-160. Quicke, D. L. J. & R. A. Kruft. 1995. Species of Yelicones (Hymenoptera: Braconidae: Rogadinae) in North America with descriptions of two new species. Ann. Entomol. Soc. Am., 88: 129— 138. Quicke, D. L. J., M. J. K. Chishti, & H. H. Basibuyuk. 1996. A revision of the Yelicones species (Hymenoptera: Braconidae: Rogadinae) from Central America, with descriptions of sixteen new species. Zool. Meded. Leiden, 70: 17-61. Quicke, D. L.J. & M.J.K. Chishti. 1997. A revision of the Yelicones species (Hymenoptera: Bra¬ conidae: Rogadinae) from Africa and the Arabian Peninsula, with descriptions of four new species. African Entomol., 5: 77-91. Quicke, D. L. J., M. J. K. Chishti, X. Chen, & R. A. Kruft. 1997. Revision of Yelicones (Hymenoptera: Braconidae: Rogadinae) from the East Palaearctic and Oriental regions with description of four new species. J. Nat. Hist., 31: 779-797. Quicke, D. L. J., A. D. Austin, & M. J. K. Chishti. 1998. Revision of Yelicones (Hymenoptera: Braconidae: Rogadinae) from the Australasian region. Invert. Taxon., 12: 897-928. Shaw, M. R. 1998. The surprising discovery of the genus Yelicones Cameron (Hymenoptera: Bracon¬ idae) in Western Europe. Br. J. Ent. Nat. Hist., 11: 15-16. Shaw, S. R. 1997. Subfamily Rogadinae s.s. pp. 403-412. In Wharton, R. A., Marsh, P. M. & Sharkey, M. J. (eds.). Manual of the New World genera of the family Braconidae (Hymenoptera). Special publication of the International Society of Hymenopterists, Number 1, Washington, D.C. Shenefelt, R. D. 1975. Braconidae 8, Exothecinae and Rogadinae. Part 12. pp. 1115-1262. In J. van der Vecht & R. D. Shenefelt (eds.). Hymenoptera Catalogus, Junk, The Hague. Togashi, I. 1980. Discovery of the genus Yelicones Cameron (Hymenoptera, Braconidae) from Japan. Kontyu, 48: 571-520. Received 11 July 2001; Accepted 23 November 2001. PAN-PACIFIC ENTOMOLOGIST 78(1): 23-33, (2002) THE SPIDER FAUNA ASSOCIATED WITH LITTER UNDER WOODRAT MIDDENS IN SOUTHERN CALIFORNIA (ARACHNIDA: ARANEAE) Richard S. Vetter & Thomas R. Prentice Department of Entomology, University of California, Riverside, California 92521 Abstract .—Litter from under wood rat ( Neotoma sp.) middens from southern California, was sampled, primarily between 1977-1985, in search of latridiid beetles. As an ancillary by-product, spiders were collected, separated and labeled with collection data but most have remained un¬ determined until now. In this paper, we determine the spider species associated with the rat middens, predominantly from five southern Californian counties, ranging in elevation from 200 to 1900 m. This study yielded 316 specimens representing 42 species, 34 genera and 20 families of spiders. The family Linyphiidae was represented here with the greatest number of genera, species and specimens. The most frequently collected species, Tapinocyba dietrichi Crosby & Bishop (Linyphiidae), contributed 37.3% of the specimen total. Spirembolus erratus Millidge (Linyphiidae) and Zanomys californica (Banks) (Amaurobiidae) were the second and third most predominant species, contributing 16.5% and 5.1%, respectively, of the specimen total. Many of the specimens identified in this study are members of minute species (1-3 mm), which can easily be overlooked unless samples are carefully scrutinized. Some of the more interesting and un¬ common species include Trogloneta paradoxum Gertsch (Mysmenidae), Gertschanapis shantzi (Gertsch) (Anapidae), several Zanomys spp., and members from the families Hahniidae, Capon- iidae and Oonopidae. Specimens collected here will contribute to the description of three new species (2 amaurobiids, 1 linyphiid) and the male of a second linyphiid species. Key Words .—Arachnida, Neotoma, rat middens, species list, Araneae, spiders. Woodrats of the genus Neotoma create large middens composed of vegetation and other materials culled from the surrounding environment (English 1923). Mid¬ dens can reach massive sizes, primarily through occupancy by successive gener¬ ations with each occupant adding material. In so doing, the rats create a micro¬ habitat that often differs from the immediate surrounding area in various biotic and abiotic aspects (Vestal 1938, Linsdale & Tevis 1951). Dimensions of 301 middens in northern California averaged 118 cm in height and 152 cm in basal diameter; the average volume of 572 middens was 0.713 m 3 (Vestal 1938). Active rat middens are strewn with fresh food cuttings and copious fecal pellets (Linsdale & Tevis 1951). Considering their structural features, middens provide ideal har¬ borage for a plethora of animals including spiders (Linsdale & Tevis 1951). Fossil midden remains have offered a wealth of data on the historical plant and arthropod components of the regions of occupancy (Hall et al. 1989, Elias et al. 1992, Clark & Sankey 1999) although, in these studies, arachnids were represented solely by the hard-bodied ticks, scorpions and pseudoscorpions. From 1977 to 1985, Ken Cooper of the University of California-Riverside (UCR) harvested arthropod inhabitants from the litter under Neotoma nests in search of beetles of the family Latridiidae. This litter was collected primarily from five southern Californian counties in a variety of habitats ranging from the Los Angeles-San Diego basins (ca. 300 m elevation) to the surrounding inland and coastal mountains (up to 1900 m). An ancillary by-product of the collections resulted in an accumulation of spiders which, until now, have languished in the 24 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1) UCR Entomology museum, mostly as undetermined specimens. We have identi¬ fied these spiders, the majority of which were less than 3 mm in body length, and present our findings here. Minute spiders pose several problems that impede identification. Size alone creates difficulties in manipulation of the specimens as well as in distinguishing the pertinent diagnostic characters. Most of the minute species determined in this study are members of the Linyphiidae which contains a cumbersome number of genera and species that are often very difficult to identify. Relatively few generic keys and current taxonomic revisions are available for this family; those that do exist often include only regional fauna. One of the most current and definitive identification guides for North American spiders (Roth 1993) presents keys to genera for every family except the Linyphiidae which are simply listed alpha¬ betically by genus for both the linyphiine (45 genera) and erigonine (117 genera) spiders. What we hope to accomplish here is to assist future investigators of Neotoma biology, add to our knowledge of Californian linyphiid distribution, and encourage eager arachnologists to pursue taxonomic studies on the little-known genera. Materials and Methods All information regarding the history of the spiders from the Neotoma middens was derived from personal communications with the collector (K. Cooper), the processor (F. Andrews) of the midden material, and the former curator (S. From- mer) of the Entomology museum collection (all former associates of UCR). Ap¬ proximately 100 Neotoma (mostly N. fuscipes with a few N. lepida ) middens were sampled. The litter under each nest was shoveled into canvas bags, brought back to the laboratory and processed in approximately 12 gallon loads, for up to 10 days, in Berlese funnels until no additional arthropods were collected. Although most of the spiders were part of the undetermined UCR Entomology Research Museum spider collection, it was not known how many spiders were actually identified and incorporated as determined species. Consequently, we searched the entire general spider collection (> 1700 vials) excepting the orb-weaver families Araneidae, Tetragnathidae, Uloboridae, the adults of which we felt would not likely be using the litter under rat middens as refugia. As a result of our search, we believe that this study represents an accurate inventory of the material recov¬ ered by the researchers. Spiders were present in samples from at least 50 Neotoma middens collected by Cooper in southern California. Also included are data from one midden from San Bernardino County, California collected by Andrews, Hardy & Eichlin in 1985. Elevations not present on the collection labels were approximated to the nearest 25 meters with topographic maps or through recent communications with K. Cooper and should be accurate to within 50 meters. Spiders are listed in Table 1 in decreasing frequency by family to show the importance of their ecological association with Neotoma nests and then alphabetically by genus within each family; families with equal numbers of specimens are grouped alphabetically. All specimens have been incorporated into the reference collection of the UCR Entomology Research Museum. Table 1. List of the spiders identified from Neotoma rat nests. Family/Genus Species Male Fem Imm County Date Elevation (meters) Locale Linyphiidae Ceratinops inflatus 2 2 Riverside 14-Jan-83 1800 0.8 km E of James Reserve, San Jacinto Mts Ceratinopsis interventa 1 Kern 23-Sep-81 800 Red Rock Cyn campsite “Ceratinopsis” palomara 1 Riverside 30-Apr-81 750 Bautista Cyn, 27.4 km SE of Valle Vista palomara 2 Riverside 29-Apr-82 300 Clinton Keith Rd, W Murrieta near entrance to cyn palomara 1 5 Riverside 14-Jan-83 1825 0.8 km E of James Reserve, San Jacinto Mts Linyphantes aeronautica 1 Riverside 27-Dec-79 400 Whitewater Cyn aeronautica 1 Riverside 25-Mar-82 500 Hemet, Motte Reserve laguna 1 San Diego 13-Mar-81 1075 San Felipe microps 1 Riverside 25-Mar-82 500 Hemet, Motte Reserve Spirembolus demonologicus 2 San Diego 13-Mar-81 1075 San Felipe erratus 1 Riverside 13-Jan-77 1100 Bautista Cyn, 27.4 km SE of Valle Vista erratus 1 Riverside 13-Feb-77 600 Harford Preserve, Gavilan Hills erratus 9 Riverside 20-Mar-77 600 Gavilan Hills erratus 10 14 1 Riverside 27-Dec-79 400 Whitewater Cyn erratus 1 Riverside 18-Nov-83 200 Vail Lake erratus 1 1 Riverside 18-Nov-83 200 Vail Lake erratus 1 Riverside 18-Nov-83 200 Vail Lake “nest 2” erratus 1 9 San Bernardino 26-Feb-77 525 E of Mentone erratus 1 San Bernardino 13-May-82 700 Baldy Mesa, Phelan & Transmission Line Rd erratus 1 Santa Barbara 11-Aug-83 1075 Figueroa Mt Rd hibernus 1 Riverside 24-Aug-83 950 El Carrizo Oaks hibernus 1 San Diego 13-Mar-81 1075 San Felipe redondo 1 San Bernardino 2-May-68 325 1.6 km E of Summit on Rt 138 Tapinocyba dietrichi 8 7 Los Angeles 11-Nov-80 1000 0.8 km N of Jet. Vasquez & Bouquet Cyns dietrichi 6 6 Los Angeles 14-Aug-81 1000 Bouquet Cyn nr. SE arm Bouquet Reservoir dietrichi 1 Los Angeles 20-Apr-82 1000 Bouquet Cyn Rd at E. end Bouquet Cyn Reservoir dietrichi 6 Los Angeles 20-Apr-82 1000 3.2 km E toward Palmdale on Bouquet Cyn Rd dietrichi 1 Los Angeles 20-Apr-82 1000 Bouquet Cyn nr. entrance Vasquez Rd dietrichi 2 24 Riverside 14-Jan-83 1825 James Reserve Lk Fulmor dietrichi 1 1 Riverside 4-Jun-83 750 7 km SE of Sage on R3 dietrichi 1 Riverside 24-Aug-83 950 El Carrizo Oaks 2002 VETTER & PRENTICE: SPIDER FAUNA OF WOOD RAT MIDDENS Table 1. Continued. to o\ Family/Genus Species Male Fem Imm County Date Elevation (meters) Locale dietrichi 3 San Bernardino 2-May-68 325 1.6 km E of Summit on Rt 138 dietrichi 5 5 San Bernardino 3-Jun-81 975 18.6 km from Hwy 15 on Rt 138 nr Little Horse- thief Ranch dietrichi 3 San Bernardino 1-May-85 1100 Summit Vly under cottonwood dietrichi 4 16 San Diego 25-Jul-79 1250 William Heise Co. Park, Pine Hills S of Julian dietrichi 1 2 San Diego 13-Mar-81 1075 San Felipe dietrichi 1 1 San Diego 13-Mar-81 725 San Felipe @ S2 dietrichi 5 8 Santa Barbara 1 l-Aug-83 1075 Figueroa Mt Rd, 1.6 km W Rngr Sta sp. #1 1 Riverside 17-Jun-77 1300 Covington Flat sp. #1 7 Riverside 9-Dec-81 1325 Joshua Tree Natl Pk, Hidden Valley sp. #1 4 San Bernardino 21-May-82 975 6.4 km N of Yucca Valley Amaurobiidae Amciurobius latescens 1 San Bernardino 26-Feb-77 525 E of Mentone Zanomys calif ornica 1 Los Angeles 11-Nov-80 1000 0.8 km N of Jet. Vasquez & Bouquet Cyns californica 1 Riverside 13-Jan-77 1100 Bautista Cyn, 27.4 km SE of Valle Vista calif ornica 4 Riverside 14-Jan-83 1825 James Reserve Lk Fulmor californica 3 Riverside 14-Jan-83 1825 0.8 km E of James Reserve, San Jacinto Mts californica 1 Riverside 18-Nov-83 200 Vail Lake “nest 2” californica 1 San Bernardino 2-May-68 325 1.6 km E of Summit on Rt 138 californica 1 San Bernardino 3-Jun-81 975 18.6 km from Hwy 15 on Rt 138 nr Little Horse- thief Ranch californica 1 San Bernardino 1-May-85 1100 Summit Valley californica 1 San Bernardino 14-Apr-85 1850 Wrightwood californica 1 San Diego 25-Jul-79 1250 William Heise Co. Park, Pine Hills S. Julian californica 1 San Diego 25-Apr-80 1075 San Felipe ochra 1 San Bernardino 15-May-7 8 1900 Bums Cyn above Rimrock ochra 2 San Bernardino 13-May-82 700 Baldy Mesa, Phelan & Transmission Line Rd ultima 3 Santa Barbara 1 l-Aug-83 1075 Figueroa Mt Rd sp. #1 4 7 Riverside 9-Dec-81 950 Joshua Tree Natl Pk THE PAN-PACIFIC ENTOMOLOGIST Vol. Table 1. Continued. Family/Genus Species Male Fem Imm County Date Elevation (meters) Locale Genus #1 1 Riverside 7-Dec-77 unknown Boyd Desert Center, Coyote Creek Anapidae Gertschcincipis shantzi 3 2 2 Los Angeles 17-Nov-80 1000 Bouquet Reservoir shantzi 3 6 Los Angeles 14-Aug-81 1000 Bouquet Cyn nr. SE arm Bouquet Reservoir Oonopidae Oonops sp. #1 1 Riverside 5-Nov-80 400 Whitewater Cyn Orchestina moaba 1 Riverside 13-Feb-77 600 Gavilan Hills moaba 1 1 Riverside 25-Dec-80 325 Box Springs Mts nr UCR moaba 2 Riverside 29-Nov-81 450 3.2 km S of Valle Vista on Rt 74, under live oak moaba 1 Riverside 9-Dec-81 950 Joshua Tree Natl Pk moaba 1 Riverside 18-Nov-83 200 Vail Lake Scaphiella hespera 1 Riverside 25-Dec-80 325 Box Springs Mts nr UCR hespera 1 1 Riverside 9-Dec-81 950 Joshua Tree Natl Pk Dictynidae Dictyna cholla 1 Riverside 27-Dec-79 400 Whitewater Cyn cholla 1 Riverside 25-Dec-80 325 Box Springs Mts nr UCR Tivyna moaba 1 Riverside 5-Nov-80 400 Whitewater Cyn moaba 1 San Bernardino 25-May-82 550 Twentynine Palms Yorima angelica 1 San Bernardino 26-Feb-77 525 E of Mentone angelica 1 Riverside 29-Apr-82 325 Clinton Keith Rd, W of Murrieta near entrance to cyn imm 1 Los Angeles 17-Nov-80 1000 Bouquet Reservoir Scytodidae Scytodes undescr. sp. 3 Riverside 13-Nov-79 400 Whitewater Cyn undescr. sp. 4 Riverside 5-Nov-80 400 Whitewater Cyn to 2002 VETTER & PRENTICE: SPIDER FAUNA OF WOOD RAT MIDDENS Table 1. Continued. to oo Family/Genus Species Male Fem Imm County Date Elevation (meters) Locale Caponiidae Orthonops imm 1 Imperial 25-Mar-80 100 Indian wash, 30 km SE of Glamis imm 3 Los Angeles 17-Nov-80 1000 Bouquet Reservoir imm 1 Riverside 25-Mar-82 500 Hemet, Motte Reserve zebra 1 Riverside 13-Feb-77 600 Gavilan Hills Corinnidae Trachelas pacificus 2 Riverside 18-Nov-83 200 Vail Lake pacificus 1 San Bernardino 28-Sep-83 425 Afton Cyn pacificus 1 Santa Barbara 11-Aug-83 1075 Figueroa Mt Rd, 1.6 km W of Rngr Sta Gnaphosidae Herpyllus propinquus 1 Riverside 17-Feb-7 8 1100 Bautista Cyn propinquus 1 San Diego 13-Mar-81 725 San Felipe @ S2 Micaria imm 1 Riverside 25-Dec-80 325 Box Springs Mts nr UCR pasadena 1 San Bernardino 1-May-85 1100 Summit Vly under cottonwood Hahniidae Hahnia sanjuanensis 2 Riverside 24-Aug-83 950 El Carrizo Oaks sanjuanensis 1 1 San Diego 25-Apr-80 1075 San Felipe Liocranidae Phrurotimpus mateonus 1 1 Riverside 18-Nov-83 200 Vail Lake Scotinella kastoni 1 San Diego 25-Apr-80 1075 San Felipe Cybaeidae Cybaeota nan a 1 Riverside 15-Nov-81 1825 James Reserve Lk Fulmor nana 1 Riverside 14-Jan-83 1800 0.8 km E of James Reserve, San Jacinto Mts Mysmenidae Trogloneta paradoxum 2 Los Angeles 14-Aug-81 1000 Bouquet Cyn nr. SE arm Bouquet Reservoir THE PAN-PACIFIC ENTOMOLOGIST Vol. Table 1. Continued. Family/Genus Species Male Fem Imm County Date Elevation (meters) Locale Plectreuridae Plectreurys deserta 1 Riverside 9-Dec-81 950 Joshua Tree Natl Pk imm 1 Riverside 22-May-80 275 Corn Springs, SW Desert Ctr Salticidae Neon pixii 1 Riverside 20-Mar-77 600 Gavilan Hills Sitticus dorsatus 1 Riverside 25-Dec-80 325 Box Springs Mts nr UCR Theridiidae Euryopis spinigera 1 Riverside 9-Dec-81 1350 Joshua Tree Natl Pk, Jumbo Rocks Cmpgrd spinigera 1 Riverside 9-Dec-81 950 Joshua Tree Natl Pk Cyrtaucheniidae Aptostichus imm 1 Riverside 10-Jan-81 325 Riverside Filistatidae Kukulcania utahana 1 Riverside 10-Jan-81 325 Riverside Lycosidae Pardosa California 1 San Bernardino 5-Jun-79 950 4.8 km S of Hesperia Pholcidae Psilochorus acanthus 1 Los Angeles 14-Aug-81 1000 Bouquet Cyn nr. SE arm Bouquet Reservoir TOTALS 88 206 22 to vo 2002 VETTER & PRENTICE: SPIDER FAUNA OF WOOD RAT MIDDENS 30 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1) Results and Discussion A total of 316 spiders representing 20 families, 34 genera and 42 species were identified in this study of Neotoma midden litter (Table 1). The linyphiid family provided the greatest number of genera (6), species (12) and overall specimens (204), garnering 65.2% of the specimen total. Tapinocyba dietrichi Crosby & Bishop (Linyphiidae) was the most frequently collected species, contributing 118 specimens (37.3% of the specimen total). T. dietrichi was found in midden debris in five California counties (Los Angeles, Riverside, San Bernardino, San Diego, Santa Barbara), ranging in approximate elevation from 325 to 1825 m but pre¬ dominantly found at higher elevations. From the 14 middens in which it was found, the average number of T. dietrichi per midden was 8.4, with a range of 1—26 specimens. The other Tapinocyba species is believed to be undescribed. The linyphiid genus Spirembolus contributed four species and the second most com¬ monly collected species, S. erratus Millidge (16.5% of specimen total). The amau- robiid genus, Zanomys, also contributed four species and the third most prevalent species, Z. californica (Banks) (5.1% of specimen total) with one new species to be described as a result of this study. Although we expected that the linyphiids would comprise the bulk of midden spider inhabitants, we were somewhat surprised to find Tapinocyba dietrichi in southern California, let alone its overwhelming contribution to the study’s spec¬ imen total. To our knowledge, the only other California specimens of this species were taken in Alpine and Alameda Counties in central California (Crosby & Bishop 1933, Boe, unpublished data). This species was not discovered during faunal studies of coastal sage scrub in either San Diego or Riverside Counties (Prentice et al. 1998, 2001) nor has it been found in montane oak ( Quercus spp.) leaf duff in southern California (Vetter, unpublished data). Spirembolus erratus was previously known only from samples collected in sycamore litter (Millidge 1980), grass litter in coastal sage scrub and in oak leaf litter (Riverside County) (Prentice & Vetter, unpublished data). In this study, S. erratus was collected most often at elevations of 200 to 700 m with two specimens taken near 1100 m. Millidge (1980) states that virtually nothing is known of the natural history of Spirembolus species in general, including our other listed Spir¬ embolus species, S. redondo (Chamberlin & Ivie), S. hibernus Millidge, and S. demonologicus (Crosby). Zanomys californica has previously been collected from Neotoma middens by J. Linsdale at the Hastings Reserve in central California (specimens at California Academy of Sciences, examined). In our study, Z. californica was more prevalent at high elevations (predominantly 1000—1850 m). Leech (1972) records both Z. californica and Z. ochra Leech (holotype) from dry leaf duff, the latter also taken from a rat midden in Juab County, Utah. Z. californica is a common inhabitant of montane oak leaf duff (moist or dry) in southern California (Vetter, unpublished data). In addition, the concomitant collection of several females of “ Ceratinopsis ” palomara Chamberlin along with the male (which is undescribed) in rat nest litter in the San Jacinto Mountains in corroboration with similar recent contempora¬ neous collections of both sexes in oak leaf duff in the same mountain range (Vetter 2002 VETTER & PRENTICE: SPIDER FAUNA OF WOOD RAT MIDDENS 31 & Prentice, unpublished data) lead to the conclusion that this species does not belong in the genus. Members of many of the spider families identified from our southern Califor¬ nian pack rat midden study are commonly found within (and possibly restricted to) the leaf litter strata. In several ecosystems, especially those within the desert, Neotoma middens represent a drastic vegetative change from the immediate sur¬ roundings and may indeed be “oases” of increased survival potential. Occupied middens contain fresh, nocturnally-harvested vegetation and copious amounts of fecal pellets (Vestal 1938, Linsdale & Tevis 1951), both of which may attract and support potential spider prey in the middens. Active and abandoned middens alike provide refugia (for the local inhabitants) that are structurally more stable and offer protection from environmental extremes than many of the other niches with¬ in the surrounding environment. Vorhies & Taylor (1945) showed that over a year’s time, temperatures inside an Arizona midden were consistently 11 to 17° C lower than soil surface temperatures. Platnick (1995) states that rat middens may provide a “vestige refuge” for the araneophagous Orthonops spp. due to habitat destruction in various regions of southern California. However, in an undisturbed site in the Colorado Desert south of Joshua Tree National Park (400 m elevation), the caponiids Orthonops iceno- glei Platnick and Tarsonops sp. were frequently collected from pitfall traps but not from the remains of a Neotoma midden at the site (Vetter, unpublished data). It may be that, in undisturbed areas, caponiids are not restricted to rat middens nor to leaf duff. Several of the species listed here were previously taken from Neotoma middens during various studies. Gertsch (1960) discovered both Gertschanapis shantzi (Gertsch) and Trogloneta paradoxum Gertsch while Platnick & Forster (1990) recorded only G. shantzi during their examination of Linsdale’s wood rat material from the Hastings Reserve. Ryckman & Lee (1956) sampled Neotoma middens for reduviid bugs ( Triatoma spp.), primarily in Riverside and San Bernardino Counties and recorded the arachnids found therein. However, for the spiders, they failed to provide frequency data which would have allowed percent species com¬ position comparisons with our study. Species listed in both their study and ours include the following: Zanomys californica, Trachelas pacificus Chamberlin & Ivie, Dictyna cholla Gertsch & Davis, Herpyllus propinquus (Keyserling), Liny- phantes laguna Chamberlin & Ivie, Pardosa californica Keyserling, and Orches- tina moaba Chamberlin & Ivie. Assuming that their determinations were based on mature specimens, the average size of the species sampled was larger than that of the species that we examined which is probably due to sampling differ¬ ences of whole nests by Ryckman and Lee in comparison to litter under the nests by Cooper. Vestal (1938) observed general aspects of Neotoma and their nests in Berkeley, California and although most of his article focused on Neotoma behavior, he did mention some arthropod associates. The only arachnids listed are mites ( Histios - toma sp.) and a pseudoscorpion (. Apocheiridium fumeroides ). Walters and Roth ( 1950 ) sampled 30 nests in Oregon, recording a myriad of arthropod inhabitants, but list only two genera and four families of spiders, Orchestina (Oonopidae), Calymmaria (Hahniidae), Linyphiidae, and Lycosidae. Linsdale and Tevis (1951) allocated 80 pages of their comprehensive book to animal associates of Neotoma 32 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1) nests at the Hastings Reserve in central California. However, they merely state that “spiders occur everywhere within the house except in the used nest” (“house” meaning the structure and “used” meaning the occupied portion of the structure). Vorhies & Taylor (1940) investigated middens of N. albigula albigula in Arizona and found that, within 100 nests, opilionids were the most common arachnid encountered (74% of the nests), followed by miscellaneous spiders (46%), black widow spiders (12%), scorpions (6%) and one tarantula (1%). In our work here, we have documented the partial diversity of the Araneae in Neotoma middens. Members of many of the families that were sampled here are infrequently collected, possibly because of their rarity, secretive habits, and/or very small size. Three of the species on our list are believed to be previously undescribed taxa, one belonging to an undescribed genus in the family Amauro- biidae (D. Ubick, personal communication). To our knowledge, two of the listed linyphiid species, Ceratinops inflatus (Emerton) and Ceratinopsis interventa Chamberlin, constitute new records for the state of California. Collecting material from underneath wood rat middens should provide a rewarding experience for the interested arachnologist by producing spiders that are rarely seen or that possess bizarre somatic features (such as Trogloneta and Gertschanapis ) and, thus, would be a fruitful challenge. Acknowledgment We thank K. Cooper for the tremendous effort spent in collecting the Neotoma nests, and F. Andrews and S. Frommer for providing information regarding the history of the spiders used in this collection. This project was funded in part by Humbug Mountain Engineering Services P-62 (RSV). Literature Cited Clark, W. H. & J. T. Sankey. 1999. Late Holocene Sonoran desert arthropod remains from a packrat midden, Catavina, Baja California Norte, Mexico. Pan-Pac. Ent., 75: 183-199. Crosby, C. R. & S. C. Bishop. 1933. American spiders: erigoneae, males with cephalic pits. Ann. Entomol. Soc. Am., 26: 105-182. Elias, S. A., J. I. Mead & L. D. Agenbroad. 1992. Late Quaternary arthropods from the Colorado plateau, Arizona and Utah. Great Basin Natur., 52: 59-67. English, P. F. 1923. The dusky-footed wood rat (Neotoma fuscipes). J. Mammal., 4: 1-9. Gertsch, W. J. 1960. Descriptions of American spiders of the family Symphytognathidae. Amer. Mus. Novit., 1981: 1-40. Hall, W. E., C. A. Olson & T. R. Van Devender. 1989. Late Quaternary and Modern arthropods from the Ajo Mountains of Southwestern Arizona. Pan-Pac. Entomol., 65: 322-347. Leech, R. 1972. A revision of the nearctic Amaurobiidae (Arachnida: Araneida). Mem. Entomol. Soc. Can., 84: 1-182. Linsdale, J. M. & L. P. Tevis, Jr. 1951. The dusky-footed wood rat. Univ. Calif. Press, Berkeley, California, 664 pp. Millidge, A. F. 1980. The erigonine spiders of North America. Part 2. The genus Spirembolus Cham¬ berlin (Araneae: Linyphiidae). J. Arachnol., 8: 109-158. Platnick, N. I. 1995. A revision of the spider genus Orthonops (Araneae, Caponiidae). Amer. Mus. Novit., 3150: 1-18. Platnick, N. I. & R. R. Forster. 1990. On the spider family Anapidae (Araneae, Araneoidea) in the United States. J. New York Entomol. Soc., 98: 108-112. Prentice, T. R., J. C. Burger, W. R. Icenogle & R. A. Redak. 1998. Spiders from Diegan coastal sage scrub (Arachnida: Araneae). Pan-Pac. Entomol., 74: 181-202. Prentice, T. R., J. C. Burger, W. R. Icenogle & R. A. Redak. 2001. Spiders from Riversidian coastal 2002 VETTER & PRENTICE: SPIDER FAUNA OF WOOD RAT MIDDENS 33 sage scrub with comparisons to Diegan scrub fauna (Arachnida: Araneae). Pan-Pac. Entomol., 77: 90-122. Roth, V. D. 1993. The Spider Genera of North America (3rd ed.). Amer. Arachnological Society, Gainesville, Florida, 203 pp. Ryckman, R. E. & R. D. Lee. 1956. Spiders and phalangids associated with mammals ( Citellus and Neotoma) in southwestern United States and northern Mexico. Ann. Entomol. Soc. Amer., 49: 406-409. Vestal, E. H. 1938. Biotic relations of the wood rat ( Neotoma fuscipes) in the Berkeley Hills. J. Mammal., 19: 1-36. Vorhies, C. T. & W. P. Taylor. 1940. Life history and ecology of the white-throated wood rat, Neotoma albigula albigula Hartley, in relation to grazing in Arizona. Univ. Arizona Agric. Exp. Sta. Tech. Bull., 86: 455-529. Vorhies, C. T. & W. P. Taylor. 1945. Water requirements of desert animals in the Southwest. Univ. Arizona Agric. Exp. Sta. Tech. Bull., 107: 487-525. Walters, R. D & V. D. Roth. 1950. Faunal nest study of the woodrat, Neotoma fuscipes monochroura Rhoads. J. Mammal., 31: 290-292. Received 30 June 2000; Accepted 4 September 2001. PAN-PACIFIC ENTOMOLOGIST 78(1): 34-42, (2002) REDESCRIPTION OF GONATOCERUS ATRICLAVUS GIRAULT (HYMENOPTERA: MYMARIDAE), WITH NOTES ON OTHER EGG PARASITOIDS OF SHARPSHOOTERS (HOMOPTERA: CICADELLIDAE: PROCONIINI) IN NORTHEASTERN MEXICO Serguei V. Triapitsyn 1 , Larry G. Bezark 2 , & David J. W. Morgan 2 department of Entomology, University of California, Riverside, California 92521 integrated Pest Control Branch, California Department of Food and Agriculture, Sacramento, California 95814 Abstract.— The mymarid wasp Gonatocerus atriclavus Girault, NEW STATUS, described orig¬ inally from Trinidad as a variety of G. triguttatus Girault, is redescribed and illustrated based on specimens reared from an egg mass of the sharpshooter Oncometopia clarior (Walker), col¬ lected in Ciudad Victoria, Tamaulipas, Mexico. This is the first known host record for this parasitoid species. Proconiine leafhopper host associations are also reported for three other eco¬ nomically important Gonatocerus species from the ater species group in North America: G. ashmeadi Girault, G. morrilli (Howard), and G. triguttatus Girault. Key Words .—Insecta, Cicadellidae, Proconiini, Mymaridae, Gonatocerus, egg parasitoid, Mex¬ ico. The glassy-winged sharpshooter, Homalodisca coagulata (Say), is native to the southeastern United States and northeastern Mexico. This species recently invaded California and has become established in the southern part of the State. In 2000, several sharpshooter populations were detected north of Kern County. In order to control populations of this xylem-feeding species, an integrated program employ¬ ing several control options is needed. One cornerstone of an integrated pest man¬ agement program will be classical biological control. Egg parasitoids of H. coagulata were discovered through survey activities con¬ ducted in California and the southeastern United States in 1996 and 1997 (Triap¬ itsyn et al. 1998). Levels of parasitism from one dominant species, Gonatocerus ashmeadi Girault (Hymenoptera: Mymaridae), have been high in southern Cali¬ fornia, but only on the summer and fall generations of the host. The level of parasitism by other local parasitoids, such as Gonatocerus morrilli (Howard) and Ufens spp. (Hymenoptera: Trichogrammatidae), has been generally very low. It is clear that parasitoids attacking the spring generation of H. coagulata need to be imported, screened through quarantine, reared, registered and released through the appropriate permit process, and monitored, in order to impact this pest. Several field expeditions to locate natural enemies of H. coagulata and its close relatives were undertaken, including two trips to the Mexican states of Nuevo Leon and Tamaulipas in early March and April 2000. Preliminary results of these activities were reported by Morgan et al. (2000); this paper provides taxonomic notes on the parasitoid species discovered through our survey and also indicates their host associations, most of which are new records. Members of the tribe Proconiini (Homoptera: Cicadellidae: Cicadellinae), to which Homalodisca Stal belongs, often are referred to as sharpshooters. Distri- 2002 TRIAPITSYN ET AL.: GONATOCERUS ATRICLAVUS REDESCRIPTION 35 Figure 1. Gonatocerus atriclavus Girault. Antenna, female. bution of the 55 genera and numerous species that comprise this tribe is restricted to the Western Hemisphere (Young 1968); only a few of those occur in the United States, most of them are Neotropical. In our survey, we searched for egg parasitoids of the species in two sharp¬ shooter genera, Homalodisca and Oncometopia Stal, based on data obtained dur¬ ing the previous year (Triapitsyn & Phillips 2000). All collections in Mexico were made under a permit issued to our collaborator, E. Ruiz Cancino, Universidad Autonoma de Tamaulipas at Ciudad Victoria. We searched for parasitized sharp¬ shooter egg masses mainly in parks, citrus orchards, and irrigated private gardens. Adult sharpshooters were collected directly in 70% ethanol for further identifi¬ cation and association with the egg masses on their host plants. All parasitized egg masses were sent to the University of California, Riverside (hereafter UCR), quarantine facility under a USD A permit, where emerged parasitoids were screened, identified, and propagated (Morgan et al. 2000). Investigative responsibilities were divided between authors so that S.V.T. iden¬ tified mymarid egg parasitoids and worked on taxonomic aspects of this study as well as on conclusive remarks in the “Discussion”, L.G.B. coordinated foreign exploration efforts, L.G.B. and S.V.T. collected parasitized egg masses of Pro- coniini in Mexico, and D.J.W.M. processed and reared the material in quarantine. Terminology for morphological features used in the description is that of Huber (1988); we use the abbreviation FI, F2, etc., to represent the first, second, etc. funicular segments of the females, and the first, second, etc. flagellar segments of the males. Measurements are given in micrometers (|mm) as length or, if necessary, as length/width. Abbreviations for depositories of specimens are as follows: CSCA, California State Collection of Arthropods, Sacramento; UATM, Univer¬ sidad Autonoma de Tamaulipas, Ciudad Victoria, Mexico; UCRC, Entomology Research Museum, University of California, Riverside; USNM, National Museum of Natural History, Washington, D.C. Gonatocerus atriclavus Girault, 1917, NEW STATUS (Figs. 1-4) Gonatocerus triguttatus atriclavus Girault, 1917: 19 (as a new variety). Gonatocerus triguttatus atriclavus Girault; Huber, 1988: 57. Type Locality. —Mitan, Trinidad. Types .—Lectotype $ on point [USNM], here designated in order to maintain stability of usage of the name, labeled: 1. “Reared from egg-mass of leaf hopper”; 36 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1) Figure 2. Gonatocerus atriclavus Girault. Forewing, female. 2. “Mitan BWI Jan. 1915”; 3. “F. W. Urich Collector”; 4. “53-41”; 5. “ Gon¬ atocerus triguttatus atriclavus Gir 8 $ types.”; 6. “Type No. 20098 U.S.N.M.”. Girault (1917) described this species as a variety of Gonatocerus triguttatus Gi¬ rault without designating a holotype, from several “types”, Catalogue No. 20098. The type series now consists of the lectotype $ mentioned above and 1 2 and 1 2002 TRIAPITSYN ET AL.: GONATOCERUS ATRICLAVUS REDESCRIPTION 37 4 Figure 4. Gonatocerus atriclavus Girault. Basal segments of antenna, male. S paralectotypes on points, here designated, labeled same as the lectotype except label 1 is lacking and label 6 with “Paratype” instead of “Type” [USNM]. Other Material Examined .—MEXICO. TAMAULIPAS, Ciudad Victoria, Tamatan, ex. parasitized egg mass of Oncometopia clarior (Walker) in hibiscus leaf collected 7 Mar 2000, L. G. Bezark and S. V. Triapitsyn; 2 $ $ and 1 S parasitoids emerged at UCR quarantine 20 Mar 2000 [UCRC], 3 9 2 and 1 3 labeled: 1. “MEXICO, interc.[epted] S. Antonio, Tex. VI-10-1959 Johnston”; 2. “59-21950 3325 on palm leaves”; 3. “ Gonatocerus triguttatus atriclavus Girault det. J. T. Huber 1984” [USNM], Original Description .—The original description is inadequate as it is limited to a single sentence: “Similar to typical form but the antenna concolorous except the club” (Girault 1917). Huber (1988) was first to notice that G. atriclavus “is probably a good species, not a subspecies of triguttatus”. Redescription. — Female. — Coloration: Head pale except upper face and ocellar triangle brown, trabeculae and occiput dark brown, eyes and ocelli dusky. Antenna with scape yellow to light brown; pedicel, F1-F3, F4 (basally) and F7 brown; F4 (distally), F5 and F6 light brown; F8 dark brown; club black. Neck pale, pronotum pale brown with darker spots; mesoscutum orange-brown anteriorly and yellowish posteriorly, parapsidal furrows black; anterior scutellum light brown to brown; axilla brown with darker spot at middle; posterior scutellum yellowish-orange-brown; dorsellum, propodeum, pro-, meso-, and metapleura brown; lateral panels of metanotum and propodeal carinae dark brown. Legs yellowish brown except all tarsi, middle and hind tibiae brown. Wings hyaline; venation brown to dark brown. Petiole dark brown; gaster pale to light yellow with dark brown bands on terga; ovipositor plates brown. Head: Width 432. Antenna (Fig. 1) with radicle 0.2X as long as scape, scape very long and wide for the species group, 3. OX as long as wide, very finely longitudinally striate (inner side) or almost smooth (outer side), with several rows of strong setae; pedicel short, smooth; FI short and without sensilla; F2-F5 subequal in length but each slightly wider than preceding article; F6 subquadrate, shorter than F5, F7 slightly wider than long, F8 much wider than long; F2 with 1 or 2 and F3-F8 each with 2 longitudinal sensilla; all funicle segments densely setose; club long (length/width 2.5:1), slightly wider than scape, and about as long as combined length of F1-F4, with 8 longitudinal sensilla, its ventral surface covered with numerous minute, short setae and placoid sensilla, its dorsal surface densely covered with longer setae. Mesosoma: Pronotum divided medially, each lobe with 1 dorsal and 1 lateral strong seta. Mesoscu¬ tum with a pair of strong adnotaular setae. Dorsellum rhomboidal (as in Fig. 3). Propodeal spiracle kidney-shaped; lateral carinae well-developed, submedial carinae almost parallel except anteriorly; propodeum (as in Fig. 3) otherwise smooth. Legs: foretibia with 6-7 conical sensilla. Forewing (Fig. 2) 3.55 X as long as wide; marginal cilia very short, longest fringe seta about V 5 maximum wing width. Forewing blade bare immediately distal to submarginal vein, with 9 microtrichia behind marginal and stigmal veins, cubital row of setae complete; remainder of blade densely setose. Submarginal vein length 263, with 2 hypochaetae, marginal vein length 216, with 7 microchaetae between proximal and distal macrochaetae, stigmal vein length 58. Hind wing blade bare except for complete rows of mi¬ crotrichia along margins and several scattered discal setae at apex. Metasoma: Petiole about as wide as long. Ovipositor about 3 4 length of gaster, barely exserted beyond its apex. Outer plates of ovipositor each with 1 basal and 1 apical seta. Measurements (n = 1).— Body: head: 67; mesosoma: 648; petiole: 72; gaster: 783; ovipositor: 603. Antenna: radicle: 66; scape: 329; pedicel: 88; FI: 47; F2: 80; F3: 84; F4: 80; F5: 80; F6: 65; F7: 51; 38 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1) F8: 43; club: 303. Forewing: 1503/423; longest fringe cilia: 88. Hind wing: 1107/69; venation: 513; longest fringe cilia: 106. Legs (given as coxa, femur, tibia, tarsus): fore: 198, 306, 351, 369; middle: 126, 270, 450, 364; hind: 201, 405, 531, 441. Male. — Coloration: Antenna brown to dark brown except base of scape yellow; face brown; ocellar triangle and occiput dark brown, remainder of vertex and gena light brown; eyes and ocelli pinkish brown. Neck light brown; pronotum, mesoscutum (except light brown edges of lateral lobes), axilla, anterior and posterior scutellum, dorsellum, and propodeum shining brown; mesosomal pleura light brown. All legs light brown except middle tibia and middle and hind tarsi slightly darker, hind tibia brown. Petiole brown, gaster light brown with several dark cross-bands on terga. Otherwise similar to female except for sexually dimorphic characters, as follows: antenna with scape and radicle fused (Fig. 4); basal flagellomeres relatively wide, all flagellomeres with numerous longitudinal sensilla. Measurements (n = 1). —Total body length (dry specimen): 1321; Antenna: scape + radicle: 153; pedicel: 54; FI: 160; F2: 182; F3: 171; F4: 186; F5: 193; F6: 186; F7: 187; F8: 172; F9: 176; F10: 173; FI 1: 233. Forewing: 1884/532. Genitalia: 303. Diagnosis .—This species is easily distinguished from all other described Ne- arctic species of the ater species group ( ater subgroup) of Gonatocerus that have the forewing with the cubital row of microtrichia complete, extending to base of marginal vein (Huber 1988) (i.e., G. ashmeadi Girault, G. fasciatus Girault, and G. triguttatus Girault) by the transverse F8 in the female antenna. If using the key by Huber (1988), G. atriclavus would be separated from G. fasciatus by having the forewing hyaline, without fascia, thus keying to couplet 5 together with G. ashmeadi and G. triguttatus. Besides the shape of F8 mentioned above, G. atriclavus can also be easily separated from these two species by the dilated scape, F5 and F6 lighter colored than other funicle segments, and a very long, black club of the female antenna. In both G. ashmeadi and G. triguttatus, the funicle of the females is uniformly dark brown to black. Distribution .—Mexico and Trinidad. Although we collected this species in Ciudad Victoria, Tamaulipas, which is near the border between the Nearctic and Neotropical regions, it appears to be a mainly Neotropical species. Hosts .—The type series was reared from a “leaf-hopper egg mass” in Trinidad (Girault 1917). Our three specimens from Tamaulipas were reared from a sharp¬ shooter egg mass on a hibiscus plant which was almost certainly laid by Oncom- etopia clarior (Walker). Upon emergence in quarantine, all the wasps were fed with honey, and the single male was given time to mate with the females. The females were then exposed to freshly laid egg masses of H. coagulata in citrus leaves several times; despite the fact that they attempted to parasitize those eggs, we failed to obtain any progeny. Eventually, wasps were killed in 70% ethyl alcohol to serve as voucher specimens. Notes on Other Egg Parasitoids of Proconiine Leafhoppers in Northeastern Mexico Gonatocerus ashmeadi Girault. This common North American species was redescribed and illustrated in detail by Huber (1988), who also indicated its known hosts: Cuerna costalis (F.), Homalodisca coagulata, H. lacerta (Fowler), and On- cometopia orbona (F.). Oncometopia clarior is added here to that list. Individuals from northeastern Mexico appeared identical to specimens collected in southern California, except for two rearings of darker-colored individuals from Tamaulipas that were not propagated. We attribute such differences in body col¬ oration to intraspecific variability, which is possibly host-induced. 2002 TRIAPITSYN ET AL.: GONATOCERUS ATRICLAVUS REDESCRIPTION 39 Under quarantine laboratory conditions (22—25° C, 40—50% RH), the Ciudad Victoria population of G. ashmeadi successfully parasitized H. coagulata eggs laid in a variety of plants including chrysanthemum ( Dendranthema sp.), crape myrtle {Lagerstroemia indica L.), grape {Vitis vinifera L.), sweet orange {Citrus sinensis (L.) Osbeck), toyon {Heteromeles arbutifolia (Aiton) M. Romer), and Verbascum sp. Eggs laid by H. lacerta in chrysanthemum, crape myrtle, and sweet orange were also accepted for parasitism. Viable offspring emerged from all ma¬ terial tested. Further studies were then conducted on the biology of G. ashmeadi as well as of the two other Gonatocerus species discussed below; the results will be reported elsewhere. Material Examined.—Gonatocerus ashmeadi: MEXICO: NUEVO LEON, Monterrey, UANL cam¬ pus, ex. parasitized host egg masses coll. 6 Mar 2000, L. G. Bezark and S. V. Triapitsyn, 18 $2,7 3 3 em. 21 Mar 2000 at UCR quarantine (ex. Oncometopia clarior (Walker) on leaves of Fraxinus sp.). TAMAULIPAS: Ciudad Victoria, 14 Feb 2000, S. N. Myartseva, 16 $ 2, 2 33 (ex. sharpshooter eggs on citrus leaf); Ciudad Victoria, Tamatan, ex. parasitized host egg masses coll. 7 Mar 2000, L. G. Bezark and S. V. Triapitsyn, 19 2 2,7 3 3 parasitoids em. in UCR quarantine 14-23 Mar 2000 (ex. Oncometopia clarior (Walker) on hibiscus leaves); Llera de Canales, ex. parasitized sharpshooter egg mass (on hibiscus leaf) coll. 8 Mar 2000, L. G. Bezark and S. V. Triapitsyn, 12,13 parasitoids em. in UCR quarantine 23 Mar 2000; Municipio Hidalgo, nr. Ejido Benito Juarez, garden in Hotel Hacienda Santa Engracia, ex. parasitized sharpshooter egg mass (on orange leaf) coll. 10 Apr 2000, L. G. Bezark and S. V. Triapitsyn, 7 2 2,3 3 3 em. 14 Apr 2000 at UCR quarantine; 5 km N of Valle Hermoso, ex. parasitized host egg masses coll. 13 Apr 2000, L. G. Bezark, 10 2 2, 11 3 3 em. 20 Apr 2000 at UCR quarantine (ex. Homalodisca coagulata (Say) on hibiscus leaves) [UATM, UCRC], Oncometopia clarior. MEXICO. NUEVO LEON: 6 km SE of Allende, Sanatorio Naturista de Canoas, 10 Apr 2000, L. G. Bezark and S. V. Triapitsyn (on citrus); Monterrey, UANL campus, 6 Mar 2000, L. G. Bezark and S. V. Triapitsyn (on crape myrtle). TAMAULIPAS, Ciudad Victoria, Tamatan, 7 Mar 2000, L. G. Bezark and S. V. Triapitsyn (on hibiscus) [CSCA], Gonatocerus morrilli (Howard). This species is common in southern U.S.A. and Mexico according to Huber (1988). The only previously known host of G. morrilli, H. coagulata, was first indicated by Turner & Pollard (1959) and later by Triapitsyn et al. (1998). We observed a female of this species ovipositing in a sharpshooter egg mass, probably that of O. sp. nr. nigricans (Walker), on a hibiscus plant in the garden of Hotel Rancho Mariposa, near Santander Jimenez in Tamaulipas. Another leafhopper host of G. morrilli that we discovered in Ta- maulipas is O. clarior. A colony of G. morrilli, individuals of which were morphologically similar to those collected in southern California, was established at UCR quarantine using wasps originating from Ciudad Victoria, Tamaulipas. They were successfully reared on both H. coagulata and H. lacerta eggs laid in host plants as described previously for G. ashmeadi. Material Examined .— MEXICO. TAMAULIPAS: Ciudad Victoria, Tamatan, ex. parasitized host egg mass coll. 7 Mar 2000, L. G. Bezark and S. V. Triapitsyn, 9 2 2,1 3 parasitoids em. in UCR quarantine 20-21 Mar 2000 (ex. Oncometopia clarior (Walker) on hibiscus leaf); Llera de Canales, ex. parasitized sharpshooter egg masses (on citrus and hibiscus leaves) coll. 8 Mar 2000, L. G. Bezark and S. V. Triapitsyn, 14 2 2,2 33 parasitoids em. in UCR quarantine 14-23 Mar 2000; same location and collectors, ex. parasitized sharpshooter egg mass (on orange leaf) coll. 11 Apr 2000, 2 2 2,1 3 parasitoids em. in UCR quarantine 17 Apr 2000; nr. Santander Jimenez, Hotel Rancho Mariposa, ex. parasitized sharpshooter egg masses (on hibiscus leaves) coll. 9 Mar 2000, L. G. Bezark and S. V. Triapitsyn, 7 2 2,3 3 3 parasitoids em. in UCR quarantine 15-20 Mar 2000; same location, 22 Apr 40 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1) 1999, S. Triapitsyn, 2 $ $ (on hibiscus, by sweeping); 5 km N of Valle Hermoso, 10 Mar 2000, S. Triapitsyn, 1 8 (on hibiscus, by sweeping) [UATM, UCRC]. Gonatocerus triguttatus Girault. This species was redescribed and illustrated by Huber (1988) based on the type series from Trinidad and additional specimens from Texas. Adult specimens of G. triguttatus were reared in northern Tamaulipas during April 1999 from the egg masses of its only known host at that time, H. coagulata, laid in citrus and peach leaves (Triapitsyn & Phillips 2000). In early spring of 2000, we dissected dead specimens of G. triguttatus from an egg mass of a sharpshooter on a citrus (orange) leaf in Gomez Farias, which is in the tropical part of Tamaulipas. These specimens apparently could not emerge from the egg mass and had died not long before we found them. From the same orange tree, we collected an adult male leafhopper, later identified by Raymond J. Gill as Paraulacizes thunbergi (Stal); that species thus can be considered as a probable host for G. triguttatus. Paraulacizes thunbergi, previously known from southern Mexico (Young 1968), is a new addition to the list of proconiine leafhoppers in Tamaulipas that feed and oviposit on citrus plants (Coronado-Bianco et al. 2000). Additional adult specimens of P. thunbergi were collected in the Nearctic part of Mexico in Nuevo Leon and Tamaulipas. Other apparent hosts of G. triguttatus are O. clarior and O. sp. (see “Material examined” below). A colony of G. triguttatus was established at UCR quarantine (Morgan et al. 2000) from adults that emerged from the egg masses of H. coagulata in hibiscus near Valle Hermoso, Tamaulipas. This colony was being reared on H. coagulata eggs laid in chrysanthemum and orange leaves. All plants tested with G. ashmeadi were also accepted by G. triguttatus. When G. triguttatus females were offered eggs of the smoke-tree sharpshooter, H. lacerta, which is native to California, they readily parasitized them and produced viable offspring. Material Examined.—Gonatocerus triguttatus: MEXICO. NUEVO LEON, 6 km SE of Allende, Sanatorio Naturista de Canoas, ex. parasitized host egg mass coll. 10 Apr 2000, L. G. Bezark and S. V. Triapitsyn, 1 $ em. 14 Apr 2000 in UCR quarantine (ex. Oncometopia clarior (Walker) on orange leaf). TAMAULIPAS: Gomez Farias, 8 Mar 2000, S. N. Myartseva, L. G. Bezark and S. V. Triapitsyn, 1 $ (dissected from a sharpshooter egg mass on orange leaf); nr. Santander Jimenez, Hotel Rancho Mariposa, ex. parasitized host egg mass coll. 9 Mar 2000, L. G. Bezark and S. V. Triapitsyn, 2 $ $ parasitoids em. in UCR quarantine 15 Mar 2000 (ex. Oncometopia sp. on hibiscus leaf); 5 km N of Valle Hermoso, 10 Mar 2000, S. Triapitsyn, 1 8 (on hibiscus, by sweeping); same location, ex. parasitized host egg masses coll. 13 Apr 2000, L. G. Bezark, 14 $ $, 15 8 8 em. 20-27 Apr 2000 at UCR quarantine (ex. Homalodisca coagulata (Say) on hibiscus leaves) [UATM, UCRC], Paraulacizes thunbergi. —MEXICO. NUEVO LEON, 6 km SE of Allende, Sanatorio Naturista de Canoas, 10 Apr 2000, L. G. Bezark and S. V. Triapitsyn (on Malva sp.). TAMAULIPAS: Gomez Farias, 8 Mar 2000, S. N. Myartseva, L. G. Bezark and S. V. Triapitsyn, 1 8 (on orange); Municipio Hidalgo, nr. Ejido Benito Juarez, garden in Hotel Hacienda Santa Engracia, 7 Mar 2000, S. N. Myart¬ seva, L. G. Bezark and S. V. Triapitsyn (on Malva sp.) [CSCA], Ufens sp. This species first emerged from a single sharpshooter egg mass laid in an orange leaf collected at Llera de Canales, Tamaulipas. It was later reared in Nuevo Leon from the eggs of O. clarior. It is a gregarious species: 76 wasps emerged from a clutch of 15 sharpshooter eggs (3—7 emergences per host egg). Morphologically, Ufens sp. is very similar to, but probably distant from, one taken from unspecified leafhopper eggs on elm and jojoba in southern California; H. lacerta is known to be a common associate to the latter plant there. It is also different from the other two Ufens species known from Homalodisca eggs in 2002 TRIAPITSYN ET AL.: GONATOCERUS ATRICLAVUS REDESCRIPTION 41 southern California. All species involved are almost certainly undescribed (J. D. Pinto, personal communication). An attempt was made to initiate a colony of the Mexican Ufens sp. at UCR quarantine. In the first generation, wasps were offered egg masses of H. coagulata laid in leaves of chrysanthemum (14 egg clutches), toyon (2), grape (2), and orange (6). Wasps emerged from only one clutch, laid in an orange leaf, after 16 days, a longer developmental period than was observed for any of the three Gon- atocerus species evaluated (D.J.W.M., unpublished data). The second generation was offered host eggs on Verbascum sp. (4), crape myrtle (6), orange (17), and chrysanthemum (8). No wasps emerged from these leaves. We suggest that a combination of conditions and strong host plant preferences are responsible for the failure to propagate this trichogrammatid species successfully. Material Examined. —MEXICO. NUEVO LEON, 6 km SE of Allende, SanatorioNaturistade Canoas, ex. parasitized host egg mass coll. 10 Apr 2000, L. G. Bezark and S. V. Triapitsyn, 37 $ $, 6 8 8 em. 17 Apr 2000 at UCR quarantine (ex. Oncometopia clarior (Walker) on orange leaf). TAMAULIPAS: Llera de Canales, ex. parasitized sharpshooter egg mass collected 8 Mar 2000, L. G. Bezark and S. V. Triapitsyn, 62 $ 9, 14 8 8 parasitoids em. in UCR quarantine 14-15 Mar 2000 [UATM, UCRC], Discussion Considering the great diversity of the proconiine leafhoppers, which are among the largest leafhoppers known, and whose eggs are laid in clusters in plant tissue (Young 1968), it is surprising how little is generally known about their biology and natural enemies beyond a few economically important species in the United States. The work by Turner & Pollard (1959) had been practically the only one available on this subject until recently, when the establishment of H. coagulata in California prompted the interest in sharpshooter investigations, including stud¬ ies of their egg parasitoids (Triapitsyn et al. 1998). Most of the reported egg parasitoids of Cuerna, Homalodisca and Oncometopia species are members of Gonatocerus. All known North American species parasitic on these leafhopper genera belong to the ater species group of Gonatocerus as defined by Huber (1988), and we believe that this might be the case for all proconiines in the New World. The amazing diversity of ater group species of Gonatocerus in Malaise trap samples from Central and South America correlates very well with the even greater diversity of the sharpshooters from these areas. Further research is needed to demonstrate the validity of this apparent correlation. It is unlikely that species of the ater group of Gonatocerus in the New World are species-specific to their leafhopper hosts; rather, they may be narrowly to broadly oligophagous, i.e., able to parasitize species of a number of closely-related host genera within the tribe Proconiini. Some species of parasitoids, however, may display a preference for certain sharpshooter species. Most likely, however, that they discriminate their hosts based on the size of hosts’ eggs and the habitat. Some sharpshooter species, like H. coagulata, are able to feed upon many plant species, but prefer certain ones for oviposition. To be successful in finding host egg masses, female parasitoids must concentrate their searching activity on those plants. As a result, different parasitoid species may have become more host plant specific than insect host specific, like for instance the common North American mymarid Anagrus nigriventris Girault (Al-Wahaibi & Walker 2000). Thus, for a classical biological control program to be successful in California against H. coa- 42 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1) gulata, whose females oviposit on a great number of different plants, introduction of many different species, as well as various biotypes from any given species, may be warranted. Acknowledgment We thank Enrique Ruiz Cancino, Svetlana N. Myartseva and Vladimir A. Trja- pitzin (Universidad Autonoma de Tamaulipas, Ciudad Victoria, Mexico) for fa¬ cilitating our collecting efforts in Mexico; Walker A. Jones (USDA-ARS, Wes¬ laco, Texas) for assistance in collecting and logistical support, Raymond J. Gill (California Department of Food and Agriculture, Sacramento) for the sharpshooter identifications, John D. Pinto (University of California, Riverside) for identifica¬ tion of Ufens species, and John T. Huber (Canadian Forestry Service, Natural Resources Canada, Ottawa) for valuable advice on the identity of Gonatocerus species. Vladimir Berezovskiy (University of California, Riverside) point- and slide-mounted the parasitoids and made line drawings. This study was sponsored in part by a grant to Mark S. Hoddle and Serguei V. Triapitsyn (University of California, Riverside) who gratefully acknowledge the California Department of Food and Agriculture for financial support. Literature Cited Al-Wahaibi, A. K. & G. P. Walker. 2000. Searching and oviposition behavior of a mymarid egg parasitoid, Anagrus nigriventris, on five host plants species of its leafhopper host, Circulifer tenellus. Entomol. Exp. Appl., 96: 9-25. Coronado-Bianco, J. M., E. Ruiz-Cancino & S. V. Triapitsyn. 2000. Chicharritas de la tribu Proconiini (Homoptera: Cicadellidae) asociadas a citricos en Tamaulipas, Mexico. Acta Zool. Mex. (n. s.), 81: 133-134. Girault, A. A. 1917. Descriptiones stellarum novarum. Privately printed, 22 pp. Huber, J. T. 1988. The species groups of Gonatocerus Nees in North America with a revision of the sulphuripes and ater groups (Hymenoptera: Mymaridae). Mem. Entomol. Soc. Canada, 141: 1-109. Morgan, D. J. W., S. V. Triapitsyn, R. A. Redak, L. G. Bezark & M. S. Hoddle. 2000. Biological control of the glassy-winged sharpshooter: current status and future potential, pp. 167-171. In Hoddle, M. S. (ed.). [Proceedings] California Conference on Biological Control, held July 11- 12, 2000 at the historic Mission Inn, Riverside, California, 205 pp. Triapitsyn, S. V., R. F. Mizell, III, J. L. Bossart & C. E. Carlton. 1998. Egg parasitoids of Homalodisca coagulata (Homoptera: Cicadellidae). Florida Entomol., 81 (2): 241-243. Triapitsyn, S. V. & P. A. Phillips. 2000. First host record of Gonatocerus triguttatus (Hymenoptera: Mymaridae) from eggs of Homalodisca coagulata (Homoptera: Cicadellidae), with notes on the distribution of the host. Florida Entomol., 83 (2): 200-203. Turner, W. F. & H. N. Pollard. 1959. Life histories and behavior of five insect vectors of phony peach disease. Tech. Bull. U.S. Dep. Agric. 1188, 28 pp. Young, D. A. 1968. Taxonomic study of the Cicadellinae (Homoptera, Cicadellidae). Part 1. Proconiini. United States Nat. Mus. Tech. Bull. 261, 287 pp. Received 8 December 2000; Accepted 13 September 2001. PAN-PACIFIC ENTOMOLOGIST 78(1): 43-55, (2002) COPULATION DURATION IN THREE SPECIES OF ANTHOCORIS (HETEROPTERA: ANTHOCORIDAE) AT DIFFERENT TEMPERATURES AND EFFECTS ON INSEMINATION AND OVARIAN DEVELOPMENT David R. Horton, Tamer a M. Lewis, & Tonya Hinojosa USDA-ARS, 5230 Konnowac Pass Road, Wapato, Washington 98951 Abstract .—We compared duration of copulation among three species of predatory bugs: Antho- coris tomentosus Pericart, A. whitei Reuter, and A. nemoralis (Fabricius). Copulation duration at room temperatures (22-24° C) was longest in A. whitei ( x = 89.3 min), of intermediate length in A. tomentosus (. x = 40.0 min), and shortest in A. nemoralis ( x = 12.7 min). By interrupting mating pairs we showed that long duration copulations were more likely than short copulations to result in insemination and in ovarian maturation in the female. Probability of ovarian devel¬ opment following a shortened copulation was often lower than probability of insemination, sug¬ gesting that insemination alone was not always enough to prompt ovarian development. Size of the sperm reservoir in the female increased with increasing duration of copulation. Males of all species transferred seminal products for most of the copulation period, suggesting that the longer copulations noted for A. whitei and A. tomentosus (relative to duration in A. nemoralis ) were not due to post-insemination mate-guarding by A. whitei or A. tomentosus. Copulation duration was longer at 15° C (range: 25-181 min) than at 25° C (range: 13-102 min). We interrupted mating pairs at both temperatures in all species to determine how the combined factors of temperature and copulation duration affect insemination rates and probability of oocyte matu¬ ration. For a given copulation duration, probability of insemination and oocyte maturation were higher at 25° C than at 15° C. Species’ differences in copulation duration may reflect differences among species in how rapidly males transfer sperm, and that lower temperatures may make it physically more difficult for the male to force seminal products through the aedeagus. Key Words .—Insecta, Anthocoridae, mating behavior, sperm transfer, copulation. Predatory bugs in the genus Anthocoris are important sources of biological control in natural and agricultural ecosystems (Lattin 1999). Despite their impor¬ tance, however, these species are poorly studied in certain aspects of basic biology. Our laboratory has recently begun to study the reproductive biology of several North American species of Anthocoris, with goals ultimately to improve our un¬ derstanding of mating behavior and reproductive biology in these insects. In the present study, we explored how copulation duration varies among three species of Anthocoris that inhabit the Pacific northwest region of the United States: A. tomentosus Pericart, A. whitei Reuter, and A. nemoralis (Fabricius). Copulation duration varies extensively among insect taxa (Thornhill & Alcock 1983), to the extent that even closely related species may exhibit large divergence in this trait (Thornhill & Alcock 1983, Cordero 1990, Lachmann 1997). There are potentially a variety of costs and benefits associated with short- or long- duration couplings (Alcock 1994). Here, we address whether artificially shortened copulations in these species affect probability of insemination, probability of ovar¬ ian development, and amount of seminal material transferred to the female. The first hypothesis tested here is that females that experience artificially shortened copulations would be less likely than undisturbed females to be inseminated, as shown in other insect species (Farias et al. 1972, Lew & Ball 1980). We also determined whether these artificially shortened copulations resulted in decreased 44 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1) probability of ovarian development in the female. Maturation of ovaries in An- thocoris requires mating (Anderson 1962, Shimizu 1967). However, it is not known whether the mating act itself is sufficient to prompt ovarian development, or whether insemination is required. If the mating act alone is enough to prompt ovarian development, then we expect that severely shortened copulations, i.e., those resulting in a lack of insemination, would still prompt ovarian development. Our second major objective was to determine whether the longer copulations of A. whitei and A. tomentosus (relative to copulation duration in A. nemoralis [see below]), was due to mate-guarding by males of these two species. Many insect species prolong copulation beyond that necessary to complete insemination, as a male strategy to delay remating by the female and to ensure paternity (Alcock 1994). Females of A. tomentosus and A. whitei will mate multiple times under laboratory conditions (unpublished data). Here, we interrupted copulating pairs at several time intervals between intromission and the end of copulation, and esti¬ mated quantity of seminal products that had been transferred by the male at each duration. If males prolonged copulation beyond the time required to inseminate the female (i.e., if the male engaged in mate-guarding behavior), we would expect that the amount of seminal material transferred in artificially shortened copulations would not differ from the amount transferred in uninterrupted matings. Finally, we determined whether temperature mediates the interaction between copulation duration and probability of insemination. In Anthocoris spp., sperm are transferred by the male through a thin membranous tube in the female (cop- ulatory tube; Carayon 1953) directly into her sperm pouch. Elsewhere, we sug¬ gested that males in certain species of Anthocoris experience physical difficulties in forcing seminal materials through the aedeagus and female copulatory tube (Horton et al. 2001). Here, we hypothesized that these difficulties would be am¬ plified with decreasing temperatures, resulting in longer copulations at a lower temperature than a higher temperature. Moreover, we hypothesized that probabil¬ ity of insemination would decrease with decreasing temperature for a copulation of a given duration. This last hypothesis was tested by interrupting mating pairs at specified time intervals for matings conducted at different temperatures. Materials and Methods Source of Insects. —Laboratory cultures of the three species were begun from field-collected insects. Nymphs and adults of A. tomentosus were collected from Salix sp. growing west of Tieton, Washington. Anthocoris whitei was collected from antelope bitterbrush, Purshia trident at a (Pursh), growing in rangeland west of Tieton. Assays with A. tomentosus and A. whitei were done using offspring of field-collected insects. The third species, A. nemoralis, is native to Europe but was released in western North America to control pear psylla, Cacopsylla pyricola (Foerster) (McMullen 1971). Nymphs and adults of A. nemoralis were collected in Richmond, California from Acacia longifolia Willdenow, where it occurs in association with a psyllid pest (Dreistadt and Hagen 1994), and from an uniden¬ tified shrub species located in Golden Gate Park, San Francisco. Insects from the two sites were combined into a single culture. Assays used a mix of first-gener¬ ation (offspring of field-collected insects) and second-generation insects. Insects were reared on pear seedlings infested with pear psylla at 22° C under long-day conditions (16:8 h [L:D]). Offspring from the parental cultures were 2002 HORTON ET AL.: ANTHOCORIS COPULATION 45 collected as older nymphs and placed singly in glass petri dishes lined with filter paper. Psylla-infested leaves were added daily to each dish. Petri dishes were checked daily for eclosion of new adults. Date of eclosion and sex of each bug were recorded. All assays used previously unmated insects 1 to 6 days of age. Voucher specimens of each species have been deposited in the M.T. James Entomology Museum at Washington State University, Department of Entomology, Pullman. Study 1. Copulation Duration, Insemination, and Oocyte Maturation. —Copu¬ lation duration was quantified for the interval beginning with intromission and ending with the male’s withdrawal of the aedeagus. Withdrawal from the female was always followed immediately by physical separation of the two sexes in all species. Matings were done at room temperature (22—24° C) under fluorescent lighting. Plastic petri dishes (60 mm in diameter) were used as mating arenas. Preliminary trials were conducted to determine copulation duration in pairs al¬ lowed to mate without interference. Based upon these trials, we established a range of copulation durations to be tested. Pairs were manually separated after the appropriate time interval by gently prodding the insects with a small paint brush. The test durations are (in min from intromission to manual interruption): A. tomentosus (0 [virgin females], 10, 20, 25, 30, 40, and uninterrupted); A. whitei (0, 10, 20, 40, 60, 90, and uninterrupted); and A. nemoralis (0, 2, 4, 8, 12, and uninterrupted). Probability of sperm transfer and ovarian maturation were recorded as a func¬ tion of copulation duration for the manipulated and uninterrupted pairs. Imme¬ diately after copulation, females were either dissected to determine whether in¬ semination had occurred, or set aside to allow oocyte development (see below). In Anthocoris, sperm are transferred by the male directly into the female’s mem¬ branous sperm pouch (Carayon 1953); there is no spermatophore. To obtain an indication about the amount of seminal material transferred by the male at dif¬ ferent copulation durations, we measured size of the sperm pouch in females allowed to copulate for specified time intervals. After mating, the female was killed by crushing her head and thorax. The abdomen was dissected away from the rest of the body in a drop of saline by using two insect pins. The sperm pouch was carefully removed from the other tissues and submerged in a drop of saline on a microscope slide. The pouch was situated on the slide so that the entrance of the copulatory tube was at the base of the pouch (Fig. 1). The pouch was then measured immediately at 50X under a dissecting microscope equipped with an ocular micrometer. Two measurements were recorded (Fig. 1): maximum length— measured from the base of the membranous pouch to the top of the pouch; max¬ imum width—measured perpendicular to the length measurement. Virgin females were included as controls. After measuring the pouch, we checked for the presence of sperm by placing a cover slip over the pouch and pressing on the slip until the pouch ruptured. Pouch contents were then examined for the presence of sperm at 100—400X under a compound microscope. Females that were not dissected were set aside to record whether ovaries ma¬ tured. Following copulation in both interrupted and uninterrupted pairs, females were separated from the male and put singly on pear seedlings infested with pear psylla. Seedlings and insects were placed in environmental chambers at 22° C and 46 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1) Figure 1. Methods used for estimating maximum length and width of sperm pouch in virgin and mated females of A. tomentosus, A. whitei, and A. nemoralis. Pouch shapes are highly variable both within and between species; illustration provides more-or-less typical shape for A. tomentosus. a 16:8 h photoperiod. Females were then dissected 8—12 (A. tomentosus) or 4—6 (A. nemoralis and A. whitei) days later; the preoviposition period (i.e., the number of days between mating and onset of egglaying) in these species is ca. 8, 4—5, and 3-4 days in A. tomentosus, A. whitei, and A. nemoralis, respectively (unpub¬ lished data). Dissected females were scored as reproductive or as non-reproduc- tive. To be scored as reproductive, the female had to have at least one ovariole containing a mature egg. Appearance of the ovaries is quite different in repro¬ ductive and non-reproductive females of these species. The mature terminal oo¬ cyte in a reproductive female is approximately 1.5 (A. nemoralis), 2.0 (A. whitei), and 4.0 (A. tomentosus) times as long as the terminal oocyte in an unmated female of the same age (unpublished data), and, moreover, has the characteristic appear¬ ance of the deposited egg. Probability that the female contained at least one mature oocyte was modeled as a logistic function of copulation duration (PROC CATMOD; SAS Institute 1987). We tested whether copulation duration affected probability of insemination using x 2 tests. To compare frequency of females scored as reproductive to fre¬ quency of females that were successfully inseminated, we conducted 2X2 con¬ tingency tests. To determine whether size of the sperm pouch depended upon 2002 HORTON ET AL.: ANTHOCORIS COPULATION 47 copulation duration, we first conducted a principal components analysis (PROC PRINCOMP; SAS Institute 1987) on the width and length measures to create a single size variable. Analysis of variance was then used to compare mean principal component scores among different copulation durations. Sample sizes are provid¬ ed in the RESULTS. Study 2. Effects of Temperature .—Two temperatures were compared for effects on mating and insemination: 15° C and 25° C. Assays were conducted under fluorescent lighting in controlled environmental rooms set to the appropriate tem¬ perature. Petri dishes containing the adult insects were placed in the rooms 1 h before the assay was conducted to allow the insects to acclimate. After 1 h, pairs (1 male and 1 female) were moved to mating arenas (plastic petri dishes 60 mm in diameter) and allowed to mate. Pairs that had not initiated copulation within 30 min of being placed in the arenas were discarded. Copulation duration was manipulated at both temperatures by separating pairs at two specified time intervals, producing for each species copulations of three durations (in min): A. tomentosus —10, 30, and uninterrupted controls; A. whitei — 20, 40, and uninterrupted controls; A. nemoralis —2, 8, and uninterrupted controls. The female from each pair was collected at the end of the mating bout. One-half of the females were then dissected to determine whether insemination had oc¬ curred. Females that were not dissected were set aside to monitor ovarian matu¬ ration, using the methods described above. Sample sizes are provided in the RE¬ SULTS. Mean copulation duration in uninterrupted pairs was compared between tem¬ peratures with a two-sample /-test (PROC TTEST; SAS Institute 1987). Contin¬ gency tests were used to test whether probability of insemination or probability of oocyte maturation in interrupted matings were affected by temperature and copulation duration. For each species, a 2 X 2 X 2 (temperature [15° C versus 25° C] X duration [short interrupted versus long interrupted] X status [inseminated versus not inseminated]) model was fitted to the insemination data and analyzed using log-linear methods (PROC CATMOD; SAS Institute 1987). The analyses were repeated replacing frequency of insemination with frequency of oocyte mat¬ uration (i.e., female containing at least one mature oocyte versus no mature oo¬ cytes). Results Study 1. Copulation Duration, Insemination, and Oocyte Maturation .—Mean copulation duration in uninterrupted pairs differed among the three species (Fig. 2: solid circles having standard error bars). There was also substantial variation within species, as indicated by the range of values noted in uninterrupted pairs: A. tomentosus, range = 7—64 min, x = 40.0 min, n = 17; A. whitei, range = 39— 138 min, x = 89.3 min, n = 13; A. nemoralis, range = 7.7-20.9 min, x = 12.7 min, n = 20. Probability of oocyte maturation in females from interrupted pairs increased with increasing duration of copulation (Fig. 2; solid circles and solid lines). Probability of ovarian maturation was 82-100% in uninterrupted females (solid circles with error bars in Fig. 2). In some interrupted females that were scored as reproductive, oocyte maturation was limited to a subset of the ovarioles, rather than in all ovarioles as seen in females from undisturbed matings. Probability of insemination increased with increasing copulation duration in all % of Females Having at Least One Mature Oocyte (•) 48 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1) 0 10 20 30 40 50 0 20 40 60 80 100120 Copulation Duration (Minutes) Figure 2. Effects of copulation duration on probability that the female had at least one mature oocyte at dissection (solid circles) and on probability of insemination (open circles; data not collected for A. tomentosus at duration = 25 min). Solid circles having standard error bars are results for females % of Females With Sperm (o) 2002 HORTON ET AL.: ANTHOCORIS COPULATION 49 three species (Fig. 2, open circles; by x 2 tests, P < 0.01 in all three species). Percentage of females having sperm was very high in the uninterrupted pairs: A. tomentosus, 100% (n = 12); A. whitei, 91.7% ( n — 12); A. nemoralis, 100% ( n — 17). Probability of insemination was often higher than probability of oocyte maturation, particularly at the shorter copulation durations (Fig. 2: compare paired open and filled circles), which may indicate that presence of sperm in the female was not always sufficient to prompt oocyte maturation. Size of the sperm pouch increased linearly with increasing duration of copu¬ lation in all three species (Fig. 3; P < 0.005 for all three species); quadratic effects were non-significant. Study 2. Effects of Temperature .—Mean copulation duration in uninterrupted pairs was significantly longer at 15° C than 25° C for all three species (Fig. 4; P < 0.01 for all species [by /-tests]). Copulations averaged 79, 32, and 12 min longer at 15° C than 25° C for A. whitei, A. tomentosus, and A. nemoralis, re¬ spectively (Fig. 4). Percentage of females that were inseminated or were characterized as having at least one mature oocyte increased with increasing temperature and duration of copulation (Fig. 5; statistical tests summarized in caption). For A. tomentosus, females that were paired at 15° C invariably failed to mature their ovaries if interrupted before finishing copulation (Fig. 5). Uninterrupted copulations tended to result in insemination and oocyte maturation at both temperatures for all three species. Discussion While it is apparent that copulation duration is highly variable in the Insecta (Thornhill & Alcock 1983, Eberhard 1996), the significance of this variation is not well understood (Cook 1994, Eberhard 1996). If the function of copulation were no more than a means to deliver sperm to the female, then one would expect selection to favor brevity (Cook 1994, Eberhard 1996). That is, time spent in copulation is not available for other activities, including functions such as feeding, egglaying, or search for additional mates (Eberhard 1996). However, the fact that copulation in many species occurs for time periods longer than necessary to trans¬ fer sperm (Cordero 1990, Alcock 1994) suggests that sperm transfer is not the sole function of copulation. Long-duration copulation may be favored for any of a number of reasons, including as a means to increase certainty of paternity, to 80% in A. tomentosus, A. whitei, and A. nemoralis, respectively (Fig. 2). To reach the point at which 75% or more of females contained sperm in the sperm pouch required copulation of approximately 4, 20, and 40 min in duration for A. nemoralis, A. tomentosus, and A. whitei, respectively. Moreover, measurements of sperm pouch size suggested that males transferred seminal prod¬ ucts for most of the time that the insects were paired. That is, for all 3 species, there was a significant linear increase in sperm pouch size with copulation du¬ ration for inseminated females (Fig. 3). This result suggests that the longer cop¬ ulations of A. whitei and A. tomentosus relative to duration in A. nemoralis were not due to mate-guarding activities by male A. whitei and A. tomentosus. Instead, species’ differences in duration apparently are due to differences in how rapidly males of each species could transfer sperm, or in species’ differences in the amount of seminal materials transferred. By manipulating copulation duration, we showed that shortened copulations had major effects on fitness. Probability of insemination and ovarian maturation decreased with decreasing duration of copulation (Fig. 2), and quantity of seminal material transferred by the male was reduced at the shorter durations (Fig. 3). Other studies have monitored the effects of artificially shortened copulations on fitness measures in insects. In several species, interrupted copulations have been shown to result in reduced probability of insemination (Farias et al. 1972, Lew & Ball 1980, Lachmann 1997), results similar to those reported here. Studies have also shown that longer copulations may result in more sperm being transferred (Yamagishi & Tsubaki 1990). We infer, based upon measurements of sperm pouch size (Fig. 3), that number of sperm transferred to the female by males of Antho¬ coris increased with increasing duration of copulation. Mating is necessary to prompt oocyte maturation in Anthocoridae, and unmated females in this family deposit few or no eggs (Anderson 1962, Shimizu 1967, Carayon 1970). Almost no research has been done to determine what factors <— Sample sizes are (see also Fig. 2): 9-14 (A tomentosus), 9-12 (A nemoralis), and 12-18 (A whitei). Sample sizes tend to be smaller than those provided in Fig. 2 because damage to the sperm pouch during dissection occasionally prevented measurement of the pouch. Open symbols are results for uninterrupted controls. Effects of copulation duration (by ANOVA; excluding uninterrupted controls): A. tomentosus (F = 5.5; df = 4,48; P = 0.001 [linear: P = 0.004; quadratic: P = 0.13]); A. whitei (F = 8.8; df = 5,82; P < 0.001 [linear: P < 0.001; quadratic: P = 0.18]); A. nemoralis (F = 16.2; df = 4,51; P < 0.001 [linear: P < 0.001; quadratic: P = 0.11]). 52 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(1) tn Q. •*- O 0) o> ra •*-> c a) o « CL $ & & 4* £ & ri > a'o ** n. oj V 10 larvae) were placed in plastic bags with double paper towel beneath the vegetation containing immatures for rearing. Small col¬ lections (<10 larvae) were placed in plastic vials with tissue paper for rearing. Some larvae and pupae were preserved in hot water and stored in 70% ethanol. Some pupal shells were saved for examination by SEM. Type Depositions .—The type series are deposited in the following institutions: CNC—Canadian National Collection, Agriculture Canada, Biosystematics Re- 134 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(2) search Centre, K. W. Neatby Bldg., C.E.F. Ottawa, Ontario K1A OC6, Can¬ ada LACM—Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007, U.S.A. NTNU—Department of Biology, National Taiwan Normal University, Taipei, Taiwan 116, R.O.C. SDNHM—San Diego Natural History Museum, San Diego, California 92112, U.S.A. UCB—Essig Museum of Entomology, University of California, Berkeley, Cal¬ ifornia 94720, U.S.A. USNM—Department of Entomology, Entomology, U.S. National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Embola Embola Walsingham 1909: 3, REVISED STATUS Type Species—Embola xanthocephala Walsingham 1909, by monotypy Diagnosis.—Embola is characterized by very long and slender saccus, phallus and ductus bursae (Hsu 1995, Hsu & Powell in press). The other genera with similar genital features have flattened antennae, while Embola possesses cylin¬ drical antennae. There is a row of long, bristle-like scales behind eyes. Corpus bursae has double signa, one dorsally immediate anterior to the junction with ductus bursae, the other on ventral wall. Embola powelli Hsu, NEW SPECIES (Figs. 1-3) Types. —Holotype, male: USA. CALIFORNIA, SAN DIEGO Co.: 2 mi (3 km) NE of Lakeside, 400' (122 m), 16 March 1994, reared from Mirabilis californica, emgd. 18 April 1994, JAP 94C54 (Y. F. Hsu, H. H. Chuah, UCB) 2002 HSU: A NEW SUN MOTH IN SOUTHERN CALIFORNIA 135 Fig. 2. Male genitalia of Embola powelli. Fig. 3. Female genitalia of Embola powelli. Paratypes (21 males, 20 females).—USA, ARIZONA, COCHISE Co.: 1 fe¬ male, 3 mi NW Chiricahua, 5 August 1991 (Y. F. Hsu & J. A. Powell, UCB). GRAHAM Co.: 1 female, Aravaipa Cyn., Wild Turkey Cr., MVL, 26 July 1989 (B. & J. F. Landry, CNC). PIMA Co.: 1 male, T19S, R16E, S18, 10 October 1960 (R. W. Hodges, USNM). CALIFORNIA, IMPERIAL Co.: 1 male, 2 mi E Moun¬ tain Springs, 28 April 1993, reared from Mirabilis tenuiloba, emgd. 18 May 1993, JAPN 93D41.1 (Y. F. Hsu, UCB); 1 male, same locality, 18 March 1994, reared from M. tenuiloba, emgd. 12 April 1994, JAP 94C62 (Y. F. Hsu & H. H. Chuah, NTNU); 2 males, same locality, 20 July 1994, reared from M. tenuiloba, emgd. 136 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(2) 25 August 1994, JAP 94G25 (Y. F. Hsu, UCB). KERN Co.: 1 male, 4 mi N Red Rock Cyn., 3 May 1968, on Haplopappus cooperi (Powell, UCB). LOS ANGELES Co.: 2 males, 3 females, San Clemente Is., Seal Cove, Rock Wall Cyn. area, 14 April 1980, associated with Mirabilis laevis (= californica) (Powell, D. Faulkner, SDNHM, UCB); 1 female, same locality, West Cove, associated with Mirabilis laevis (= californica ), 15 April 1980 (Powell, Faulkner, UCB); 1 female, Whittier, 12 December 1910, on flowers of Encelia californica (P. H. Timberlake, USNM). RIVERSIDE Co.: 1 male, 4 mi E Elsinore, R. R. Cyn., 17 April 1965 (Powell, UCB). SAN BERNARDINO Co.: 1 male, 7 mi SE Kelso, Vulcan Mine Rd., 26 April 1977 (Powell, UCB). SAN DIEGO Co.: 4 males, 1 female, 1 mi E Cardiff, 24/31 March 1974 (Powell, SDNHM, UCB); 1 female, La Jolla, 18 June 1963 (Powell, UCB); 1 male, 2 mi NE Lakeside, 400', 30 March 1961, 1 male, same locality, 13 March 1963; 1 male, 1 female, same locality, 24/25 March 1993 (all Powell, UCB); 1 female, 27 April 1993, reared from M. californica, emgd. 11 June 1993, JAP 93D39 (Y. F. Hsu, UCB); 1 female, same locality, 16 March 1994, reared from M. californica, emgd. 17 April 1994, JAP 94C54 (Y. F. Hsu, H. H. Chuah, UCB). SANTA BARBARA Co.: 1 female, Santa Cruz Island, Lower Central Valley, 24 May 1984 (J. F. Landry, CNC). VENTURA Co.: 1 male, 1 female, Santa Susana Mtns., Tapo Cyn., 16/19 April 1939, on M. californica (L. M. Martin, LACM). MEXICO, BAJA CALIFORNIA NORTE: 1 female, 4 mi SW La Zapopita, Valle de Trinidad, 16 April 1961 (F. S. Tmxal, LACM). BAJA CALIFORNIA SUR: 1 female, 30 km E La Ribera, Rancho Las Barracas, 21/24 March 1982 (M. Irwin, E. Schlinger, UCB). CHIHUAHUA: 1 female, 12 mi W Hidalgo Del Parral, 6200', 14 July 1964 (Powell, UCB); DURANGO: 1 male, 26 mi S La Zarca, 16 July 1964, on Encelia (J. A. Chemsak, UCB). Description. —MALE. FW length 2.8-5.4 mm (3.58 mm ± 0.49 mm, n = 21). Head: Frons, vertex metallic gray tinged with blue. Scaling behind eyes cream-white. Antenna metallic dark gray. Labial palpus metallic gray with basal segment cream-white. Thorax: Metallic dark gray tinged with blue. Legs metallic gray tinged with blue. Profemur and mesotibia with distal ends cream-white. Metatibia with a whorl of white scales adjacent to spurs; black scaling in front of the white whorl. Medial spurs of metatibia with inner one 2.2X longer than outer. Forewing: Metallic chrome or flame orange with distal margin metallic gray tinged with blue; 3 costal and 1 dorsal metallic gray spots tinged with blue. A transverse band of same color located at % from base; Cl proximal, C2 and C3 distal to transverse band; spots and band edged with black scaling. Extensive black scaling along costa and dorsal margin in some specimens. Fringe gray tinged with orange. Hindwing: Metallic pale gray tinged with blue. Fringe gray tinged with orange, turning cream-white toward tornus. Abdomen: Metallic black banded with silver, with creamy yellow terminal end. Genitalia: As in Fig. 2 (drawn from EME slide 3646, Riverside Co., CA; n = 8). Tegumen cone-shaped, attenuate to up-curved, blunt distal end. Socii elongate, dilated at base, rod-like with a blunt, down-curved distal tip, 0.65X tegumen length. Saccus 3.15X tegumen length Valva broad, elongate with basal portion narrowed. Phallus very narrow, slightly down-curved distally, 1.2X longer than tegumen + saccus. Cornuti a cluster of wart¬ like protuberances at distal end of aedeagus. FEMALE. FW length 2.8-4.8 mm (3.55 mm ± 0.59 mm, n = 17). Color pattern as described for male but lacking cream-yellow terminal scaling on abdomen. Genitalia: As in Fig. 3 (drawn from YFH slide 1044, Graham Co., AZ; n = 7). Medial, sclerotized, band of apophyses anteriores oval or somewhat rectangular. Ventral signum elongate, irregularly bordered, forming a deeply invaginated band; dorsal signum oval or an elongate, slightly depressed band, length variable, ranging from half long to nearly as long as dorsal signum. Early Stages .—larva cylindrical, cream colored, with two S V setae on A9 (Hsu & Powell in press); pupa (Fig. 7) cylindrical, brown, with short lateral bristles present on weak lateral ridges on abdomen. 2002 HSU: A NEW SUN MOTH IN SOUTHERN CALIFORNIA 137 Distribution. —USA. (California, Arizona); Mexico (Baja California Norte, Baja California Sur, Chihuahua, Durango). Voltinism. —Evidently a multivoltine species, as moths have been collected in all seasons. Etymology. —This species is named in honor of Dr. Jerry A. Powell who col¬ lected the first series of this insect from the type locality, and for his significant contribution to the knowledge of North American microlepidoptera fauna. Diagnosis. —Within Embola, E. powelli is unique in having it socii dilated at base and abruptly narrowed toward the distal end. While all the other known Embola species have the ventral and dorsal signa different in shape (Hsu & Powell in press), the two signa of E. powelli are similar. E. powelli is also the only Embola with a metallic gray transverse band on the forewing. Biology. —Larval hosts are Mirabilis californica (JAP 93D39, 94C54) and M. tenuiloba (JAP93D41.1, 94C62) (Nyctaginaceae) in southern California. The lar¬ va is a stem borer that enters the stem by boring a hole at any position on the stem; frass is deposited in the canal made by larva. Pupation occurs in the stem, and the adult emerges from the stem through a hole made in the larval stage. The adult raises its hindlegs in repose. Discussion The discovery of biology of Embola powelli suggests that the endophagous behavior of the larva may be a general feeding strategy shared by all Embola species. Monte (1934) and Costa Lima (1936, 1945) reported larvae of South American E. obolarcha (Meyrick) as borers in cecidomyid galls on Piper species (Piperaceae). The identity of the moths they observed has yet to be confirmed, but it will confirm the endophagous behavior of Embola larvae if their moths were true E. obolarcha. Besides Embola species, Lamprolophus is the only heliodinid that is known to have larva feeding as a stem borer. In that species, however, the behavior is facultative. Busck (1900) indicated that larvae of L. lithella use young stems and eject frass through a hole on the stem (Pig. 4). I further observed larvae of that species leaving a stem to bore into another when the old one deteriorated, and a group of larvae were seen in a single large cavity in a soft, young stem (JAP 94D98). Moreover, the pupal morphology of L. lithella is typical of heliodinids, with the body flattened dorsoventrally and with prominent lateral ridges and long, lateral bristles (Pig. 6). In contrast, the larvae of Embola are obligate borers, making a linear gallery in the stem and depositing frass within the gallery (Pig. 5). Pupae of E. powelli show modifications related to the life style as obligate borers. The body is cylindrical, lateral ridges of the abdomen are greatly reduced, nearly obsolete, and the lateral bristles on abdominal segments are considerably shortened (Fig. 7). Phylogenetic analysis indicates Embola to be more closely related to genera such as Scelorthus, Lithariapteryx, and Aetole, which are either leaf skeletonizer or leaf miners during larval stages, than to Lamprolophus (Hsu 1995, Hsu & Pow¬ ell in press). As a result, the stem-boring strategy found in Lamprolophus and Embola is unrelated and not homologous. Obligate stem boring behavior is found exclusively only in Embola, and is hypothesized to be a synapomorphy of mem¬ bers of Embola as well as a uniquely derived larval feeding behavior in the 138 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(2) Fig. 4. Feeding hole and extruded frass of Lamprolophus lithella on Pisonia aculeata (Arrow indicates frass). Fig. 5. Feeding gallery containing deposited frass by a larva of Embola powelli in a stem of Mirabilis californica (Arrow indicates frass). Fig. 6. SEM of pupal shell of Lamprolophus lithella. Fig. 7. SEM of pupal shell of Embola powelli. 2002 HSU: A NEW SUN MOTH IN SOUTHERN CALIFORNIA 139 evolutionary history of Heliodinidae, with morphological modifications associated with the boring feeding strategy present only in the Embola lineage. Acknowledgment This paper is dedicated to Jerry A. Powell, my major professor at UCB. He supported my investigation of heliodinid moths from all possible ways, and I have gained the majority of my knowledge of microlepidoptera from him. I also thank Terry L. Harrison, Department of Entomology, University of Illinois for his com¬ ments on the manuscript, Jean-Frangois Landry, Canadian National Collection, Julian P. Donahue, Natural History Museum of Los Angeles County, Ronald W. Hodges, formerly with the Systematic Entomology Lab, USDA, U.S. National Museum of Natural History, for the loan of specimens. Literature Cited Busck, A. 1900. New American Tineina. Jour. N. Y. Entomol. Soc. 8: 234-248. Brown, J. W. & J. A. Powell 1991. Systematics of the Chrysoxena group of genera (Lepidoptera: Tortricidae: Euliini). U. Calif. Publ. Entomol. Ill: 1-87. Costa Lima, A. 1936. Terceiro Catalogo dos Insetos que Vivem nas Plantas do Brasil. Rio de Janeiro. Costa Lima, A. 1945. Insetos do Brasil. 5° Tomo. Lepidopteros, l a parte. Esc. Nac. Agron. Serie Didatica 7: 1-379. Diakonoff, A. & Y. Arita. 1979. Three new species of the so-called Glyphipterigidae auctorum from Japan (Lepidoptera). Zool. Meded. 54: 95-100. Fal’kovich, M. I. 1990. Heliodinidae. pp. 699-700. In Medvedev, G. S. (ed.), Keys to the Insects of the European Part of the USSR. Volume IV. Lepidoptera, Part II. English Edition. E. J. Brill. Floater, G. J. 1995. Seed predation and flower visiting by Epicroesa sp. (Lepidoptera: Heliodinidae) on a rare Seychelles tree. Phelsuma 3: 31-36. Harrison, T. & S. Passoa. 1995. Mirabilis-fceding Heliodines (Lepidoptera: Heliodinidae) in central Illinois, with description of a new species. Proc. Entomol. Soc. Wash 97: 63-70. Heppner, J. B. & B. Landry. 1994. A new sun moth from Galapagos Islands (Lepidoptera: Heliodi- nidae). Tropical Lepidoptera 5: 126-128. Hsu, Y. F. 1995. Systematics of moths formerly assigned to Heliodines Stainton and phylogenetic relationships within Heliodinidae (Lepidoptera: Heliodinidae). Ph.D. dissertation, University of California, Berkeley. Hsu, Y. F. & J. A. Powell, (in press) Phylogenetic relationships within Heliodinidae and systematics of moths formerly assigned to Heliodines Stainton (Lepidoptera: Yponomeutoidea). U. Calif. Publ. Entomol. Klots, A. B. 1970. Lepidoptera. pp. 115-130. In Tuxen, S. L. (ed.), A Taxonomists’ Glossary of Genitalia in Insects (2nd ed.) Munksgard, Copenhagen. Meyrick, E. 1914. Lepidoptera Heterocera Fam. Heliodinidae. Genera Insectorum 165: 1-29. Monte, O. 1934. Borboletas que vivem em plantas cultivadas. Seer. Agr. Est. Minas Gerais 21: 1-219. Powell, J. A. 1980. Evolution of larval food preferences in microlepidoptera. Ann. Rev. Entomol. 25: 133-159. Powell, J. A. 1991. A review of Lithariapteryx (Heliodinidae), with description of an elegant new species from coastal sand dunes in California. J. Lepid. Soc. 45: 89-104. Smithe, F. B. 1975. Naturalist’s Color Guild. The American Museum of Natural History, New York. Walsingham, T. 1909. Embola. Biol. Centr.-Amer., Heter. 4: 3-4. Wester, C. 1956. Comparative bionomics of two species of Heliodines on Mirabilis. Proc. Entomol. Soc. Wash. 58: 43-36. Received 8 December 2000; Accepted 9 January 2002 PAN-PACIFIC ENTOMOLOGIST 78(2): 140-150, (2002) SPIDERS (ARANEAE) AS POTENTIAL PREDATORS OF LEAFROLLER LARVAE AND EGG MASSES (LEPIDOPTERA: TORTRICIDAE) IN CENTRAL WASHINGTON APPLE AND PEAR ORCHARDS Eugene R. Miliczky and Carrol O. Calktns Yakima Agricultural Research Laboratory, United States Department of Agriculture—Agricultural Research Service, 5230 Konnowac Pass Road, Wapato, Washington 98951 Abstract .—Eleven species of arboreal, hunting spiders, common in central Washington apple and pear orchards, were evaluated as potential predators of the tortricid leafrollers, Pandemis pyru- sana Kearfott and Choristoneura rosaceana (Harris), pests in Pacific Northwest orchards. All species fed on leaf roller larvae established on apple and pear seedlings or branches during small cage tests. Cheiracanthium mildei L. Koch was the most effective predator in these tests, con¬ suming 65% of larvae. C. mildei was also the most effective predator among six species used in tests where leafroller larvae were established on small, caged apple trees. Twelve species of arboreal, hunting spiders were tested as predators of C. rosaceana egg masses. C. mildei was the most effective egg mass predator and 35 of 112 individuals consumed part or all of an egg mass. In addition, Oxyopes scalaris Hentz and Cheiracanthium inclusum (Hentz) exhibited some feeding on eggs. Key words .—Arachnida, Araneae, biological control, Cheiracanthium mildei, leafrollers, or¬ chards, predation, spiders. Spiders are important predators of insects in most terrestrial habitats (Foelix 1996). Apple orchards heavily treated with synthetic, broad-spectrum insecticides have few spiders, but in orchards where these chemicals are not used spider numbers may be much higher (Mansour et al. 1980a, Madsen & Madsen 1982). Even limited use of synthetic, broad-spectrum insecticides in apple orchards re¬ sults in lower spider numbers compared to orchards where they are not used (Knight et al. 1997, Miliczky et al. 2000). Spiders have been observed feeding on a variety of apple and pear pest insects (Dondale 1956, Wisniewska and Prokopy 1997, ERM, personal observations) and evidence of their importance in control of some pest species has been presented. MacLellan (1973) showed that spiders were valuable predators of the light brown apple moth, Epiphyas postvittana Walker, in Australia and Mansour et al. (1980c) found that spider predation on the Egyptian cotton leafworm, Spodoptera littoralis (Boisduval), reduced damage to apple in Israel. The use of synthetic broad-spec¬ trum insecticides in central Washington apple and pear orchards has been decreas¬ ing in recent years as new pest control technologies, such as pheromone based mating disruption for codling moth, Cydia pomonella L., and specific insecticides, have been adopted (Knight 1995). As a result, spiders and other natural enemies may assume a greater role in orchard pest management programs. The leafrolling caterpillars, Pandemis pyrusana Kearfott and Choristoneura rosaceana (Harris) (both Tortricidae), are pests of apple, pear, and other fruit trees in eastern Washington. The bionomics of the two insects are similar as they have two generations per year and overwinter as second and third instar larvae. Both species lay compact masses of up to 300 eggs and, as larvae, roll leaves to form 2002 MILICZKY & CALKINS: SPIDER PREDATION ON LEAFROLLERS 141 protected feeding sites (Schuh & Mote 1948, Beers et al. 1993). Larvae feed primarily on foliage but also damage fruit by surface feeding (Beers et al. 1993). We report here the results of tests designed to evaluate the potential for pre¬ dation on larvae of P. pyrusana and C. rosaceana of 11 species of hunting spiders. Twelve species of hunting spiders were also tested as predators of C. rosaceana egg masses. Hunting spiders do not construct webs for prey capture but either actively seek prey or wait for prey to come to them (Wise 1993, Foelix 1996). All tested species occur in south-central Washington apple and pear orchards and are frequently found in the tree canopy where there is potential for contact with leaf rollers (Miliczky et al. 2000). Materials and Methods Greenhouse and laboratory tests were conducted at the USDA-ARS Yakima Agricultural Research Laboratory near Wapato, Washington. Tests were also con¬ ducted in an outdoor screenhouse and in small, experimental orchards at the USDA-ARS research farm, 26 km east of Yakima, Washington. Spiders used in the tests were collected in Yakima Co., Washington from apple and pear orchards and adjacent habitat. Spiders not used within two or three days of capture were maintained in the laboratory on live insects including field-col¬ lected Lygus spp. and laboratory reared Drosophilidae and larval codling moth. Spiders were not starved prior to use in a test. Some individuals were used in more than one test and some were transferred to a second or third cage during a given test. Small Cage Tests .—Small cage tests were conducted with various plant ma¬ terial, but in all cases infestation with leafroller larvae and introduction of spiders were as follows. Three to five leafroller larvae were placed on leaves of a test plant. Larval size was adjusted according to size of the spiders used in a given experiment because spiders generally take prey smaller than themselves (Wise 1993). Third instars were used in tests with small to medium size spiders; fourth to early fifth instars were used in tests with larger spiders. The plant was then enclosed in one of two types of cage. Cylindrical cages made of flexible, plastic screen (seven mesh per cm) were used in greenhouse, screenhouse, and orchard tests. Cages were 48 cm long and 16 cm in diameter. One end was equipped with a sewn-in, circular piece of screen with an opening for leafroller and spider in¬ troduction. The second type of cage, used in laboratory tests with potted seedlings, was constructed of clear, plastic sheeting (Mylar) formed into a cylinder 45 cm tall and 17.5 cm in diameter. The ends of the cylinder were inserted into the tops of paper ice cream cartons. The center of the carton forming the top of the cage was removed and replaced with fine mesh, organdy gauze for ventilation. Leaf- roller larvae were given one to three days to feed on host plants and establish leafrolls prior to spider introduction. Just before introduction of a spider, a plant was examined to determine the number of established leafroller larvae. Eleven species (five to 50 individuals per species) were evaluated in small cage tests. Evidence of predation was scored at two to four day intervals after spider introduction. Plants were examined and dead larvae, when found, were inspected under a microscope at 6X-50X to determine cause of death. Larvae killed by spiders showed one or more of the following features: 1) Head capsule and/or pronotum pierced one or more times by the spider’s chelicerae, 2) Body markedly 142 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(2) chewed up, shrunken, and discolored, 3) Larva torn into two pieces at some point along its length with one piece often missing, 4) Large wound(s) present with internal fluid/tissue exuding. Test larvae fed on by spiders in the laboratory showed the same features. Dead larvae that did not exhibit one or more of these features were assumed to have died from some other, usually unknown, cause. Often, little remained of a spider-killed larva except the exoskeleton and undi¬ gested plant material in the gut. Cast exoskeletons were readily distinguishable from spider-killed larvae. Tests ended when all larvae had been killed, died of other causes, remained unconsumed for several days, or pupated (predation on pupae was originally thought to be unlikely but did occur occasionally—see be¬ low). Maximum head capsule width was measured with an ocular micrometer for 109 spider-killed leaf roller larvae that were recovered (five P. pyrusana and 104 C. rosaceana). Head capsule widths of third instar and last instar larvae of P. pyrusana and C. rosaceana were measured for comparison with head capsules of spider-killed larvae. Twenty-four small cage tests were conducted using a variety of host plants and plant sizes. Twelve tests were conducted in the laboratory on pear seedlings grow¬ ing in 9 cm X 9 cm X 9 cm plastic pots and housed in the clear, plastic cages. One test was conducted in the greenhouse using potted apple seedlings (9 cm X 9 cm X 9 cm pots) covered with flexible screen cages. Six tests were conducted on small (2 m tall) apple trees (varieties Red and Golden Delicious) growing in large, plastic pots (55 cm diameter X 45 cm deep) in an outdoor screenhouse. Branches of appropriate length were enclosed in flexible, screen cages. Two tests were conducted in small, apple orchards (varieties Fuji and Golden Delicious) and three were conducted in a small, pear orchard (variety Bartlett). Branches of appropriate length were enclosed in flexible screen cages. Five to 20 cages were used per test depending on availability of spiders or potted plants. Pandemis pyrusana larvae were used in four laboratory and three field tests on pear and in two screenhouse tests on apple. Choristoneura rosaceana larvae were used in eight laboratory tests on pear and in one greenhouse, four screenhouse and two field tests on apple. Both species were from laboratory colonies main¬ tained on artificial diet. Control cages, seeded with leafroller larvae but without spiders, were main¬ tained in some tests to verify that larvae constructed leafrolls, fed on foliage, and developed normally under the experimental conditions. Prey presentation in small cage tests was designed to approximate the condi¬ tions a spider would encounter in the orchard in that larvae were allowed to establish leafrolls on live plants before spider introduction. However, the small volume of the cage and the high density of leafrollers on the short length of test branch may have made it more likely that a spider would encounter and attack a leafroller than would be the case under field conditions. Large Cage Tests .—These tests were conducted in an orchard of small Red Delicious apple trees and were designed to more closely approximate actual field conditions than the small cage tests. Trees were lightly pruned, if necessary, to fit inside 1.8 m X 1.8 m X 1.8 m screen cages supported by tubular, metal frames. One wall of a cage was zippered to allow access. Soil was piled around the bottom perimeter of each cage to help prevent escape of spiders and leafrollers. Prior to leafroller introduction, trees were beat with a stiff rubber hose to dislodge poten- 2002 MILICZKY & CALKINS: SPIDER PREDATION ON LEAFROLLERS 143 tial predators onto a 0.45 m 2 tray. Predators were then removed from the cage. Thirty-five C. rosaceana larvae (third and fourth instar) were seeded onto each of four trees used in a test. Larvae were placed on young, still-growing leaves. Two to four days later the number of established larvae was counted and spiders were introduced. Trees were inspected at two to four day intervals thereafter to determine the number of surviving leafrollers and to look for evidence of pre¬ dation: dead larvae and empty leafrolls. These were examined at 6X-50X. Tests were continued until surviving C. rosaceana had pupated. Spiders used in large cage tests were species judged to have the greatest po¬ tential as leafroller predators based on small cage test results. Three large cage tests were conducted. The first was run from 6 to 24 July 2000 and used females or large immatures of five species: Oxyopes scalaris Hentz (Oxyopidae) and four species of Salticidae, Eris militaris (Hentz), Phidippus audax (Hentz), P. clarus Keyserling, and P. comatus Peckham & Peckham. One individual of each species was placed in each of two cages. Two control cages received leafroller larvae only. The second test (31 July—22 August 2000) employed four female P. clarus in one cage, four female P. comatus in one cage, two female and two large immature Cheiracanthium mildei L. Koch (Clubionidae) in one cage, and one cage was a control. Five immature (ca. one-half grown) C. mildei were used in each of three cages for the third test (30 August-18 September 2000). The fourth cage was a control. Egg Mass Predation Tests .—Egg masses of C. rosaceana were used in all tests. Egg masses were presented to spiders in three ways. Initially, egg masses laid on wax paper were obtained from the rearing facility at the Yakima laboratory. Egg masses on wax paper were attached to pear leaves with double-sided tape in a pear orchard at the experimental farm. Branches with egg-bearing leaves were enclosed in flexible, screen cages and spiders introduced. Seven to ten individuals of three species were tested: Pelegrina aeneola (Curtis) (Salticidae), Misumenops lepidus (Thorell) (Thomisidae), and Philodromus cespitum (Walckenaer) (Phil- odromidae). Caged female moths were allowed to lay eggs on potted apple or pear seedlings for a second series of tests. Egg-bearing leaves were clipped and the petiole inserted through a hole in the cap of a water-filled, 145 ml plastic vial. The leaf blade was enclosed in a second vial attached above the water vial. The spider was housed in the upper vial, the end of which was screened for ventilation. Tests were continued until the eggs were consumed, reached the black-head stage, or hatched. Five to 100 individuals of the following species were tested: P. aeneola, P. comatus, P. clarus, P. audax, E. militaris, O. scalaris, C. mildei, Cheiracan¬ thium inclusum (Hentz), Xysticus cunctator Thorell (Thomisidae), and Anyphaena pacifica (Banks) (Anyphaenidae). A final series of tests utilized 30 cm tall, potted pear seedlings whose leaves bore one or more egg masses. Seedlings were enclosed in cylindrical plastic cages (described above) and a spider was introduced. Tests were continued as before. Twelve C. mildei were tested. Results Small Cage Tests .—Both leafroller species constructed normal leafrolls on apple and pear. Percent of larvae in control cages that survived to the end of the exper¬ iments was, however, about twice as high on apple as it was on pear (Table 1). 144 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(2) Seven of 11 spider species were tested on both apple and pear. Because pre¬ dation rates by these species were similar on both plants, results for all small cage tests are combined in Table 1. Predation rates on leafroller larvae among the 11 species ranged from 5% to 65% based on spider-killed larvae that were recovered and examined microscopically (Table 1). Table 1 also indicates that some larvae were unaccounted for at the ends of experiments based on the number of larvae that established leafrolls prior to spider introduction. Some of these larvae prob¬ ably abandoned their leafrolls and escaped from the cages or were otherwise unaccounted for. Some larvae, however, may have been killed by spiders but their remains were not recovered. If this was the case then predation rates may have been somewhat higher than those given in Table 1. Cheiracanthium mildei was the most effective predator of leafroller larvae. Predation rates by this species were the same on apple and pear (65%). Only two of 37 individuals failed to consume at least one larva and both escaped and were lost during testing. On the other hand, one adult female consumed 14 of 20 leaf rollers during the course of three separate tests. She ate larvae, including last instars, a pupa, and an emerged adult. A second adult female consumed all five larvae in one cage within three days and four of five larvae in a second cage within four days. Adult female C. mildei seemed especially voracious predators. Cheiracanthium inclusum was less effective, consuming 35% of larvae. Among the other species, the four largest salticids ( E. militaris and the three species of Phidippus) and the lynx spider, O. scalaris, consumed 28-51% of larvae. The remaining four species each consumed fewer than 20% of the larvae. Large Cage Tests. —Results of the large cage tests are summarized in Table 2. Leafroller establishment in the first test was high in all four cages, but predation by the spiders was low; one larva killed in one cage and two in the second. Low levels of parasitism by Colpoclypeus floras (Walker) (Hymenoptera: Eulophidae) occurred in all cages. Only four of 28 larvae were killed by P. comatus during the second large cage test and no predation by P. clarus was observed. Both species had performed much better in small cage tests. C. mildei, however, destroyed 10 of 27 larvae, 16 others were unaccounted for, and only one survived to the end of the test. Predation by C. mildei during the third, large cage test ranged from 12% to 29% and several larvae were unaccounted for in each cage. All larvae not killed by the spiders or unaccounted for were parasitized by C. floras and no leafrollers survived to the end of the test, including those in the control cage. Sizes of leafroller larvae preyed upon by four spider species and by different sizes of individuals within a species overlapped considerably (Table 3). Mean head capsule width of third instar P. pyrusana larvae was 0.55 mm (range: 0.47— 0.58 mm; n = 23) and mean width of last instars was 1.66 mm (range: 1.41-1.84 mm; n = 19). Corresponding dimensions for C. rosaceana were slightly larger: 0.59 mm (range: 0.57-0.61 mm; n — 4) for third instar and 1.77 mm (range: 1.46-2.00 mm; n = 6) for last instar larvae. All species and size classes of spiders, except male P. comatus, preyed on leafroller larvae that were within the size ranges of last instar P. pyrusana and C. rosaceana. Earlier instars were also preyed upon (Table 3). These four spider species were among the largest tested and most individuals were adults or large immatures (>Vi grown) when used in the experiments. Table 1. Small cage test results. Spiders tested for feeding propensity on two species of leafroller larvae (LR) in laboratory, greenhouse, field cage, and field situations on apple and pear foliage. Spider species Spider family No. individuals tested No. LRs available % LRs eaten % LRs not accounted for % LRs survived Cheiracanthium mildei 3 Clubionidae 37 204 65 25 10 Eris militaris 3 Salticidae 15 69 51 23 26 Phidippus comatus 3 Salticidae 50 193 41 35 24 Oxyopes scalaris a Oxyopidae 11 47 40 13 47 Phidippus clarus 3 Salticidae 45 157 36 28 36 Cheiracanthium inclusum 3 Clubionidae 14 62 35 23 42 Phidippus audax 3 Salticidae 5 18 28 56 17 Philodromus californicus 0 Philodromidae 7 33 18 45 36 Phanias sp. b Salticidae 7 31 13 42 45 Pelegrina aeneola 0 Salticidae 19 43 7 44 49 Anyphaena pacified 0 Anyphaenidae 9 38 5 50 45 Control (apple) — — 51 — 18 82 Control (pear) — — 20 — 55 45 a Species tested on both apple and pear. b Species tested on pear only. c Species tested on apple only. 2002 MILICZKY & CALKINS: SPIDER PREDATION ON LEAFROLLERS 145 Table 2. Large cage test results. Fates of leafroller larvae on small, caged apple trees when exposed to different species of spiders. See text for details. Contents of cage No. LRs established % LRs eaten % LRs not accounted for % LRs parasitized % Other mortality % LRs survived Spider 1 31 3 Large Cage Test #1 26 3 3 65 Spider 2 31 6 0 6 6 81 Control 1 31 — 10 13 10 68 Control 2 28 — 11 4 0 86 P. comatus 28 14 Large Cage Test #2 18 0 7 61 P. clarus 26 0 0 0 0 100 C. mildei 27 37 59 0 0 4 Control 22 -—• 50 0 0 50 C. mildei 1 31 26 Large Cage Test #3 26 48 0 0 C. mildei 2 25 12 28 60 0 0 C. mildei 3 31 29 16 55 0 0 Control 18 — 17 83 0 0 <1 oo 146 THE PAN-PACIFIC ENTOMOLOGIST Vol. 2002 MILICZKY & CALKINS: SPIDER PREDATION ON LEAFROLLERS 147 Table 3. Head capsule widths of leafroller larvae consumed by four spider species and different size classes within spider species during predation experiments. Spider species Spider stage (No. tested) Mean spider carapace width 1 Mean LR head width Range in LR head widths No. LRs measured C. mildei female (4) 2.56 mm (10) 1.41 mm 1.1-1.7 mm 7 C. mildei sub-female 2 (2) — 1.44 mm 1.0-1.8 mm 5 C. mildei immatures 3 (8) — 1.20 mm 0.8-1.7 mm 28 C. inclusum immatures (4) _4 1.14 mm 0.8-1.7 mm 6 P. clarus female (1) 3.49 mm (10) 1.40 mm 0.7-1.7 mm 5 P. clarus sub-female (2) — 1.65 mm 1.5-1.8 mm 4 P. clarus immatures (10) — 1.29 mm 1.0-1.8 mm 22 P. comatus sub-female (6) 2.60 mm (2) 5 1.52 mm 1.0-1.9 mm 10 P. comatus male (3) 2.94 mm (10) 1.10 mm 1.0-1.2 mm 7 P. comatus sub-male (7) 1.95 mm (1) 1.33 mm 0.9-1.6 mm 13 P. comatus sub-sub-male (2) — 1.55 mm 1.4-1.7 mm 2 1 Number of specimens measured is given in parentheses. 2 Sub-female and sub-male spiders are one molt from reaching adulthood. Sub-male status deter¬ mined by enlarged but undifferentiated, terminal segment of the pedipalp. Sub-female status deter¬ mined by subsequent rearing. 3 Immature spiders of all species were approximately one-half grown or larger but do not include sub-adults. 4 Mean carapace width of adult female C. inclusum = 2.28 mm (n = 4). Immatures used here were approximately one-half grown. 5 Probable sub-females. Mean width of adult female = 3.23 mm (n = 10). Egg Mass Predation. —No predation was observed on egg masses laid on wax paper and taped to leaves of pear trees by the three species tested in this way. Also, seven of ten species did not feed when housed in small, plastic vials with single apple or pear leaves on which an egg mass had been laid. However, two of 25 O. scalaris and one of seven C. inclusum consumed part or all of an egg mass during single leaf tests. C. mildei was the most effective egg predator in single leaf tests and 29 of 100 individuals fed on egg masses, often consuming them in their entirety within 24 hours. Eighteen of the 100 C. mildei were tested twice. Eleven failed to feed either time, six fed once, and one fed on eggs in both tests. Unconsumed eggs in a fed-upon mass often developed normally. Six of 12 C. mildei also fed on eggs laid on leaves of 30 cm tall pear seedlings. Two of these spiders consumed three masses each and a third individual consumed two masses and part of a third. Discussion Several spider species that occur in central Washington orchards preyed on leafroller larvae during these tests. C. mildei was the most effective predator in both small and large cage tests. This species is a native of the Mediterranean region and was probably introduced into North America (Edwards 1958). It is widely distributed in central Washington orchards (Miliczky et al. 2000, E.R.M. personal observation). C. mildei is a long-legged, swift running, nocturnal hunter that reaches an adult body length of 10 mm (Dondale & Redner 1982). Spiders were often found in silken retreats on leaves during daytime cage inspections. Other studies have indicated the importance and versatility of C. mildei as a predator of pest insects. It was the dominant spider in an unsprayed apple orchard 148 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(2) in Israel where it was an effective predator of the Egyptian cotton leaf worm, Spodoptera littoralis (Boisduval), (Mansour et al. 1980a). Specialized predation on cryptic larvae of the leaf miner, Phyllonorycter blancardella (F.), was reported by Corrigan & Bennett (1987). They observed small holes in the lower surface of leaf mines, from which larvae were absent, and attributed the holes and absence of larvae to predation by C. mildei. Our unpublished observations showed that C. mildei employed specialized behaviors when attacking leafroller larvae includ¬ ing the cutting of small holes in rolled leaves. Wise (1993) speculated that C. mildei, because of its abundance and effective hunting techniques, may make important contributions to pest suppression in a number of agroecosystems. C. inclusum, a species native to North America (Edwards 1958), was found less frequently in central Washington orchards than C. mildei (Miliczky et al. 2000, ERM personal observation). The two species are similar in size and ap¬ pearance. C. inclusum is a common spider in western Oregon apple orchards (Bajwa & AliNiazee 2001) and is also abundant in central California vineyards (Costello & Daane 1999). Peck & Whitcomb (1970) described C. inclusum as “an indiscriminate and voracious feeder” that accepted a wide range of insect prey in laboratory trials including larvae of several species of Lepidoptera. It may be of importance as a predator of citrus leafminer, Phyllocnistis citrella Stainton, in Florida citrus groves (Amalin et al. 2001). C. inclusum' s predation rate in small cage tests (35%) was about half that recorded for C. mildei. Several spider species showed high predation rates on leafroller larvae in small cage tests but consumed few, if any, larvae in large cage tests. These spiders, the salticids, P. comatus, P. clarus, P. audax, and E. militaris, and the oxyopid, O. scalaris, are all diurnal hunters with good eyesight and visually oriented hunting strategies (Wise 1993). Phidippus spp. are heavy-bodied, hairy, often colorful spiders whose adult body lengths frequently exceed 10 mm (Kaston 1978). E. militaris and O. scalaris are smaller. Observations of predatory behavior in these species showed little tendency to invade leafrolls and extract larvae. Rather, they appeared opportunistic in their predatory behavior and were quick to snatch ex¬ posed larvae. In small cages where prey densities were high, these spiders may have had frequent opportunities for predation on exposed or partially exposed larvae. Similar opportunities may have been infrequent in the large cages where leafroller densities were lower and the trees presented a much greater area to be searched for prey. Spider predation on insect eggs appears to be quite common; members of the Clubionidae, Oxyopidae, Salticidae, Lycosidae, and Anyphaenidae are the most frequently reported egg predators; and eggs of Lepidoptera are most commonly preyed upon (Nyffeler et al. 1990). Predation on an egg mass of the eastern spruce budworm, Choristoneura fumiferana (Clemens), by the jumping spider, Pelegrina flavipedes (G. & E. Peckham), was reported by Jennings & Houseweart (1978). C. mildei was quite an effective predator of leafroller egg masses in these tests where search area was limited to a single leaf or a small seedling. Some individ¬ uals, however, failed to feed on eggs even after four or five days in close proximity to them. Mansour et al. (1980b) noted that C. mildei spiderlings fed on infertile, conspecific eggs within the egg sac, a behavior also noted for C. inclusum (Peck & Whitcomb 1970). One of seven C. inclusum fed on a leafroller egg mass in our tests, and the species has also been found to prey on eggs of the tobacco 2002 MILICZKY & CALKINS: SPIDER PREDATION ON LEAFROLLERS 149 budworm, Heliothis virescens (Fabricius), (McDaniel & Sterling 1982) and the velvetbean caterpillar, Anticarsia gemmatalis Hubner (Buschman et al. 1977). The tests described above show that among several species of hunting spiders found in central Washington orchards a range of abilities as predators of leafroller larvae exists. Some species showed little potential whereas C. mildei was quite effective, a conclusion supported by the consistent performance of numerous in¬ dividuals during small cage tests and C. mildeV s markedly better performance in the large cage tests compared to other species. C. mildei may also be quite an effective egg mass predator. Although leaf area searched during egg predation tests was tiny compared to that of an entire tree, the belief that egg mass predation by C. mildei may occur in the field is strengthened by the fact that 35 of 112 individuals consumed eggs whereas only three individuals among the other 11 species (149 total individuals) did so. In orchards where C. mildei and some of the other species are present, spiders may contribute substantially to natural con¬ trol of leafroller pests. Acknowledgment We would like to thank Kathie Johnson and Jeanine Jewet for supplying leaf- roller larvae and egg masses. Debee Broers supplied apple and pear seedlings. Jerry Gefre and John Harvey helped get the large cages into working order. Mer- ilee Bayer provided able technical assistance. Alan Knight provided the flexible screen cages. G. B. Edwards identified Phidippus comatus. Dan Mayer, Alan Knight, and two anonymous reviewers read earlier versions of the manuscript and we thank them for their constructive comments. Partial funding for this project was provided by the Washington Tree Fruit Research Commission. Literature Cited Amalin, D. M., J. E. Pena & R. McSorley. 2001. Predation by hunting spiders on citrus leafminer, Phyllocnistis citrella Stainton (Lepidoptera: Gracillariidae). J. Entomol. Sci. 36: 199-207. Bajwa, W. I. & M. T. AliNiazee. 2001. A survey of spiders in apple trees in the Willamette Valley of Oregon. J. Entomol. Sci. 36: 214-217. Beers, E. H., J. F. Brunner, M. J. Willett & G. M. Warner, (eds.). 1993. Orchard pest management: a resource book for the Pacific Northwest. Good Fruit Grower, Yakima, Washington. Buschman, L. L., W. H. Whitcomb, R. C. Hemenway, D. L. Mays, N. Ru, N. C. Leppla & B. J. Smittle. 1977. Predators of velvetbean caterpillar eggs in Florida soybeans. Environ. Entomol. 6; 403-407. Corrigan, J. E. & R. G. Bennett. 1987. Predation by Cheiracanthium mildei (Araneae, Clubionidae) on larval Phyllonorycter blancardella (Lepidoptera, Gracillariidae) in a greenhouse. J. Arach- nol. 15: 132-134. Costello, M. J. & K. M. Daane. 1999. Abundance of spiders and insect predators on grapes in central California. J. Arachnol. 27: 531-538. Dondale, C. D. 1956. Annotated list of spiders (Araneae) from apple trees in Nova Scotia. Can. Entomol. 88: 697-700. Dondale, C. D. & J. H. Redner. 1982. The insects and arachnids of Canada. Part 9. The sac spiders of Canada and Alaska (Araneae: Clubionidae and Anyphaenidae). Agric. Can. Publ. No. 1724. Edwards, R. J. 1958. The spider subfamily Clubioninae of the United States, Canada, and Alaska (Araneae: Clubionidae). Bull. Mus. Comp. Zool. 118: 365-436. Foelix, R. F. 1996. Biology of spiders (2nd ed.) Oxford University Press. New York. Jennings, D. T. & M. W. Houseweart. 1978. Spider preys on spruce budworm egg mass. Entomol. News. 89: 183-186. Kaston, B. J. 1978. How to know the spiders (3rd ed.). Wm. C. Brown Co. Dubuque, Iowa. 150 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(2) Knight, A. L. 1995. The impact of codling moth (Lepidoptera: Tortricidae) mating disruption on apple pest management in Yakima Valley, Washington. J. Entomol. Soc. Brit. Columbia. 92: 29-38. Knight, A. L., J. E. Turner & B. Brachula. 1997. Predation on eggs of codling Moth (Lepidoptera: Tortricidae) in mating disrupted and conventional Orchards in Washington. J. Entomol. Soc. Brit. Columbia. 94: 67-74. MacLellan, C. R. 1973. Natural enemies of the light brown apple moth, Epiphyas postvittana, in the Australian Capital Territory. Can. Entomol. 105: 681-700. Madsen, H. F. & B. J. Madsen. 1982. Populations of beneficial and pest arthropods in an organic and a pesticide treated apple orchard in British Columbia. Can. Entomol. 114: 1083-1088. Mansour, F., D. Rosen & A. Shulov. 1980a. A survey of spider populations (Araneae) in sprayed and unsprayed apple orchards in Israel and their ability to feed on larvae of Spodoptera littoralis (Boisd.). Acta Oecologica/Oecol. Applic. 1: 189-197. Mansour, F., D. Rosen & A. Shulov. 1980b. Biology of the spider Chiracanthium mildei (Arachnida: Clubionidae). Entomophaga. 25: 237-248. Mansour, F., D. Rosen, A. Shulov & H. N. Plaut. 1980c. Evaluation of spiders as biological control agents of Spodoptera littoralis larvae on apple in Israel. Acta Oecologica/Oecol. Applic. 1: 225-232. McDaniel, S. G. & W. L. Sterling. 1982. Predation of Heliothis virescens (F.) eggs on cotton in east Texas. Environ. Entomol. 11: 60-66. Miliczky, E. R., C. O. Calkins & D. R. Horton. 2000. Spider abundance and diversity in apple orchards under three insect pest management programmes in Washington State, U.S.A. Ag. For. Entomol. 2: 203-215. Nyffeler, M., R. G. Breene, D. A. Dean & W. L. Sterling. 1990. Spiders as predators of arthropod eggs. J. Appl. Entomol. 109: 490-501. Peck, W. B. & W. H. Whitcomb. 1970. Studies on the biology of a spider, Chiracanthium inclusum (Hentz). Bull. Univ. Arkansas Agric. Exp. Stn. 753. Schuh, J. & D. C. Mote. 1948. The oblique-banded leaf roller on red raspberries [ Archips rosaceana (Harris)]. Oregon Agric. Exp. Stn. Tech. Bull. 13. Wise, D. H. 1993. Spiders in ecological webs. Cambridge University Press, Cambridge, U.K. Wisniewska, J. & R. J. Prokopy. 1997. Do spiders (Araneae) feed on rose leafhopper ( Edwardsiana rosae\ Auchenorrhyncha: Cicadellidae) pests of apple trees? Eur. J. Entomol. 94: 243-251. Received 10 April 2001; Accepted 6 March 2002 PAN-PACIFIC ENTOMOLOGIST Information for Contributors See volume 74: 248-255, October 1997, for detailed general format information and the issues thereafter for examples; see below for discussion of this journal’s specific formats for taxonomic manuscripts and locality data for specimens. 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Blackman, R. L., P. A. Brown & V. F. Eastop. 1987. Problems in pest aphid taxonomy: can chromosomes plus morphometries provide some answers? pp. 233-238. In Holman, J., J. Pelikan, A. G. F. Dixon & L. Weismann (eds.). Population structure, genetics and taxonomy of aphids and Thysanoptera. Proc. international symposium held at Smolenice Czechoslovakia, Sept. 9-14, 1985. SPB Academic Publishing, The Hague, The Netherlands. Ferrari, J. A. & K. S. Rai. 1989. Phenotypic correlates of genome size variation in Aedes albopictus. Evolution, 42: 895-899. Sorensen, J. T. (in press). Three new species of Essigella (Homoptera: Aphididae). Pan-Pacif. Entomol. Illustrations. — Illustrations must be of high quality and large enough to ultimately reduce to 117 X 18 1 mm while maintaining label letter sizes of at least 1 mm; this reduction must also allow for space below the illustrations for the typeset figure captions. 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Authors and affiliations are listed in the last, left indented paragraph of the note with the affiliation underscored. Page Charges. — PCES members are charged $35.00 per page, for the first 20 (cumulative) pages per volume and full galley costs for pages thereafter. Nonmembers should contact the Treasurer for current nonmember page charge rates. Page charges do not include reprint costs, or charges for author changes to manuscripts after they are sent to the printer. Contributing authors will be sent a page charge fee notice with acknowledgment of initial receipt of manuscripts. I Volume 78 THE PAN-PACIFIC ENTOMOLOGIST April 2002 Number 2 Contents POTAPOV, M. & M. CULIK-A new species of Folsomia (Collembola: Isotomidae) from Brazil, with notes on foil-setae in the Fimetaria group - 69 HSU, L.-P. & C.-S. CHEN-A new species of Ugandatrichia (Trichoptera: Hydroptilidae) from Taiwan- 74 GE, S.-Q., S.-Y. WANG, X.-K. YANG, & W.-Z. LI-A revision of the genus Agrosteella Medvedev (Chrysomelidae: Chrysomelinae) -- 80 GILBERT, A. J. & F. G. ANDREWS—Studies on the Chrysomelidae (Coleoptera) of the Baja California Peninsula: the genus Dysphenges Horn (Galerucinae: Alticini)- 88 BRENNER, G. J„ P. T. OBOYSKI, & P. C. BANKO-Parasitism of Cydia spp. (Lepidoptera: Tortricidae) on Sophora chrysophylla (Fabaceae) along an elevation gradient of dry subalpine forest on Mauna Kea, Hawaii - 101 BRAILOVSKY, H.—Two new species of Mictis Leach (Heteroptera: Coreidae: Mictini) from Sulawesi- 110 XU, X., C.-M. YIN, & C. E. GRISWOLD-A new species of the spider genus Macrothele from the Gaoligong Mountains, Yunnan, China (Araneae: Hexathelidae) - 116 DODDS, K. J. & D. W. ROSS-Relative and seasonal abundance of wood borers (Buprestidae, Cerambycidae) and Cucujidae trapped in Douglas-Fir beetle pheromone- baited traps in northern Idaho - 120 HSU, Y.-F.—Larval and pupal biology of a new sun moth in southern California; novel host use strategy in the evolution of Heliodinidae (Lepidoptera: Yponomeutoidea). - 132 MILICZKY, E. R. & C. O. CALKINS—Spiders (Araneae) as potential predators of leafroller larvae and egg masses (Lepidoptera: Tortricidae) in central Washington apple and pear orchards - 140 The PAN-PACIFIC ENTOMOLOGIST Volume 78 July 2002 Number 3 Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES (ISSN 0031-0603) The Pan-Pacific Entomologist EDITORIAL BOARD R. M. Bohart J. T. Doyen J. E. Hafernik, Jr. Warren E. S a vary Published quarterly in January, April, July, and October with Society Proceedings usually appearing in the following October issue. 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See the back cover for Information-to-Contributors, and volume 73(4): 248-255, October 1997, for more detailed information. Information on format for taxonomic manu¬ scripts can be found in volume 69(2): 194-198. Refer inquiries for publication charges and costs to the Treasurer. The annual dues, paid in advance, are $25.00 for regular members of the So¬ ciety, $26.00 for family memberships, $12.50 for student members, or $40.00 for institutional subscriptions or sponsoring members. Members of the Society receive The Pan-Pacific Entomologist. Single copies of recent numbers or entire volumes are available; see 72(4): 247 for current prices. Make checks payable to the Pacific Coast Entomological Society. J. Y. Honda, Editor R. E. Somerby, Book Review Editor Robert Zuparko, Treasurer Pacific Coast Entomological Society OFFICERS FOR 2002 Katherine N. Schick, President Vincent F. Lee, Managing Secretary Rolf L. Aalbu, President-elect Richard M. Brown, Recording Secretary Robert L. Zuparko, Treasurer THE PAN-PACIFIC ENTOMOLOGIST (ISSN 0031-0603) is published quarterly for $40.00 per year by the Pacific Coast Entomological Society, % California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599. Periodicals postage is paid at San Francisco, CA, and additional mailing offices. POSTMASTER: Send address changes to the Pacific Coast Entomological Society, % California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599. This issue mailed 8 November 2002 The Pan-Pacific Entomologist (ISSN 0031-0603) PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, U.S.A. © The paper used in this publication meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PAN-PACIFIC ENTOMOLOGIST 78(3): 151-167, (2002) OBITUARY AND BIBLIOGRAPHY OF KENNETH S. HAGEN (1919-1997), DEDICATED ENTOMOLOGIST AND TEACHER Robert L. Zuparko Essig Museum of Entomology, 201 Wellman Hall, University of California, Berkeley, California 94720 Kenneth Sverre Hagen, professor emeritus of Entomology at the University of California, Berkeley, and past president of the Pacific Coast Entomological So- 152 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) Figure 1. Collecting trip in Sunset Valley, near Santa Barbara, California in 1939, with (L-R) Edwin Van Dyke, Burdett White, Bill Barr & Ken Hagen. ciety, died suddenly of a ruptured aortic aneurism on 10 Jan 1997. He is survived by his wife Maxine, his son Kent, and his brother Paul. Worldwide he was re¬ garded as an authority on biological control and insect nutrition. Those who worked with him, also regarded him as the first one to go to for an identification, a literature source, or a cup of coffee. Ken was born in Oakland, California on 26 Nov 1919. His parents were from Norway, and his father was a seaman, serving on such ships as the Balclutha, and eventually reached the rank of first mate. Ken’s mother passed away when he was a teenager, and he and his brother had to fend for themselves when their father was away on a voyage. As a boy, Ken enjoyed chemistry and natural history, and collecting insects was a favorite pastime, but it was a favored teacher in the 9th grade at Lockwood Junior High School who recognized his talents and stimulated him to focus on science as a career. Ken attended Fremont High School in Oakland, where he played football and set a high jump record in track. He graduated from Fremont High in 1938, and then enrolled in San Francisco State College. During this time, his love of natural history led him to the California Academy of Sciences, first as a part-time preparator of insects, and later as an assistant caretaker in the Steinhart Aquarium. He went collecting with Edwin van Dyke, and climbed Mt. Whitney with his fellow coleopterist, Bill Barr. At San Francisco State he contin¬ ued to play football, and received his A.A. degree in 1942. Ken then attended U.C. Berkeley, where he was offered a football scholarship. However, Ken did not play football at Cal, but concentrated on his studies (taking up to 24 units a term) to earn his B.S. in entomology in 1943. He then went to 2002 ZUPARKO: HAGEN OBITUARY 153 Officer Candidate School at Columbia University, where was commissioned in the U.S. Navy as one of the “90 day wonders”. Ken was then given a brief leave, which he used to return to Oakland and marry his fiancee, Maxine White, on 1 Dec 1943. A week later he went to Norfolk, Virginia to attend Amphibious Train¬ ing School, and then was shipped out to Europe. During the war he served on the USS Anne Arundel, as a lieutenant in charge of a landing craft section, and saw action in the Neptune Invasion at Omaha Beach in Normandy, and the Dragon Invasion in the south of France in 1944. In 1945, he participated in the landings at Okinawa (where the fierce fighting strand¬ ed his boat on the beach overnight), and later helped transport Chinese troops. He developed quite a reputation among his comrades for his entomological in¬ terests—en route for the Pacific, Ken was seen leaning over the rail with a net, sweeping the vegetation while passing through the Panama Canal. In 1946, Ken came back to California and was hired as the supervising ento¬ mologist for the West Side Alfalfa Pest Control Association in California’s Central Valley, responsible for overseeing 10,000 acres of alfalfa, and becoming the first supervised control entomologist in California. This position played a key role in the development of integrated pest management, and was the predecessor of to¬ day’s pest control advisor. Ken then returned to Berkeley as a graduate student, working as a technician in the Division of Biological Control. He received his M.S. there in 1948, and his Ph.D. in 1952, under the direction of Richard Doutt. This was a particularly rich time to be at Berkeley, as Ken studied under such luminaries as E. O. Essig, E. G. Linsley, R. L. Usinger and A. E. Michelbacher. He also spent a year in Oahu working on the oriental fruit fly with Robert van den Bosch, and worked in the statewide Department of Biological Control under Harry Scott Smith, whose signed photograph was one of Ken’s treasured posses¬ sions. He was appointed Junior Entomologist in the Division of Biological Con¬ trol, Agricultural Experiment Station (at the Gill Tract in Albany, California) in 1952, and advanced to Entomologist in 1965, and to Professor of Entomology in 1969. Ken took a special leave of absence from 1961 to 1963 to work for the government of Greece to advise and develop culturing techniques for the olive fly. He officially retired in 1990, but continued to work at the Gill Tract until the day of his death. It was remarked that the way you knew Ken was retired was that he only worked half a day on Saturday. Biological control was his passion as well as his profession. Biocontrol has been subdivided into the three tactics of importation, conservation and augmen¬ tation. Evidence of Ken’s solid training and great command of the field was that he was well-versed in all three. Besides publishing on the history of biological control, Ken was involved in the importation of the natural enemies of pear psylla, acacia psyllid, spotted alfalfa aphid, blue alfalfa aphid, pea aphid, walnut aphid, plum aphid, european asparagus aphid, iceplant scales, Egyptian alfalfa weevil and walnut husk fly. He was familiar with the conservation of the natural enemies through his work on population monitoring and reducing insecticide usage for Colias caterpillars and aphids in alfalfa. However, it was in the area of augmentation of natural enemies, coupled with insect nutrition, that Ken made his most important contributions to science. He was the first to develop an “artificial egg” for the mass-rearing of Chrysoperla, and helped develop artificial diets for mass-rearing Trichogramma, coccinellids 154 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) Figure 2. Some pioneers of modern biological control in Palm Canyon, Riverside County, Cali¬ fornia in 1948 (L-R) (front row) Stan Flanders, Glen Finney, Charles Fleschner, P. H. Timberlake, Everett Dietrick & Paul DeBach; (second row) Blair Bartlett, Ken Hagen, Prof. Harry Smith, Dave Lloyd, Harold Compere & Ed Steinhaus; (third row) Carl Huffaker, Ken Hughes (kneeling), Ted Fisher & A. J. Baisinger. and tephritids. His innovative work, with Richard Tassan, on food sprays for predators was a major breakthrough in biological control and continues to serve as a basic example for augmentation of field populations of entomophagous in¬ sects. Ken considered that his most significant research contribution was presented in a 1986 paper, wherein he hypothesized that the occurrence of amino acids in honeydew helped protect honeydew producers from ant predation, and presented data showing that chrysopids were attracted to a combination of plant volatiles and kairomones from honeydew, but the attraction varied with the age of the crop. Ken was truly a scientist of international stature and experience. He engaged in collaborative research in Mexico, Central America, Brazil, Greece, Kenya and China, but his travels also extended through Europe to India, Malaysia, Australia, New Zealand and Chile. Of the 22 visiting scientists and postdoctoral students he hosted in his lab, 18 were from other countries, and of the 28 graduate students he supervised, eight were from other countries, while he was an external examiner of dissertations of another ten students from outside the United States. Ken’s research interests extended beyond biological control, including aquatic Hymenoptera and the immature stages of Hymenoptera, but especially the bio- systematics of Hymenoptera (Encyrtidae) and Coleoptera (Coccinellidae and An- thicidae). His work with the Coccinellidae included documenting the complex migratory behavior of the convergent ladybeetle, which involved the use of hot air balloons and scoops fitted onto fixed wing aircraft to sample airborne beetles. This work led to an article in the National Geographic (1970) entitled “Following 2002 ZUPARKO: HAGEN OBITUARY 155 Figure 3. At Gill Tract, Albany, California in 1985: (L-R) Chuck Kennett, Carl Huffaker, Dick Doutt & Ken Hagen. the ladybug home”. Ken was particularly pleased with that issue, since it also included an article on his ancestors, the Vikings. He codescribed Karpinskiella paratomicobia (Hymenoptera: Pteromalidae) (Hagen & Caltagirone 1968), and had the following patronyms named in his honor: Notoxus hageni (Coleoptera: Anthicidae) (Chandler 1982), Gnathoweisea hageni (Coleoptera: Coccinellidae) (Gordon 1985), Olla hageni (Coleoptera: Coc- cinellidae) (Vandenberg 1992), Meleoma kennethi (Neuroptera: Chrysopidae) (Tauber 1969), Metaphycus hageni (Hymenoptera: Encyrtidae) (Daane & Calta¬ girone, 1999), and the Hagen glands in Braconidae (Hymenoptera) (Buckingham & Sharkey 1988). Ken was a member of the Entomological Society of America (president of the Pacific Branch in 1979 and fellow), American Entomological Society, Entomo¬ logical Society of Canada (fellow), Pacific Coast Entomological Society (president 1968-69 and Honored Member), Entomological Society of Washington, Kansas Entomological Society, Hawaiian Entomological Society, Georgia Entomological Society, Society of Systematic Zoology, the Coleopterists Society, American As¬ sociation for the Advancement of Science (fellow), American Institute of Biolog¬ ical Sciences, International Society of Hymenopterists, and the International Or¬ ganization of Biological Control (president 1980-84). He was honored at the 1989 national meeting of the Entomological Society of America with a symposium entitled “Native and Introduced Predaceous Cocci¬ nellidae: A Tribute to Kenneth S. Hagen for His Contributions to Coccinellid Biology”. In 1990 he was the recipient of the prestigious Berkeley Citation pre¬ sented by the University of California, Berkeley, for outstanding service to the 156 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) University, and honored by the California State Senate Rules Committee Reso¬ lution #2513. In 1992 he received the Distinguished Service Award by the As¬ sociation of Applied Insect Ecologists and the Lifetime Excellence in Entomology from the Hawaiian Entomological Society, and in 1993, the Distinguished Service Award (Honored Member) by the Pacific Coast Entomological Society. In 1995 the International Organization of Biological Control presented Ken with the Dis¬ tinguished Biological Control Science Award, and he presented an invitational talk on the Chemical Ecology of Chrysopidae at the IOBC’s conference honoring him. In 1998, a review of forage alfalfa pest management was dedicated to Ken (Summers 1998). Irrespective of these many scientific honors, Ken Hagen was probably best known among his colleagues for several personal traits. First, he always kept a pot of coffee going in his lab, and this served as a focal point for staff and visitors to drop in and discuss entomology. Second, he had a virtual encyclopedic knowl¬ edge of entomology and biological control. At the Gill Tract, it was generally understood that if you had a question, your first stop should be Hagen’s office. And if he didn’t immediately know the answer to the question, as often as not, he was able to swivel around in his chair, and from his immense reprint collection pick out the appropriate reference. Finally, he was extremely generous with his time and knowledge. No matter who approached him, be it a professor, graduate student, staff personnel, farmer or member of the general public, Ken would be happy to lay aside whatever he was working on, and give that person his full attention until he got the answer, or could refer the person to the correct authority. And if the search dragged on, it did no good to tell Ken to forget it—he just “hung in there” and kept looking for your answer. Ken was also popular with the local elementary school teachers, taking out the young students to the Gill Tract’s alfalfa field and showing them how to sweep for insects. To Ken, this commitment to teach others about entomology was as natural as can be, possibly reflecting his own debt of gratitude to those teachers who helped him, and he willed his substantial entomological library to the Division of Biological Control. Ken liked working with wood and was a fine carpenter. He was also interested in sailing, stamp collecting, astronomy, and (due to his studies of anthicids) sand dunes. However, outside of entomology, Ken’s greatest interest was book col¬ lecting. He was a keen bibliophile, and would bind his own books. His book and journal collection eventually outgrew his house, and when the house next to his came up for sale, Ken and Maxine ended up buying it, largely to use the garage as a storage space for his overflowing library. A tireless researcher, a loyal and dedicated member of the University of Cali¬ fornia faculty, an enthusiastic teacher, a helpful and stimulating colleague, and a generous human being, Ken Hagen was, in every sense of the word, a true gen¬ tleman. Bibliography Of K.S. Hagen I have attempted to include all of Ken’s writings to indicate the breadth of his interests. This list includes governmental reports, abstracts and non-scientific works which may not qualify as “published scientific articles”. Except for those indicated with by asterisk (*), all items have been checked against the originals. 2002 ZUPARKO: HAGEN OBITUARY 157 1. 1946. The occurrence of Ceutorhynchus assimili (Paykul) in California. Pan- Pacif. Ent. 22: 73. 2. 1949. Two new ichneumonid host records. Pan-Pacif. Ent. 25: 25. 3. 1949. (R. L. Doutt & K.S.H.) Periodic colonization of Chrysopa californica as a possible control of mealybugs. J. Econ. Ent. 42: 560. 4. 1949. (R. L. Doutt & K.S.H.) Baker mealybug use of green lacewing in control studied. Calif. Agric. 3 (3): 7. 5. 1950. (R. L. Doutt & K.S.H.) Biological control measures applied against Pseudococcus maritimus on pears. J. Econ. Ent. 43: 94-96. 6. 1950. Fecundity of Chrysopa californica affected by synthetic foods. J. Econ. Ent. 43: 101-104. 7. 1950. (with R. L. Doutt) Brontispa yoshinoi Barber, a description of adult and immature stages. Ann. ent. Soc. Am. 43: 311-319. 8. 1950. (with G. L. Finney) A food supplement for effectively increasing the fecundity of certain tephritid species. J. Econ. Ent. 43: 735. 9. 1952. Influence of adult nutrition upon fecundity, fertility, and longevity of three tephritid species. Ph.D. Thesis, University of California, Berkeley, Cal¬ ifornia. 10. 1953. A premating period in certain species of the genus Opius. Proc. Ha¬ waii ent. Soc. 15: 115-116. 11. 1953. (S. Maeda, K.S.H. & G. L. Finney) Artificial media and the control of microorganisms in the culture of tephritid larvae. Proc. Hawaii ent. Soc. 15: 177-185. 12. 1953. Influence of adult nutrition upon the reproduction of three fruit fly species, pp. 72-76. In California Senate, 3rd special report Joint Legislative Committee on Agricultural Livestock Problems on Control of the Oriental Fruit Fly. 13. *1953. (S. Maeda, K.S.H. & G. L. Finney) Role of microorganisms in the culture of fruit fly larvae, pp. 84-86. Ibid. 14. 1954. (R. van den Bosch, E. J. Dietrick & K.S.H.) Foes of California crop pests imported. Western Grower and Shipper 25(8): 40-41. 15. 1956. Aquatic Hymenoptera. pp. 289-292. In Usinger, R. L. (ed.). Aquatic insects of California. University of California Press, Berkeley, California. 16. 1956. (R. F. Smith & K.S.H.) Enemies of the spotted alfalfa aphid. Calif. Agric. 10(4): 8-10. 17. 1957. (C. S. Davis, A. S. Deal, J. E. Dibble, R. C. Dickson, E. J. Dietrick, G. L. Finney, H. Graham, K.S.H., I. M. Hall, Jr., J. K. Hollaway, L. G. Jones, J. D. Paschke, B. Puttier, H. T. Reynolds, E. I. Schlinger, R. F. Smith, E. H. Stanford, V. M. Stern, E. S. Sylvester & R. van den Bosch) The spotted alfalfa aphid and its control in California. University of California Agricul¬ tural Extension Service Manuscript. 18. *1957. Notes on the convergent lady beetle, Hippodamia convergens. Pest Control Review. Jan. 1957: 4-5. 19. 1958. (with J. K. Holloway, F. E. Skinner & G. L. Finney) Aphid parasites established. Calif. Agric. 12(2): 3, 15. 20. *1958. (with R. F. Smith) How many lady beetles are necessary to control aphids in alfalfa? Pest Control Review. March 1958: 3-4. 158 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) 21. *1958. (with E. E. Stevenson & R. F. Smith) Summary of spotted alfalfa aphid control. Pest Control Review. June 1958: 3-4. 22. 1958. (with R. F. Smith) Chemical and biological methods of pest control. Agric. chem. 13: 30-32, 98-92. 23. 1958. Honeydew as an adult fruit fly diet affecting reproduction. Proc. 10th Int. Cong. Entomol. 3: 25-30. 24. 1959. (R. van den Bosch, E. I. Schlinger, E. J. Dietrick, K.S.H. & J. K. Holloway) The colonization and establishment of imported parasites of the spotted alfalfa aphid in California. J. econ. Ent. 52: 136-141. 25. 1959. (V. M. Stern, R. F. Smith, R. van den Bosch & K.S.H.) The integration of chemical and biological control of the spotted alfalfa aphid. I. The inte¬ grated control concept. Hilgardia 29: 81-101. 26. 1959. (R. F. Smith & K.S.H.) The integration of chemical and biological control of the spotted alfalfa aphid. II. Impact of commercial insecticide treatments. Hilgardia 29: 131-154. 27. 1959. (R. F. Smith & K.S.H.) Integrated control programs in the future of biological control. J. econ. Ent. 52: 1106-1108. 28. 1960. Biological control with lady beetles, pp. 28-35. In Handbook on bi¬ ological control of plant pests. Plants & Gardens 16(3) (Special printing). 29. 1960. (with E. I. Schlinger) Imported Indian parasite of pea aphid estab¬ lished in California. Calif. Agric. 14(9): 5-6. 30. 1960. (V. M. Stern, R. F. Smith, R. van den Bosch & K.S.H.) Effectiveness of integrated control programs against pests on agricultural crops. Calif. Agric. 14(9): 7-8. 31. 1960. (E. I. Schlinger, K.S.H. & R. van den Bosch) Imported French parasite of walnut aphid established in California. Calif. Agric. 14(11): 3-4. 32. 1962. Biology and ecology of predaceous Coccinellidae. Ann. Rev. Ent. 7: 289-326. 33. 1962. (R. van den Bosch, E. I. Schlinger & K.S.H.) Initial field observations in California on Trioxys pallidus (Haliday) a recently introduced parasite of the walnut aphid. J. econ. Ent. 55: 857-862. 34. 1963. (with L. Santas & A. Tsecouras) A technique of culturing the olive fly, Dacus oleae Gmel., on synthetic media under xenic conditions, pp. 333- 356. In Radiation and radioisotopes allied to insects of agricultural impor¬ tance. International Atomic Energy Agency, Vienna. 35. 1964. (O. G. Bacon, V. E. Burton, A. S. Deal, K.S.H., C. S. Koehler, H. T. Reynolds, R F. Smith, V. M. Stern, J. E. Swift & R. van den Bosch) Pest and disease control program for alfalfa hay. Calif. Agric. Exp. Sta. Leaflet #85. 36. 1964. Developmental stages of parasites, pp. 168-246. In DeBach, P. (ed.). Biological control of insect pests and weeds. Reinhold, New York. 37. 1964. Nutrition of entomophagous insects and their hosts, pp. 356-380. Ibid. 38. 1964. (P. DeBach & K.S.H.) Manipulation of entomophagous species, pp. 429-458. Ibid. 39. 1965. (R. F. Smith & K.S.H.) Modification of the natural regulation of aphids by local climates in California, pp. 372-374. In Freeman, P. (ed.). Proc. Xllth Int. Cong. Entomol., London. 2002 ZUPARKO: HAGEN OBITUARY 159 40. 1966. (with R. L. Tassan) A method of providing artifical diets for Chrysopa larvae. J. econ. Ent. 58: 999-1000. 41. 1966. (R. van den Bosch & K.S.H.) Predacious and parasitic arthropods in California cotton fields. Calif. Agric. Exp. Sta. Bull. 820. 42. 1966. Dependence of the olive fly, Dacus oleae, larvae on symbiosis with Pseudomonas savastanoi for the utilization of olive. Nature 209: 423-424. 43. 1966. (D. W. Walker, A. Alemany, V. Quintana, F. Padovani & K.S.H.) Improved xenic diets for rearing the sugarcane borer in Puerto Rico. J. econ. Ent. 59: 1-4. 44. 1966. (with R. L. Tassan) The influence of protein hydrolysates of yeasts and chemically defined [diets] upon the fecundity of Chrysopa carnea Ste¬ phens. Vest. Csl. zool. Spol. 30: 219-227. 45. 1966. (with R R Sluss) Quantity of aphids required for reproduction by Hippodamia sp. in the laboratory, pp. 47-59. In Hodek, I. (ed.). Ecology of aphidophagous insects Czechoslovakia Academy of Sciences, Prague. 46. 1966. (with R L. Tassan) Artificial diet for Chrysopa carnea Stephens, pp. 83-87. Ibid. 47. 1966. (with R. L. Tassan) A method of coating droplets of artifical diets with paraffin for feeding Chrysopa larvae, pp. 89—90. Ibid. 48. 1966. Coccinellid aggregations, pp. 131-133. Ibid. 49. 1966. Suspected migratory flight behaviour of Hippodamia convergens. pp. 135-136. Ibid. 50. 1966. (with R. R. Sluss) Factors influencing the dynamics of walnut aphid populations in northern California, pp. 243-248. Ibid. 51. 1966. (R. F. Smith & K.S.H.) Natural regulation of alfalfa aphids in Cali¬ fornia. pp. 297-315. Ibid. 52. *1967. (with R. van den Bosch et al.) Biological control of the bollworm. pp. 9-15. In University of California, Division of Agriculture Sciences Pro¬ gress Report on Research 1966. 53. 1967. (J. T. Shimizu & K.S.H.) An artifical oviposition site for some Het- eroptera that insert their eggs into plant tissue. Ann. ent. Soc. Am. 60: 1115— 1116. 54. 1968. (with R. van den Bosch) Impact of pathogens, parasites and predators on aphids. Ann. Rev. Ent. 13: 325-384. 55. 1968. (with L. E. Caltagirone) A new nearctic species of Karpinskiella (Hy- menoptera: Pteromalidae). Pan-Pacif. Ent. 44: 241-248. 56. 1970. (with R L. Tassan) the influence of food wheast and related Sacca- romyces fragilis yeast products on the fecundity of Chrysopa carnea (Neu- roptera: Chrysopidae) Can. Ent. 102: 806-811. 57. 1970. (D. R. Laing & K.S.H.) A xenic, partially synthetic diet for the oriental fruit moth, Grapholitha molesta (Lepidoptera: Oleuthreutidae). Can. Ent. 102: 25-252. 58. 1970. (R. L. Tassan & K.S.H.) Culturing green lacewings in the home and school. University of California Extension Service, One Sheet Answers 246. 59. 1970. Notes on the convergent lady beetle ( Hippodamia convergens). Uni¬ versity of California Extension Service, One Sheet Answers 247. 60. 1970. Collecting and handling the convergent lady beetle ( Hippodamia con- 160 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) vergens). University of California Extension Service, One Sheet Answers 248. 61. 1970. Following the ladybug home. National Geographic Magazine 137: 542-553. 62. 1970. (with R. L. Tassan & E. F. Sawall, Jr.) Some ecophysiological rela¬ tionships between certain Chrysopa, honeydews and yeasts. Boll. Lab. Ent. Agr. Filippo Silvestri 28: 113-134. 63. 1971. (with R. van den Bosch & D. L. Dahlsten) The importance of natu¬ rally-occurring biological control in the western United States, pp. 253-293. In Huffaker, C. B. (ed.). Biological Control. Plenum Press, New York. 64. 1971. (R. van den Bosch, T. F. Leigh, L. A. Falcon, V. M. Stern, D. Gonzales & K.S.H.) The developing program of integrated control of cotton pests in California, pp. 377—394. Ibid. 65. 1971. (with E. F. Sawall & R. L. Tassan) The use of food sprays to increase effectiveness of entomophagous insects. Proc. Tall Timb. Conf. Ecol. Anim. Cont. Habit. Manage. 2: 59-81. 66. 1972. (I. Hodek, K.S.H. & H. F. van Emden) Methods for studying effec¬ tiveness of natural enemies, pp. 147-188. In van Emden, H. F. (ed.). Aphid technology. Academic Press, London. 67. 1972. (with R. L. Tassan) Exploring nutritional roles of extracellular sym¬ biotes on the reproduction of honeydew feeding adult chrysopids and te- phritids. pp. 323-351. In Rodriguez, J. R. (ed.). Insect and mite nutrition. North-Holland, Amsterdam. 68. 1973. (with J. M. Franz) A history of biological control, pp. 433-476. In Smith, R. F., T. E. Mittler & C. N. Smith (eds.). History of Entomology. Annual Reviews Inc., Palo Alto, California. 69. 1974. The significance of predacious Coccinellidae in biological and inte¬ grated control of insects. Entomophaga, Mem. Hors-Serie. 7: 25-44. 70. 1974. (G. F. Rajendra & K.S.H.) Trichogramma oviposition into artifical substrates. Environ, ent. 3: 399-401. 71. 1974. (with R. Hale) Increasing natural enemies through use of supplemen¬ tary feeding and non-target prey. pp. 170-181. In Maxwell, F. E. & F. A. Harris (eds.). Proceedings of the Summer Institute on Biological Control of Plant Insects and Diseases. University Press of Mississippi. 72. 1974. (Book review) Biology of Coccinellidae by I. Hodek. Science 183: 299. 73. 1975. (P. Neuenschwander, K.S.H. & R. F. Smith) Predation on aphids in California’s alfalfa fields. Hilgardia 43: 53-78. 74. 1976. Role of nutrition in insect pest management. Proc. Tall Timb. Conf. Ecol. Anim. Cont. Habit. Manag. 6: 221-261. 75. 1976. (with P. Greany, E. F. Sawall, Jr. & R. L. Tassan) Tryptophan in artificial honeydew as a source of an attractant for adult Chrysopa carnea. Environ, ent. 5: 458-468. 76. 1976. (H. F. van Emden & K.S.H.) Olfactory reactions of the green lace¬ wing, Chrysopa carnea, to tryptophan and certain breakdown products. En¬ viron. ent. 5: 469-473. 77. 1976. (W. R. Bowen, V. E. Burton, K.S.H., V. M. Stern, C. G. Summers & 2002 ZUPARKO: HAGEN OBITUARY 161 N. C. Toscano) Insect control guide for alfalfa hay. University of California, Department of Agricultural Sciences Leaflet #2763. 78. 1976. (with S. Bombosch & J. A. McMurtry). The biology and impact of predators, pp. 93-142. In Huffaker, C. B. & P. S. Messenger (eds.). Theory and practice of biological control. Academic Press, New York. 79. 1976. (with G. A. Viktorov, K. Yasumatsu, & M. F. Schuster) Biological control of pests of range, forage and grain crops, pp. 397-442. Ibid. 80. 1977. (D. A. Nordlund, W. J. Lewis, R. L. Jones, H. R. Gross, Jr. & K.S.H.) Kairomones and their use for management of entomophagous insects. VI. An examination of the kairomones for the predator Chrysopa carnea Ste¬ phens at oviposition sites of Heliothis zea (Boddie). J. Chem. Ecol. 3: 507- 511. 81. *1977. (W. C. Mitchell, C. O. Andrew, K.S.H., R. A. Hamilton, E. J. Harris, K. L. Maehler, & R. H. Rode. The Mediterrean fruit fly and its economic impact on Central American countries and Panama. UC/AID Report on Pest Management and Related Environmental Protection Project. 82. *1977. Biological and integrated control of insect pests in Brazil. Graduate Training, Accomplishments and Future. Brazil/Michigan State PEAS Report. 83. 1977. (D. S. Chandler & K.S.H.) New synonymy of North American No- toxus (Coleoptera: Anthicidae). Pan-Pacif. Ent. 53: 230—232. 84. 1977. (T. Finlayson, T. & K.S.H.) Final-instar larvae of parasitic Hymenop- tera. Pest Management Paper #10, Simon Fraser University, Burnaby, British Columbia. 85. 1978. (D. E. Pinnock, K.S.H., D. V. Cassidy, R. J. Brand, J. E. Milstead & R. L. Tassan) Integrated pest management in highway landscapes. Calif. Agric. 32(2): 33-34. 86. 1978. Aquatic Hymenoptera. pp. 233—243. In Merritt, R. W. & K. W. Cum¬ mins (eds.). An introduction to the aquatic insects of North America. Ken¬ dall/Hunt, Dubuque, Iowa. 87. 1978. (S. A. Hassan & K.S.H.) A new artificial diet for rearing Chrysopa carnea larvae. Z. agnew. Ent. 86: 315—330. 88. 1978. (K.S.H., E. F. Sawall, Jr. & R. L. Tassan) (Abstract) The attraction of Chrysopa carnea adults to comercially available predator food sprays. Ab¬ stracts of submitted papers Pacific Branch ent. Soc. Am. 62nd Annual Meet¬ ing, Scottsdale, Arizona, June 19-22. 1978: 26. 89. 1979. (R. L. Tassan, K.S.H. & E. F. Sawall, Jr.) The influence of field food sprays on the egg production rate of Chrysopa carnea. Environ, ent. 8: 81— 85. 90. 1979. (W. J. Lewis, M. Beevers, D. A. Nordlund, H. R. Gross, Jr. & K.S.H.) IX. Investigations of various kairomone-treatment patterns for Trichogram- ma spp. J. Chem. Ecol. 5: 673-680. 91. 1979. (with J. A. McMurtry) Natural enemies and predator-prey ratios, pp. 28-40. In Davis, D. W., S. C. Hoyt, J. A. McMurtey, & M. T. AliNiazee (eds.) Biological control and insect pest management. University of Cali¬ fornia, Division of Agricultural Sciences, Priced Publication #4096. 92. 1979. (with G. W. Bishop) Use of supplemental food and behavioral chem¬ icals to increase the effectiveness of natural enemies, pp. 49-60. Ibid. 162 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) 93. 1979. (M. T. AliNiazee, K.S.H., & S. C. Hoyt) Current trends and future outlook, pp. 96-102. Ibid. 94. 1979. (G. J. Tsiropoulos & K.S.H.) Ovipositional response of the walnut husk fly, Rhagoletis completa, to artificial substrates. Z. agnew. Ent. 88: 547-550. 95. 1980. (P. Neuenschwander & K.S.H.) Role of the predator Hemerobius pa- cificus in a non-insecticide treated artichoke field. Environ, ent. 9: 492—495. 96. 1980. (W. J. Lewis, K.S.H., W. L. Roelofs & L. M. Schoonhoven) Status and potential use of behavioral chemicals in pest management. FAO Plant Protection Bulletin 28: 121—128. 97. 1981. (A. P. Gutierrez, J. U. Baumgaertner & K.S.H.) A conceptual model for growth, development, and reproduction in the ladybird beetle, Hippo- damia convergens (Coleoptera: Coccinellidae). Can. Ent. 113: 21—33. 98. 1981. (P. Greany & K.S.H.) Prey selection, pp. 121-135. In Nordlund, D. A., R. L. Jones & W. J. Lewis (eds.). Semiochemicals—their role in pest control. Wiley, New York. 99. 1981. (J. B. Johnson & K.S.H.) A neuropterous larva uses an allomone to attack termites. Nature 289: 506-507. 100. 1981. (with W. W. Allen & R. L. Tassan) Mediterranean fruit fly: the worst may be yet to come. Calif. Agric. 35(3 & 4): 5-7. 101. 1981. IOBC Presidential address. Int. Org. Biological Control Newsletter #19-20: 1-4. 102. 1981. (R. L. Tassan, K.S.H. & D. V. 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Report to State of California Department of Parks and Recre¬ ation. 121. 1986. (S. H. Dreistadt, K.S.H. & D. L. Dahlsten) Predation by Iridomyrmix humilis [Hym.: Formicidae] on eggs of Chrysoperla carnea [Neu.: Chry- sopidae] released for inundative control of Illinoia liriodendri [Horn.: Aphi- didae] infesting Liriodendron tulipifera. Entomophaga 31: 397-400. 122. 1987. Nutritional ecology of terrestrial insect predators, pp. 533-577. In Slansky, F. Jr. & J. G. Rodriguez (eds.). The nutritional ecology of insects, mites, and spiders. John Wiley and Sons, New York. 123. 1987. (R. Garcia & K.S.H.) Summer dormancy in adult Agabus disintegra¬ te (Crotch) (Coleoptera: Dytiscidae) in dried ponds in California. Ann. ent. Soc. Am. 80: 267-271. 164 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) 124. 1987. (G. J. Tsiropoulos & K.S.H.) Effect of nutritional deficiencies, pro¬ duced by antimetabolites, on the reproduction of Rhagoletis completa Cres- son (Dipt., Tephritidae). Z. agnew. Ent. 103: 351-354. 125. 1987. (S. H. Dreistadt, D. L. Dahlsten & K.S.H.) Integrated pest manage¬ ment of tuliptree aphids, pp. 35-37. In Minimal Maintenance Landscaping. A report to the Elvenia J. Slosson Fund for Ornamental Horticulture: 1983- 1986. University of California, Division of Agriculture and Natural Re¬ sources. 126. 1987. (P. Neyman & K.S.H.) (Abstract). Toxoptera aurantii on camellias in a large urban garden: abundance and control by natural enemies. Abstracts of submitted and invited papers. Pacific Branch ent. Soc. Am. 71st Annual Meeting, Portland, Oregon, June 23—25 1987: 13. 127. 1987. (T. Wipperfurth, K.S.H. & T. E. Mittler) Egg production by the coc- cinellid Hippodamia convergens fed on two morphs of the green peach aphid, Myzus persicae. Entomologia exp. appl. 44: 195—198. 128. 1988. (Abstract) (R. Garcia & K.S.H.). Ecological adaptations for survival in Agabus disintegratus (Crotch) (Coleoptera: Dytiscidae). Proc. XVIIIth Int. Cong. Ent., Vancouver, B.C., Canada: 34. 129. 1988. (Abstract). Physiological interactions in host-parasite systems and their implications for biological control. Proc. XVIIIth Int. Cong. Ent., Van¬ couver, B.C., Canada: 98. 130. 1990. (A. P. Gutierrez, K.S.H. & C. K. Ellis) Evaluating the impact of natural enemies: a multitrophic perspective, pp. 81-109. In Mackauer, M., L. E. Ehler & J. Roland (eds.). Critical issues in biological control. Intercept Ltd. Andover, Hants, U.K. 131. 1990. (R. Garcia, K.S.H., W. G. Voight) Life history, termination of summer diapause, and other seasonal adaptations of Agabus disintegratus (Crotch) (Coleoptera: Dytiscidae) in the Central Valley of California. Quaest. ent. 26: 139-149. 132. 1990. Biological control of insect pests in alfalfa hay. pp. 74-83. In Pro¬ ceedings of the 20th California Alfalfa Symposium, Visalia, California. Uni¬ versity of California Cooperative Extension Service, Davis, California. 133. 1990. (with S. H. Dreistadt) First California record for Anthocoris nemoralis (Fabr.) (Hemiptera: Anthocoridae), a predator important in the biological control of psyllids (Homoptera: Psyllidae). Pan-Pacif. Ent. 66: 323-324. 134. 1991. (M. Y. Hussein & K.S.H.) Rearing of Hippodamia convergens on artifical diet of chicken liver, yeast and sucrose. Ent. exp. appl. 59: 197— 199. 135. 1991. (C. G. Summers, K.S.H. & V. M. Stern) U. C. IPM pest management guidelines: Alfalfa. University of California Department of Agriculture and Natural Resources Publication #3339. 136. 1992. (P. G. da Silva, K.S.H. & A. P. Gutierrez) Functional response of Curinus coeruleus (Col.: Coccinellidae) to Heteropsylla cubana (Horn.: Psyllidae) on artificial and natural substrates. Entomophaga 37: 555-564. 137. 1992. (K. M. Daane, G. Y. Yokota, R. F. Gill, L. E. Caltagirone, K.S.H., D. Gonzalez, P. Stary, & W. E. Chaney) Imported parasite may help control European asparagus aphid. Calif. Agric. 46 (6): 12-14. 138. *1992. (L. F. Schultz & K.S.H.) Report of the evaluation mission on the 2002 ZUPARKO: HAGEN OBITUARY 165 collaborative project on the biological control of tsetse and crop pests by the International Center of Insect Physiology and Ecology and the Depart¬ ment of Entomology, Vageningen Agricultural University. 139. 1993. (Y. Zheng, K. M. Daane, K.S.H. & T. E. Mittler) Influence of larval dietary supply on food consumption, food utilization efficiency, growth and development of the lacewing Chrysoperla carnea. Ent. exp. appl. 67: 1-7. 140. 1993. (Y. Zheng, K.S.H., K. M. Daane & T. E. Mittler) Influence of larval food consumption on the fecundity of the lacewing Chrysoperla carnea. Ent. exp. appl. 67: 9-14. 141. *1993. (with L. E. Caltagirone) Biological control of the pepper tree psyllid. Final report to State of California Department of Transportation, RT56G062. 142. 1993. (K. M. Daane, G. Y. Yokota, Y. D. Rasmussen, Y. Zheng & K.S.H.) Effectiveness of leafhopper control varies with lacewing release methods. Calif. Agric. 47 (6): 19-23. 143. *1994. (Abstract) Stethorus histrio, a predator of tetranychid mites from Australia established in California. 78th Annual Meeting Pacific Branch Ent. soc. Am. 144. 1994. (S. H. Dreistadt & K.S.H.) European elm scale (Homoptera: Eriococ- cidae) abundance and parasitism in northern California. Pan-Pacif. Ent. 70: 240-252. 145. 1994. (S. H. Dreistadt & K.S.H.) Classical biological control of the acacia psyllid, Acizzia uncatoides (Homoptera: Psyllidae), and predator-prey-plant interactions in the San Francisco Bay area. Biol. Cont. 4: 319-327. 146. 1995. (K. M. Daane, G. Y. Yokota, K.S.H. & Y. Zheng) Field evaluation of Chrysoperla spp. in augmentative release programs for the variegated grape leafhopper, Erythroneura variabilis. pp. 105-120. In Nicoli, G., M. Benuzzi, & N. C. Leppla (eds.). Proceedings of the 7th Workshop of the IOBC work¬ ing group: Quality control of mass reared arthropods. Rimini, Italy. 147. 1995. (J. A. McMurtry, L. A. Andres, T. S. Bellows, Jr., S. C. Hoyt & K.S.H.) A historical overview of regional research project W-84. pp. 3-5. In Nechols, J. R., L. A. Andres, J. W. Beardsley, R. D. Goeden, & C. G. Jackson (eds.). Biological control in the western United States. University of California, Division of Agriculture and Natural Resources Publication #3361. 148. 1995. (J. W. Beardsley, K.S.H., J. R. Leeper, & R. L. Tassan) Acacia psyllid. pp. 91-92. Ibid. 149. 1995. (T. R. Unruh, P. H. Westigard, & K.S.H.) Pear psylla. pp. 95-100. Ibid. 150. 1995. (K. M. Daane, K.S.H., D. Gonzalez, & L. E. Caltagirone) European asparagus aphid, pp. 120-122. Ibid. 151. 1995. (D. Gonzalez, K.S.H., P. Stary, G. W. Bishop, D. W. Davis, & K. S. Pike) Pea aphid and blue alfalfa aphid, pp. 129-135. Ibid. 152. 1995. (M. T. AliNiazee & K.S.H.) Walnut aphid. Part 1. pp. 140-141. Ibid. 153. 1995. (C. E. Kennett, K.S.H. & K. M. Daane) Citricola scale, pp. 148-149. Ibid. 154. 1995. (R. L. Tassan & K.S.H.) Iceplant scales, pp. 150-154. Ibid. 155. 1995. (L. K. Etzel, K.S.H., D. Gonzalez & J. J. Ellington) Egyptian alfalfa weevil, pp. 176-179. Ibid. 166 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) 156. 1995. (with R. L. Tassan., M. Fong & M. T. AliNiazee) Walnut husk fly. pp. 224-227. Ibid. 157. 1995. (S. H. Dreistadt, K.S.H. & L. G. Bezark) Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), first western United States record for this Asi¬ atic lady beetle. Pan-Pacif. Ent. 71-135-136. 158. 1996. Aquatic Hymenoptera. pp. 474-483. In Merritt, R. W. & K. W. Cum¬ mins (eds.). An introduction to the aquatic insects of North America. 3rd ed. Kendall/Hunt, Dubuque, Iowa. 159. 1996. (K. M. Daane, G. Y. Yokota, Y. Zheng & K.S.H.) Inundative release of common green lacewings (Neuroptera: Chrysopidae) to suppress Ery- throneura variabilis and E. elegantula (Homoptera: Cicadellidae) in vine¬ yards. Environ, ent. 25: 1224—1234. 160. 1998. (K. M. Daane, K.S.H. & N. J. Mills) Predaceous insects for insect and mite management, pp. 62-115. In Ridgway, R. L., M. P. Hoffmann, M. N. Inscoe, and C. S. Glenister (eds.). Mass-reared natural enemies: appli¬ cation, regulation, and needs. Thomas Say Publications in Entomology. En¬ tomological Society of America, Lanham, Maryland. 161. 1999. (with N. J. Mills, G. Gordh, & J. A. McMurtry) Terrestial arthropod predators of insect and data mite pests, pp. 383-503. In Bellows, T. S., T. W. Fisher, L. E. Caltagirone, D. L. Dahlsten, C. Huffaker & G. Gordh (eds.). Handbook of biological control. Academic Press. San Diego, California. 162. 1999. (S. N. Thompson & K.S.H.) Nutrition of entomophagous insects and other arthropods, pp. 594-652. Ibid. 163. 2000. (M. J. Tauber, C. A. Tauber, K. M. Daane & K.S.H.) Commerciali¬ zation of predators: recent lessons from green lacewings (Neuroptera: Chry¬ sopidae: Chrysoperla). American Entomologist 46: 26-38. 164. 2000. (K. M. Daane, M. S. Barzman, L. E. Caltagirone & K.S.H.) Meta- phycus anneckei and M. hageni (Hym.: Encyrtidae): two discrete species parasitic on the black scale, Saissetia oleae. BioControl 45: 269—284. 165. *2001. (K. M. Daane & K.S.H.) An evaluation of lacewing releases in North America, pp. 398-407. In McEwen, P. K., T. R. New, & A. Whittington (eds.). Lacewings in the crop environment. Cambridge University Press, London. Acknowledgment I wish to thank Ken’s many colleagues who shared their memories with me. I am especially grateful to Leo Caltagirone and Charlie Summers for their editorial suggestions and help in describing Ken’s career. I thank the two anonymous re¬ viewers for their suggested improvements to the manuscript. I also thank Maxine Hagen, who graciously provided photographs and led me through the events of Ken’s early days. This paper was partially funded by a grant from the C. P. Alexander fund from the Pacific Coast Entomological Society. Literature Cited Buckingham, G. R. & M. J. Sharkey. 1988. Abdominal exocrine glands in Braconidae (Hymenoptera). pp. 199-242. In Gupta, V. K. (ed.). Advances in Parasitic Hymenoptera Research, 1988. Chandler, D. S. 1982. A revision of North American Notoxus with a cladistic analysis of the New World species. Entomography 1: 333-438. 2002 ZUPARKO: HAGEN OBITUARY 167 Daane, K. & L. E. Caltagirone. 1999. A new species of Metaphycus (Hymenoptera: Encyrtidae) parasitic on black scale, Saissetia oleae (Olivier) (Homoptera: Coccidae). Pan-Pacif. Ent. 75: 13-17. Gordon, R. D. 1985. The Coleoptera (Coccinellidae) of America north of Mexico. J. New York Ent. Soc. 93: 1-912. Hagen, K. S. 1970. Following the ladybug home. National Geographic Magazine 137: 542-553. Hagen, K. S. & L. E. Caltagirone. 1968. A new nearctic species of Karpinskiella (Hymenoptera: Pteromalidae). Pan-Pacif. Ent. 44: 241-248. Summers, C. G. 1998. Integrated pest management in forage alfalfa. Integrated Pest Management Reviews 3: 127-154. Tauber, C. A. 1969. Taxonomy and biology of the lacewing genus Meleoma (Neuroptera: Chrysopi- dae), Univ. Calif. Publ. Ent. 58: 1-94. Vandenberg, N. J. 1992. Revision of the New World lady beetles of the genus Olla and description of a new allied genus (Coleoptera: Coccinellidae). Ann. Ent. Soc. Am. 85: 370-392. Received 1 March 2001; Accepted 13 September 2001 PAN-PACIFIC ENTOMOLOGIST 78(3): 168-176, (2002) NICKEL ACCUMULATION IN SERPENTINE ARTHROPODS FROM THE RED HILLS, CALIFORNIA Michael A. Wall* and Robert S. Boyd Department of Biological Sciences, 101 Rouse Life Sciences Building, Auburn University, Auburn, Alabama 36849-5407 U.S.A. Abstract. —Serpentine soils are characterized by high levels of heavy metals (e.g., Ni, Fe, Cr), and low levels of important plant nutrients (e.g., P, Ca, N). Due to these inhospitable edaphic conditions, serpentine soils are typically home to a very specialized flora. Although much is known about the serpentine flora, almost no research has investigated the arthropods of serpentine areas. In this study, we sampled the arthropods associated with Streptanthus polygaloides (Gray), a Ni hyperaccumulator, and the arthropod community of the surrounding serpentine area in the Red Hills of California. Arthropods were then analyzed for Ni content to investigate Ni transport within the serpentine ecosystem. Arthropods associated with S. polygaloides contained signifi¬ cantly higher concentration of Ni than those collected in the surrounding community. One insect associated with S. polygaloides, Melanotrichus boydi Schwartz and Wall (Hemiptera: Miridae), accumulated 770 |xg Ni/g. Key Words. —Insecta hyperaccumulation, serpentine arthropods. Whereas some heavy metals are essential nutrients and are crucial to the sur¬ vival of most organisms, in excessive doses they also can be toxic. At many sites heavy metal concentrations in the environment are artificially elevated due to anthropogenic influences such as mining and metal smelting. Because it is pos¬ sible in some cases to know when human-caused metal contamination began (Brooks 1998), many authors have been able to conduct research on rates of adaptation of organisms to metal contamination (e.g., Wu et al. 1975, Posthuma 1990). This research is not only interesting from a microevolutionary standpoint but also has revealed physiological and genetic mechanisms by which organisms adapt to high concentrations of metal in the environment. However, the relatively recent history of anthropogenic metal contamination at mines and smelters pre¬ cludes studying the effects of heavy metals on these human-impacted ecosystems over longer time spans. Serpentine soils provide an opportunity to study long-term effects of heavy metals on ecosystems. Distributed around the globe, serpentine soils are high in heavy metals such as Ni, Fe, and Cr but are low in important plant nutrients like Ca and P (Brooks 1987). Within North America, there are extensive areas of serpentine soils in California and Oregon (Kruckeberg 1984). Whereas Coleman (1967) suggested that these serpentinized areas of California and Oregon have been exposed since the Tertiary, a more recent hypothesis by Raven and Axelrod (1978) suggests a much younger origin for these sites (3 to 24 million years ago). In either case, plant and animal species in these areas have had millions of years in which to adapt to the unique edaphic conditions provided by serpentine soil. The serpentine flora of California has been the subject of botanical research since the late 1800’s (Kruckeberg 1984). Much of this research has focused on * Currently Department of Ecology and Evolutionary Biology, 75 North Eagleville Road, U-43, Uni¬ versity of Connecticut, Storrs, Connecticut 06269-3043 U.S.A. 2002 WALL & BOYD: NICKEL IN SERPENTINE ARTHROPODS 169 the adaptations of plants to the unique edaphic conditions of serpentine soils (Brooks 1987). Of particular interest has been the accumulation of Ni by plants. Serpentine plant species commonly accumulate higher levels of Ni than nonser¬ pentine species (Brooks 1987). Moreover, some species of serpentine plants, termed “hyperaccumulators,” contain over 1000 fxg Ni/g (Brooks et al. 1977). These concentrations of Ni have been found to defend plants against many types of herbivores (see reviews by Boyd and Martens 1998, Boyd 1998). The high Ni concentration in Ni hyperaccumulators provides a unique environment to which herbivores must adapt in order to utilize these plants as a food source. Many authors have hypothesized that there should be a unique fauna associated with the serpentine flora (Proctor and Woodell 1975, Kruckeberg 1984, Brooks 1987), but little research has specifically addressed this idea. The purpose of this study was to assess interspecific differences in the Ni concentration of arthropods associated with a serpentine community with an emphasis on insects associated with metal hyperaccumulating plants. Materials and Methods Study Site. —This study was conducted in the Red Hills Management Area in Tuolumne County, California. The entire management area is underlain with ser¬ pentine soils (Franklin et al. 1997). Much of the community-level sampling in this study took place in a large (ca. 2000 m 2 ) area along Red Hills Road in the Red Hills Management Area. The vegetation was dominated by Ceanothus cu- neatus (Hook.) Nutt., Clarkia biloba (Durand) Nels & Macbr., Calycadenia mul- tiglundulosa ssp. bicolor (Greene) Keck, and Streptanthus polygaloides Grey. Species-specific sampling took place throughout the Red Hills Management Area, but was concentrated at the location described above. Community-level Sampling. —We sampled arthropods within the general ser¬ pentine community of the Red Hills Management Area via both pitfall-trapping and black-lighting. In June 1996 and 1997, 21 pitfall traps were placed at the study site. Traps were arranged in an approximately 30 m by 70 m grid. Traps were placed approximately 10 m apart. A trap consisted of a 14 cm diameter by 12 cm deep plastic cup set into the ground so that the lip of the cup was level with the soil surface. Cups contained approximately 100 m of a 50:50 mixture of ethylene glycol and water. Cups were covered by a 22 cm diameter plastic plate to reduce evaporation of preservative. A 2-3 cm space was left between the cover and the soil surface to allow arthropods to easily enter traps. For black lighting, on three occasions over a one week period in June of 1996 five black-lights were spaced at approximately 50 m intervals throughout a site (ca. 2000 m 2 ) in the Red Hills Recreational Area. Lights were turned on around 9 pm and allowed to shine against aim 2 white sheet. We sampled lights at 10 pm, 11 pm, and 12 am. During a collecting bout, arthropods were arbitrarily collected for approximately 5-10 minutes at each light. Species-Specific Sampling : Streptanthus Polygaloides. —Accompanying the broad community-level sampling of the Red Hills, we also specifically sampled the arthropods associated with S. polygaloides, the only known Ni hyperaccu¬ mulator in the Red Hills. Streptanthus polygaloides is endemic to serpentine bar¬ rens of the foothills of the western part of the Sierra Nevada from Fresno County, California north to Butte County, California (Kruckeberg 1984). Containing an 170 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) average of 9750 fxg Ni/g (Reeves et al. 1981, Kruckeberg & Reeves 1995), S. polygaloides is an annual that often grows in relatively dense stands. Streptanthus polygaloides was sampled via both sweep-netting and visual in¬ spection. Sampling took place during June of 1996, 1997, and 1998 while the plants were in flower. Dense pure stands of S. polygaloides were targeted for sweep-netting in order to reduce inadvertent sampling of other plant species. Apis mellifera L. and Bombus vandykkei (Frison) were used to compare Ni concentrations found within members of the same species that occur both on and off serpentine soils. Both species were collected from S. polygaloides in the Red Hills and from Heteromeles arbutifolia (Lindley) Roemer, a shrub growing on a nonserpentine site >15 km from the Red Hills, for comparison. Elemental Analysis. —Specimens were sorted according to morphotype and rep¬ resentatives of each morphotype were pinned and labeled for later identification to the lowest taxonomic level that could be readily attained. Individual specimens were air-dried for at least 72 h at 67° C and weighed. Individuals of the same morphotype weighing less than 50 mg were combined in order to create samples of at least that mass for analysis. Specimens were then analyzed for Ni concen¬ tration as described below. In order to sample variation in Ni values adequately, we analyzed at least three samples of each morphotype to generate means (± SD). Many of our morphotypes could not be analyzed for Ni concentration, due to their low mass. Morphotypes with insufficient biomass to create three samples for analysis are not included in the data presented here. Nickel concentration was determined with an atomic absorption spectrophotom¬ eter (Instrumentation Laboratory, IL 251). Samples were digested in borosilicate glass test tubes using 3-5 ml of concentrated nitric acid at 110° C for 6-8 h, after which time most of the liquid had evaporated. The residue was then redissolved in 3-5 ml of 1 M hydrochloric acid at 110° C for 2-4 h. The solutions were then diluted with distilled water to a volume of 10 or 25 ml, depending on the original mass of the dried sample. Reagent blanks were made and processed with every batch of samples in order to correct for any contamination generated by the tech¬ nique. All metal values are reported as p,g metal/g on a dry weight basis. Specimens containing unusually high levels of Ni (> 300 |xg Ni/g) were also analyzed for Cr in order to rule out contamination by soil. Chromium levels are several orders of magnitude higher in serpentine soils than in the plants growing on serpentine soils. Unusually high levels of Ni accompanied by high levels of Cr indicate the potential of soil contamination (Brooks 1987). Concentrations of Cr were determined via inductively coupled argon plasma spectrophotometry (Jar¬ rell-Ash, ICAP 9000). Data Analysis. —Nickel concentrations in arthropod tissues were analyzed by one-way analysis of variance (ANOVA) in order to determine if association with S. polygaloides influenced specimen Ni concentrations. Nickel concentrations were log-transformed in order to satisfy the assumptions of ANOVA (Zar 1984). Log-transformed Ni concentrations of hemipteran herbivores collected on S. po¬ lygaloides were also analyzed via one-way ANOVA. In this case, post-hoc mean separations were performed using Fisher’s Protected Least Significant Difference (PLSD) test (SAS Institute 1998) in order to compare Ni concentrations between pairs of hemipteran species. Vouchers of analyzed specimens were deposited in 2002 WALL & BOYD: NICKEL IN SERPENTINE ARTHROPODS 171 Table 1. Nickel concentration (mean ± SD) of insect species or morphospecies associated with Streptanthus polyaloides in the Red Hill Recreational Area, California. Order Family Morphospecies or species Mean ± SD n Coleoptera Bruchidae Ac antho sc elides seminulum Horn 55 ± 96 3 Melyridae Mel-1 129 ± 18 3 Diptera Otitidae Oti-1 58 ± 8 3 Hemiptera Miridae Melanotrichus boydi Schwartz & Wall 777 ± 162 8 Lygus hesperus Knight 131 ± 126 3 Pentatomidae Thyanta pallidovirens (Stal) 40 ± 28 3 Rhopalidae Liorhyssus hyalinus (Fabicus) 48 ± 50 4 Hymenoptera Apidae Apis mellifera L. 43 ± 24 10 Bombus vandykei (Frison) 38 ± 34 10 Halictidae Dialictus sp. 51 ± 27 3 Lepidoptera Lycaenidae Everes amyntula (Boisduval) 36 ± 34 3 Overall Mean 128 ± 218 the University of Connecticut insect collection with voucher labels with the prefix, “wall-boyd-CA” followed by the morphospecies names listed in Tables 1 and 2. Results We collected a total of 110 morphotypes of arthropods. Almost half of these arthropods (50) were collected in association with S. polygaloides. The remaining arthropods (60) were collected by black-lighting and pitfall-trapping. Of all the morphotypes collected, only 33 were collected in great enough number to allow us to analyze three replicates. Eleven of the 33 analyzed morphotypes were col¬ lected in association with S. polygaloides. The arthropods analyzed contained an average of 65 ± 132 pig Ni/g. However, one plant bug associated with S. poly¬ galoides, Melanotrichus boydi Schwartz and Wall, contained an average of 111 ± 162 p,g Ni/g (Table 1) (also see Schwartz and Wall 2001). Melanotrichus boydi contained almost no Cr (1 ± 2 pig Cr/g, n = 3). If samples of M. boydi are excluded, the average Ni content of the arthropods sampled decreases to 43 ± 34 Rg Ni/g. Nickel content of arthropods associated with S. polygaloides (Table 1) was significantly higher than the Ni content of arthropods collected via black-lighting and pitfall-trapping (Table 2, ANOVA: F = 11.45; df = 1, 31; P = 0.002). Even when M. boydi is excluded from this analysis, there is still significantly more Ni in arthropods associated with S. polygaloides than in arthropods collected via black-lighting and pitfall-trapping (ANOVA: F = 8.69; df = 1, 30; P = 0.006). Melanotrichus boydi contained more Ni than other hemipteran herbivores found feeding on S. polygaloides. Other than M. boydi, three other hemipteran herbi¬ vores were collected from S. polygaloides in great enough numbers to analyze: Lygus hesperus Knight (Heteroptera: Miridae), Thy ant a pallidovirens (Stal) (Het- eroptera: Pentatomidae), and Liorhyssus hyalinus (F.) (Heterptera: Rhopalidae). 172 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) Table 2. Nickel concentrations (mean ± SD) of insect species or morphotypes collected via pitfall traps and black lights in the Red Hills Recreational Area, California. Order Family Morophospecies or species Mean ± SD n Coleoptera Elateridae Ela-1 41 ± 69 3 Eucnemidae Euc-1 23 ± 15 3 Euc-2 19 ± 15 3 Tenebrionidae Eleodes sp. 75 ± 71 3 Lepidoptera Geometridae Geo-1 7 ± 7 3 Geo-2 8 ± 14 3 Geo-3 13 ± 23 3 Geo-4 62 ± 80 3 Geo-5 13 ± 16 3 Geo-6 11 ± 13 3 Geo-7 13 ± 12 3 Noctuidae Noc-1 40 ± 59 3 Noc-2 32 ± 24 3 Pyralidae Pyr-1 12 ± 21 3 Pyr-2 44 ± 69 3 Pyr-3 12 ± 22 3 Neuroptera Corydalidae Cor-1 32 ± 21 3 Myrmeleontidae Myr-1 13 ± 7 3 Orthoptera Acrididae Malanopus sp. 40 ± 15 5 Gryllacrididae Ceuthophilus sp. 51 ± 28 3 Gryllidae Gryllus assimilis (Fabricius) 133 ± 77 3 Tettigoniidae Arethea sp. 55 ± 28 4 Overall Mean 34 ± 29 All of these insects were observed feeding on the young stems and leaves of S. polygaloides in the field. Also, the presence of nymphs of L. hesperus, M. boydi, and T. pallidovirens on S. polygaloides supports the hypothesis that S. polyga¬ loides can be a host for these herbivores. These herbivores varied significantly in Ni content (ANOVA: F = 8.36; df = 3, 15; P = 0.002). Although all these herbivores appeared to be feeding on S. polygaloides, M. boydi contained signif¬ icantly more Ni than the other three species of hemipterans (Fisher’s PLSD test: P = 0.001, 0.008, and 0.002 for pairwise comparisons of M. boydi with L. hes¬ perus, T. pallidovirens, and L. hyalinus respectively). Apis mellifera collected while visiting flowers of S. polygaloides contained significantly higher levels of Ni (46 ± 8 pug Ni/g, n = 10) than those collected from Heteromeles arbutifolia (16 ± 8 pug Ni/g, n = 10) in a non-serpentine environment (ANOVA: F = 7.88; df = 1, 18; P = 0.012). Like A. mellifera, Bombus vandykei collected from S. polygaloides contained significantly more Ni (38 ± 15 pig Ni/g, n = 5) than those collected from H. arbutifolia (12 ± 14 pig Ni/g, n = 5); (ANOVA: F = 8.60; df = 1, 8; P = 0.019). Discussion Historically, little attention has been paid to accumulation of Ni by arthropods in terrestrial environments (see Hopkin 1989). This may be due to the typically 2002 WALL & BOYD: NICKEL IN SERPENTINE ARTHROPODS 173 low levels of Ni found in arthropods from metal-contaminated sites (e.g., Helio- vaara and Vaisanen 1990, Heliovaara et al. 1990, Bagatto and Shorthouse 1996). On the other hand, a study of arthropods collected from serpentine sites in Zim¬ babwe reported very high levels of Ni accumulation (Wild 1975). Termites col¬ lected from these sites contained 5000 fxg Ni/g and 1500 fxg Cr/g (Wild 1975). Wild (1975) hypothesized that these high levels of Ni and Cr were the result of accumulation from ingested plant material. In plant material, however, high levels of Cr are typical of soil contamination on specimens (Brooks 1987). As with Wild’s (1975) work with termites from the serpentine exposures of Zimbabwe, there is the potential that residual plant material in the gut and/or dust on speci¬ mens may have artificially elevated Ni concentrations in Melanotrichus boydi. Soil/dust contamination is unlikely unless Cr concentrations in a sample exceed 100 (jig Cr/g (Brooks 1987). Levels of Cr in M. boydi, averaging only 1 p,g Cr/ g, are extremely low and thus soil/dust contamination seems unlikely to be the source of Ni in the specimens analyzed. There are several alternatives that may explain the disparity in Ni content found amongst herbivores of Streptanthus polygaloides. In one scenario, metal content varies between tissues and organs of the plant and the observed differences in herbivore metal content simply reflect the differences in the plant tissue or organ type upon which was fed. In this study, the herbivores with the three lowest Ni values, Thyanta pallidovirens, Liorhyssus hyalinus, and Acanthoscelides semi- mulum, were collected on the developing fruits of S. polygaloides. Indeed, the developing fruits (1100-5230 fxg Ni/g) of S. polygaloides are known to contain less Ni than leaves (3300-14,800 fxg Ni/g) or flowers (2860-16,400 |Jig Ni/g) (Reeves et al. 1981). On the other hand, both mirid species, M. boydi and Lygus hesperus, feed on the same plant organs, young developing leaves and flowers. Thus it is difficult to explain the disparity in metal content of the two mirids, as being due to differences in organ metal content. These differences could, however, be due to variation in metal content between tissues. Boyd and Martens (1999) found that aphids were able to feed upon S. polygaloides without accumulating significant amounts of Ni. The implication being that the aphids contained little Ni because the phloem sap contains little Ni. Unlike aphids, no mirids are known to tap into sieve elements and feed directly on phloem sap (Wheeler 2001). While feeding on phloem sap is unlikely to explain the low metal content in L. he perns relative to M. boydi, the two species could target tissues of differing metal content within S. polygaloides. Unfortunately, nothing is known about the distribution of metal between tissues in S. polygaloides. The alternative scenario is that these species feed on similar tissues but vary in their ability to accumulate and/or excrete metal. If the insects are unable to avoid metal, then excretion and storage-detoxification are the two major strategies for terrestrial invertebrates dealing with toxic levels of heavy metal (Dallinger 1993). To effectively distinguish which strategy is employed by which species would require laboratory-based studies with artificial diets as opposed to our own field-based studies. It is interesting to note, however, that previous work has shown M. boydi to be a specialist on S. polygaloides (Wall 1999) and L. hesperus is widely known to be extremely polyphagous (Schwartz & Footit 1998). Thus differences in strategy may represent different “trade-offs” by specialist versus polyphagous herbivores (Futuyma & Moreno 1988). 174 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) We found that arthropods associated with a Ni hyperaccumulator, S. polyga- loides, contained more Ni than arthropods collected from the general serpentine community. Arthropods collected from S. polygaloides also contained more Ni than arthropods reported in the literature that were collected from sites contami¬ nated with Ni (e.g., Heliovaara and Vaisanen 1990, Heliovaara et al. 1990). The higher levels of Ni found in arthropods associated with S. polygaloides probably reflect the degree to which hyperaccumulators accumulate Ni, as well as the bio¬ availability of that Ni. Although plants from Ni-contaminated sites typically con¬ tain relatively low levels of Ni (e.g., Koricheva and Haukioja 1995), S. polyga¬ loides contains an average of 9750 [xg Ni/g (Reeves et al. 1981). Often, Ni con¬ tamination at polluted sites is the result of aerial deposition and therefore is only available in an inorganic form (Berthelsen et al. 1995). Within hyperaccumulators, Ni appears to be complexed with an organic acid (Lee et al. 1978) and is poten¬ tially more available to arthropods. The variation in metal content that we observed within insect orders highlights the need to separate taxa to finest taxonomic level that material allows. In their study on metal content of insects associated with Scottish serpentine soils, Dav¬ ison et al. (1999) pool their specimens together at the ordinal level for metal analysis. This does not allow exploration of intraordinal variation in metal content, which our study suggests, can be significant. For instance, mean values of Ni within the Hemiptera ranged from 40 ± 28 |xg Ni/g in T. pallidovirens to 777 ± 162 [xg Ni/g in M. boydi. From this perspective, our own study would have been more informative had we the biomass to analyze different body parts in order to isolate Ni localization within the body. The variation of Ni content that we observed in this study offers a unique system upon which comparative studies can be built. For instance, both an en¬ demic specialist (i.e., M. boydi ) and cosmopolitan generalists (e.g., T. pallidovi¬ rens, and L. hesperus ) feed on S. polygaloides. These two groups differ signifi¬ cantly in metal content (Table 1), yet the function of elevated Ni content in M. boydi and the physiological mechanisms by which it sequesters Ni remain unclear. Research that contrasts the physiological mechanisms of metal tolerance in M. boydi with those of generalist herbivores could address both of these issues. Acknowledgment We would like to thank T. Henry, R Naskrecki, M. Schwartz, and D. Rider for assisting in identification of specimens. Furthermore, we thank M. Davis, A. Teem and E. Watkins for assistance in the field and laboratory. Literature Cited Bagatto, G. & J. D. Shorthouse. 1996. Accumulation of Cu and Ni successive stages of Lymantria dispar L. (Lymantriidae, Lepidoptera) near ore smelters at Sudbury, Ontario, Canada. Env. Poll., 92: 7-12. Berthelsen, B. O., E. Steinnes, W. Solberg & L. Jingsen. 1995. Heavy metal concentrations in plants in relation to atmospheric heavy-metal deposition. J Environ. Qual., 24: 1018-1026. Boyd, R. S. 1998. Hyperaccumulation as a plant defensive strategy, pp. 181-201. In R. R. Brooks (ed.). Plants that hyperaccumulate heavy metals: their role in phytoremediation, microbiology, archaelogy, mineral exploration and phytomining. CAB International, Wallingford. Boyd, R. S. & S. N. Martens. 1998. The significance of metal hyperaccumulation for biotic interac¬ tions. Chemoecology, 8: 1-7. 2002 WALL & BOYD: NICKEL IN SERPENTINE ARTHROPODS 175 Boyd, R. S. & S. N. Martens. 1999. Aphids are unaffected by the elemental defense of the Ni hyperaccumulator Streptanthus polygaloides (Brassicaceae). Chemoecology, 9: 1-7. Brooks, R. R. 1987. Serpentine and its vegetation: a multi-disciplinary approach. Dioscorides Press, Portland. Brooks, R. R. 1998. Phytoarchaeology and hyperaccumulators, pp. 153-180. In R. R. Brooks (ed.). Plants that hyperaccumulate heavy metals: their role in phytoremediation, microbiology, ar¬ chaeology, mineral exploration, and phytomining. CAB International, Wallingford. Brooks, R. R., J. Lee, R. D. Reeves & T. Jaffre. 1977. Detection of nickeliferous rocks by Analysis of herbarium specimens of indicator plants. J. Geochemical Exloration, 7: 49-57. Coleman, R. G. 1967. Low temperature reaction zones and alpine ultramafic rocks of California, Oregon, and Washington. U.S.G.S. Bull. 1247: 1-49. Dallinger, R. 1993. Strategies of metal detoxification in terrestrial invertebrates, pp. 245-289. In R. Dallinger & P. S. Rainbow (eds.). Ecotoxicology of metals in invertebrates. Lewis Publishers, Boca Raton. Davison, G., C. L. Lambie, W. M. James, M. E. Skene & K. E. Skene. 1999. Metal content in insects associated with ultramafic and non-ultramafic sites in the Scottish Highlands. Ecol. Entomol., 24: 396-401. Franklin, A., I. Noell, M. Wakabayashi, R. Wood, K. Shaffer, J. Haas, C. Knowles & J. Knowles. 1997. Red Hills of California: partnership for ecosystem preservation, pp. 173-174. In T. Jaffre, R. D. Reeves & T. Becquer (eds.). The ecology of ultramafic and metalliferous areas. ORSTOM, Noumea. Futuyma, D. J. & G. Moreno. 1988. The evolution of ecological specialization. Annu. Rev. Ecol. Syst., 19: 207-233. Heliovaara, K. & R. Vaisanen. 1990. Heavy-metal content of Bupalus piniarius (Lepidoptera: Geo- metridae) and Panolis flammea (Lepidoptera: Noctuidae) near an industrial source. Environ. Entomol., 19: 481-485. Heliovaara, K., R. Vaisanen, E. Kemppi & M. Lodenius. 1990. Heavy metal concentration in males and females of three pin diprionids (Hymenoptera.). Entomol. Fennica, 1: 175-179. Hopkin, S. P. 1989. Ecophysiology of metals in terrestrial invertebrates. Elsevier Applied Science, London. Koricheva, J. & E. Haukioja. 1995. Variations in chemical composition of birch foliage under air- pollution stress and their consequences for Eriocrania miners. Environ. Pollut., 88: 41-50. Kruckeberg, A. R. 1984. California serpentines: flora, vegetation, geology, soils, and management problems. University of California Press, Berkeley. Kruckeberg, A. R., and R. D. Reeves. 1995. Nickel accumulation by serpentine species of Streptanthus (Brassicaceae): field and greenhouse studies. Madrono 42: 458-469. Lee, J., R. D. Reeves, R. R. Brooks & T. Jaffre. 1978. The relationship between nickel and citric acid in some nickel-accumulating plants. Phytochemistry, 17: 1033-1035. Posthuma, L. 1990. Genetic differentiation between populations of Orchesella cincta (Collebola) from heavy-metal contaminated sites. J. Appl. Ecology, 27: 609-622. Proctor, J. & S. R. J. Woodell. 1975. The ecology of serpentine soils. Adv. Ecol. Research, 9: 255- 266. Raven, P. H. & D. I. Axelrod. 1978. Origin and relationship of the California flora. Univ. Calif. Publ. In Botany, 72: 1-134. Reeves, R. D., R. R. Brooks & R. M. MacFarlane. 1981. Nickel uptake by California Streptanthus and Caulanthus with particular reference to the hyperaccumulator S. polygaloides Gray (Bras¬ sicaceae). Am. J. Bot., 68: 708-712. SAS Institute. 1998. StatView: StatView Reference. Second Edition. SAS Institution, Cary. Schwartz, M.D. & R.G. Foottit. 1998. Revision of the Nearctic Species of the Genus Lygus Hahn with a Review of the Palaeartic Species (Heteroptera: Miridae). Associated Publishers, Gainsville. Wall, M. A. 1999. Nickel accumulation in serpentine arethropods with emphasis on a species of Melanotrichus (Heteroptera: Miridae). M.S. Thesis, Auburn University, Alabama. Wheeler, A. G. Jr. 2001. Biology of the plant bugs (Hemiptera: Miridae): pest, predators, opportunists. Cornell University Press, Ithaca. Wild, H. 1975. Termites and the serpentines of the Great Dyke of Rhodesia. Trans. Rhod. Sci. Assoc., 57: 1-11. Wu, L., A. D. Bradshaw & D. A. Thurman. 1975. The potential for evolution of heavymetal tolerance 176 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) in plants III. The rapid evolution of copper tolerance in Agrostis stolonifera. Heredity, 34: 165— 187. Zar, J. H. 1996. Biostatistical analysis. Prentice-Hall, Englewood Cliffs. Received 28 December 2000; Accepted 20 June 2002. PAN-PACIFIC ENTOMOLOGIST 78(3): 177-183, (2002) OVIPOSITION PREFERENCES OF SCIKTOTHRIPS PERSEAE NAKAHARA (THYSANOPTERA: THRIPIDAE) IN SOUTHERN CALIFORNIA AVOCADO ORCHARDS Mark S. Hoddle Department of Entomology, University of California, Riverside, California 92521, U.S.A. Abstract .—A survey was conducted over the period July 1998-1999 to determine the oviposition preferences of female Scirtothrips perseae in a southern California avocado orchard. Female thrips oviposited into the undersides and topsides of immature avocado leaves, small fruit, and petioles from immature fruit. A significant oviposition hierarchy was determined with immature fruit being most preferred for oviposition followed by undersides of immature leaves, immature fruit petioles, and the topsides of immature leaves. Immature leaf petioles and stems were not used for oviposition. Of field collected fruit, small fruit 25-54 mm in length were most preferred for oviposition as fruit in this size range, on average, had the greatest mean numbers of S. perseae larvae emerging from them. The results of this work have important applications for the development of integrated pest management (IPM) programs using carefully timed natural enemy releases and pesticide applications to reduce low-density S. perseae populations before fruit of a size vulnerable to thrips feeding damage is set on trees. Key Words .—Insecta Scirtothrips perseae, Per sea americana, avocado, oviposition preference. Scirtothrips perseae Nakahara (Thysanoptera: Thripidae) is a new pest of av¬ ocados ( Persea americana Mill. [Lauraceae]) in California USA, and at time of discovery this insect was a species new to science (Nakahara 1997). Populations of S. perseae were first found in June 1996 damaging avocado fruit and foliage in Saticoy and Oxnard, Ventura County, and later around Irvine, Orange County, both in California. By July 1997, infestations of S. perseae were north of the initial discovery areas into San Luis Obispo County and south into San Diego County (Hoddle & Morse 1997). This pest is native to Mexico and Guatemala, and in California, S. perseae has only been found feeding on avocados suggesting that this thrips has a highly restricted host range (Hoddle et al. 2002). Scirtothrips perseae builds to high densities on immature avocado foliage and cumulative feeding damage by larvae and adults can induce premature defoliation by mid to late spring. Thrips larvae and adults feeding on immature fruit are the primary cause of economic damage to avocados in California. Feeding damage results in brown scarring to fruit skin as it matures. Heavily infested orchards in Ventura County experienced 50-80% crop damage in 1997, and much of the damaged fruit was either unmarketable or downgraded in packinghouses. In 1998, crop loses due to damaged fruit that were downgraded and increased production costs due to insecticide use to control S. perseae, cost the California avocado industry approximately $7.6-13.4 million (US) (Hoddle et al. 1998, 1999). Little is known about the developmental and reproductive biology, and field ecology of S. perseae in its native range or California. The purpose of this work was to determine what substrates are most preferred for oviposition by S. perseae in avocado orchards. Improved understanding of the egg-laying choices by fe¬ males may assist in timing natural enemy releases or pesticide applications to protect the most preferred oviposition substrates. Optimizing treatment timing 178 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) may maximize control impact and reduce the number of spray applications needed to prevent thrips from causing economic damage. Materials and Methods Study Site .—A commercial 40 ha ‘Hass’ avocado orchard (85% of fruit pro¬ duction in California is from the ‘Hass’ cultivar) in Bonsall, San Diego County, California, USA (33°16.45 N, 117°13.09 W, elevation 124 m) was selected for this study. This orchard had a very heavy S. perseae infestation when surveys were conducted over the period July 1998-July 1999. No sprays were applied for thrips control over this time period. The orchard was located in plant climate zone 2S (southern coastal valley [Kimball & Brooks 1959]) and subject to a moderating marine influence. Surveying Potential Oviposition Sites .—Potential oviposition substrates used by S. perseae were investigated by collecting % expanded avocado leaves, young green twigs from terminal branches, petioles from % expanded leaves, immature fruit, and fruit petioles from the study site. Plant parts were measured, placed on foam pads saturated with water in stainless steel trays, and held in temperature cabinets at 25° C under long days (L:D 14:10). Glass cells (2.8 cm diameter, 1.5 cm height) with the top opening covered with polyester mesh (95 micron open¬ ings) were adhered to upper leaf and lower leaf surfaces with Duco® Stik-Tak (Devcon Consumer Product, Illinois, USA) to trap emerged larvae. Plant parts were examined daily, and numbers of emerged S. perseae larvae were recorded for seven consecutive days following field collection. Emergence of Larvae from Immature Avocado Leaves .—Mean numbers of lar¬ vae emerging from immature leaves and percentage infested leaves were deter¬ mined by making weekly collections of 20 leaves at the study site from July 1998-March 1999. Leaves were placed upper side down on water-saturated foam pads in stainless steel trays and held at 25° C under long days (L:D 14:10) in temperature controlled cabinets. Larval emergence per leaf was recorded daily for seven days. Determining Avocado Lruit Size Preferences for Oviposition .—Substantial off bloom over the summer of 1998 supplemented normal fruit production in spring 1999 and resulted in significant numbers of fruit in all size categories being present over the course of this survey. Immature avocados were picked weekly from fruit bearing trees at the study site. A total of 1066 fruit were collected from 29 January 1999 to 12 July 1999. Harvested fruit were numbered, and per fruit lengths (mm) and diameters (mm) were recorded. Fruit were adhered to the bot¬ toms of stainless steel pans with Duco® Stik-Tak, and pans were partially filled with water to prevent S. perseae larvae leaving fruit from which they had emerged. Fruit in pans were placed in temperature controlled cabinets at 25° C under long days (L:D 14:10) and numbers of emerged larvae per fruit were re¬ corded and removed with a moistened paint brush daily for seven consecutive days. Fruit were placed in one of 15 size categories based on length. The mean number of emerged larvae, and percentage of fruit infested with S. perseae in each size category were calculated. Statistical Analyses. —Numbers of S. perseae larvae emerging from potential oviposition substrates were compared using Log-likelihood Ratio Test (i.e., G- test) to determine if substrate preferences for oviposition existed. Pair-wise com- 2002 HODDLE: SCIRTOTHRIPS PERSEAE OVIPOSITION IN AVOCADO 179 Table 1. Total number of emerged Scirtothrips perseae larvae from potential oviposition substrates. Collected plant material was incubated at 25°C for seven days. Numbers followed by italicized Roman numerals are significantly different from each other. Oviposition substrate Size (cm) ± SE n No. emerged larvae Young leaf petioles 5.95 ± 0.12 a 40 0 % Expanded avocado leaves (tops) 52 4 i % Expanded avocado leaves (bottoms) 51 78 a Thin green stems 0.75 ± 0.06 b 40 0 Thick green stems 1.47 ± 0.10 b 40 0 Immature fruit 2.85 ± 0.17 b 15 43 Hi Immature fruit petioles 9.90 ± 0.54 a 15 5 iv a Mean length. b Mean diameter. parisons between substrates from which larvae emerged were made using x 2 as sample sizes were large (Sokal and Rohlf 1995). Numbers of S. perseae larvae emerging per fruit in each size category were log-transformed and mean numbers of emerged larvae were compared across size categories using ANOVA in SAS (SAS 1990) with Tukey’s Studentized Range Test at the 0.05 level of significance being used for means separation. Results Oviposition Preferences.—Scirtothrips perseae larvae emerged from the tops and bottoms of immature avocado leaves, immature fruit, and the petioles attached to collected fruit. Significant differences in larval emergence from different ovi¬ position substrates were observed (x 3 2 = 131.28; P < 0.001). Significantly more S. perseae larvae emerged from immature fruit (xi 2 = 10.17; P = 0.001) followed by emergence from the undersides of immature leaves (xi 2 = 17.25; P < 0.001) > immature fruit petioles (xi 2 = 4.63; P — 0.031) > topsides of immature leaves. Immature leaf petioles and stems were not used for oviposition (Table 1). Larval Emergence Patterns from Immature Leaves. —Larval emergence from immature leaves declined to very low levels from August to mid November 1998. Emergence rates peaked at the end of February 1999 at 33.18 ± 10.05 emerged larvae (range 0-189 emerged larvae per leaf) per leaf and declined by approxi¬ mately 50% to 15.58 ± 2.73 larvae per leaf in early April 1999 (Fig. 1A). During periods of very low larval thrips emergence, percentage of infested leaves reached zero on only four occasions, twice in September and November 1998 (Fig. IB). Fruit Size Preferences for Oviposition.—Scirtothrips perseae larvae emerged from 55.66% of 1066 fruit that were 4-96 mm in length. Significant differences in mean numbers of larvae emerging from each fruit size category (F (14 1045) = 45.75; P < 0.0005) (Fig. 2A). The largest mean number (34.63 larvae per fruit ± 5.27 [SE]) of larvae per fruit emerged from fruit in the 40-44 mm length category and the lowest number of thrips larvae emerged from fruit >75 mm in length (0.05 larvae per fruit ± 0.04 [SE]) were observed (Fig. 2A). The highest percentage of infested fruit were 40-44 mm in length (94.37%) and the least infested fruit category (4.11%) were >75 mm in length (Fig. 2B). 180 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) Date Leaves Harvested Figure 1. (A) Mean number (± SE) of emerged Scirtothrips perseae larvae per leaf and (B) percentage of leaves from which S. perseae larvae emerged for leaves collected at Bonsall, California. Discussion Scirtothrips perseae females preferentially oviposited into immature avocado fruit, with the undersides of immature avocado leaves being the next highly pre¬ ferred oviposition site when these two substrates were simultaneously available. Upper surfaces of immature leaves and immature fruit petioles were the least favored oviposition substrates and no larvae were recovered from immature leaf 2002 HODDLE: SCIRTOTHRIPS PERSEAE OVIPOSITION IN AVOCADO 181 0) 0) 0> 0) 0) o> in CM w 7 in (O N 1 ID 1 in ■ in in ■ in ■ in A T- CM in 10 o O) o (0 o d) N (/) T3 d) 't-* d> 100 90 80 70 60 50 40 30 20 10 0 B 0) 0) 0) 0) 0) 0) in CM cn in (0 1 10 B in i in i in ■ in 8 in A CM cn * in (0 Fruit Size Category (Length mm) Figure 2. (A) Mean number (± SE) of emerged Scirtothrips perseae larvae per fruit length cat¬ egory, and (B) percentage of fruit in each length category from which S. perseae larvae emerged. petioles and green twigs indicating that egg-laying females do not utilize these structures. Significantly more S. perseae larvae emerged from field-collected fruit 25-54 mm in length than other size categories. Avocados spontaneously abort —90% of fruit <10 mm in length (Yee et al. 2001a) and selection of fruit by female S. perseae in the size range 25-54 mm is probably under strong selection pressure 182 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) as the feeding resource selected for larvae by females needs to be mature enough to remain on trees, yet tender enough to permit oviposition and sufficient larval feeding time for immature thrips to complete development before fruit skin is too thick to feed on (i.e., fruit >55 mm in length). Field observations indicate that S. perseae is most commonly found on fruit 20-40 mm in length, and most economic scarring occurs over a two week period when fruit 5-14 mm retained by trees are used for feeding by adult and immature thrips (Yee et al. 2001 ab). Emergence of S. perseae larvae from field collected leaves over July-August and November-March suggests that large founding populations of thrips could be accidentally imported into the USA on smuggled plant material. Work on Seri- cothrips staphylinus Haliday (Thysanoptera: Thripidae) used for the biological control of Ulex europaeus (L.), a noxious weed in New Zealand, has demonstrated that 33% of carefully managed releases of just 10 adult thrips into a permissive environment can result in establishment and proliferation (Memmott et al. 1998). On average, more than 10 larval S. perseae per leaf emerged over October-March in this study suggesting that individual leaves may harbor enough thrips eggs to found incipient populations. As part of an IPM program being developed for S. perseae in California, mon¬ itoring of low-density pest populations and application of carefully timed insec¬ ticide sprays with high selectivity towards thrips on immature foliage during the pre-bloom period is being investigated. The results of this oviposition preference study suggest that proactive pesticide applications or natural enemy releases (e.g., Franklinothrips orizabensis Johansen [Thysanoptera: Aeolothripidae] [Hoddle et al. 2000, 2001a, b]) on trees with immature leaves in spring prior to fruit set may help to selectively protect the most preferred oviposition and larval feeding sites from S. perseae. Acknowledgments This work was supported in part by funds from the California Avocado Com¬ mission. Jennifer Jones assisted with fieldwork and data entry. Mr. David Hedrick kindly provided unlimited access to Rancho Camargo. Literature Cited Hoddle, M. S. & J. G. Morse. 1997. Avocado thrips: a serious new pest of avocados in California. California Avocado Society Yearbook, 81: 81-90. Hoddle, M. S., J. G. Morse, P. A. Phillips & B. A. Faber. 1998. Progress on the management of avocado thrips. California Avocado Society Yearbook, 82: 87-100. Hoddle, M. S., J. G. Morse, W. L. Yee & P. A. Phillips. 1999. Further progress on avocado thrips biology and management. California Avocado Society Yearbook, 83: 105-126. Hoddle, M. S., L. Robinson, K. Dreshcer & J. Jones. 2000. Developmental and reproductive biology of a predatory Franklinothrips n. sp. (Thysanoptera: Aeolothripidae). Biol. Cont., 18: 27-38. Hoddle, M. S., S. Nakahara & P. A. Phillips. (2002). Foreign exploration for Scirtothrips perseae Nakahara (Thysanoptera: Thripidae) and associated natural enemies on avocado {Persea amer- icana Miller). Biol. Cont. 24: 251-265. Hoddle, M. S., K. Oishi & D. Morgan. 2001a. Pupation biology of Franklinothrips orizabensis (Thy¬ sanoptera: Aeolothripidae) and harvesting and shipping of this predator. Fla Entomol. 84: 272- 281. Hoddle, M. S., J. Jones, K. Oishi, D. Morgan & L. Robinson. 2001b. Evaluation of diets for the development and reproduction of Franklinothrips orizabensis (Thysanoptera: Aeolothripidae). Bull. Entomol. Res., 91: 273-280. 2002 HODDLE: SCIRTOTHRIPS PERSEAE OVIPOSITION IN AVOCADO 183 Kimball, M. H. & F. A. Brooks. 1959. Plantclimates of California. Cal. Ag., 13: 7-12. Memmott, J. Fowler, S. V. & R. L. Hill. 1998. The effect of release size on the probability of estab¬ lishment of biological control agents: gorse thrips (Sericothrips staphylinus ) released against gorse (Ulex europaeus ) in New Zealand. Biocontrol Science and Technology, 8: 103-115. Nakahara, S. 1997. Scirtothrips perseae (Thysanoptera: Thripidae), a new species infesting avocado in southern California. Insecta Mundi, 11: 189-192. SAS Institute. (1990). SAS/STAT User’s Guide: Statistics Version 6. SAS Institute, Cary, North Car¬ olina. Sokal, R. R. & Rohlf, F. J. 1995. Biometry: the principles and practice of statistics in biological research. Third Edition. W.H. Freeman and Company, New York. Yee, W. L., P. A. Phillips, J. L. Rodgers & B. A. Faber. 2001a. Relationships between Scirtothrips perseae (Thysanoptera: Thripidae) populations on avocado leaves, fruit, and scarring damage on fruit. Environ. Entomol., 30: 932-938. Yee, W. L., P. A. Phillips, J. L. Rodgers & B. A. Faber. 2001b. Phenology of arthropod pests and associated natural predators on avocado leaves, fruit, and in leaf litter in Southern California. Environ. Entomol., 30: 892-898. Received 16 October 2001; Accepted 20 June 2002. PAN-PACIFIC ENTOMOLOGIST 78(3): 184-187, (2002) THE DISCOVERY OF THE GENUS GNAMPTODON HALIDAY (HYMENOPTERA: BRACONIDAE) IN CHINA, WITH DESCRIPTION OF ONE NEW SPECIES Xuexin Chen 1 ’ 2 , J. B. Whitfield 2 , & Junhua He 1 institute of Applied Entomology, Zhejiang University, Hangzhou 310029, China department of Entomology, University of Illinois, 320 Morrill Hall, 505 S. Goodwin Ave, Urbana, Illinois 61801, U.S.A. Abstract. —Two species of Gnamptodon are reported in this paper from China including one new species, Gnamptodon chinensis sp. nov. It represents the first record of the genus Gnamptodon as well as the subfamily Gnamptodontinae in China. Key Words. —Insecta, Hymenoptera, Braconidae, Gnamptodontinae, Gnamptodon, Braconidae, new species, China. The genus Gnamptodon Haliday contains some of the smallest Braconidae, usually scarcely longer than 1 mm, which are exclusively parasitoids of the mining caterpillars of Nepticulidae (Lepidoptera). Thirty seven species, i.e., 15 Palaearc- tic, 7 Nearctic, 3 Oriental, 8 Australian, 3 Afrotropical and 1 Neotropical, have been described worldwide at present (van Achterberg 1983, 1988; Belokobylskij 1987; Narendran & Rema 1996; Papp 1996, 1997; Tobias & Saidov 1997). There were no species recorded in China before this study, although several species have been reported from adjacent countries, such as G. orientalis van Achterberg from Thailand, G. nepalicus Fischer from Nepal, G. indicus Narendran & Rema and G. malabaricus Narendran & Rema from India, and Gnamptodon georginae van Achterberg from the Russian Far East. The species of this genus seem in general to be rarely collected. Only five specimens were found during this study after the first author examined all of the most important collections in China, including the Parasitic Hymenoptera Collection in Zhejiang University (which started in the 1920s and contains about 0.5 million pinned specimens, and as many specimens in alcohol) and the insect collections of Academia Sinica in Beijing and Shanghai. Two species of Gnamptodon are recognized in this paper from China, including one new species, Gnamptodon chinensis sp. nov., from the Oriental part of the country. It represents the first record of the genus Gnamptodon as well as of the subfamily Gnamptodontinae in China. Specimens of the two Chinese species were collected by sweep net, and therefore there is no host record. The subfamily Gnamptodontinae contained 4 genera originally, i.e., Gnamp¬ todon Haliday, 1837 (cosmopolitan), Pseudognaptodon Fischer, 1965 (New World), Gnaptogaster Tobias, 1976 (Palaearctic) and Liparophleps Enderlein, 1920 (Neotropical) (van Achterberg 1983). Liparophleps has subsequently been determined to represent a junior synonym of Semirhytus Szepligeti (Doryctinae), and thus has been removed from Gnamptodontinae (Marsh 1976, Wharton 1997). Recently Belokobylskij (1999) described another monotypic genus, Neognamp- todon Belokobylskij, 1999 of the subfamily from Madagascar. Only the biology of Gnamptodon and Pseudognaptodon is known; both are parasitoids of nepticulid 2002 CHEN: NEW GNAMPTODON 185 Figures 1-2. Gnamptodon chinensis sp. nov., holotype. 1, wings. 2, first-third metasomal tergites, dorsal view. larvae, but it is not yet definitively demonstrated whether they are endo- or ec- toparasitoids. For the identification of the genus Gnamptodon Haliday, see van Achterberg (1983). For the terminology used in this paper, see van Achterberg (1993) and Chen & He (1997). The type specimen is deposited in the Hymenoptera Collec¬ tion, Zhejiang University, Hangzhou, China (ZJU). Gnamptodon Chinensis Chen & Whitfield, NEW SPECIES Gnamptodon chinensis sp. nov. (Figs. 1 and 2) Description-. —Female: body length 1.5 mm, fore wing length 1.5 mm. Head: Antennal segments 19, length of third segment 1.2 times fourth segment, length of third, fourth and penultimate segments 3.3, 2.8 and 2.6 times their width respectively; length of eye 2.3 times temple in dorsal view; POL: OD: OOL = 4:2:9; frons virtually flat and distinctly granulate; vertex concave and smooth; face distinctly convex and smooth, with long setae; length of malar space 1.6 times basal width of mandible. Mesosoma: Length of mesosoma 1.6 times its height; mesosoma smooth; mesoscutal lobes nearly glabrous, without medial depression; scutellar sulcus narrow and finely crenulate. Wings: Fore wing, r: 3-SR: SRI = 4:12:48; 1-CU1: 2-CU1 = 2:16; 2-SR: 3-SR: r-m = 16:12:10; length of pterostigma 0.7 times vein 1-R1; length of distance between apex of wing and apex of marginal cell 0.2 times vein 1-R1; pterostigma robust; vein SRI nearly straight. Legs: Length of femur, tibia, and basitarsus of hind leg 3.6, 7.0 and 4.0 times their width respec¬ tively. Metasoma: Length of first tergite equal to its apical width, its surface distinctly longitudinally rugose, with dorsal carinae developed; curved transverse elevation of second tergite distinct and smooth, second tergite behind the transverse elevation distinctly longitudinally rugose, apical margin smooth; median length of elevation of second tergite 0.36 times median length of rest of tergite; second intersegmental suture of metasoma distinct, crenulate, with no additional grooves; third tergite basally longitudinally striate, rest of third tergite and following tergites smooth; ovipositor slightly curved downwards, with nodus subapically; length of sheath 0.08 times fore wing, 0.8 times hind basitarsus. Color: Head yellowish brown, vertex darker; antenna brown, basal four segments yellowish; palpi yellow; mesosoma reddish brown, mesoscutum, scutellum and propodeum darker reddish brown; legs brownish yellow, tarsi and hind tibia yellow; metasoma darker reddish brown, ventrally, tergites 2-4 laterally and fifth and its following tergites brownish yellow. Wing membrane hyaline, pterostigma and veins brown. Male: Unknown. 186 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) Material examined: holotype female, China: Zhejiang, Mt. Gutian, 1990.vii—viii, Ma Yun, no.905760. Diagnosis .—This species is from the Oriental part of China and is similar to G. orientalis van Achterberg, 1983, but can be separated from the latter by having the frons distinctly granulate; scutellar sulcus finely crenulate; length of distance between apex of wing and apex of marginal cell of fore wing 0.2 times vein 1- Rl; pterostigma robust; and first and second tergites distinctly longitudinally ru¬ gose. It is also similar to G. pumilio (Nees), but can be readily distinguished from the latter by the sculpture of the metasomal tergites. It also can be easily separated from the other known species from China, G. georginae van Achterberg, 1983, by having the much longer vein 1-R1 of the fore wing. Gnamptodon georginae van Achterberg, 1983 Gnamptodon georginae van Achterberg, 1983, Tijdscrift voor Entomologie, 126(2): 33. Material examined: 1 female, China: Liaoning, Dalian, 1991.ix.4, Lou Juxian, no.975981. Distribution: China (Liaoning); Russia Far East, Algeria, Bulgaria, Switzerland, Hungary. Note: This species is new to China. Acknowledgment We thank Dr C. van Achterberg (Leiden, the Netherlands) for his comments on the first draft. The project was partly supported by a National Scientific Foun¬ dation of China (NSFC: 39970099) to the first author. Literature Cited Achterberg, C. van. 1983. Revisionary notes on the subfamily Gnaptodontinae, with descriptions of eleven new species (Hymenoptera: Braconidae). Tijdschr. v. Entomol., 126(2): 25-57. Achterberg, C. van. 1988. A new species of the genus Gnamptodon from Italy (Hymenoptera: Bra¬ conidae). Entomol. Bericht., Amsterdam, 48(10) 1988: 159-161. Belokobylskij, S. A. 1987. Subfamily Gnaptodontinae (Hymenoptera: Braconidae) in the USSR Far East. pp. 78-83. In Kapustina, O. G. (ed.). Taxonomy of insects of Siberia and USSR Far East. Vladivostok, 1987: 1-132. Belokobylskij, S. A. 1999. New genera of the subfamilies Rhyssalinae, Exothecinae and Gnampto- dontinae from the Old World (Hymenoptera: Braconidae). Zoosystem. Ross., 8(1): 155-169. Chen, Xuexin & H, Junhua. 1997. Revision of the subfamily Rogadinae (Hymenoptera: Braconidae) from China. Zool. Verhand. Leiden, 308: 1-187. Fischer, M. 1987. Hymenoptera. Opiinae 3—aethiopische, orientalische, australische und ozeanische Region. Tierreich, 104: ix-xv, 1-734. Marsh, R M. 1976. Pars 13. Braconidae 9, Doryctinae. pp. 1331. In Vecht, J. vander & R. D. Shenefelt (eds.). Hymenopterorum Catalogus (Nova Editio). Dr. W. Junk, The Hague. Narendran, T. C. & C. G. Rema. 1996. Three new species of Braconidae (Hymenoptera) from India. J. Ecobiol., 8(2), 1996: 135-142. Papp, J. 1996. Braconid wasps from the Cape Verde Islands (Hymenoptera, Braconidae) 1. Cheloninae, Exothecinae, Homolobinae, Microgastrinae, Rogadinae. Bol. Mus. Munic. Funch., 48: 197-216. Papp, J. 1997. New braconid wasps (Hymenoptera, Braconidae) in the Hungarian Natural History Museum, 5. Ann. Hist.-Nat. Mus. Nat. Hung., 89: 157-175. Tobias, V. I. & N. Sh. Saidov. 1997. Two new species of braconid wasps (Hymenoptera, Braconidae) from Tajikistan. Entomol. Oboz., 76(1): 210-212, 236. 2002 CHEN: NEW GNAMPTODON 187 Wharton, R. A. 1997. Subfamily Gnamptodontinae. pp. 256-259. In Wharton, R. A., P M. Marsh & M. J. Sharkey (eds.). Manual of the new world genera of the family Braconidae (Hymenoptera). Spec. Pub. Intern. Soc. Hymenopt., No. 1. Washington, D.C. Received 20 February 2002; Accepted 8 May 2002. PAN-PACIFIC ENTOMOLOGIST 78(3): 188-196, (2002) TWO NEW SPECIES OF BETELGEUSE FROM MEXICO (HYMENOPTERA: BRACONIDAE: EUPHORINAE) Scott Richard Shaw U.W. Insect Museum, Department of Renewable Resources, University of Wyoming, Laramie, Wyoming 82071-3354 Abstract .—Two new species of euphorine Braconidae from Mexico, Betelgeuse piceus, NEW SPECIES and Betelgeuse variabilis, NEW SPECIES, are described and illustrated. A key to the three known species is included. The male of B. variabilis is described, and represents the first record of males for this genus. Previously all genera in the tribe Dinocampini were thought to reproduce via thelyotokous parthenogenesis, with males being absent or extremely rare. Unusual variation of the fore wing vein Rs+M is documented and discussed. Key Words. —Insecta, Braconidae, Euphorinae, Dinocampini, Betelgeuse, Mexico, new species. The genus Betelgeuse (pronounced “beetle-juice”) was described by Shaw (1988) to include one Mexican species placed in the tribe Dinocampini. The genus is named after the star Betelgeuse in the constellation Orion (a sword-bearing hunter in Greek mythology), because the female wasp has a conspicuous sword¬ like ovipositor (Fig. 14). Despite its distinctive appearance and ease of recogni¬ tion, specimens of Betelgeuse are extremely rare in collections. The purpose of this paper is to describe two new species of Betelgeuse based on material from the Canadian National Collection (Ottawa) and the California Academy of Sci¬ ences (San Francisco). Like the type-species of the genus, Betelgeuse aztecus, both new species also have females with serrate antennae and are known only from Mexico. Males are described for one of the new species, B. variabilis, this being the first record of males for the genus. Sexual dimorphism of the antenna is documented (Figs. 7 and 8). The same species also exhibits unusual variation of the fore wing vein Rs + M (Figs. 10—13). Materials and Methods Betelgeuse species can be identified as members of the subfamily Euphorinae using the keys of Shaw (1995) or Sharkey (1997). Diagnosis of the genus Betel¬ geuse follows that of Shaw (1988). Specimens can be determined as Betelgeuse using the key provided by Shaw (1997). The genus is easily distinguished by its very coarse head (Fig. 1) and mesosomal sculpture, large propodeal tubercles (Fig. 14), and females with serrate antenna (Figs. 3-5). Members of this genus are the only known braconid wasps with serrate antennae. The morphological terminology used here follows that of Sharkey and Wharton (1997). Fore wing venation terminology is illustrated in Fig. 9. Microsculpture terminology follows that of Harris (1979). Since specimens have the metasoma bent in different positions, body length was measured by adding the combined lengths of the head, mesosoma, and metasoma (excluding the ovipositor). Key to Female Specimens of Betelgeuse la. Antenna with 13 flagellomeres (Fig. 14); apical flagellomere about 2X longer than preceding flagellomere; head and mesosoma mostly orang- 2002 SHAW: NEW BETELGEUSE FROM MEXICO 189 ish brown; forewing with stigma dark brown; second subdiscal cell of forewing mostly clear or faintly infumate . Betelgeuse aztecus Shaw lb. Antenna with 9 flagellomeres (Fig. 7); apical flagellomere at least 3X longer than preceding flagellomere (Fig. 6); head and mesosoma mostly very dark brown or black; forewing with stigma nearly black; second subdiscal cell of forewing with a deeply infumate, very distinct, darkly pigmented patch (Figs. 9 and 10) . 2 2a. Head and mesosoma black; cross-vein lcu-a of forewing angled poste¬ riorly towards wing base (Fig. 9) ... Betelgeuse piceus, NEW SPECIES 2b. Head and mesosoma dark brown infumated with black; cross-vein lcu-a of forewing vertical or angled slightly away from wing base (Fig. 10) . Betelgeuse variabilis NEW SPECIES Betelgeuse piceus Shaw, NEW SPECIES (Figs. 1-6, 9) Types. —Holotype, female; data: MEXICO. Chis (= Chiapas), San Cristobal (de las Casas), 2200 m, 26-27 May 1990, H. Howden, B. Gill, FIT [deposited at Canadian National Collection, Ottawa]. Paratype: 1 female, same data as holotype [deposited at University of Wyoming Insect Museum, Laramie]. Description of Holotype Female .—Body length (excluding ovipositor) 6.0 mm; forewing length 3.3 mm. Head transverse, in dorsal view 2.1 X broader than long; surface sculpture coarsely and evenly rugose (Fig. 1); eye elongate oval, not bulging anteriorly beyond face; eyes in anterior view distinctly converging ventrally; shortest inter-ocular distance 1.6X clypeus width; eye apparently glabrous; me¬ dian frontal carina absent, obscured by rugose sculpture; inter-antennal distance 2.4X socket width; scrobes very slightly protuberant; scape elongate, gradually curved, gradually wider apically; scape length 4.4X width at apex (Fig. 2); pedicel somewhat globose (Fig. 3); flagellum 9-segmented, con¬ siderably shorter than body length; flagellomeres 1-5 longer than wide (Figs. 3-5), F1-F4 of similar width, F5 slightly less wide, F1-F5 somewhat flattened, forming serrations antero-laterally, each ser¬ ration terminating apically in a point with a single large seta; FI 2.OX longer than apical width; F2- F5 each relatively shorter than preceding flagellomere; F6-F8 each compact, about as long as wide (Figs. 5 and 6); apical flagellomere (F9) 3.8X longer than wide, apically pointed (Fig. 6); ocellar triangle small, distance between lateral ocellus and compound eye 2.5X distance between lateral ocelli; occipital carina complete dorsally and somewhat crenulate, ventrally absent and not meeting hypos- tomal carina; malar space short, slightly greater than % eye height; malar suture absent; facial setae minute, not obscuring face; lower clypeal margin truncate; mandibles very sharply pointed, when closed overlapping for about 3 4 mandible length; maxillary palpus 5-segmented; labial palpus 2-seg- mented. Mesosoma with surface sculpture entirely coarsely rugose to rugo-punctate; notaulus and sternaulus indistinct from general rugose sculpture; scutellar furrow 8-foveate; anterior margin of scutellar furrow weakly carinate; scutellum large, triangular, and projecting posteriorly as a distinct conical point; propodeum with deep postero-median impression, as deep as basal width of petiole (metasomal tergum 1); postero-lateral corners of propodeum developed as broad lateral tubercles; petiolar notch short, not extending past metacoxal cavity; hind coxa rugose, remainder of leg coarsely granular; metafemur length 3.9X maximum width; tarsal claw simple. Fore wing with pterostigma large, nearly semi-circular; forewing vein 1M gradually and evenly curved; vein RS+Ma absent basally but present apically as very short stub (about Vi length of vein r); vein RS+Mb long and curved; vein 3RS entirely tubular and distinct, reaching wing margin well before wing apex; marginal cell shorter than pterostigma length (about 0.8X pterostigma length); vein r-m absent; vein M+CU slightly curved at middle; tubular portion of vein 2M about equal in length to vein r; vein lCUa angled posteriorly at 110 degrees relative to vein M+CU; vein 2CU slightly longer than m-cu, curved apically; vein lcu-a postfurcal by distance slightly greater than its length, angled posteriorly toward wing margin; vein 2-1A short and straight, about equal in length to lCub. Hind wing with 3 hamuli; vein RS absent basally but apically indicated by infumate line nearly reaching wing tip; vein 2M 190 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) Figures 1-6, scanning electron micrographs of Betelgeuse piceus, NEW SPECIES, 200X magnifi¬ cation, anterior view. Figure 1. Coarse facial sculpture below antennal insertions. Figure 2. Antennal scape showing coarse sculpture. Figure 3. Pedicel and first antennal flagellomere. Figure 4. Second and third flagellomeres. Figure 5. Fourth to sixth flagellomeres. Figure 6. Seventh to ninth flagello- meres (antennal apex). indicated by infumation nearly reaching lower wing margin; vein cu-a curved, meeting vein 1A posteriorly in a tubular joint, without obvious bulla or wing fold. Metasoma with petiole not fused ventrally, apex 2.9X broader than base, about 0.6X as long as metasoma beyond petiole (excluding ovipositor); tergum 1 rugose over basal 3 4, smooth over apical X A\ glymma absent; dorsope absent; petiolar spiracles just beyond middle of petiole, barely prominent; syntergum 2+3 smooth and highly polished, about % as long as petiole; lateral fold of syntergum 2+3 present; suture between terga 2+3 present laterally; terga 4-8 exposed beyond syntergum 2+3, all smooth and highly polished; tergum 8 short and strongly compressed to form a vertical slit above ovipositor; ovipositor (fully exserted) slightly longer than head and mesosoma combined; sheaths narrower, shorter, and curled. Color: Body mostly black, except scape, pedicel, apical flagellomere, mandible, palpi, and metasomal sclerites very dark reddish brown; flagellomeres 6-8, tarsi apically, ovipositor, and sheaths much lighter yellowish brown; eye and ocelli silvery white; metasomal membranes laterally and ventrally white; forewing 2002 SHAW: NEW BETELGEUSE FROM MEXICO 191 with stigma nearly black; second subdiscal cell of forewing with a deeply infumate, very distinct, darkly pigmented patch. Variation .—Single paratype female as in holotype except body length 5.6 mm; fore wing length 3.2 mm; vein RS+Ma absent basally but present apically as a short branch (only slightly shorter than vein r). Diagnosis .—This species can be distinguished from Betelgeuse aztecus by the antenna with only 9 flagellomeres, forewing with the stigma nearly black, and second subdiscal cell of forewing with darkly pigmented patch. Betelgeuse piceus is more similar to Betelgeuse variabilis, NEW SPECIES but can be distinguished by its entirely black head and mesosoma, and by cross-vein lcu-a of the fore wing, which is strongly angled posteriorly towards wing base. Distribution .—Known only from the type-locality in Chiapas, Mexico. Remarks .—The arrangement of antennal placodes in five distinct ranks on fla- gellomere 9 (Fig. 6) indicates that this elongate apical flagellomere is probably formed from the fusion of five flagellomeres. Therefore, the presence of 13 fla¬ gellomeres (as in Betelgeuse aztecus ) is probably the primitive condition for the genus. The elongated 9 th flagellomere is interpreted here as a synapomorphy in¬ dicating the close phylogenetic relationship of the two new species described in this paper. Etymology .—The species epithet piceus is derived from the Latin for “pitch- black,” in reference to the predominant black body color of this species. Betelgeuse variabilis Shaw, NEW SPECIES (Figs. 7 and 8, 10-13) Types .—Holotype, female; data: MEXICO. Hgo. (= Hidalgo), Hwy. 105 2.7 mi N. Tlanchinol, 5000', 15 June 1983, C. W. & L. O’Brien & G. B. Marshall [deposited at California Academy of Sciences, San Francisco]. Paratypes: 1 male, same data as holotype; 1 male, same data as holotype except collected 2 August 1982, C. W. & L. O’Brien & G. Wibner, collectors [deposited at California Acad¬ emy of Sciences, San Francisco and University of Wyoming Insect Museum, Laramie]. Description of Holotype Female .—Body length 4.5 mm; forewing length 3.6 mm. Head transverse, in dorsal view 2.6X broader than long; surface sculpture coarsely and evenly rugose; eye elongate oval, not bulging anteriorly beyond face; eyes in anterior view distinctly converging ventrally; shortest inter-ocular distance 1.4X clypeus width; eye apparently glabrous but with scattered minute setae; median frontal carina absent, rugose facial sculpture interrupted medially by a vertical groove; inter- antennal distance 2.5 X socket width; scrobes very slightly protuberant; scape elongate, gradually curved, gradually wider apically, somewhat flattened dorso-ventrally; scape length 5.OX width at apex; pedicel somewhat globose, except dorsally with a short serration; flagellum 9-segmented, considerably shorter than body length; flagellomeres 1-4 longer than wide, F5 about as long as wide, F1-F4 of similar width, F5 slightly less wide, F1-F5 somewhat flattened, forming serrations antero-laterally, each serration terminating apically in a point with a single large seta; FI 1.8X longer than apical width; F2-F5 each relatively shorter than preceding flagellomere; F6-F8 each compact, about as long as wide; apical flagellomere (F9) 3.8X longer than wide, apically pointed; ocellar triangle small, distance between lateral ocellus and compound eye 2.9X distance between lateral ocelli; occipital carina complete dorsally and somewhat crenulate, ventrally absent and not meeting hypostomal carina; malar space short, about 1/6 eye height; malar suture absent; facial setae minute, not obscuring face; lower clypeal margin truncate; mandibles very sharply pointed; maxillary palpus 5-segmented; labial palpus 2-segmented. Mesosoma with surface sculpture entirely coarsely rugose to rugo-punctate; no- taulus and sternaulus indistinct from general rugose sculpture; scutellar furrow 14-foveate; anterior margin of scutellar furrow weakly carinate; scutellum large, triangular, and projecting posteriorly as 192 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) Figures 7-8, antenna of Betelgeuse varibilis, NEW SPECIES, anterior view. Figure 7. Female. Figure 8. Male. a distinct conical point; propodeum with deep postero-median impression, as deep as basal width of petiole (metasomal tergum 1); postero-lateral corners of propodeum developed as lateral tubercles; petiolar notch short, not extending past metacoxal cavity; hind coxa rugose, remainder of leg coarsely granular; metafemur length 4.5X maximum width; tarsal claw simple. Fore wing with pterostigma large, nearly semi-circular; forewing vein 1M only very slightly curved, nearly straight; vein RS+Ma absent basally but present apically as short branch about !4 to Vi as long as vein r (short branch of vein RS+Ma longer in left wing than in right wing); vein RS+Mb long and slightly curved; vein 3RS entirely tubular and distinct, reaching wing margin well before wing apex; marginal cell about equal to pterostigma length; vein r-m absent; vein M+CU slightly curved at middle; tubular portion of vein 2M about equal in length to vein r; vein lCUa angled posteriorly at 110 degrees relative to vein M+CU; vein 2CU slightly longer than m-cu, curved apically; vein lcu-a postfurcal by distance 1.5X greater than its length, vertical or angled slightly away from wing base; vein 2-1A short and straight, about equal in length to lCub. Hind wing with 3 hamuli; vein RS indicated by infumate line nearly reaching wing tip; vein 2M spectral; vein cu-a curved, meeting vein 1A posteriorly in a tubular joint, without obvious bulla or wing fold. Metasoma with petiole not fused ventrally, apex 3.5X broader than base, about 0.75X as long as metasoma beyond petiole (excluding ovipositor); tergum 1 weakly longitudinally rugose over basal %, smooth over apical Va\ glymma absent; dorsope absent; petiolar spiracles just beyond middle of petiole, not prominent; syntergum 2 + 3 smooth and highly polished, about 3 4 as long as petiole; lateral fold of syntergum 2+3 present; suture between terga 2+3 present laterally; terga 4-8 exposed beyond syntergum 2+3, all smooth and highly polished; tergum 8 short and strongly compressed to form a vertical slit above ovipositor; ovipositor (fully exserted) slightly longer than head and mesosoma combined; sheaths narrower, shorter. Color: Body mostly dark reddish brown, except scape, pedicel, mandible, fore and middle coxae, ovipositor, and sheaths lighter yellowish brown; flagellomeres 6-8 light yellowish white; eye and ocelli silvery white; meta¬ somal membranes laterally and ventrally white; forewing with stigma dark brown; second subdiscal cell of forewing with a deeply infumate, very distinct, darkly pigmented patch. Variation, Paratype Males .—Aside from primary sexual differences, similar to female except body 2002 SHAW: NEW BETELGEUSE FROM MEXICO 193 10 Figures 9-10, fore wing of Betelgeuse females. Figure 9. Betelgeuse piceus, NEW SPECIES. Figure 10. Betelgeuse varibilis, NEW SPECIES. 11 12 13 Figures 11-13, medial area of fore wing of male Betelgeuse varibilis, NEW SPECIES showing variation of vein RS+Ma, first submarginal, and first discal cell. Figure 11. Right wing of paratype (August 2) with vein RS+Ma partly present. Figure 12. Left wing of paratype (June 15) with vein RS+Ma absent. Figure 13. Right wing of paratype (June 15) with vein RS+Ma entirely present. 194 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) Figure 14. Habitus of Betelgeuse aztecus Shaw. somewhat larger and more robust; body length 5.4-6.0 mm; fore wing length 4.1-4.2 mm; head not as broad and narrow as in female, in dorsal view 2.3-2.4X broader than long; eyes smaller and face wider than in female, shortest inter-ocular distance 2.2X clypeus width; rugose facial sculpture not interrupted medially by a vertical groove; inter-antennal distance wider than in female, 3.OX socket width; antenna longer and more slender than in female, not serrate (Fig. 8); scape cylindrical in cross- section, not flattened dorso-ventrally as in female; scape length 4.7 X width at apex; pedicel globose; flagellum slender, flagellomeres 6-11 shorter and wider than Fl-5 giving the flagellum a slightly clavate appearance; FI 9.3X longer than wide; F2 6.7X longer than wide; F3 6.OX longer than wide; F4 5.3X longer than wide; F5 4.3X longer than wide; F6-7 each 2.OX longer than wide; F8-10 each 1.4X longer than wide; Fll (apical flagellomere) 3.5X longer than wide; distance between lateral ocellus and compound eye much wider than in female, equal to 3.5X distance between lateral ocelli; malar space very broad, about % eye height; scutellar furrow 10-12-foveate; mesonotum with antero- submedial areas and postero-lateral areas smoother and slightly depressed, as compared with female; metafemur length 4.8X maximum width; forewing vein 1M gradually and evenly curved; vein RS+Ma varying from completely present, to partly present as a short branch about equal in length to vein r, to completely absent (one specimen has vein RS+Ma absent in the left wing but completely present in the right wing); hind wing vein RS absent basally but apically indicated by infumate line nearly reaching wing tip; vein 2M indicated by infumation nearly reaching lower wing margin; petiolar spiracles situated more posteriorly than in female, about % distance from base of petiole; petiolar sculpture more coarse than in female, extending further posteriorly; metasoma beyond petiole shorter 2002 SHAW: NEW BETELGEUSE FROM MEXICO 195 and broader than in female; tergum 8 not compressed to form a vertical slit; genitalia short and barely exposed; parameres tapering to rounded, setose tips. Diagnosis.—Betelgeuse variabilis, NEW SPECIES can be distinguished from Betelgeuse aztecus by the antenna with only 9 flagellomeres and second subdiscal cell of forewing with darkly pigmented patch. Betelgeuse variabilis is similar to Betelgeuse piceus NEW SPECIES but can be distinguished by its reddish brown head and mesosoma, and by the cross-vein lcu-a of the fore wing, which is vertical or slightly angled away from the wing base (Fig. 10). Distribution .—Known only from the type-locality in Hidalgo, Mexico. Remarks .—This description represents the first record of males for this genus. Previously all genera in the tribe Dinocampini were thought to reproduce via thelyotokous parthenogenesis, with males being absent or extremely rare. Since this species is only known from three specimens, it is uncertain if this record of males is a rare occurrence (as in other dinocampines) or if males are common in this species. The males differ most notably from females by the antennal flagellum being slender and gradually wider apically (Fig. 8), not serrate as in females of this genus. However, the very coarse sculpture of the mesosoma and propodeum with large tubercles remain diagnostic for the genus, regardless of sex. The range of variation in the form of fore wing vein RS+Ma in this species is quite remarkable, and merits special discussion. In most Braconidae this vein is either present or absent, and its presence or absence is often used as a diag¬ nostic character for genera. There are few published cases in the family Bracon¬ idae where a single species exhibits so much variation for wing venation (Konig 1972), and it’s remarkable that only three specimens should show so much vari¬ ation. In the female holotype vein RS + Ma is present as an apical short branch (Fig. 10), but this varies in length between the left and right wing of the same specimen. The male paratype dated 2 August 1982 is presumably “normal” in that the wings are symmetrical, with both wings having an apical branch of vein RS + Ma that extends half-way across the combined first submarginal and discal cells (Fig. 11). The male paratype dated 15 June 1983 is presumably “abnormal” in that the wings are asymmetrical: vein RS+Ma is entirely absent from the left wing (thus the first submarginal and discal cells are combined) (Fig. 12), while it is more or less entirely present in the right wing (thus the first submarginal and discal cells are clearly separated) (Fig. 13). In the right wing of this specimen the basal % of vein RS+Ma is not tubular, but it is clearly indicated by very dark pigmentation. The apical % of the vein is clearly tubular and strongly indicated. In my previous paper (Shaw 1988) the significance of the absence of this vein in Betelgeuse aztecus was discussed (under the older Muesebeckian system the same vein was termed the “first segment of the cubitus”). Since other genera of the tribe Dinocampini (e.g., Dinocampus, Ropalophorus, Centistina ) have this vein present, its absence in Betelgeuse was regarded as a convergence with eu- phorine section 3 tribes (which mostly lack this vein). Nevertheless, the placement of Betelgeuse in the tribe Dinocampini is clearly supported by the elongated scape and labial palpus reduced to 2 segments. The present discovery that two new species of Betelgeuse have the vein at least partly present (and that it is a highly plastic character) lends further support to the interpretation of its total loss in some Betelgeuse species as an evolutionary convergence. 196 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) Etymology .—The species epithet variabilis is derived from the Latin for “changeable,” in reference to the extreme variation of fore wing vein RS + Ma exhibited by this species. Acknowledgments Thanks are given to Michael Sharkey (University of Kentucky, formerly Ca¬ nadian National Collection, Ottawa) and to Robert Zuparko (California Academy of Sciences, San Francisco) for initially recognizing these specimens and calling them to my attention for study. Kathleen M. Horton, Managing Editor of Pysche, kindly granted permission to reprint the beautiful habitus illustration done by Kathy Brown-Wing, which was first published in my 1988 paper. Teresa Williams (Western Research Institute), assisted with the environmental scanning electron microscope. Literature Cited Harris, R. A. 1979. A glossary of surface sculpturing. Occasional Papers in Entomology, 28: 1-31. Konig, R. 1972. Zur systematik, faunistik, phanologie un okolgie mitteleleuropaischer Braconiden (Hymenoptera) (1.) Faunistisch-Oekologisch Mitteilungen, 4(3): 85-106. Sharkey, M. J. 1997. Key to the New World subfamiles of the family Braconidae. pp. 39-63. In Wharton, R. A., P. M. Marsh and M. J. Sharkey (eds.). Manual of the New World genera of the family Braconidae (Hymenoptera). Special Publication of the International Society of Hy- menopterists, Number 1. Sharkey, M. J. & R. A. Wharton. 1997. Morphology and terminology, pp. 19-37. In Wharton, R. A., P. M. Marsh and M. J. Sharkey (eds.). Manual of the New World genera of the family Bracon¬ idae (Hymenoptera). Special Publication of the International Society of Hymenopterists, Num¬ ber 1. Shaw, S. R. 1988. A new Mexican genus and species of Dinocampini with serrate antennae (Hyme¬ noptera: Braconidae: Euphorinae). Psyche, 95: 289-297. Shaw, S. R. 1995. Braconidae. pp. 431-463. In Hanson, P. E. and I. D. Gauld (eds.). The Hymenoptera of Costa Rica. Oxford University Press, Oxford. Shaw, S. R. 1997. Subfamily Euphorinae. pp. 234-254. In Wharton, R. A., P. M. Marsh and M. J. Sharkey (eds.). Manual of the New World genera of the family Braconidae (Hymenoptera). Special Publication of the International Society of Hymenopterists, Number 1. Received 12 April 2002; Accepted 1 August 2002. PAN-PACIFIC ENTOMOLOGIST 78(3): 197-214, (2002) REVIEW OF THE GENUS HESPEROBAENUS LECONTE (COLEOPTERA: MONOTOMIDAE) OF AMERICA, NORTH OF MEXICO Yves Bousquet Eastern Cereal and Oilseed Research Centre, Agriculture and Agri-Food Canada, Ottawa, Ontario K1A 0C6 Abstract .—The genus Hesperobaenus LeConte in North America, north of Mexico, is reviewed. One new species is described, H. constricticollis, NEW SPECIES (type locality: Sabal Palm Grove Sanctuary, near Brownsville, Texas) and a new combination, H. unicolor (Casey), NEW COMBINATION is proposed. Hesperobaenus arizonicus Casey, NEW SYNONYM, is placed for the first time in synonymy with H. abbreviatus (Motschulsky). A key is provided for the discrimination of the species along with distributional maps and illustrations of the most impor¬ tant character states. Key Words .—Insecta, Coleoptera, Monotomidae, Hesperobaenus, new species, new combination, North America. The genus Hesperobaenus was described by John L. LeConte in 1861 for two species, Monotonia rufipennis LeConte of North America, now a junior synonym of Hesperobaenus abbreviatus (Motschulsky), and Rhizophagus capito Fairmaire of Honolulu, Hawaii. Since then, only a few species have been added to the genus, all from North America and Central America. The genus has never been revised and species discrimination is very difficult with the existing literature. The purpose of this work is to provide a taxonomic review of the species of Hesperobaenus occurring in Canada and the United States. Materials and Methods This review is based on the study of about 1600 specimens of Hesperobaenus. The material was borrowed from the following institutions referred to in the text by their acronyms. Names of curators follow the institutional addresses. AMNH: Department of Entomology, American Museum of Natural History, Central Park West at 79 th Street, New York, NY 10024, U.S.A. Lee H. Herman. BMNH: The Natural History Museum, Cromwell Road, London SW7 5BD, England. Malcolm Kerley. CAS: Department of Entomology, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118, U.S.A. David H. Kavan- augh. CDAE: California State Collection of Arthropods, Department of Food and Agriculture, 1220 N Street, Sacramento, California 95814, U.S.A. Fred G. Andrews. CMNH: The Carnegie Museum of Natural History, 4400 Forbes Avenue, Pitts¬ burgh, Pennsylvania 15213-4080, U.S.A. Robert L. Davidson. CNC: Canadian National Collection of Insects, Agriculture and Agri-Food Canada, Ottawa, Ontario K1A 0C6. 198 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) CUIC: Department of Entomology, Cornell University, Ithaca, New York 14850, U.S.A. James K. Liebherr. FMNH: Field Museum of Natural History, Roosevelt Road at Lake Shore Drive, Chicago, Illinois 60605, U.S.A. Alfred F. Newton, Jr. FSCA: Florida State Collection of Arthropods, Florida Department of Agri¬ culture and Consumer Services, P.O. Box 147100, Gainesville, Florida 32614, U.S.A. Michael C. Thomas. INHS: Section of faunistic surveys and insect identification, Illinois Natural History Survey, 607 East Peabody Drive, Champaign, Illinois 61820, U.S.A. Kathryn C. McGiffen. LSUC: Louisiana State University Insect Collection, Department of Entomol¬ ogy, Louisiana State University, Baton Rouge, Louisiana 70803, U.S.A. Vicky L. Moseley. MCZ: Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138, U.S.A. David G. Furth. NHDE: Entomological Museum, Department of Zoology, University of New Hampshire, Durham, New Hampshire 03824, U.S.A. Donald S. Chan¬ dler. OSUC: Department of Entomology, Ohio State University, 1735 Neil Avenue, Columbus, Ohio 43210, U.S.A. Charles A. Triplehorn. TAMU: Department of Entomology, Texas A&M University, College Station, Texas 77843, U.S.A. Edward G. Riley. USNM: National Museum of Natural History, Smithsonian Institution, Wash¬ ington, D.C. 20560, U.S.A. Gloria N. House. The following measurements were made on some specimens using an ocular micrometer in a stereoscopic microscope at 80 X: maximum width of head, in¬ cluding eyes (WH); maximum width of pronotum (WP); length of pronotum along midline (LP); length of elytra from posterior extremity of scutellum to tip of right elytron (LE). Genus Hesperobaenus LeConte, 1861 Hesperobaenus LeConte 1861: 86. Type species: Monotoma rufipennis LeConte, 1858 (= Rhyzophagus abbreviatus Motschulsky, 1845), PRESENT DESIG¬ NATION [the designation of Hesperobaenus abbreviatus by Sharp (1900: 565) is invalid since the species was not originally included in the genus]. Horn (1879a: 262); Blatchley (1910: 667); Casey (1916: 91); Arnett (1962: 768); Sen Gupta (1988: 17, 44); Downie and Arnett (1996: 985, 988). Recognition .—The following character states distinguish members of Hespe¬ robaenus from those of other genera of Nearctic Monotomidae. Head without antennal grooves. Antenna with 3-segmented club (seemingly 2-segmented). Pro- notal disc with impunctate median zone. Elytral disc with setigerous punctures arranged in longitudinal rows; indexed part of elytron with 4—5 rows of punctures (punctures of medial rows are more or less confused in most species). Fore coxae rounded. Sen Gupta (1988) provided a detailed description of the genus. Habitat. —Very little is known about the habitat requirements of the species of Hesperobaenus. Label information attached to specimens studied suggest that they are associated with yucca and sotol plants (family Liliaceae) or are found under 2002 REVIEW OF HESPEROBAENUS 199 the bark of dead trees. The species are probably fungus feeders. Lawrence (1991) reported that Hesperobaenus species have been taken in fruiting bodies of Hy¬ po xylon and Daldinia (Ascomycetes: Xylariaceae). Discussion .—Beside the species treated in the present work, four other names are associated with the genus Hesperobaenus (see Hetschko 1930): capito Fair- maire, 1850 (originally described as a member of Rhizophagus ) reported from Tahiti and the Hawaiian Islands; humeralis Reitter, 1873 listed by Hetschko (1930) as a junior synonym of capito ; lineellis Reitter, 1873 (originally described as a member of Europs ) reported with doubt from North America; and stipes Sharp, 1900 reported from Guatemala. According to Sharp (1900: 565), capito belongs to the genus Europs. I have studied the type specimen of stipes (BMNH); it differs from other Hesperobaenus species in having only three rows of setigerous punc¬ tures on the inflexed part of the elytron. Its generic position remains uncertain but I doubt that it belongs to the genus Hesperobaenus. Key to Nearctic Species of Hesperobaenus 1. Elytral intervals 3 and 5 with setigerous punctures at least over most of anterior half. Male last visible sternite with large, oval and shallow me¬ dian depression . 2 — Elytral intervals 3 and 5 without setigerous punctures or at most with 1— 4 punctures at base. Male last visible sternite without depression . 3 2. Scutellum without setigerous punctures. Eyes convex, temples at least % longitudinal diameter of eyes (Fig. 2) . H. alternatus Schaeffer — Scutellum with 1-4 short setigerous punctures. Eyes less convex, temples distinctly shorter, less than % longitudinal diameter of eyes (Fig. 3) . . . . H. unicolor (Casey) 3. Metacoxal bead, on first visible abdominal sternite, triangularly produced, with or without linear prolongation. 4 — Metacoxal bead, on first visible abdominal sternite, not triangularly pro¬ duced, at most somewhat thickened, without linear prolongation. 6 4. Scutellum with setigerous punctures. Pronotum elongate (LPAVP = 1.06- 1.14). Prosternal apophysis with isodiametric microsculpture .... . H. fenyesi Van Dyke — Scutellum without setigerous punctures. Pronotum transverse to subquad¬ rate (LPAVP = 0.90—1.05). Prosternal apophysis without microsculpture or with micro sculpture near apex only. 5 5. Pronotum subquadrate (LPAVP = 0.96-1.05) with punctures subcontig- uous laterally. Temples long, more than half longitudinal diameter of eyes (Fig. 6). Anterior half of elytra distinctly paler than posterior half in most specimens. Anterior angles of pronotum laterally slightly pro¬ duced in most specimens (Fig. 6) . H. abbreviatus (Motschulsky) — Pronotum slightly transverse (LPAVP = 0.90-0.96) with punctures not subcontiguous laterally. Temples shorter, half longitudinal diameter of eyes or less (Fig. 7). Elytra more or less uniformly colored. Anterior angles of pronotum not produced laterally (Fig. 7). . . H. rufipes LeConte 6. Pronotum only slightly narrowed basally (Fig. 4). Temples long, more than half longitudinal diameter of eyes (Fig. 4) . H. subtestaceus Reitter 200 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) - Pronotum markedly narrowed basally (Fig. 8). Temples shorter, less than half longitudinal diameter of eyes (Fig. 8) . H. constricticollis n.sp. HESPEROBAENUS ALTERNATUS Schaeffer, 1910 Hesperobaenus alternatus Schaeffer, 1910: 213. Type locality: ARIZONA. Huachuca Mts. Type Material .—Schaeffer (1910) described H. alternatus from an unspecified number of specimens collected in the Huachuca Mountains in Arizona. The USNM contains two specimens of that species, a male and a female, in the general collection, both labelled as “Type”. These syntypes bear the following labels: “Huach Mts. Ariz./TYPE/alternatus Schaef. [handwritten]/Hesperobaenus alter¬ natus Schaef. [handwritten]/C. Schaeffer Collection R. 11.II.36 [partly handwrit- ten]/Nevermann Collection 1940”. Description. —Habitus (Fig. 1). Body length: 2.9-3.5 mm. Coloration. —Dorsal surface red-brown, disk of elytra slightly paler than pronotum in many specimens. Microsculpture. —Prosternal apophysis without microsculpture. Head. —Wider in males (WH/WP = 0.97-1.02; x = 0.99; n = 10) than in females (WH/WP = 0.90-0.95; x = 0.92; n = 10). Eye convex, longitudinal diameter 1.5-1.6X length of antennomere I. Temple moderately long, 0.6-0.7X longitudinal diameter of eye, rounded posteriorly, not produced (Fig. 2). Antennomere IX about as wide as long, subequal in width to antennomere X. Prothorax. Pronotum slightly elongate (LP/WP = .99-1.08; x = 1.04; n = 20), with maximal width slightly before apex (Fig. 2); anterior angle rounded, not produced; punctures narrowly spaced laterally but not subcontiguous; disc slightly convex, with narrow median impunctate area. Hypomeron not rugose. Elytra. —Proportionally short (LE/LP = 1.73-1.93; x = 1.83; n = 20), with short, vague, shallow oblique impression on anterior third near suture in many specimens. Third and fifth intervals with numerous setigerous punctures mostly on anterior two-thirds; scutellum without setigerous punc¬ tures. Abdomen. —First visible sternite with coxal bead rounded, not triangularly produced. Male last visible sternite with shallow, oval, median depression. Male Genitalia. —Aedeagus as in Fig. 9. Diagnosis .—Distinguished from other species of Hesperobaenus by the pres¬ ence of setigerous punctures on the third and fifth elytral intervals in combination with the absence of setigerous punctures on the scutellum. Distribution .—This species is known from southeastern Arizona and Texas (Fig. 16). Beside the specimens listed below I have seen seven specimens labelled “Florida: Hillsborough Co. Dover, 11.11.1987, J. Felty palm flowers ex Texas” in Florida State Collection of Arthropods, Gainesville, Florida. Habitat .—Label data indicate that the species is associated with yucca plants. Material Examined. —ARIZONA. COCHISE Co.: Chiric[ahua] Mts (20, USNM). ChiricahuaMtns., Rucker Cny. (2, CNC) [at light]. Chiricahua Mtns. near Portal (6, FSCA) [Yucca], 5 mi W Portal (1, CDAE) [Yucca sp. Lep. frass], W Stronghold (3, FSCA). Stronghold, Dragoon Mtns. (3, MCZ) [ex leaf axils of Yucca with dead flowers in them], Huach[ucha] Mts. (2, USNM). SANTA CRUZ Co.: Santa Rita Mts. (19, MCZ, USNM). Santa Rita Mts., Madera Cyn. (5, FSCA) [1—dead Sotol], TEX¬ AS. BASTROP Co.: Bastrop State Park (1, CNC) [light], BEXAR Co.: San Antonio (19, CNC). BOWIE Co.: Maud (1, OSUC). CAMERON Co.: county record only (17, USNM) [fallen fruit Yucca treculeana Carr.]. Brownsville (8, USNM) [Yucca blossom], GILLESPIE Co.: Fredericksburg (1, CNC). HUD¬ SPETH Co.: Eagle Flat (1, USNM) [ Yucca macrocarpa]. KERR Co.: Kerrville (8, CNC) [Yucca flowers/on yucca], KLEBERG Co.: Kingsville (3, CUIC). SUTTON Co.: Sonora (3, TAMU). “Sinton, Welder Wildlife Fd.” (1, TAMU). Hesperobaenus unicolor (Casey), 1916, NEW COMBINATION Europs unicolor Casey, 1916: 95. Type locality: “TEXAS”. Type Material .—Casey’s collection in USNM contains a single specimen, a 2002 REVIEW OF HESPEROBAENUS 201 Figure 1. Hesperobaenus alternatus Schaeffer (c?), habitus. Scale bar — 1 mm. 202 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) Figures 2-8. Head and pronotum, dorsal view. Figure 2. Hesperobaenus alternatus Schaeffer (c?); Figure 3. H. unicolor Casey (3). Figure 4. H. subtestaceus Reitter (d). Figure 5. H. fenyesi Van Dyke ($). Figure 6. H. abbreviatus Motschulsky ($). Figure 7. H. rufipes LeConte ($). Figure 8. H. constricticollis Bousquet (3). Scale bar =1.0 mm. 2002 REVIEW OF HESPEROBAENUS 203 Figure 9-11. Aedeagus. Figure 9. Hesperobaenus alternatus Schaeffer. Figure 10. H. unicolor Casey. Figure 11 . H. subtestaceus Reitter. Scale bar = 0.2 mm. 204 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) male, labelled: “Tex./Casey bequest 1925/Type USNM 49192/unicolor Csy [handwritten]”. Description .—Same character states as H. alternatus except for the following. Body length: 3.0- 3.8 mm. Head. —Proportionally narrower (WH/WP = 0.83-0.91; x = 0.86; n — 10 in males; WH/ WP = 0.80-0.84; x = 0.83; n = 10 in females). Eye longer, longitudinal diameter about twice length of antennomere I; temple shorter, about 0.2-0.3X longitudinal diameter of eye, and slightly produced posteriorly (Fig. 3). Prothorax .—Pronotum with punctuation smaller, punctures more distantly sepa¬ rated; disc flat to slightly depressed; sides more regularly rounded. Elytra. Scutellum with 1-4 setig- erous punctures. Male Genitalia. —Aedeagus as in Fig. 10. Diagnosis. —Most similar to H. alternatus but differs readily by larger eyes, shorter temples and presence of setigerous punctures on the scutellum. Distribution. —This species is known from southern Arizona, New Mexico and southwestern Texas (Fig. 17). Habitat. —Label data suggest that this species is associated with sotol plants (Dasylirion sp.). Discussion. —This species is the adelphotaxon (i.e., sister species) of H. alter¬ natus. The presence of setigerous punctures on the third and fifth elytral intervals and the presence of an oval, median depression on the last visible abdominal sternite of the male are synapomorphies for the two species. Material Examined .—ARIZONA. PIMA Co.: Santa Catalina Mts., Molino Basin, (11, CNC, FSCA) [ex sotol], Baboquivari Mts. (3, MCZ). SANTA CRUZ Co.: Santa Rita Mts. (4, MCZ, USNM) [2-in dying Dasylirion ]. NEW MEXICO. DONA ANA Co.: Organ Mts., Solidad Can. (3, MCZ). LINCOLN Co.: 10 mi. E Carrizozo, Valley of Fire (5, CDAE). TEXAS. BREWSTER Co.: Big Bend Nat. Pk„ Green Gulch (18, CNC) [on sotol, Dasylirion leiophyllum ]. Alpine (1, USNM). CULBERSON Co.: Guadalupe Mountains Nat. Pk., McKittrick Canyon (1, CNC). Hesperobaenus subtestaceus Reitter, 1876 Phyconomus subtestaceus Reitter, 1876: 299. Type locality: “MEXICO”. Phyconomus subtestaceus var. discoideus Reitter, 1876: 299. Type locality not stated. Synonymy established by Sharp (1900: 565). Hesperobaenus subtestaceus : Sharp (1900: 565). Type Material .—Reitter (1876) described H. subtestaceus and his variety dis¬ coideus from an unspecified number of specimens. I have not seen syntypes of these taxa which are probably deposited in the Museum d’Histoire Naturelle de Paris. However, I have seen a male and female identified by Sharp in BMNH which he compared to the types of H. subtestaceus (see Sharp 1900: 565-566). Description .—Body length: 2.9 mm. Coloration. —Dorsal surface red-brown, with area around scu¬ tellum and propygidium darker. Microsculpture .—Prosternal apophysis without microsculpture. Head .—Slightly wider than pronotum (WH/WP = 1.03). Eye convex, longitudinal diameter about 1.2X length of antennomere I. Temple moderately long, about 0.7 X longitudinal diameter of eye, rounded posteriorly and somewhat bulbous (Fig. 4). Antennomere IX slightly wider than long, about as wide as antennomere X. Prothorax .—Pronotum slightly transverse (LP/WP = 0.97) with sides slightly convergent in posterior half; anterior angle rounded, not produced (Fig. 4); punctures narrowly spaced laterally, not subcontiguous; disc flat, with moderately wide, median impunctate area. Hypom- eron not rugose. Elytra .—Moderately long (LE/LP = 1.92), with very small, shallow oblique impres¬ sion on anterior third near suture. Third and fifth intervals with 0-1 setigerous puncture at base; scutellum with 1 setigerous puncture. Abdomen .—First visible sternite with coxal bead not triangularly produced, without longitudinal extension. Male last visible sternite without depression. Male Genita¬ lia. —Aedeagus as in Fig. 11. 2002 REVIEW OF HESPEROBAENUS 205 Diagnosis .—Distinguished from other species by features given in the key to species. Superficially most similar to H. alternatus but readily differentiated by the absence of setigerous punctures on the third and fifth elytral intervals. Distribution .—This species is at present known only from southwestern Texas and central Mexico. Habitat .—No data available. Material Examined. —TEXAS. 1.8 mi W McDonald Observatory road on Hwy 118, JeffDavis Co., 9. VIII. 1992, W. Godwin & E. Riley (1(5, TAMU). I have also seen 2 specimens from Guanajuato, Mexico (BMNH). HESPEROBAENUS FENYESI VAN DYKE, 1945 Hesperobaenus fenyesi Van Dyke, 1945: 102. Type locality: CALIFORNIA. Pas¬ adena. Type Material. —The holotype, a male housed in CAS, is labelled: “Pasadena Cal./Mar./[small yellow round label]/A. Fenyes Collection/Holotype Hesperoba¬ enus fenyesi Van Dyke [handwrittenj/California Academy of Sciences Type No. 5436”. Description .—Body length: 2.6-3.1 mm. Coloration. —Dorsal surface red-brown, elytra in most specimens slightly paler than forebody. Microsculpture .—Prosternal apophysis with isodiametric mi¬ crosculpture. Head .—Not wider in males (WH/WP = 0.96-1.01; x = 0.98; n — 8) than in females (WH/WP = 0.93-1.00; x = 0.98; n — 10). Eye convex (slightly more than in H. abbreviatus), hemispherical, longitudinal diameter about 1.5X length of antennomere I. Temple moderately long, 0.4-0.6X longitudinal diameter of eye, slightly produced posteriorly (Fig. 5). Antennomere IX about as long as wide, slightly narrower than antennomere X. Prothorax .—Pronotum elongate (LP/WP = 1.06-1.14; x = 1.10; n = 18); anterior angle slightly produced anterolaterally in most specimens (Fig. 5); punctures very narrowly spaced laterally, subcontiguous; disc more or less flat to slightly convex, with narrow, median impunctate area. Elytra .—Moderately long (LE/LP = 1.90-2.04; x = 1.97; n = 18), with short, vague, shallow oblique impression on anterior third near suture in many specimens. Third and fifth intervals with 0-2 setigerous punctures at base; scutellum with 2-5 setigerous punc¬ tures. Abdomen .—First visible sternite with coxal bead triangularly produced, with longitudinal ex¬ tension. Male last visible sternite without depression. Male Genitalia .—Aedeagus as in Fig. 12. Diagnosis .—Distinguished from other Hesperobaenus treated by the expanded microsculpture on the prosternal apophysis. Distribution .—This species is known only from southern California (Fig. 17); it may also occur in Arizona. Habitat. —One specimen seen was collected in “decaying yucca”. Material Examined. —ARIZONA. “Ariz” (1, INHS). CALIFORNIA. “Cal.” (5, INHS, MCZ, USNM). “S.Cal.” (2, CAS, INHS). KERN Co.: 2 mi E Caliente (1, CDAE). Walker Pass (2, CAS). LOS ANGELES Co.: Claremont (1, CDAE). Santa Monica (3, INHS). Pomona (2, MCZ) [decaying yucca]. Sierra Madre (2, CAS). Pasadena (11, CAS, CUIC, MCZ, USNM). SAN BENITO Co.: 6 mi E Los Gatos Creek Road (1, CDAE) [antifreeze pit trap]. SAN BERNARDINO Co.: Arrowhead (4, MCZ). SAN DIEGO Co.: Poway (1, CAS). SANTA BARBARA Co.: Carpinteria (1, CAS). Hesperobaenus abbreviatus (Motschulsky), 1845 Rhyzophagus abbreviatus Motschulsky, 1845: 371. Type locality: «CALIFOR- NIE». Monotoma rufipenne LeConte, 1858: 64. Type locality: CALIFORNIA. San Jose. Synonymy established by Horn (1879: 262). 206 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) Figures 12-15. Aedeagus. Figure 12. Hesperobaenus fenyesi Van Dyke. Figure 13 . H. abbreviatus Motschulsky. Figure 14. H. rufipes LeConte. Figure 15. H. constricticollis Bousquet. Scale bar = 0.2 mm. 2002 REVIEW OF HESPEROBAENUS 207 Figure 16. Collection localities for Hesperobaenus alternatus Schaeffer. Rhizophagous corpulentus Reitter, 1873: 35, Type locality: «AMER.». Synonymy established by Horn (1879b: 331) Hesperobaenus abbreviatus: Horn (1879a: 262); Hatch (1962: 253). Hesperobaenus arizonicus Casey, 1916: 92. Type locality: «ARIZONA». NEW SYNONYMY. Type Material .—Motschulsky (1845) described H. abbreviatus from an un¬ specified number of specimens. I have not seen syntypes of this species which are probably housed in the Zoological Museum, Moscow University, Moscow, Russia. LeConte (1858) described H. rufipennis from an unspecified number of speci¬ mens. His collection (in MCZ) contains six specimens. The first one, a male, is labelled “Type 7044/Hesperobaenus rufipennis Lee. Monotoma Lee. S. Jose [handwritten]”. The next three specimens have no labels. The next one is labelled “15 [handwritten]” and the last one “Van.”. Probably only the first one is part of the type series. Reitter (1873) described R. corpulentus , which he credited to “Motsch i. litt.” from an unspecified number of specimens. The location of the syntype(s) is un¬ known to me. Casey’s collection in USNM contains one specimen of H. arizonicus, a male, labelled: “Ari/Casey bequest 1925/Type USNM 49190/ arizonicus Csy. [hand¬ written]”. 208 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) (#)■ Description. —Body length: 2.0-2.8 mm. Coloration. Dorsal surface red-brown with basal half of elytra conspicuously paler, yellow to red (a few specimens seen with uniformly pale coloration or with uniformly dark elytra). Microsculpture. —Prosternal apophysis with isodiametric microsculpture at apex. Head. —Wider in males (WHAVP = 1.00-1.04; x = 1.02; n = 10) than in females (WHAVP = 0.89-0.99; x = 0.95; n — 10). Eye convex, longitudinal diameter 1.5-1.6X length of antennomere I. Temple moderately long, 0.5-0.7X longitudinal diameter of eye, and slightly produced posteriorly (Fig. 6). Antennomere IX slightly longer than wide, slightly narrower than antennomere X. Protho¬ rax. —Pronotum subquadrate (LP/WP = 0.96-1.05; x = 1.01; n = 20), with maximal width at apex or before (apical 4/5); anterior angle slightly produced laterally in most specimens (Fig. 6); punctures very narrowly spaced laterally, subcontiguous; disc slightly convex, with narrow median impunctate area. Elytra. —Shorter than in H. rufipes (LE/LP = 2.04-2.17; x = 2.11; n = 20), with oblique depression on anterior third near suture. Strial punctures finer and shallower than in H. rufipes. Third and fifth intervals with 1-4 setigerous punctures at base. Scutellum without setigerous punctures. Abdomen .—First visible sternite with coxal bead triangularly produced, with longitudinal extension. Male last visible sternite without median depression. Male Genitalia .—Aedeagus as in Fig. 13. Diagnosis. —Distinguished from other Hesperobaenus studied by the bicolored elytra (in most specimens) and the laterally produced anterior angles of the pron¬ otum. Most similar to H. feneysi, especially specimens with uniformly colored elytra, but distinguished by less expanded microsculpture on the prosternal apoph¬ ysis. Synonymy. —I have compared the syntype of H. arizonicus with several spec¬ imens identified as H. abbreviatus from California, Washington and British Co¬ lumbia. I was unable to find any consistent structural differences between the specimens notwithstanding Casey’s (1916: 92) statement. 2002 REVIEW OF HESPEROBAENUS 209 Distribution .—This species ranges from southern British Columbia south to southern California, east to Idaho, Colorado and New Mexico (Fig. 18). The record from Dallas, Texas, is suspect. Habitat .—Found mainly under the bark of dead trees. Discussion .—The coloration on the dorsal surface of the body varies for this species. While most specimens (at least 90%) have a red-brown forebody with bicolored elytra, some have the dorsal surface more or less entirely red-brown (mostly specimens from the northern part of the species distribution) and others have the dorsal surface, or the elytra only, entirely pale, flavous (some specimens from the southern part of the species distribution). Material Examined. —CANADA. BRITISH COLUMBIA. Vancouver (7, CAS, INHS, MCZ) [3— under Alnus bark], Paxton Valley (1, CAS). Creston (9, CNC, CUIC, MCZ, USNM) [4—excordwood], Oliver (2, CNC). 7 mi N Oliver (1, CNC). Salmon Arm (3, CNC). Enderby (1, CNC). Robson (1, CNC). Duncan (1, CNC). UNITED STATES OF AMERICA. ARIZONA. APACHE Co.: Chuska Mts. (1, MCZ) [under bark of Quercus]-, idem, Wagon Wheel Forest Cp. (1, FSCA). COCHISE Co.: 5 mi W Portal (1, CDAE) [blacklight], Chiricahua Mts. (3, FSCA, MCZ); idem, W of Portal (1, FSCA) [bark], COCONINO Co.: Grand Canyon Nat. Pk. (1, CDAE) [ex bark-ground cover], GILA Co.: Payson (7, CAS) [under bark of rotten oak stump], Pinal Mts. (2, USNM). SANTA CRUZ Co.: Santa Rita Mts. (3, CAS); idem, Madera Cyn. (5, FSCA). Pajarito Mts., Sycamore Cyn. (3, FSCA) [under bark oak]; idem, Pena Blanca (1, FSCA) [under bark oak], PIMA Co.: Santa Catalina Mts., Redington Pass (2, FSCA) [under bark hackberry]; idem, Bear Canyon (7, FSCA) [under bark oak/pine]; idem, Peppersauce Cyn. (1, FSCA) [under bark oak], “Graham Mts., Wet Cyn.” (1, FSCA). CALIFORNIA. “Cal.” (37, AMNH, CAS, INHS, MCZ, USNM). ALAMEDA Co.: county record only (11, CAS, FMNH, MCZ). Berkeley (61, CAS, CUIC). Alameda (3, CAS, FMNH). Oakland (2, USNM). Tracy 210 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) (1, USNM) [under bark]. Redwood Canyon (1, CAS). BUTTE Co.: Chico (1, CDAE) [ex raisin trap]. Paradise (1, CAS). CALAVERAS Co.: 4.8 km S West Point (1, CAS). Mokelumne Hill (16, CAS). COLUSA Co.: 3 mi S Lodoga (3, CDAE) [ex Polyporus sulphureus]. CONTRA COSTA Co.: Vine Hill (22, CAS). Mt. Diablo (5, CAS) [ Quercus dingsolepis]. EL DORADO Co.: county record only (1, USNM) [under bark of Pinus sabiniana ]. 3 mi W Grizzly Flat (4, CDAE) [under oak bark/reared from polypore on Pinus sp.]. 2 mi. S Kyburz (1, CDAE). 5 mi E Volcanoville (4, CDAE). 2 mi NE Auburn (60, CDAE) [under bark of standing, burn killed Pinus /Quercus], 6 mi SW Ice House (1, CDAE) [conifer bark]. 3 mi S Somerset (1, TAMU). 1.2 mi W Stumpy Meadows Lake (2, CDAE) [under bark Abies concolor]. Polock Pines (1, OSUC). Pine Hill nr Rescue (1, OSUC). Lake Edson (3, CDAE) [under Pinus bark], FRESNO Co.: county record only (1, CAS). Clovis (1, USNM). 3 mi NE Auberry (1, CDAE) [under bark of Quercus]. Huntington Lake (1, CAS). 10 mi N Parkfield (2, CDAE) [under Pinus bark], HUMBOLDT Co.: county record only (2, USNM). 5 mi NW Garberville, (2, CDAE) [malaise trap]. Green Point (1, CAS). Fieldbrook (1, USNM). Hydesville (1, CAS). KERN Co.: county record only (2, USNM). LAKE Co.: county record only (2, CAS). Clearlake Oaks (3, CAS, CDAE) [ex Digger pine]. Lucerne (2, CAS). LASSEN Co.: Pine Cr. (6, CAS). Facht (1, CAS). LOS ANGELES Co.: county record only (6, CAS, USNM). Sierra Madre (1, CAS). San Gabriel Canyon (2, CDAE). Los Angeles (8, CAS, CUIC). Pasadena (21, AMNH, CAS, CUIC, FMNH, MCZ). Long Beach (2, CAS) [on lichens], San Dimas Exp. For. (3, CAS). 2 mi E Three Points (13, CDAE) [under oak bark]. Pomona (4, INHS). Jackson Lake (1, MCZ). MADERA Co.: Placer Ran. St. (1, CAS) [ Alnus rhombifolia]. Anderson Valley (1, CAS) [ Quercus californica]. MARIN Co.: county record only (10, CAS, CUIC). Mill Valley (2, CAS) [rotting tomato], Novato (2, CAS) [ultraviolet light]. Taylorville (3, CUIC). Lagunitas (1, CAS). Muir Woods (1, CAS). Bon Tempe Lake (1, CAS). Inverness (3, CAS). Stinson Beach (1, CAS). Alpine Dam (1, CAS). L. Lagunitas Rd. (2, CAS). Lagunitas (1, MCZ) [ex Lenzites betulina ]. Carson Ridge (1, MCZ) [Polystictus versicolor on Umbellularia], MARIPOSA Co.: 1.5 mi NE Darrah (1, CDAE) [under bark Pinus]. Mariposa (4, CDAE) [ex Plantanus], Anderson Valley (3, CAS) [ Libocedrus decurrens ]. Yosemite Nat. Pk. (1, CAS). MENDOCINO Co.: county record only (1, CAS). 6 mi N Willits (1, MCZ) [ex Polyporus sulfureus]. MERCED Co.: 2 mi E Cressey (7, CDAE) [under bark Populus tremuloides]. Los Banos Valley (15, CDAE) [under Populus/ SVz/Y/Cottonwood bark]. MONTEREY Co.: 4 mi SE Notleys Landing (3, CDAE) [ex polypore fungus]. Soledad (1, CDAE) [on Pleurotus ostreatus], 2.4 mi N Parkfield (10, CDAE) [fungus under bark of Populus], Bradley (1, CDAE). Jamesburg (2, AMNH). Carmel (5, CAS). Big Sur (5, CAS). NAPA Co.: Capella Cr. (6, CDAE). 3 mi NW Lake Berryessa (22, CDAE) [under bark standing fire killed Pinus sabiniana], 2 mi NNE Angwin (24, CAS) [stump of Quercus kelloggii/on rotting apple/under bark of Pinus ponderosa/under pile of weeds]. Pope Valley (1, OSUC). NEVADA Co.: Sagehen Cr. (7, FSCA, OSUC). Truckee (1, USNM). PLACER Co.: county record only (3, CAS). Folsom Lake (2, CDAE) [ex Pleurotus ostreatus], 5 mi S Auburn (1, CDAE) [berlesed from oak duff], PLUMAS Co.: county record only (1, USNM) [pine bark], 6 mi N.W. Chester (1, USNM). RIVERSIDE Co.: 2 mi NW Gilman Hot Springs (1, CDAE). Hemet (2, CDAE). Glenlvy Hot Spr. (2, CAS). Riverside (2, CAS, CUIC). Hempt Res. (1, CAS). SACRAMENTO Co.: county record only (1, USNM). Sacramento (4, CDAE). Sacramento River (1, CDAE) [barking Riparian Woodland], SAN BERNARDINO Co.: county record only (4, OSUC). Colton (3, USNM). San Antonio Creek (4, CDAE). N Ceder Springs (3, CDAE). Oak Glen (2, CDAE). Chino Hills, Carbon Canyon (1, FSCA) 1. SAN DIEGO Co.: country record only (2, CDAE, FMNH) [1—peach]. Warner Springs (2, AMNH). Poway (2, CAS). Banner (3, CAS). Burnt Rancheria PC. (1, USNM). Mts. near Clairemont (3, FMNH, MCZ). SAN FRANCIS¬ CO Co.: San Francisco (17, CAS, CUIC, USNM). SAN LUIS OBISPO Co.: 1 mi SW Cholame (4, CDAE) [under bark, oak stump], 2.8 mi S Atascadero (1, CDAE). SAN MATEO Co.: Crystal Springs L. (1, CAS). Stanford (5, CAS). SANTA BARBARA Co.: Santa Barbara (1, CAS). Santa Cruz Isl. (4, FSCA) [1— Quercus agrifolia]. SANTA CLARA Co.: county record only (1, USNM). Stevens Ck. (1, CAS). Los Gatos (10, CAS, USNM). Alum Rock Park (1, USNM). Gilroy Hot Spg (1, CAS). SANTA CRUZ Co.: 9 mi NE Big Basin (12, CAS). Ben Lomond (1, CAS). Felton (4, USNM) [under bark], SHASTA Co.: county record only (7, FMNH). Redding (1, CDAE). SISKIYOU Co.: county record only (3, CAS). Soda Springs (1, CAS). Dorris (2, AMNH) [pine stump], McCloud (1, CAS). Antelope Cr. (1, CAS). Lava Beds National Monument (Mammouth Crater) (2, AMNH) [under bark]. SONOMA Co.: county record only (5, MCZ, USNM). Duncan Mills (1, CAS). Cazadero (1, CUIC). Sobre Vista (1, CAS). Santa Rosa (3, AMNH, CDAE). SUTTER Co.: country record only (1, CDAE) [collected from peach], TEHAMA Co.: Red Bluff (6, CAS, OSUC). 12 mi SW Red Bluff (4, OSUC). TRINITY Co.: county record only (1, CAS). Waeverville (1, CDAE). Carrville (1, CAS). 12 mi SE Hyampom 2002 REVIEW OF HESPEROBAENUS 211 (1, CAS). TULARE Co.: Ash Mountain (3, CDAE). Clough Caves (1, CAS). Kaweah (5, CAS). Colony Mill (1, CAS). VENTURA Co.: Ojai (1, MCZ). YUBA Co.: Spenceville Wildlife Area (10, CDAE) [under oak bark], 15 mi E Marysville (5, CDAE) [antifreeze pit trap]. “Clayton” (2, CAS). “Santa Cruz Mts.” (20, CAS, FMNH, MCZ, USNM). “Norval Flats” (1, CAS). “Alder, San Antonio Can., San Berdo. Mts.” (2, CAS). “Warners” (2, CAS). “Malibou Beach” (6, CAS). “Northfork” (9, CUIC). COLORADO. ROUTT Co.: Steamboat Springs (1, CAS) [under aspen bark], IDAHO. ADA Co.: Barber Park (3, FSCA). CANYON Co.: Parma (1, USNM). KOOTENAI Co.: Coeur d’Alene (10, AMNH, MCZ, USNM). NEBRASKA. DAWES Co.: Pine Ridge (1, USNM). NEVADA. State record only (3, AMNH, USNM). LYON Co.: Dayton (8, AMNH, MCZ) [bark dead Populus ]. STOREY Co.: 6 mi Canyon, Virginia City (2, TAMU). WASHOE Co.: Reno (2, AMNH). NEW MEXICO. TAOS Co.: San Juan Valley (3, MCZ). OREGON. “Or” (7, CMNH, FMNH, INHS, MCZ). BAKER Co.: Dooley Mt. (4, AMNH). BENTON Co.: Monroe (2, USNM) [in flight], 2 mi W Corvallis (13, AMNH) [under dead oak bark], GRANT Co.: John Day (2, AMNH). HOOD RIVER Co.: Hood River (4, USNM). JACKSON Co.: Medford (1, CAS). Shady Cove (1, AMNH) [under oak bark]. JOSEPHINE Co.: Illinois River (1, AMNH). KLAMATH Co.: Klamath Falls (12, AMNH) [under bark juniper stump], 15 mi NW Bly (3, AMNH) [on yellon pine]. Upper Klamath Lake (17, AMNH) [dead poplar bark], Merrill (1, AMNH) [tree litter]. Hildebrand (6, AMNH) [pine bark], 7 mi W Keno (1, AMNH) [under bark], YAMHILL Co.: county record only (2, CAS, USNM). TEXAS. Dallas Co.: DALLAS (1, MCZ). UTAH. SUMMIT Co.: Park City (1, USNM). UTAH Co.: Mt. Timpanogos (1, CAS). East Utah Lake (7, TAMU). WASHINGTON. “Wash” (2, OSUC). GRAYS HARBOR Co.: Hoquiam (1, USNM). OKANOGAN Co.: Omak (1, MCZ). SPOKANE Co.: 9 mi N Spokane (1, FMNH). THURS¬ TON Co.: Olympia (6, MCZ). Tenino (2, USNM). WHHMAN Co.: Wawawai Cyn (2, LSUC). Pullman (4, USNM). HESPEROBAENUS RUFIPES LeCONTE, 1863 Hesperobaenus rufipes LeConte, 1863: 65. Type locality: «southern states». Horn (1879: 263); Blatchley (1910: 669); Downie & Arnett (1996: 988). Type Material. —LeConte (1863) described H. rufipes from an unspecified num¬ ber of specimens. His collection (in MCZ) contains two specimens. The first one, a male, is labelled “[orange disc]/Type 7045/H. rufipes Lee, [handwritten]”. The second, also a male, has an orange disc only. Description. —Body length: 2.0-2.8 mm. Coloration. —Dorsal surface uniformly red-brown. Micro- sculpture. -Prosternal apophysis without microsculpture. Head. —Wider in males (WH/WP = 0.98- 1.05; x = 1.02; n — 10) than in females (WH/WP = 0.91-0.98; x = 0.94; n — 10). Eye convex, longitudinal diameter 1.4-1.5 X length of antennomere I. Temple short, about 0.4 X longitudinal diameter of eye, and slightly produced posteriorly (Fig. 17). Antennomere IX slightly wider than long, slightly narrower than antennomere X. Prothorax. —Pronotum transverse (LP/WP = 0.90-0.96; x = 0.93; n = 20), with maximal width before apex, at apical 4/5; anterior angle rounded, not produced (Fig. 17); punctures narrowly spaced laterally but not subcontiguous; disc slightly convex, with rather wide median impunctate area. Hypomeron not rugose. Elytra. —Proportionally long (LE/LP = 2.17— 2.33; x = 2.24; n — 20), with oblique depression on anterior third near suture. Strial punctures rather coarse and deep. Third and fifth intervals with 0-2 setigerous punctures at base. Scutellum without setigerous punctures. Abdomen. —First visible sternite with coxal bead triangularly produced, without or with short longitudinal extension. Male last visible sternite without median depression. Male Gen¬ italia. —Aedeagus as in Fig. 14. Diagnosis. —Distinguished from other Hesberobaenus treated by features given in the key to species. Distribution. —This species occurs from Maryland to Kansas, south to Florida and Texas (Fig. 19). Habitat. —Found under the bark of oak and maple trees. Blatchley (1928: 66) reported that this species occurs frequently, around Dunedin, Florida, “beneath the close fitting bark of dead water-oak”. 212 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) Figure 19. Collection localities for Hesperobcienus rufipes LeConte. Material Examined. —ALABAMA. “Ala” (1, MCZ). DEKALB Co.: De Soto State Park (2, CAS). JEFFERSON Co.: Birmingham (1, FMNH). LAUDERDALE Co.: county record only (1, CNC). LEE Co.: county record only (2, LSUC). MOBILE Co.: Mobile (1, MCZ). TUSCALOOSA Co.: county record only (3, CNC). “Spring Hill” (5, OSUC, USNM). ARKANSAS. HEMPSTEAD Co.: Hope (2, MCZ). JOHNSON Co.: Ozone (3, TAMU). LOGAN Co.: Mt. Magazine Lookout (1, CNC) [sifting deciduous leaf litter], WASHINGTON Co.: county record only (6, INHS). Fayetteville (10, INHS). DISTRICT OF COLUMBIA. “DC” (2, MCZ). Washington (1, USNM). FLORIDA. “Fla” (1, INHS). ALACHUA Co.: county record only (5, CUIC). San Felasco Hammock (8, FSCA). Gainesville (2, FSCA). “At Levy Co. line SR 24” (6, FSCA) [under bark of Quercus sp.]. COLLIER Co.: Choko- loskee (4, CUIC, MCZ). COLUMBIA Co.: county record only (1, FSCA) [under bark of dead Quercus laevis ]. DUVAL Co.: county record only (11, FSCA) [under bark of Quercus ]. MARION Co.: Rainbow Springs (34, FSCA). Ocala (1, FSCA). OKALOOSA Co.: Fort Walton Beach (6, CNC). Niceville (1, CNC). Nr. Deerland (14, FSCA) [turkey oak bark]. “0.3 mi N jet US 90 & CR 345” (40, FSCA) [under bark of Quercus laevis ]. PINELLAS Co.: Dunedin (5, AMNH, MCZ, USNM). GEORGIA. “Geo” (1, MCZ). CHATHAM Co.: Savannah (1, MCZ) [under bark and in fungi], HABERSHAM Co.: Cornelia (1, USNM) [Quercus], JEFFERSON Co.: Louisville (1, USNM) [under bark]. LAMAR Co.: Bamesville (2, MCZ). ILLINOIS. LA SALLE Co.: Starved Rock (1, FMNH). UNION Co.: Anna (1, INHS). VERMILION Co.: Muncie (5, USNM). KANSAS. LEAVENWORTH Co.: Leavenworth (2, CAS, CNC). KENTUCKY. HENDERSON Co.: Henderson (1, CAS). LOUISIANA. CADDO Parish: parish record only (1, LSUC). MADISON Parish: Tallulah (1, USNM). “Vowell’s Mill” (18, MCZ). “Bay Sara” (4, USNM). MARYLAND. BALTIMORE Co.: Catonsville (5, USNM) [under bark of hickory log], MONTGOMERY Co.: 3 mi S Colesville (1, MCZ) [under bark maple]. PRINCE GEORG¬ ES Co.: county record only (1, USNM). Hyattsville (1, USNM). MISSISSIPPI. FORREST Co.: Hat¬ tiesburg (3, AMNH). GEORGE Co.: Lucedale (77, CUIC). GREENE Co.: Avera (5, CUIC). MIS¬ SOURI. “Mo” (2, CAS, USNM). St. Louis City: St. Louis (2, USNM). NORTH CAROLINA. “N.C.” (11, MCZ, OSUC, USNM). CLEVELAND Co.: Kings Mountain (1, TAMU). FRANKLIN Co.: county record only (1, FSCA). MOORE Co.: Southern Pines (3, USNM). POLK Co.: Tryon (6, USNM) 2002 REVIEW OF HESPEROBAENUS 213 [Hicoria], WAKE Co.: Raleigh (5, FSCA). OHIO. DELAWARE Co.: county record only (1, FSCA). HOCKING Co.: county record only (4, OSUC). OKLAHOMA. CADDO Co.: county record only (1, CAS). CHEROKEE Co.: 5 mi NE Qualls (1, CAS). LATIMER Co.: county record only (29, FSCA, NHDE, USNM) [2—under oak bark], 5 mi W Red Oak (3, FSCA). PAYNE Co.: Stillwater (1, MCZ). PITTSBURG Co.: McAlester Army Ammunition Plant (1, OSUC). PENNSYLVANIA. ALLEGHENY Co.: Pittsburgh (2, CMNH). SOUTH CAROLINA. “SC” (2, MCZ). PICKENS Co.: Clemson (19, CAS, TAMU) [under oak bark/under bark]. Rocky Bottom (3, USNM). TENNESSEE. CUMBERLAND Co.: 8 km NW Rockwood (3, USNM). TEXAS. “Tex” (6, MCZ, USNM). CHAMBERS Co.: Anahuac (1, USNM). DALLAS Co.: Dallas (1, MCZ). HARRIS Co.: Katy (8, FSCA). Houston (1, USNM). KERR Co.: Kerrville (20, USNM). JEFFERSON Co.: Beaumont (2, USNM). MONTGOMERY Co.: Conroe (8, FSCA). TRAVIS Co.: Austin (4, CAS). VICTORIA Co.: Victoria (1, USNM). “Cypress Mills” (1, MCZ). VIRGINIA. CAROLINE Co.: Ladysmith (11, AMNH). PAGE Co.: Luray (1, FMNH). WEST VIRGINIA. PENDLETON Co.: Smoke Hole (1, CUIC) [light trap], HESPEROBAENUS CONSTRICTICOLLIS BOUSQUET, NEW SPECIES Type Material. —Holotype (d) labelled: “TEX: Cameron Co. Sabal Palm Grove Sanct., IV-8-1994 Coll. E.G. Riley/from Sabal Palm Grove [handwritten]/ E.G. Riley Collection/Holotype Hesperobaenus constricticollis Bousquet”, depos¬ ited in Texas A&M University, College Station, Texas. Description. —Body length: 2.6 mm. Coloration. —Head, pronotum and scutellum light red-brown, elytra paler, yellow. Microsculpture. —Prosternal apophysis without microsculpture. Head. —Wider than pronotum (WH/WP = 1.09). Eye rather large, longitudinal diameter about 1.5 X length of antennomere I. Temple moderately long, about 0.5 X longitudinal diameter of eye, not produced posteriorly (Fig. 8). Antennomere IX as wide as long, narrower than antennomere X. Prothorax .— Pronotum transverse (LP/WP = 0.88) with sides markedly narrowed in posterior half; anterior angle rounded, not produced (Fig. 8); punctures rather distantly spaced laterally, not subcontiguous; disc flat, with moderately wide, median impunctate area, widening in posterior half. Hypomeron rugose. Elytra. —Moderately long (LE/LP = 1.84), without oblique impression on anterior third. Third and fifth intervals with 0-2 setigerous puncture at base; scutellum without setigerous puncture. Abdo¬ men. —First visible sternite with coxal bead not triangularly produced but thickened, without longi¬ tudinal extension. Male first abdominal sternite with small tuft of short (but longer than adjacent ones) setae at middle. Male last visible sternite without depression. Male Genitalia. —Aedeagus as in Fig. 15. Diagnosis .—Distinguished from other Hesperobaenus treated by the markedly narrowed pronotum posteriorly. Etymology .—The specific name derives from the Latin constrictus, a, um (con¬ stricted) and collum, —i (used for pronotum); it refers to the markedly narrowed sides of pronotum toward base. Distribution .—This species is known only from the type locality. The Sabal Palm Grove Sanctuary is located in a bend of the Rio Grande along the United States-Mexico border, near Brownsville, in southeastern Texas. Habitat .—The species may be associated with Sabal Palms. Material Examined. —See Type Material. Acknowledgment I thank Henri Goulet and Serge Laplante for their constructive comments on the manuscript and Go Sato for the habitus and inking of the drawings. Literature Cited Arnett, R. H. Jr. 1962. The beetles of the United States (a manual for identification). Part V. Suborder Polyphaga (Cont.) Series Cucujiformia (Cont.) Tenebrionoidea Cucujoidea. The Catholic Uni¬ versity of America Press, Washington, D.C. Pp. 645-850. 214 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) Blatchley, W. S. 1910. An illustrated descriptive catalogue of the Coleoptera or beetles (exclusive of the Rhynchophora) known to occur in Indiana—with bibliography and descriptions of new species. The Nature Publishing Co., Indianapolis. 1386 pp. Blatchley, W. S. 1928. Notes on some Florida Coleoptera with descriptions of new species. Can. Entomol., 60: 60-73. Casey, T. L. 1916. Memoirs on the Coleoptera. VII. The New Era Printing Company, Lancaster (PA). 390 pp. Downie, N. M. & R. H. Arnett, Jr. 1996. The beetles of northeastern North America. Volume II. Polyphaga: series Bostrichiformia through Curculionoidea. The Sandhill Crane Press, Gaines¬ ville (FL). Pp. 891-1721. Fairmaire, L. 1850. Essai sur les Coleopteres de la Polynesie (suite). Revue et Magasin de Zoologie pure et appliquee (2 e serie), 2: 50-64. Hatch, M. L. 1962. The beetles of the Pacific Northwest. Part III. Pselaphidae and Diversicornia I. University of Washington Publications in Biology. Volume 16. Seattle. 503 pp. Hetschko, A. 1930. Fam. Cucujidae (Pars 109). In Junk, W. & S. Schenkling (eds.). Coleopterorum catalogus. Junk, Berlin. 122 pp. Horn, G. H. 1879a. Synopsis of the Monotomidae of the United States. Trans. Am. Entomol. Soc., 7: 257-267. Horn, G. H. 1879b. Revision of the Nitidulidae of the United States. Trans. Am. Entomol. Soc., 7: 267-336. Lawrence, J. F. 1991. Rhizophagidae (Cucujoidea) (including Monotomidae). pp. 460-462. In Stehr, F. W. (ed.). Immature Insects. Volume 2. Kendall/Hunt Publishing Co., Dubuque, Iowa. LeConte, J. L. 1858. Description of new species of Coleoptera, chiefly collected by the United States and Mexican Boundary Commission, under Major W. H. Emory, U.S.A. Proc. Acad. Nat. Sci. Phila., 10: 59-89. LeConte, J. L. 1861. Classification of the Coleoptera of North America. Prepared for the Smithsonian Institution. Part I. Smithsonian Miscellaneous Collections [No. 136]. xxiv + 214 pp. LeConte, J. L. 1863. New species of North American Coleoptera. Prepared for the Smithsonian In¬ stitution. Part I. Smithsonian Miscellaneous Collections No. 167. 86 pp. Motschulsky, V. de. 1845. Observations sur le musee entomologique de l’Universite Imperiale de Moscou. Bull. Soc. Imp. Nat. Mosc., 18: 332-388. Reitter, E. 1873. Die Rhizophaginen, monographisch bearbeitet. Verhandlungen des Naturforschenden Vereines in Briinn, 11 [1872]: 27-48. Reitter, E. 1876. Revision der Monotomidae (sensu LeConte). Dtsch. Entomol. Z., 20: 295-301. Schaeffer, C. F. A. 1910. New Clavicorn Coleoptera. J. N. Y. Entomol. Soc., 18: 210-216. Sen Gupta, T. 1988. Review of the genera of the family Rhizophagidae (Clavicornea: Coleoptera) of the world. Memoirs of the Zoological Survey of India, 17: 1-58. Sharp, D. 1900. Fam. Cucujidae. Fam. Monotomidae. pp. 499-579. In Godman, F. D. & O. Salvin (eds.). Biologia Centrali-Americana. Insecta. Coleoptera. Volume II. Part 1. Taylor & Francis, London. Van Dyke, E. C. 1945. New species of North American Coleoptera. Pan-Pac. Entomol., 21: 101-109. Received 29 June 2001, Accepted 20 May 2002. PAN-PACIFIC ENTOMOLOGIST 78(3): 215-218, (2002) Scientific Note PHYTOSEIID MITE FAUNA ON GORSE, ULEX EUROPAEUS L., IN WESTERN OREGON, USA WITH NEW RECORDS FOR PHYTOSEIULUS PERSIMILIS ATHIAS-HENRIOT AND AMPLYSEIUS GRAMINIS (CHANT) (ACARI: PHYTOSEIIDAE) Gorse, Ulex europaeus L. (Fabaceae), is a spiny evergreen shrub native to western Europe. The plant was intentionally introduced into coastal regions of southern Oregon (USA) in the late 1800s. Since its introduction, U. europaeus has escaped cultivation and aggressively invaded natural and disturbed habitats in western North America, including British Columbia, Washington, Oregon, northern California, and Hawaii. In an effort to suppress this weed, the European native spider mite Tetranychus lintearius Dufour (Acari: Tetranychidae) was in¬ troduced into gorse-dominated habitats of western Oregon in 1994 (Rees, N. E., R C. Quimby, Jr., G. L. Piper, E. M. Coombs, C. E. Turner, N. R. Spencer & L. V. Knutson. 1996. Biological Control of Weeds in the West. Western Society of Weed Science, Bozeman, Montana). Establishment, persistence, and efficacy of weed suppression by an introduced biological control agent can be affected by mortality from predators, parasites or pathogens acquired in the adventive range of the agent (Goeden, R. D. & S. M. Louda. 1976. Annu. Rev. Entomol., 21: 325-342). Predatory mites (Phytoseiidae), for instance, suppress spider mites in managed and unmanaged ecosystems world¬ wide (Helle, W., & M. W. Sabelis. 1985. Spider Mites: Their Biology, Natural Enemies and Control. Elsevier, Amsterdam). Unfortunately, the phytoseiid fauna in most natural habitats is poorly known. Therefore, we sought to determine which predatory mites are associated with the invasive weed U. europaeus and its bio¬ logical control agent T. lintearius in western Oregon. Surveys for phytoseiids were performed at six (four coastal and two inland) sites in western Oregon: near Astoria, Baker Beach, Bandon, Clackamas, Elk River, and Sutherlin (Table 1). Monthly surveys were performed at Baker Beach, Bandon, and Sutherlin from March 1998 through March 1999 and single surveys were conducted at the remaining sites. Surveys consisted of sampling U. euro¬ paeus branches every 10 m along a randomly selected 100 m transect. A total of 20 samples were collected from each transect by randomly selecting two inde¬ pendent terminal U. europaeus branches at each sampling point and excising ca. 25 cm of foliage from each branch. Each sample was placed into a polyethylene bag, transported to the laboratory, and branches were individually washed to ex¬ tract arthropods within 48 h. The extraction method entailed placing individual U. europaeus branches in separate one-liter jars and adding 300 ml of 70% ethanol (Pratt, P. D. & B. A. Croft. 2000. Environ. Entomol., 29: 1034-1040). Lids were placed on the jars and shaken manually for 30 sec, left to rest for 1 min, and then shaken again for 30 sec. Plant material was removed with forceps and slowly rinsed with 70% ethanol over jars. The ethanol and associated contents were poured into a Whatman No. 4 filter paper funnel, gravity filtrated, and examined 216 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) Table 1. Phytoseiid mites collected from Ulex europaeus. Species Life style type 1 Research site GPS coordinates 2 Amblesius andersoni (Chant) III Baker Beach 44.0915 N 124.1158 W Neoseiulus fallacis (Garman) II Baker Beach 44.0915 N 124.1158 W Typhlodromus pyri Scheuten III Sutherlin 43.3970 N 123.2974 W Amblyseius graminis (Chant) III 3 Astoria 46.2783 N 123.9970W Typhlodromus arboreus (Chant) III Baker Beach, Bandon, Elk River (see above and below) Phytoseiulus persimilis Athias-Henriot I Bandon 43.0543 N 124.4083 W Galendromus occidentalis (Nesbitt) II Clackamas 45.2391 N 122.4268 W 1 Type I = specialized predators of Tetranychus species; Type II = selective predators of tetranychid mites, particularly with those that produce copious webbing; Type III = generalist predators. 2 Global positioning system in decimal degrees (Elk River: 24.7648 N 124.4626 W). 3 Probable classification, life history studies needed to quantify life style type. within 5 min under a binocular microscope at 40X magnification. All phytoseiid mites that were washed from branches were mounted on glass slides in Hoyer’s media and identified according to morphological characters (Schuster, R. O., and A. E. Pritchard. 1963. Hilgardia, 34: 191-194). In preliminary samples, the predatory mite Phytoseiulus persimilis Athias-Hen- riot was collected at the Bandon survey site. This phytoseiid is a specialist pred¬ ator that feeds primarily on spider mites that belong to the genus Tetranychus (McMurtry, J. A. & B. A. Croft. 1997. Annu. Rev. Entomol., 42: 291-321) and it is the most common biological control agent released for suppression of pest mites in agricultural and horticultural systems throughout the world (Helle & Sabelis 1985). Because of the potential for P. persimilis to suppress the beneficial T. lintearius, we sought to assess the geographic range of this predatory mite in southern Oregon. This was done on 15 Sept. 1998 by sampling foliage of U. europaeus (as described earlier) every 1.6 km along a N-S transect radiating from the epicenter at each study site. To increase the probability of collecting P. per¬ similis and accurately measure its geographic distribution, only U. europaeus fo¬ liage containing colonies of T. lintearius was selected at each survey point. Tran¬ sects were extended north and south until three consecutive samples failed to Table 2. Dominant predatory mite species collected at each survey site. Site name Species Peak density 1 Month 2 Astoria Amblyseius graminis 0.45 (0.13) April 3 Baker Beach Typhlodromus arboreus 2.28 (0.52) May Bandon Phytoseiulus persimilis 2.75 (2.02) October Clackamas Typhlodromus arboreus 0.51 (0.28) September 3 Elk River Typhlodromus arboreus 1.50 (1.19) July 3 Sutherlin Typhlodromus pyri 4.65 (0.90) July 1 Peak densities of phytoseiid mites per sample, mean (SE). 2 Month when peak density was recorded. 3 Because only a single sample was collected from these sites, Peak density and Month many not accurately describe the population densities. 2002 SCIENTIFIC NOTE 217 TIPpTIPpTIPpTIPpTIPpTIPpTIPpTIPpTIPpTIPpTIPpTIPpTIPpTIPpTIPp 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 Census Sites Figure 1. Predator-prey ratios and age distribution for Tetranychus lint ear ius (Tl) and Phytoseiulus persimilis (Pp) along a north (15) to south (1) transect (7 = initial spider mite release site) during a 1999 survey of the invasive weed Ulex europaeus in Bandon, Oregon (U.S.A.). produce the predatory mite or T. lintearius populations were no longer present. Extraction and identification of arthropods was performed as described above. Among the Phytoseiidae collected during 1998 and 1999, 57% of the species were generalists (Type III) species, which feed on various mites, insects, and pollens (Table 1, McMurtry & Croft 1997). These generalist predators were also the dominant (most abundant) species at five of the six survey sites (Table 2). At the Bandon site, however, the specialist Type I predator of Tetranychus species, P. persimilis, was the most common natural enemy collected from T. lintearius colonies. To our knowledge, this is the first detailed survey of phytoseiid mites along the coastal regions of Oregon. Findings from this survey included the first col¬ lection of Amblyseius graminis (Chant) in North America. This predatory mite is endemic to the Old World and explanations for its adventive geographic distri¬ bution in the Pacific Northwest remain unclear. Although collected from T. lin¬ tearius colonies, attempts to establish a laboratory culture of A. graminis when held with the spider mite were unsuccessful, suggesting that the predator may not readily feed on this spider mite and thus is unlikely to interfere with biological control. The distribution of P. persimilis in southern Oregon appears to be limited to a 20.8 km transect centered in the city of Bandon. Census site 7 in Fig. 1 represents the Bandon study site, with the predatory mite distributed 11.2 km north and 8 km south. Phytoseiulus persimilis densities along the sampled transect were sim¬ ilar to those of T. lintearius ( t = 0.46, df = 24, P = 0.65), when the extreme sample locations 1 and 15 are excluded from the analysis. Predator-prey ratios and age distributions at each census site are also presented in Fig. 1. These data reflect a distribution of life stages of a predator population that was rapidly in¬ creasing over most of the range of sample sites. It should be noted that, at sites 1 and 15, there were prey mites but no predator mites. Phytoseiulus persimilis had not yet expanded into these outer limits and thus the distribution of prey life stages reflected the reproduction of the spider mite without major predation influ¬ ences. Also note that, within the area where P. persimilis is distributed, there were three sites (2, 5, and 9; Fig. 1) where prey mite populations were decreased by 218 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(3) predation to the extent that there were only adult mites, and all egg and immature life stages (preferred life stages for P. persimilis) had been eliminated. This is the first report of P. persimilis occurring in natural systems of Oregon, and possibly the first record north of the Sacramento Valley, California. The geographic distribution of P. persimilis in Oregon presently appears to be limited to the vicinity of the city of Bandon. Possible explanations for this recent occur¬ rence include introductions by horticulturalists for spider mite control in glass¬ houses or ornamental plants. An alternative, but less likely, explanation is that P. persimilis populations naturalized in the region but remained undetected until sufficient populations of prey species had increased by the recent introduction of T. lintearius. Although often considered a semitropical species, the survival of P. persimilis during recent atypically cool winters (< — 5° C) suggests that it has extended its geographic range to include coastal regions of western Oregon. In general, these findings suggest that T. lintearius has developed new asso¬ ciations with generalist and specialist predatory mites. However, association among these mites is not sufficient evidence to conclude that phytoseiids are negatively impacting T. lintearius, and thus biological control of gorse. Additional data describing prey suitability and field-based exclusion tests are needed to quan¬ tify the impacts of phytoseiids on the biological control agent T. lintearius. Records .—OREGON. CLATSOP Co. Astoria, 4 Apr 1998, E. M. Coombs, Amblyseius graminis, branches of Ulex europaeus. COOS Co. Bandon, 17 Mar 1998, P. D. Pratt, Phytoseiulus persimilis, branches of Ulex europaeus. Acknowledgment .—We thank J. A. McMurtry (of Oregon State University) for identification of the phytoseiid mites and comments on the manuscript. This re¬ search was funded, in part, by a grant from Oregon Department of Agriculture. Paul D. Pratt, USDA/ARS, Invasive Plant Research Laboratory, 3205 College Ave., Fort Lauderdale, Florida 33314, Eric M. Coombs, Oregon Department of Agriculture, 635 Capitol St. NE, Salem, Oregon 97301-2532 and Brian A. Croft, Department of Entomology, Oregon State University, Corvallis, Oregon, 97331. Received 29 June 2001; Accepted 8 May 2002. 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W. 1984. An introduction to multivariate statistical analysis (2nd ed). John Wiley & Sons, New York. Blackman, R. L., P. A. Brown & V. F. Eastop. 1987. Problems in pest aphid taxonomy: can chromosomes plus moiphometrics provide some answers? pp. 233-238. In Holman, J., J. Pelikan, A. G. F. Dixon & L. Weismann (eds.). Population structure, genetics and taxonomy of aphids and Thysanoptera. Proc. international symposium held at Smolenice Czechoslovakia, Sept. 9-14, 1985. SPB Academic Publishing, The Hague, The Netherlands. Ferrari, J. A. & K. S. Rai. 1989. Phenotypic conelates of genome size variation in Aedes albopictus. Evolution, 42: 895-899. Sorensen, J. T. (in press). Three new species of Essigellci (Homoptera: Aphididae). Pan-Pacif. Entomol. 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L.—Obituary and bibliography of Kenneth S. Hagen (1919-1997), dedicated entomologist and teacher _ 151 WALL, M. A. & R. S. BOYD—Nickel accumulation in serpentine arthropods from the Red Hills, California _ 168 M. S. HODDLE—Oviposition preferences of Scirtothrips perseae Nakahara (Thysanoptera: Thripidae) in southern California avocado orchards _ 177 X. CHEN, J. B. WHITFIELD, & J. HE—The discovery of the genus Gnamptodon Haliday (Hymenoptera: Braconidae) in China, with description of one new species _ 184 S. R. SHAW—Two new species of Betelgeuse from Mexico (Hymenoptera: Braconidae: Euphorinae) _ 188 Y. BOUSQUET—Review of the genus Hesperobaenus LeConte (Coleoptera: Monotomidae) of America, north of Mexico _ 197 SCIENTIFIC NOTE: P. D. PRATT, E. M. COOMBS, & B. A. CROFT—Phytoseiid mite fauna on gorse, Ulex earo- paeus L., in western Oregon, USA with new records for Phytoseiulus persimilis Athias- Henriot and Amblyseius graminis (Chant) (Acari: Phytoseiidae)- 115 The PAN-PACIFIC ENTOMOLOGIST Volume 78 October 2002 Number 4 Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES (ISSN 0031-0603) The Pan-Pacific Entomologist EDITORIAL BOARD R. M. Bohart J. T. Doyen J. E. Hafemik, Jr. Warren E. Savary Published quarterly in January, April, July, and October with Society Proceedings usually appearing in the following October issue. All communications regarding non-receipt of numbers should be addressed to: Vincent F. Lee, Managing Secretary; and financial communications should be addressed to: Robert L. Zuparko, Treasurer; at: Pacific Coast Entomological Society, Dept, of Entomology, California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599. 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Members of the Society receive The Pan-Pacific Entomologist. Single copies of recent numbers or entire volumes are available; see 72(4): 247 for current prices. Make checks payable to the Pacific Coast Entomological Society. R. Somerby, Editor R. E. Somerby, Book Review Editor Robert Zuparko, Treasurer Pacific Coast Entomological Society OFFICERS FOR 2002 Katherine N. Schick, President Vincent F. Lee, Managing Secretary Rolf L. Aalbu, President-elect Richard M. Brown, Recording Secretary Robert L. Zuparko, Treasurer THE PAN-PACIFIC ENTOMOLOGIST (ISSN 0031-0603) is published quarterly for $40.00 per year by the Pacific Coast Entomological Society, % California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599. Periodicals postage is paid at San Francisco, CA, and additional mailing offices. POSTMASTER: Send address changes to the Pacific Coast Entomological Society, % California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599. This issue mailed 4 March 2003 The Pan-Pacific Entomologist (ISSN 0031-0603) PRINTED BYTHEALLEN PRESS, INC., LAWRENCE, KANSAS 66044, U.S.A. © The paper used in this publication meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PAN-PACIFIC ENTOMOLOGIST 78(4): 219-229, (2002) ALLOZYME PHYLOGENY OF NORTH AMERICAN COPPERS (LYCAENINAE: LYCAENIDAE) Gordon F. Pratt and David M. Wright Department of Entomology, University of California, Riverside, California 92521 Department of Pathology, 100 Medical Campus Drive, Lansdale, Pennsylvania 19446 Abstract .—Phytogenies were created with allozyme data of 15 species and two subspecies of North American coppers. Most of the species align with the currently recognized subgenera of the subfamily Lycaeninae. These subgenera exhibit a significant level of genetic differentiation that is peihaps equivalent to genera. The subgenus Epidemia unexpectedly includes Lycaena hyllus (Cramer), which is currently assigned to a separate monotypic subgenus. The Nei Distance tree separates the North American taxa into two distinct biological groups. One group diapauses as partially grown larvae and has its closest relatives in the Palaearctic based on morphological data. The other group is endemic and diapauses in the egg stage (first instars within eggs). Divergence within the Distance Wagner tree parallels unique host shifts that have occurred several times in the North American coppers. Most shifts have originated from a Rumex feeding species. The host shift to Eriogonum has produced at least two and possibly more species. A few species have adapted to Vaccinium; these shifts appear to have occurred independently. Key Words .—Insecta, Lycaena, Epidemia, Lycaeninae, host shifts, diapause. The taxonomy of the North American coppers (subfamily Lycaeninae) has a complex history. In the past two centuries, the species have been classified under several lycaenid genera such as Polyommatus Latreille, Lycaena Fabricius, Chry- sophanus Hubner, and Heodes Dalman. Miller & Brown (1979) in an attempt to balance taxonomy with a proposed copper phylogeny resurrected genera originally named by Scudder in 1876 ( Tharsalea, Chalceria, Gaeides, and Epidemia) and also erected monotypic genera ( Hyllolycaena and Hermelycaena) for the species Lycaena hyllus and L. hermes (W. H. Edwards) respectively. Their subsequent systematic catalogue followed the same trend using a total of seven genera (Miller & Brown 1981). However, a recent checklist of California butterflies conserva¬ tively placed all coppers in the genus Lycaena with further division provided by the subgenera Lycaena, Epidemia, Chalceria, Hermelycaena, and Tharsalea (Em- mel et al. 1998). These authors did not recognize the subgenus Gaeides, because its type species is closely related to the type species of Chalceria. Chalceria has page priority over Gaeides in Scudder (1876). Miller & Brown (1979) pointed out the primitive features of Lycaena phlaeas and L. cuprea and speculated on their probable origin in the Palaearctic. The source of North American L. phlaeas populations is uncertain, but L. phlaeas has many subspecies throughout the Old World, several of which could have served as founders. Regarding L. cuprea, Klots (1936) and Sibatani (1974) independently noted a kinship between this North American species and the Eurasian species L. alciphron (Rottemburg). Strong similarities exist between these two species in both genitalia and facies. While it is tempting to link L. cuprea with L. alciphron in a unique Palaearctic genus, the proper generic assignment of alciphron is pres¬ ently uncertain. Modern workers have variably placed it in Heodes, Lycaena, and 220 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) Thersamonolycaena (Higgins 1975, Higgins & Riley 1983, Korshunov & Gor¬ bunov 1995, Tolman 1997, Tusov 2000, Gorbunov 2001). Also of great interest are recently discovered, high altitude Asian species, which display even stronger similarities to L. cuprea (Wyatt 1961, Churkin 1999). New studies are needed to resolve the relationship of L. cuprea with Palaearctic genera. Coppers as a whole have a worldwide distribution indicating these butterflies may belong to a very old lineage (Lewis 1973, Miller & Brown 1979). It has been suggested that their earliest divergence began before the continental sepa¬ ration of Pangaea in Late Cretaceous (Miller & Brown 1979). Their distribution presently extends from Eurasia to South Africa, from Asia to New Zealand (in¬ cluding Malayan Peninsula and Papuan region), and from North America to Cen¬ tral America where a single species, Lycaena ( lophanus ) pyrrhias (Godman & Salvin), resides in high elevation cloud forests. They are absent throughout South America and Australia (Clark & Dickson 1971, Miller & Brown 1979, Gibbs 1980, Higgins & Riley 1983, Korshunov & Gorbunov 1995, Gorbunov 2001). Copper larvae throughout the world, with the exception of North America, feed exclusively on plants in the family Polygonaceae. They chiefly utilize closely related members of the genera Rumex, Polygonum, and Muehlenbeckia. In North America several coppers have shifted onto unique non-polygonaceous hosts, in¬ cluding Rhamnus (Rhamnaceae), Eriogonum (Polygonaceae), Ribes (Grossulari- aceae), Vaccinium (Ericaceae), and Potentilla (Rosaceae) species. Determining what factors caused these butterflies to make host shifts is essential to understand¬ ing their evolution. In this study we produced phylogenies using allozyme analyses of 15 species of North American coppers. We surveyed the various trees for species clusters and compared these clusters for taxonomic congruence with the currently known taxa of the Lycaeninae. We also compared life history features and speculated how some North American coppers may have evolved through diapause changes and host shifts. Materials and Methods Enzyme Analysis .—Fresh or frozen butterflies were homogenized, electropho- resed on 10% starch gels, stained for enzymes, and scored following the procedure of Pratt (1994). The butterfly sample sizes and sites of collection are shown in Table 1. They were stored at —70° C. After removal of the wings the remainders were homogenized in 50 pi of buffer (0.005 M Tris-HCl pH 7.5) per butterfly. The homogenates were stored in microtiter plates at —70° C and electrophoresed on gels with a citrate-aminopropyl-morpholine continuous system (pH 8.5) (Clay¬ ton & Tretiak 1972). The enzymes aconitase (ACO-1 & ACO-2), adenylate kinase (AK-1 & AK-2), aspartate amino transferase (AAT-1 & AAT-2), alpha glycero¬ phosphate dehydrogenase (aGPD), glucose phosphate isomerase (GPI), glucose- 6-phosphate dehydrogenase (G6PD), hexokinase (HEX-1 & HEX-2), isocitrate dehydrogenase (IDH-1 & IDH-2), malic dehydrogenase (MDH-1 & MDH-2), malic enzyme (ME-1), peptidase [leucyl-glycyl-glycine (PEP-1 & PEP-2) as a substrate], phosphoglucomutase (PGM), and superoxide dismutase (SOD-1, SOD- 2, SOD-3) were stained with conventional histochemical stains (Shaw & Prasad 1970). Alleles were scored by distance from the origin. Analysis of Allelic Variation .—The allelic variations of the 22 presumptive loci 2002 PRATT & WRIGHT: LYCAENID ALLOZYME PHYLOGENY 221 Table 1. Sample sizes and locations of Lycaena populations used for enzyme analysis. Subgenus Species N Location Chalceria rubida 8 Bridgeport CA xanthoides 3 Southern CA editha 8 E Sierra Nevada, CA dione 8 Lincoln, Nebraska gorgon 8 Frazier Park, CA heteronea 8 White Mts., CA Epidemia helloides 8 Olancha, CA nivalis 13 Sonora Pass, CA mari posa 8 Cedar Lake, CA epixanthe 5 Chats worth, NJ Hyllolycaena hyllus 9 Raven wood, MD Tharsalea arota arota 6 San Gabriel Mts., CA arota nubila 4 Santa Monica Mts., CA Hermelycaena hermes 3 San Diego Co., CA Lycaena phlaeas 10 Newark, DE cuprea 2 Donner Pass, CA were analyzed as individual genotype data by BIOSYS-1 (Swofford & Selander 1989). Numerous genetic distances (Nei, Nei unbiased, Nei minimum, Nei un¬ biased minimum, Nei identities, Nei unbiased identities, Rogers, Modified Rogers, Prevosti, Cavalli-Sforza & Edwards chord, Cavalli-Sforza & Edwards arc, and Edwards “E”) were produced by BIOSYS-1. Cluster analyses were performed by the method of Sneath and Sokal (1973) using the genetic distances and the following algorithms: unweighted pair-group method with arithmetic averaging (UPGMA), weighted pair-group method with arithmetic averaging (WPGMA), single linkage (SL), and complete linkage (CL). Distance Wagner trees, utilizing the multiple addition criterion algorithm of Swofford (1981), were produced by midpoint rooting with Rogers, Modified Rogers, Prevosti, Cavalli-Sforza & Ed¬ wards chord, Cavalli-Sforza & Edwards arc, and Edwards “E” distances (Parris 1972). Results The mean number of alleles per locus, percent polymorphic loci, and hetero¬ zygosity of North American copper species are shown in Table 2. (A copy of allele frequencies is available upon request.) The mean number of alleles per locus and percent polymorphic loci ranged from 1.1 to 6.7 and 9.1 to 54.5, respectively (Table 2). Many cluster analysis trees were produced using algorithms and various genetic distances. Trees with the highest cophenetic correlation and lowest standard de¬ viation were identical in topology to the UPGMA tree created with Nei distances (Fig. 1). The Nei Distance tree groups most species into their currently recognized higher taxa. The species L. rubida (Behr), L. xanthoides (Boisduval), L. editha (Mead), L. dione (Scudder), L. gorgon (Boisduval), and L. heteronea Boisduval cluster in the subgenus Chalceria. Allied species L. gorgon and L. heteronea from a distinct separate cluster within Chalceria. The species L. epixanthe (Boisduval & Le Conte), L. hyllus, L. helloides (Boisduval) L. mariposa (Reakirt), and L. 222 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) Table 2. Mean number of alleles per locus, percent polymorphic loci, and heterozygosity. Population Mean no. of alleles per locus Mean % of loci polymorphic* Heterozygosity Direct count H-W expected** rubida 1.4 27.3 0.124 0.109 (0.1) (0.049) (0.042) xanthoides 1.2 13.6 0.045 0.079 (0.1) (0.033) (0.043) editha 6.7 50.0 0.195 0.170 (0.2) (0.058) (0.046) dione 1.6 54.5 0.239 0.197 (0.1) (0.064) (0.049) gorgon 6.6 50.0 0.186 0.164 (0.2) (0.052) (0.042) heteronea 6.6 31.8 0.157 0.132 (0.2) (0.061) (0.049) helloides 6.6 40.9 0.146 0.135 (0.2) (0.044) (0.040) nivalis 3.0 18.2 0.076 0.070 (0.0) (0.038) (0.033) mariposa 6.6 45.5 0.102 0.100 (0.2) (0.030) (0.030) epixanthe 1.4 27.3 0.127 0.118 (0.1) (0.048) (0.045) hyllus 1.5 31.8 0.095 0.105 (0.2) (0.035) (0.039) arota arota 1.3 22.7 0.106 0.107 (0.2) (0.045) (0.046) arota nubila 1.3 31.8 0.140 0.138 (0.1) (0.054) (0.048) hermes 1.1 9.1 0.030 0.042 (0.1) (0.021) (0.031) phlaeas 1.5 40.9 0.141 0.116 (0.1) (0.050) (0.037) cuprea 1.3 31.8 0.174 0.174 (0.1) (0.062) (0.061) * A locus is considered polymorphic, if more than one allele was detected. ** Unbiased estimate. nivalis (Boisduval) cluster in the subgenus Epidemia. The species L. arota (Bois- duval) and L. hermes cluster together, however they branch below the branching points of other subgeneric groups suggesting they belong to separate subgenera. Lycaena phlaeas and L. cuprea constitute a cluster that branches basally to all of the above taxa. Many Distance Wagner Trees were produced by midpoint rooting with various genetic distances. The tree with the highest cophenetic correlation and lowest percent standard deviation was created with Cavalli-Sforza & Edwards arc dis¬ tances. It is shown in Fig. 2 with host plants added to right in order to facilitate discussion of host shifts. In this phylogeny Chalceria again divides into two distinct clusters with L. gorgon and L. heteronea forming a closely allied pair. The Epidemia align in a fashion similar to the Nei Distance tree with a few subtle differences in branching sequence. Lycaena epixanthe clusters with L. hyllus, 2002 PRATT & WRIGHT: LYCAENID ALLOZYME PHYLOGENY 223 Distance 1.40 1.17 0.93 0.70 0.47 0.23 0.00 H-1-1-1-1-t-1-1-1-1-1-1-K Chalceria Epidemia Tharsalea j Hermelycaena Lycaena H-1-1-1-1-1-1-1-1-—I-1-1-h 1.40 1.17 0.93 0.70 0.47 0.23 0.00 Figure 1. Nei Distance tree. UPGMA tree of 16 North American coppers using Nei distances. Cophenetic correlation 0.925; percent standard deviation 18.896. Subgeneric designations in right margin. whereas in the Nei Distance tree each species branches independently from the main branch of the group. Lycaena helloides clusters with L. nivalis, but in the Nei Distance tree it clusters with L. mariposa. Lycaena arota and L. hermes do not cluster together in the Distance Wagner tree. Lycaena arota branches from an ancestral stem shared with Epidemia, while L. hermes branches toward the base of the subfamily just after the L. phlaeas and L. cuprea branch. Discussion Copper Taxonomy. —In our study Nei distances are selected for cluster analysis since they have been used for evolutionary estimates and phylogenies in other arthropods. Despite small sample sizes, which can diminish the overall confidence in results, all of our trees demonstrate regular branching patterns. The trees branch into distinct species clusters with notable levels of genetic differentiation. The topologies of the Nei Distance and Distance Wagner trees (Figs. 1 and 2) show remarkable congruence with the currently recognized North American copper sub- 224 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) Distance from root 0.00 0.09 0.18 0.26 0.35 0.44 0.53 -+--1-1-1-1-1- H.-» I -1-1-t-—+ rubida xanthoides -— editha — dione — gorgon - heteronea - arota arota - arota nubila - - epixanthe - hyllus - helloides - nivalis - mariposa — hermes - phlaeas - -- cuprea Rumex Eriogonum Ribes J Vaccinium 1 Rumex Polygonum 3 Vaccinium 3 Rhamnus Rumex, Oxyria H- f ■ ■ 4 -1 I I-1— ■ H- i -1-1-1-h 0.00 0.09 0.18 0.26 0.35 0.44 0.53 Figure 2. Distance Wagner tree of 16 North American coppers using Cavalli-Sforza & Edwards arc distances. Cophenetic correlation 0.912; percent standard deviation 13.159. Total length of tree = 4.537. Host plants in right margin. genera. Allozyme differentiation implies that these subgenera are perhaps com¬ patible with genera. Certain copper taxa are absent from the analysis. The addition of Lycaena ferrisi K Johnson & Balogh, L. dorcas Kirby, L. cuprea snowi (W. H. Edwards), and L. ( lophanus ) pyrrhias, as well as larger sample sizes, may improve the trees and disclose further relationships. Miller & Brown (1979) pointed out the primitive position of Lycaena phlaeas and L. cuprea relative to other North American taxa. The ancestral position of these species is supported by our allozyme phylogenies. Several subspecies of L. phlaeas and many L. cuprea-l ike species occur throughout Eurasia (Henriksen & Kreutzer 1982, Higgins & Riley 1983, Korshunov & Gorbunov 1995, Tusov 2000, Gorbunov 2001). With extant relatives in the Palaearctic, it is most likely that the ancestors of both L. phlaeas and L. cuprea originated in the Palaearctic. In our study the population of Lycaena phlaeas sampled is from eastern North America. Klots (1951) noted that eastern subspecies L. phlaeas americana Harris, corrected to L. phlaeas hypophlaeas (Boisduval) by Emmel & Pratt (1998), is 2002 PRATT & WRIGHT: LYCAENID ALLOZYME PHYLOGENY 225 morphologically similar to European L. phlaeas. Opler and Krizek (1984) sug¬ gested that hypophlaeas is adventive and was most likely introduced into North America from Scandinavia during the American Colonial period (17th—18th cen¬ tury). An alternative hypothesis is that eastern populations of hypophlaeas existed endemically in the high elevations of the White Mountains in New England and expanded their range with the introduction of Rumex acetosella. An expansion of this sort has been observed with alpine populations of L. cuprea and L. editha. Both of these species have broadened their range with the introduction of Rumex acetosella into western North America (Emmel & Pratt, personal observation). Also high altitude California L. phlaeas from 12,000 feet elevation can be ex¬ perimentally reared on Rumex crispus at 800 feet elevation (and lower), suggest¬ ing that the species has the ability to rapidly adapt to lowland conditions (Ballmer & Pratt 1989a). Oxyria digyna is the primary host plant of arctic-alpine L. phlaeas in North America (Shields and Montgomery 1966, Ferris 1974, Emmel & Pratt 1998). This plant occurs locally at high elevations on Mount Washington in New Hampshire; the possible existence of high altitude L. phlaeas colonies there and elsewhere in New England has not been studied. Allozyme evidence suggests that each of the four species at the base of the tree (L. phlaeas, L. cuprea, L. arota, L. hermes ) could belong to a separate genus or subgenus. The genetic distance between L. phlaeas and L. cuprea in the Nei Distance Tree (Fig. 1) is greater than the basal branch leading to all other sub¬ genera. Lycaena arota and L. hermes from western North America form a cluster pair in the Nei Distance tree (Fig. 1), but fail to do so in the Distance Wagner Tree (Fig. 2). Thus these species seem to require a different grouping above the species level. If they were placed in separate subgenera, two (L. arota, L. hermes) would occupy monotypic subgenera. Lycaena phlaeas and L. cuprea on the other hand belong to a polytypic Holarctic subgenus. The current assignment of L. cuprea to the Lycaena may change once comparative molecular studies with Pa- laearctic taxa have been completed. Lycaena phlaeas, the type species of Lycaena, will not change assignment. In both trees (Figs. 1 and 2), the species in Chalceria segregate into two distinct subclusters consisting of L. rubida, L. xanthoides, L. editha, and L. dione in one group and L. gorgon and L. heteronea in the other. The branch length between them in the Nei Distance tree (Fig. 1) is virtually the same as branch lengths of other subgenera. A notable shift in host plants has accompanied this split. The first group uses hosts in the plant genus Rumex, while L. gorgon and L. heteronea have shifted to Eriogonum. Ballmer and Pratt (1989b) recognized distinct differ¬ ences in the larvae of these two groups. If these two groups are eventually rec¬ ognized as separate genera or subgenera, Chalceria must be applied to the former and a new genus must be erected for L. gorgon and L. heteronea. In the Nei Distance tree (Fig. 1), the distance between L. dione and L. xan¬ thoides is greater than that between L. xanthoides and L. editha. This is consistent with the notion that L. dione is a distinct species and supports the recent elevation by Opler and Malikul (1992). Whether L. xanthoides and L. editha are fully separate species is a controversial subject (Scott 1980, Pratt et al. 1991). In this analysis L. xanthoides lies intermediate between L. editha and L. dione. Although the genetic distance between L. xanthoides and L. editha is relatively small, L. editha tentatively should retain full species status. The allozyme relationships of 226 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) this group are congruent with the phylogeny in our previous morphological study of the L. edit ha Complex (Pratt et al. 1991). It should be noted that our allozyme study analyzed only a single population of each species and did not sample in¬ termediate L. edit ha X L. xanthoides populations in northern California. We also did not examine L. ferrisi, the purported oldest member of this group. These limitations make it difficult to offer a stronger statement about the species status of L. edit ha. Most of the species within the subgenus Epidemia form a clear cluster with the surprising exception of L. hyllus, which is currently assigned to the subgenus Hyllolycaena. In the Nei Distance tree (Fig. 1), L. hyllus is more closely related to the other Epidemia species than is L. epixanthe. Lycaena hyllus and L. epix- anthe branch together In the Distance Wagner tree (Fig. 2), implying an ancestral relationship. Comparing their phenotypes, this relationship hardly seems possible. It appears that L. hyllus is either a member of Epidemia or L. epixanthe is a member of Hyllolycaena. (Alternatively, L. epixanthe could represent a separate monotypic subgenus.) Egg morphology and larval chaetotaxy reveal a close re¬ lationship between L. hyllus and the Epidemia species (Wright, personal obser¬ vation). Future analyses with the addition of L. dorcas may help determine the breadth of Epidemia and its potential inclusion of L. hyllus. Diapause Changes .—The first branch in the Nei Distance tree (Fig. 1) demar¬ cates a conspicuous split between the species pair L. phlaeas and L. cuprea and the remaining North American coppers. This branch is coincident with a signifi¬ cant biological modification in diapause. Both L. phlaeas and L. cuprea diapause as partially grown larvae, while all other North American coppers diapause in the egg stage, or more accurately as first instars within eggs (Scott 1981, Wright 1983, Pratt & Ballmer 1986). Coppers outside of North America principally dia¬ pause as partially grown larvae well beyond the first instar (Clark & Dickson 1971, Gibbs 1980, Henriksen & Kreutzer 1982, Higgins & Riley 1983). These observations suggest that the evolution of the North American species, excluding those with their closest relatives in the Palaearctic (L. phlaeas and L. cuprea ), involved a diapause change from partially grown larvae to first instar larvae within eggs. Most species of this unique group of obligate egg-diapausers are univoltine. One curious exception is L. hyllus, which has two broods. Progeny of the first brood of L. hyllus develop directly without diapause while the second brood in late summer produces eggs whose first instars enter diapause (Opler & Krizek 1984). Thus the diapause stage of this species is ultimately the same as its North American relatives. The modification of voltinism in L. hyllus appears to be a response to habitat, climate, moisture, and host availability. The precise mecha¬ nism how this species controls diapause is unknown. It is not clear in which stage multibrooded lowland L. helloides diapauses, but we speculate that it too dia¬ pauses within eggs like its congeners. This species is univoltine at high altitude where Scott (1986) reported egg hibernation. High altitude California L. helloides (> 6000 feet) when reared for three generations without diapause near sea level, entered diapause in late fall within eggs (Pratt, personal observation). Host Plant Shifts .—It is likely that the original host plant of the North American coppers was Rumex or a closely related member of the Polygonaceae. Supporting this conclusion is the observation that coppers worldwide use Polygonaceae spe- 2002 PRATT & WRIGHT: LYCAENID ALLOZYME PHYLOGENY 227 cies almost exclusively. The only continent where coppers venture onto hosts outside of the Polygonaceae is North America. Since North American coppers (excluding subgenus Lycaena ) are more derived than their Palaearctic counter¬ parts, host shifts that occurred on this continent were most likely from Polygon¬ aceae to another plant family. Also each North American subgenus with more than one species contains at least one species that feeds on either Rumex or Polygonum (Ballmer & Pratt 1989b). The major host shifts of the North American coppers align with the major branches in the allozyme phylogenies. The first branching stem (subgenus Lycae¬ na) in the Distance Wagner Tree (Fig. 2) did not switch hosts, but the following divergence (L. hermes ) saw a host shift to the plant Rhamnus crocea Nuttal in Torrey and Gray (Rhamnaceae). The branching stem of the Chalceria did not involve a host shift, but within the subgenus there occurred a split leading to the closely allied pair, L. gorgon and L. heteronea, which shifted onto Eriogonum. Although Eriogonum belongs to the Polygonaceae family, the ability to feed on this plant genus may be considered a unique host shift. Rumex feeding species of Chalceria (L. rubida, L. xanthoides, L. editha, and L. dione ) will not feed on Eriogonum in the lab, and in similar fashion L. heteronea and L. gorgon larvae will not feed on Rumex. The Rumex feeding species of Chalceria can readily switch between Rumex and Polygonum (Pratt, personal observation). These ob¬ servations suggest that the shift to Eriogonum is an actual host shift and not easily reversible. The species diversity of the Eriogonum feeders may be greater than presently appreciated. The lineage involving L. heteronea may contain two or more species. On the eastern slopes of the Sierra Nevada in western North America occur two sympatric populations of L. heteronea, one using Eriogonum umbellatum Torrey and the other Eriogonum nudum Douglas ex Bentham. Adults and larvae of these two populations differ morphologically and suggest that two species coexist (Pratt et al. 1991). Samples of these populations were not included in this analysis. The next major divergence in the Distance Wagner Tree (Fig. 2) splits the subgenera Tharsalea and Epidemia. Tharsalea is represented by a single species, which has shifted to exclusive use of Ribes (Glossulariaceae). In the subgenus Epidemia three species feed on either Rumex or Polygonum (Polygonaceae), but two species have shifted to Vaccinium (Ericaceae) (Wright 1983, Pratt & Ballmer 1986, Scott 1986). It appears that these host shifts, unlike the shift to Eriogonum, occurred independently since the two Vaccinium feeders are not closely related. The bog species L. epixanthe is more closely allied to L. hyllus, while L. mariposa is more closely related to L. helloides. We note with interest that host shifts to plants in Ericaceae have also occurred independently in the polyommatine genera Agriades, Lycaeides, and Vacciniina, especially in species adapted to the bog-like habitats (Scott 1986, Emmel & Emmel 1998). Lycaena dorcas, another bog/fen dweller in the Epidemia, has shifted to Potentilla in the Rosaceae. Acknowledgment We sincerely thank the following individuals for their services: Cecilia Pierce and Greg Ballmer helped collect many of the butterflies used in this study; Tom Emmel and John Emmel provided invaluable discussions; Tom Wood of the Uni¬ versity of Delaware and Clay Sassaman of the University of California at Riv- 228 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) erside generously provided lab space for the electrophoretic analysis; and two anonymous reviewers provided helpful comments for improvements in the man¬ uscript. Literature Cited Ballmer, G. R. & G. F. Pratt. 1989a. Instar number and larval development in Lycaena phlaeas hypophlaeas (Boisduval) (Lepidoptera: Lycaenidae). J. Lep. Soc., 43: 59-65. Ballmer, G. R. & G. F. Pratt. 1989b. A survey of the last instar larvae of the Lycaenidae (Lepidoptera) of California. J. Res. Lep., 27 (1): 1-80. Churkin, S. V. 1999. A new species of Thersamonolycaena from Tadjikistan (Rhopalocera, Lycaeni¬ dae). Atalanta, 29 (1/4): 125-129. Clark, G. C. & C. G. C. Dickson. 1971. Life histories of the South African lycaenid butterflies. Purnell & Sons, Cape Town. 272 pp. Clayton, J. W. & D. N. Tretiak. 1972. Amine-citrate buffers for pH control in starch gel electrophoresis. J. Fish. Res. Board Canada, 29: 237-251. Emmel, J. F. & T. C. Emmel. 1998. A new spedes of Agriades (Lepidoptera: Lycaenidae) from the Sierra Nevada and Trinity Alps of California, and the biology and geographic variation of Agriades podarce in California, pp. 287-302. In Emmel, T. C. (ed.). Systematics of western North American butterflies. Mariposa Press, Gainesville, Florida. Emmel, J. F., T. C. Emmel & S. O. Mattoon. 1998. A checklist of the butterflies and skippers of California, pp. 825-836. In Emmel, T. C. (ed.). Systematics of western North American But¬ terflies. Mariposa Press, Gainesville, Florida. Emmel, J. F. & G. F. Pratt. 1998. New subspecies of Lycaeninae from California and a type locality restriction for Chrysophanus cupreus W. H. Edwards (Lepidoptera: Lycaenidae). pp. 661-680. In Emmel, T. C. (ed.). Systematics of western North American butterflies. Mariposa Press, Gainesville, Florida. Farris, J. S. 1972. Estimating phylogenetic trees from distance matrices. Am. Nat., 106: 645-668. Ferris, C. D. 1974. Distribution of arctic-alpine Lycaena phlaeas L. (Lycaenidae) in North America with designation of a new subspedes. Bull. Allyn Mus., 18: 1-13. Gibbs, G. W. 1980. New Zealand butterflies. Identification and natural history. Williams Collins Pub¬ lishers Ltd., Auckland. 207 pp. Gorbunov, P. Y. 2001. The butterflies of Russia: classification, genitalia, keys for identification (Lep¬ idoptera: Hesperoidea and Papilionidae). Thesis, Ekaterinburg, 320 pp. Henriksen, H. J. & I. Kreutzer. 1982. The butterflies of Scandinavia in nature. Skandinavisk Bogforlag, Odense, Denmark, pp. 215. Higgins, L. G. 1975. The classification of European butterflies. William Collins Sons & Co., London. 320 pp. Higgins, L. G. & N. D. Riley. 1983. A field guide to the butterflies of Britain and Europe (5th ed.). Collins Clear Type Press, London. 384 pp. Klots, A. B. 1936. The interrelationships of the spedes of the genus Lycaena Fabricius (Lepidoptera, Lycaenidae). Bull. Brooklyn Ent. Soc., 31: 154-170. Klots, A. B. 1951. A field guide to the butterflies of North America, east of the Great Plains. Houghton Mifflin Company, Boston, Massachusetts. 349 pp. Korshunov, Y. & P. Gorbunov. 1995. Butterflies of the Asian part of Russia. [In Russian] Ural Uni¬ versity Press, Ekaterinburg. 202 pp. Lewis, H. L. 1973. Butterflies of the world. Follett Publishing Company, Chicago, Illinois. 312 pp. Miller, L. D. & F. M. Brown. 1979. Studies in the Lycaeninae (Lycaenidae) 4. The higher classification of the American coppers. Bull. Allyn Mus., 51: 1-30. Miller, L. D. & F. M. Brown. 1981. A catalogue/checklist of the butterflies of America north of Mexico. The Lep. Soc., Memoir No. 2: 1-280. Opler, P. A. & G. O. Krizek. 1984. Butterflies east of the Great Plains. The Johns Hopkins University Press, Baltimore, Maryland. 294 pp. Opler, P. A. & V. Malikul (illust.). 1992. A field guide to eastern butterflies. Peterson Field Guide Series. Houghton Miffin Company, New York. 396 pp. Pratt, G. F. 1994. Evolution of Euphilotes (Lepidoptera: Lycaenidae) by seasonal and host shifts. Biol. J. Linn. Soc., 51: 387-416. 2002 PRATT & WRIGHT: LYCAENID ALLOZYME PHYLOGENY 229 Pratt, G. F. & G. R. Ballmer. 1986. Clarification of the larval host plant of Epidemia mariposa (Lycaenidae) in Northern California. J. Lep. Soc., 40: 127. Pratt, G. F., D. M. Wright & G. R. Ballmer. 1991. Multivariate and phylogenetic analyses of larval and adult characters of the Editha Complex of the genus Lycaena (Lepidoptera: Lycaenidae). J. Res. Lep., 30: 175-195. Scott, J. A. 1980. Geographic variation in Lycaena xanthoides. J. Res. Lep. 18: 50-59. Scott, J. 1981. Hibernal diapause of North American Papilionoidea and Hesperioidea. J. Res. Lep., 18: 171-200. Scott, J. 1986. The butterflies of North America. A natural history and field guide. Stanford University Press, Stanford, California. 584 pp. Scudder, S. H. 1876. Synonymic list of the butterflies of North America, north of Mexico. Bull. Buffalo Soc. Nat. Sciences, 3: 98-129. Shaw, C. R. & R. Prasad. 1970. Starch gel electrophoresis of enzymes: a compilation of recipes. Biochem. Gea, 4: 297-320. Shields, O. & J. C. Montgomery. 1966. The distribution and bionomics of arctic-alpine Lycaena phlaeas subspecies in North America. J. Res. Lep., 5: 231-242. Sibatani, A. 1974. A new genus for two new species of Lycaeninae (s. str.) (Lepidoptera: Lycaenidae) from Papua New Guinea. [Appendix: a tentative scheme of higher classification of Lycaeninae (s.str.) of the world.] J. Australian Ent. Soc., 13: 95-110. Sneath, P. H. A. & R. R. Sokal. 1973. Numerical taxonomy; the principles and practice of numerical classification. W. H. Freeman, San Francisco, California. 573 pp. Swofford, D. L. 1981. On the utility of the distance Wagner procedure, pp. 25-43. In Funk, V. A. & D. R. Brooks (eds.). Advances in cladistics: proceedings of the first meeting of the Willie Hennig society. New York Botanical Gardens, New York. Swofford, D. L. & R. B. Selander. 1989. BIOSYS-1, a computer program for the analysis of allelic variation in population genetics and biochemical systematics. Illinois Natural History Survey, Champaign, Illinois. Tolman, T. 1997. Butterflies of Europe. Princeton University Press, Princeton, New Jersey. 320 pp. Tusov, V. K. (ed.) 2000. Guide to the butterflies of Russia and adjacent territories (Lepidoptera, Rhopalocera). Volume 2 Pensoft, Sofia-Moscow. 580 pp. Wright, D. M. 1983. Life history and morphology of the immature stages of the bog copper butterfly Lycaena epixanthe (Bsd. & Le C.) (Lepidoptera: Lycaenidae). J. Res. Lep., 22: 47-100. Wyatt, C. W. 1961. Additions to the Rhopalocera of Afghanistan with descriptions of new species and subspecies. J. Lep. Soc., 15 (1): 1-18. Received 20 February 2002; Accepted 8 November. PAN-PACIFIC ENTOMOLOGIST 78(4): 230-234, (2002) SYNONYMY OF DASYMUTILLA SICHELIANA (SAUSSURE) (HYMENOPTERA: MUTILLIDAE) 1 Donald G. Manley 2 and William R. Radke 3 department of Entomology, Clemson University, Pee Dee Research and Education Center, 2200 Pocket Road, Florence, South Carolina 29506-9706 3 United States Fish and Wildlife Service, San Bernardino/Leslie Canyon National Wildlife Refuges, Post Office Box 3509, 7628 North Highway 191, Douglas, Arizona 85608 Abstract.—Dasymutilla sicheliana (Saussure) and D. thera (Cameron) have been known from females only, and were synonymized by Mickel in 1965. Dasymutilla intermixta Mickel and D. thalia (Cameron) have been known from males only. Examination of the holotypes and long series of specimens of both has shown these males to be the same species. Pitfall trap collections of both males and females at the Leslie Canyon National Wildlife Refuge near Douglas, Arizona, as well as collection data from specimens collected in both the United States and Mexico, have led to the conclusion that these are male and female of the same species. The name D. sicheliana has precedence. A complete synonymy is included. Key Words .—Insecta, Hymenoptera, Mutillidae, Dasymutilla sicheliana, Dasymutilla intermixta, Dasymutilla thalia, synonymy. Dasymutilla sicheliana was first described as Mutilla sicheliana by Saussure (1867). Saussure made his description on the basis of five female specimens, all from Mexico (two from Cordilliere and three from Tehuacan), and noted three different variations. Mickel examined Saussure’s material in 1931 (unpublished notes) and designated a lectotype. That specimen is in the Geneva Museum, as are Saussure’s other specimens. Sphaerophthalma (sic.) prunotincta was described as a new species by Cock¬ erell (1895) on the basis of a female found in Guanajuato, Mexico, by Dr. A. Duges. Although Cockerell made comparisons to several other species in his description, there was no reference made to M. sicheliana. After examination of Duges’ type specimen, Andre (1898) wrote that S. prunotincta was a synonym of M. sicheliana. The type specimen for S. prunotincta has since been lost (Mickel 1928, 1965). Sphaerophthalma (sic.) thera was described as a new species by Cameron (1895) on the basis of a female found in Milpas, Mexico, by Forrer. That type was examined by Mickel and found to be a synonym for D. sicheliana (1965). The holotype of S. thera is in the British Museum. Mutilla gynaecologica is listed as a new name for S. thera by Dalle Torre (1897). The type specimens of M. sicheliana and S. thera were examined by DGM and found to be in agreement with Mickel. Dasymutilla sicheliana has been pre¬ viously known only from the female, with its distribution being Arizona and Mexico. Dasymutilla intermixta was described by Mickel (1928). The holotype for this species is in the University of Minnesota collection, and has been examined by 1 Technical contribution no. 4767 of the South Carolina Agricultural Experiment Station, Clemson University. 2002 MANLEY & RADKE: D. SICHELIANA SYNONYMY 231 DGM. It has been known only from the male, with its distribution being Arizona and New Mexico. Sphaerophthalma (sic.) thalia was described by Cameron (1895) on the basis of two male specimens found in Guerrero, Mexico by H. H. Smith. The holotype is in the British Museum, and has been examined by DGM. It has been known only from the male, with its distribution being listed as Mexico. Materials and Methods Beginning in 2000, U. S. Fish & Wildlife personnel (WRR) collected mutillids coincidentally with targeted reptiles and amphibians being live-trapped as part of a population dynamics study. The collections were made at Leslie Canyon Na¬ tional Wildlife Refuge in Cochise County, Arizona; Township 21 South, Range 28 East, Section 20 (Lat: 31°35.330' Long: 109°30.500')- Trap arrays were located at an elevation of 1419 m in various microhabitats within the canyon’s riparian corridor. Each array consisted of pitfall traps (19-liter capacity buckets buried in the ground to the rim) at the end of 7.6 m sections of tan-painted, metal drift fences 36 cm high having 2-compartmental funnel traps located at the center of the fence. The funnel traps were boxes 1.2 m long by 0.6 m wide, and 0.3 m high constructed of 2.3 mm (1/8") hardware cloth separated longitudinally by a piece of plywood such that they functioned as two parallel traps. Funnels with 5 cm entrance holes led into these traps from each end. Funnels and pitfall traps were shaded from the sun with plywood coverings, and plywood bucket trap covers were elevated approximately 4 cm from ground level to provide access to reptiles, amphibians, and invertebrates. The plywood covers made it impossible for flying invertebrates to view the contents of the pitfall traps, and additionally made it virtually impossible for flying invertebrates to exit the traps once inside. About 5 cm of loose soil served as a substrate in the pitfall and funnel traps. In 2000, four trap arrays were operated from 5 April through 7 December. In 2001, eight trap arrays were in operation from 29 March through 6 December. Traps were checked approximately every other day to remove and record the vertebrates and invertebrates that were captured. All mutillids (Hymenoptera: Mutillidae) were collected, pinned, and shipped to Clemson University (DGM) for identifi¬ cation. On 19 June 2000 one of the pitfall traps contained a single female of D. sich- eliana and a single male of D. intermixta. On 20 June 2001 one of the pitfall traps contained a single female of D. sicheliana and eight males of D. intermixta. On 2 July 2001 one of the pitfall traps contained a single female of D. sicheliana and a single male of D. intermixta. All collections were by WRR, and no other mutillid specimens were collected in those traps on those dates. Material Examined. —In addition to the type specimens mentioned above, the following material has been examined by DGM (all are females of D. sicheliana and males of D. intermixta and D. thalia): USA. ARIZONA. COCHISE Co.\ Mouth of Carr Canyon, Huachuca Mts., 10 Aug 1940, C. D. Michener, 1 $; Garden Canyon, Huachuca Mts., 1954, W. H. Mann, 1 $ (homotype, M. sicheliana)-, Cottonwood Canon, Peloncillo Mts., 25 Sep 1958, D. S. Creighton, 1 $; Portal, 8 Sep 1959, H. E. Evans, 1 <3; Skeleton Canyon, 24 Aug 1962, P. Weens, 1 $ (homotype, D. intermixta)-, Carr Canyon, Huachuca Mts., 20 Jul 1969, G. H. & D. E. Nelson, 1 $ (homotype, S. thera)\ Miller Canyon, Huachuca Mts., 3 Apr 1973, R. F. Sternitzky, 1 2; Miller Canyon, Huachuca Mts., 21 Jun 1974, E. R. Hoebeke, 1 <3; Chiricahua Mts., 9 Aug 1974, G. H. Nelson, 1 6 (homotype, D. intermixta)-, Portal, 12 Aug 1974, H. & M. Townes, 2 c CD O C 4 O 4 O o r- Q. C/) O I ^ 0 35 37.5 40.0 42.5 45.0 47.5 49.0 52.5 55.0 Temperature Treatment (x 3 hrs) Figure 2. Mean (±1 SE) concentrations of hsp70 (ng/gl) derived from head capsules of Megachile apicalis and M. rotundata for nine temperature treatments. asitoid feeding activity) at these two temperatures, respectively. None of these had advanced beyond the prepupal stage. All cocoons showed evidence of para- sitoid entry (small circular openings) and no pre-adult bees had survived parasit- oid attack after the four-week incubation period. Teneral adults had apparently succumbed to dessication after parasitoid damage to the cocoon. Trial 2: Survivorship at 50.0° C .—The 50.0° C treatment corroborated our ear- Figure 3. Percent prepupal survivorship of M. apicalis and M. rotundata compared at nine tem¬ perature regimes. 242 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) lier result that M. apicalis has higher survivorship at the highest temperature treatments in our study. Emergence of adult Megachile apicalis from brood cells after six weeks was 40.6% (39 of 96 viable brood cells) relative to 20.0% (18 of 90 viable cells) for M. rotundata. Dissection of non-emerged cocoons revealed that 68.8% (66 of 96) of M. apicalis brood cells exposed to 50.0° C either suc¬ cessfully emerged, died as teneral adults or advanced to the pupal stage. For M. rotundata , 32.2% (29 of 90) of prepupae advanced to the same stages. No para- sitoid feeding activity was observed in any cells of either species during this trial. A number of bees that emerged from cocoons showed evidence of teratogenic effects, including at least 28.2% (11 of 39) of M. apicalis and 5.6% (1 of 18) of M. rotundata. Typically, these malformations related to appendages (e.g., missing or stunted wings and legs). Trial 3: Stress Proteins in Acclimated vs Heat-shocked Larvae .—Using the same ELISA technique as described above, no significant difference in hsp70 levels was found between M. apicalis (5.25 ± 0.749 ng/pl) and M. rotundata (5.90 ± 1.067 ng/|jil) when maintained at 25.0° C {t — —0.503, df = 4, P = 0.32). However, a significant difference in levels was recorded after the three- hour exposure to 35.0° C. Megachile apicalis contained substantially lower levels of hsp70 than M. rotundata (4.15 ± 0.543 vs 7.24 ± 0.481; t = —4.269, df = 4, P < 0.01). Discussion Although both of our study species appear to tolerate high-temperature envi¬ ronments, the geographic range and nesting habits of M. apicalis suggest it has higher thermotolerance than M. rotundata (Thorp 1996, Barthell et al. 1998). Our study corroborates these observations with higher survivorship for M. apicalis exposed to 50.0° C (and above) while higher stress protein (hsp70) levels were measured in M. rotundata in response to elevated temperatures (except at 55.0° C where the most rapid mortality presumably occurred for both species). These differences are likely to be induced by heat-shock since we later failed to find a significant difference between species when samples received a four-day accli¬ mation period at 25.0° C in Trial 3. (Hsp70 levels did, however, differ significantly after the three-hour exposure to 35.0° C.) These results therefore support the conclusion that the elevated levels of hsp70 recorded in M. rotundata were a result of temperature-induced stress. It is unclear whether any species-specific developmental differences influenced our results in the experimental trials. Megachile apicalis and M. rotundata do have differing flight periods (Thorp et al. 1992) and the developmental sequence from diapausing prepupal to pupal stages just prior to adult emergence could affect hsp70 levels differentially between species. Megachile rotundata demon¬ strates variation among protein concentrations during this transition, for example, and hsp70 appears to be one of those that varies (Rank et al. 1982, 1989; Hranitz & Barthell in press). However, we exposed both study species (together) to at least a three-week cold storage period and we collected cells of M. apicalis later in the season (August) to obtain as many of the diapausing generation bees as possible (the same stage as the commercially-harvested M. rotundata used in the study). The literature is replete with evidence that elevated hsp70 levels reflect stress 2002 BARTHELL ET AL.: MEGACHILE HIGH TEMPERATURE RESPONSE 243 in organisms (Feder & Hofmann 1999). In insects this pattern has been observed in fmitfly larvae, Drosophila melanogaster (Meigen), when exposed to high tem¬ peratures in nature (Feder et al. 1997). Cold-shock stress also produces elevated levels of hsp70 in the flesh fly Sarcophaga crassipalpis Macquart (Joplin et al. 1990) . In diapausing gypsy moths, Lymantria dispar L., an elevation in stress proteins occurs in response to both high and low temperatures (Yocum et al. 1991) . Both bacterial infection and heat stress increase hsp70 production in honey bees (Severson et al. 1990, Gregorc & Bowen 1999). The significantly higher hsp70 levels we recorded in M. rotundata probably reflect lower heat tolerance in this species and is consistent with the northerly distribution of its feral popu¬ lations in California. However, since the M. rotundata specimens used in our study originated from a commercial source in Canada we still await the oppor¬ tunity to compare individuals of M. apicalis and M. rotundata sampled from sympatric populations in California. Although no thermotolerance studies exist in the literature for M. apicalis, several such studies exist for the economically important M. rotundata. Whitfield and Richards (1992) demonstrated a slightly decreased developmental rate for later instar larvae of M. rotundata in the range of 32.0° to 35.0° C. Undurraga and Stephen (1980) found no survivorship of either prepupae or pupae during either brief or extended (as long as 3 h) exposures to 50.0° C, but with high survivorship at 45.0° C. The latter study shows lower survivorship at 50.0° C than in our study for M. rotundata (0.0 vs 20.0%). However, these differences could be explained by differing acclimation conditions between studies as well as larval developmental conditions in the original, pre-study environments; the importance of this latter point is emphasized in recent contributions by Kemp and Bosch (2000, 2001). Our criteria for assessing survivorship (based upon parasitoid feeding activity) differed from these other studies as well. Nonetheless, it is clear that M. rotundata is severely compromised (> 50.0% mortality) in the prepupal stage when exposed to 50.0° C for a three hour period or less. Teratogenic effects were not specifically addressed in the mortality studies cited above, nor by Tepedino and Parker (1986), even though we observed them in both species after Trial 2. Such effects were not conspicuous among specimens that emerged from any of the temperature treatments conducted in Trial 1. Al¬ though prepupae used in Trial 2 came from the same source as those used in Trial 1 (and were maintained in cold storage), there is a possibility that bees in Trial 2 were somehow physiologically altered between trials. It may also be that other components of our experimental protocol (or these factors in combination with the treatment exposure) produced the effects we observed in Trial 2. Although no comparative studies measuring survivorship and stress proteins among solitary bee species exist in the literature, such studies have been con¬ ducted for marine molluscs. Two blue mussel species, Mytilus trossulus Gould and M. galloprovincialis Lamarck (collected from northern and southern latitudes, respectively), for example, show significant differences in hsp levels under tem¬ perature stress (Hofmann & Somero 1996). Mytilus trossulus accumulates higher levels of hsp66 and hsp70 than M. galloprovincialis after acclimation to 13.0° C for eight weeks, evidence that the more cold-adapted M. trossulus experiences greater physiological stress at temperatures above their normally encountered tem¬ perature (ca. 10.0° C). In another study, Tomanek and Somero (1999) studied four 244 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) marine snails in the genus Tegula collected from sites along the Pacific coast of the U.S.A. One of these, T. rugosa (Adams), originated from a more southerly locale than the others (Baja California) and displayed higher onset, peak and inactivation temperatures for hsp70 expression relative to three temperate species collected near Pacific Grove, California. One of the three temperate species, T. funebralis (Adams), lives in a broader range of habitats (mid- to low-intertidal zones) that can expose it to high air temperatures during low tide. It also has higher onset, peak and inactivation temperatures in comparison with the other two temperate species. Our findings are consistent with aspects of the marine invertebrate studies cited above. Megachile rotundata, a more northerly-distributed species accumulates higher hsp70 levels than M. apicalis during heat-treatments in our study. The generally high levels of hsp70 in Trial 1 may have obscured the actual peaks as well as the onset and inactivation temperatures for both species. However, a sig¬ nificant decline from the observed peaks of hsp70 occurred at a lower temperature in M. rotundata than in M. apicalis, suggesting a higher inactivation temperature for the latter species. The unexpectedly high levels of hsp70 at lower temperatures (relative to acclimated individuals in Trial 3) may reflect a heat-shock response to rapid heating rates (> 10° C in < 10 min) used in Trial 1, a condition unlike larvae would experience in nature. Lutterschmidt and Hutchison (1997) caution about such confounding effects in studies of thermotolerance. Thermotolerance in an invasive species may be an important indicator of the range that it will ultimately occupy in a newly invaded environment. Indeed, thermotolerance has been a key element in predicting the outcomes of other in¬ vasive Hymenoptera such as African honey bees (Taylor 1977). Although the original, Eurasian ranges of M. rotundata and M. apicalis are unclear from reports in the literature, these species are currently occupying ranges in the western USA that are in accord with the thermotolerance patterns recorded in our study. Me¬ gachile apicalis appears to be able to occupy a higher thermal niche relative to M. rotundata, but we still await comparisons with sympatric native species iden¬ tified in previous studies (Barthell et al. 1998, Frankie et al. 1998). Such com¬ parisons will allow us to determine if thermotolerance is a factor in the rapid range expansion observed for M. apicalis since its initial detection in southern California nearly two decades ago (Cooper 1984). Acknowledgment We thank D. Nahuliak, R. Bitner and S. Peterson (International Pollination Systems) for their advice and assistance. The management and staff of the Cos- umnes River Preserve near Galt, California, facilitated our field collections of M. apicalis specimens. In the laboratory, M. Hartless assisted in dissections and prep¬ aration of specimens for laboratory analyses. Specimen x-rays were conducted by S. Maslowski (University of California, Davis, Veterinary Medical Hospital). An earlier draft of this manuscript was provided for commentary to G. Frankie (Uni¬ versity of California, Berkeley), W. Kemp (USDA, Fogan, Utah) and W. Stephen (Oregon State University). Funding for this work was provided by the University of Central Oklahoma’s College of Graduate Studies and Research (Deans S. N. Rao and W. R. Radke). The C. P. Alexander Grant (Pacific Coast Entomological Society) provided a partial page charge waiver for this article. 2002 BARTHELL ET AL.: MEGACHILE HIGH TEMPERATURE RESPONSE 245 Literature Cited Barthell, J. F. 1992. Heterogeneity, invaders, and the population dynamics of some oak forest solitary bee communities. Ph.D. Dissertation, University of California, Berkeley. Barthell, J. F, G. W. Frankie & R. W. Thorp. 1998. 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PAN-PACIFIC ENTOMOLOGIST 78(4): 247-254, (2002) A NEW SPECIES OF GLAUCINA HULST FROM WYOMING AND COLORADO, AND DESCRIPTION OF THE FEMALE OF G. NEPHOS RINDGE (LEPIDOPTERA: GEOMETRIDAE) Clifford D. Ferris 1 ’ 3 and John S. Nordin 2 ’ 3 Research Associate, C. P. Gillette Museum of Arthropods Diversity, Colorado State University, Ft. Collins, Colorado (mailing address: P.O. Box 3351 University Station, Laramie, Wyoming 82071-3351) 2 2217 Sky View Lane, Laramie, Wyoming 82070 3 Research Associate, Florida State Collection of Arthropods, Gainesville, Florida 32608 Abstract .—A new species in the geometric! moth genus Glaucina is described from Wyoming and Colorado. The previously unknown female of Glaucina nephos Rindge is described from Wyoming specimens. Key Words .—Insecta Colorado, Geometridae, Glaucina n. sp., Insecta, Lepidoptera, Wyoming. Ultraviolet light traps placed by the first author in the Sherman Hills east of Laramie, Wyoming in late June and early July 1999 yielded two males and two females of a large and virtually unmarked gray Glaucina. Comparison of these specimens with the imagines illustrated in Rindge’s 1959 revision of the genus produced no matches. Subsequent dissections of a male and female with com¬ parison to Rindge’s plates again produced no matches. Photographs of the spec¬ imens and their genitalia were subsequently sent to Dr. Rindge at the American Museum of Natural History for his opinion. In a letter to Ferris dated 19 April 2001, he replied: “I agree that the Glaucina is an undescribed species; we have nothing to match it.” When Nordin then examined a backlog of unplaced specimens in his collection and unprepared material in his freezer, he found four specimens from Colorado and a few additional specimens from Albany Co., Wyoming. While Ferris was on extended travel in 2001, Nordin operated ultraviolet light traps in Albany Co. at sites where specimens had been taken previously. This effort generated the additional specimens included in the type series. In addition to the new species, two other large Glaucina, interrupt aria (Grote) and nephos Rindge, occur in Albany Co. The dark dorsal forewing markings of G. nephos immediately separate it from the new species. The new species is most easily confused with G. interruptaria. Fresh specimens of G. interruptaria have a distinct pattern of fine dark markings on the dorsal forewing that is absent in the new species. Worn specimens must be dissected to ascertain identity. Addi¬ tional comments follow in the Diagnosis and Discussion section. To permit easy comparison of species, the format of the ensuing descriptions is purposely modeled after the format used in the 1959 revision of Glaucina by Rindge. The new species is described from 51 males and 9 females from Wyo¬ ming and Colorado. It seems to have been missed previously because it flies early in the season when weather conditions are extremely variable, and before itinerant 248 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) collectors normally travel in the region. Additionally, it is localized and may have specialized habitat requirements. Glaucina incognitaria Ferris and Nordin, NEW SPECIES (Figs. 1-5, 7-12) Types. —Holotype, male (Figs. 1 and 2), Wyoming, Albany Co., T[ownship]15N R[ange]71 W S[ection]29, NE of Pole Mtn., south of Happy Jack Rd., 41°13.78' N 105°22.30' W, 2538 m, 23.vi.2001, leg: J. S. Nordin. Paratypes, 50 males and 9 females with data as follows (specimens leg: J. S. Nordin except as noted): WYOMING, Albany Co.: T12 N R72 W S24, 41°00.34' N 105°25.00' W, 2309 m, 2.vii.l999, If; T14 N R71 W S36, 41°08.19' N 105°17.54' W, 2355 m, 8.vii.l995, 1 m; T15 N R71 W S18, 41°15.82' N 105°23.64' W, 2500 m, 1- 2.vii.l999, 2 m, If, leg; C. D. Ferris; T15 N R71 W S29, 41°13.78' N 105°22.30' W, 2538-2544 m, 24-25.vi. 1999, If, leg: C. D. Ferris; 22.vi.2000, 4 m; 24.vi.2000, 1 m; 25.vi.2001, 14 m, 2f; 27.vi.2001, 2 m; 28.vi.2001, 8 m, 4f; 30.vi.2001, 9 m; 2.vii.01 4 m; T15 N R73 W SI, 41°17.89' N 105°31.50' W, 2277 m, 5.vii.l988, 1 m. COLORADO: Alamosa Co., Road 150, Zapata Creek, 37°41' N 105°33' W, 2380 m, 4.vi.l994, 2 m; Dolores Co., Road 532 NW slope of Cottonwood Creek, 37°40' N 108°18.5' W, 2380 m, 24.v.2000, 1 m; Rio Blanco Co., Hwy. 139 at Garfield Co. line, 39°39' N 108°48' W, 2176 m, 27.V.1990, 1 m. Holotype and a female paratype will be deposited in the collection of the American Museum of Natural History. Additional paratypes will be deposited in other public museums and in the collections of the authors. Description of Male (Fig. 1). —Head, vertex dark gray, scales very narrowly white-tipped; frons dark gray with a few scattered whitish scales, mainly dorsolaterally, dorsolateral areas swollen and clearly separated by a trough dors ally and grading into a slight ridge toward the lower margin of the frons; palpi dark gray with whitish scales basally just below eye and at their extreme tips, palpi extending beyond plane of frons by approximately two-thirds of the diameter of the eye, antenna approximately 1 cm in length or 55% of the length of the FW, stalk obscurely speckled gray and whitish; narrow white collar just at base of head and behind a broader collar of white-tipped dark gray scales at the front of the thorax. Thorax above medium gray with white-tipped scales, some scales grayish-brown or dark gray; below white at base of wings shading into pale gray distally; legs clothed with white-tipped medium gray scales. Abdomen gray to grayish brown, sprinkled with a few dark scales especially toward anterior portion of each segment with a terminal row of strongly white- tipped scales along the posterior margin of the first four segments; ventrally paler with heavy sprinkling of white scales; aggregations of dark gray scales along the midline and immediately to each side form three somewhat broken thin dark parallel lines (visible only if there is no abdominal greasing). Upper Surface of Wings: Forewings, uniform medium gray with only the slightest suggestion of dark scaling forming an indistinct pm line; under magnification a few widely scattered dark scales are visible; fringes concolorous to the naked eye, but under magnification flecked with white and with darker scales at the vein ends. Hind wings concolorous with forewings with a weak accumulation of dark scales at anal angle; fringes as in the forewings. Under Surface of Wings: Uniformly medium gray with some diffuse speckling by slightly darker scales; hind wing only very slightly lighter in color than forewing. Length of Fore wing: Holotype = 18 mm; range 16-19 mm; average (51 males) 17.75 mm. Description of Female (Fig. 3 ).—Similar to the male, except for filiform antennae and shorter and stouter abdomen. Length of Forewing: 16-19 mm; average of 9 females = 17.9 mm. Male Genitalia .—Fifteen specimens dissected. Uncus with width of base just slightly less than length of uncus, lateral margins expanded basally, the apex decurved terminally and ending in a sclerotized point; gnathos with small median enlargement and slightly bilobed apically; valves broadly rounded with slightly angulate outer margin, costa (when not flattened, see left side of Fig. 7) broadly convex and folded, enlarged medially into valve, distally slightly tapered and ending in a setose 2002 FERRIS & NORDIN: GLAUCINA FROM WYOMING AND COLORADO 249 HOLJ3TYPE Glaus ota oicogmtaria C. D. Ferns & J S. Nordm 25 June 2001 black light trap leg. J.S. Nordm WYOMING: Albany Co. T15N R71W Sect. 29 NE of Pole Mt. south of Happy Jack Rd, El. 8320 ft. 2 PARA TYPE No. 01 Glaus ma inc ogmlaria C. D. Ferris &J . S Nordin 28 June 2001 black light trap leg. J.S. NoroirT WYOMING: Albany Co. T15N R71W Sect. 29 NE of Pole Mt. south of Happy Jack Rd, El. 8320 ft. 3 4 0 III 2 cm Figures 1-6. Glaucina species. Figure 1. Glaucina incognitaria Ferris & Nordin, holotype male (dorsal). Figure 2. Specimen labels for male holotype; red holotype label. Figure 3. Glaucina incog¬ nitaria Ferris & Nordin, female paratype (dorsal). Figure 4. Specimen labels for female paratype; yellow paratype label. Figure 5. Male paratype (dorsal) of G. incognitaria showing forewing partial median band, Albany Co., WY, 30. vi. 2001, J. S. Nordin. Figure 6. Glaucina nephos female (dorsal), Albany Co., WY, 21.V.1999, C. D. Ferris. terminal protuberance; sacculus arm long (extending about 0.8 times the length of the costa) and moderately slender with slightly broader width distally, terminating in a rounded apex armed with six spines, two heavy outer spines and four smaller and less robust inner spines (Fig. 8), base of valve with sclerotized band of nonuniform width extending from basal area of inner portion of costal swell¬ ing to the sclerotized base of the sacculus; median juxta slightly longer than wide with finely pitted surface; saccus wide and broadly convex; aedeagus (Fig. 9) as long as the valves, moderately straight with diameter approximately one-sixth of the length, vesica armed with a slightly curved and slender dentate strip, which upon vesica eversion (Fig. 10) resolves into a membranous narrow band, the surface of the anterior half with triangular projections similar to the teeth of a wood rasp. 250 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) Figures 7-12. G. incognitaria genitalia. Figure 7. Male, aedeagus removed. Figure 8. Male, right valve (flattened). Figure 9. Male aedeagus, arrows indicate ends of vesica sclerotized band. Figure 10. Male, vesica everted, arrows indicate ends of vesica sclerotized band. Figure 11. Female. Figure 12. Female, detail of lamella postvaginalis. Female Genitalia (Fig. 11 ).—Two specimens dissected. Sterigma with a large asymmetrical oval (compressed at top) slightly sclerotized lamella postvaginalis (Fig. 12) bordered anteriorly and laterally by numerous lightly sclerotized folds, with antevaginalis a small medially-indented sclerotized ridge; ductus bursae very short, sclerotized, roughly cylindrical, slightly longer than wide; corpus bursae elongate with tapering sides, with virtually unsclerotized longitudinally striated short tapered neck, neck and body hardly separable, enlarging into terminal, ovoid portion of bursae; prominent signum, transverse with inward-pointing median ridge, located at approximately mid-distance between the base of the ductus bursae and the apex of the corpus bursae; ovipositor lobes typical of the genus. 2002 FERRIS & NORDIN: GLA UCINA FROM WYOMING AND COLORADO 251 Biology and Larval Host. —Unknown. Presence of fine soil particles trapped in the hairs at the tip of the abdomen of female specimens suggests oviposition close to the ground on a low-growing herbaceous plant. With a few exceptions (open prairie), this moth has been taken by black light in moderately dry coniferous environments (spruce-pine in Wyoming). Distribution .—As is shown in Fig. 15, this moth is known presently from several tightly-grouped localized areas in Albany Co., Wyoming, and from two counties in western and one in extreme south-central Colorado. Etymology .—The species name reflects the previously unknown status of this moth and is configured to be consistent with the formation of other species names in the genus. A suggested common name is “Unknown Glaucina.” Diagnosis and Discussion .—On the dorsal fore wing in nine males and one female there is the suggestion of antemedial and postmedial lines (Fig. 5), thus forming an open medial band. The only variation in the male genitalia noted was 252 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) Figures 14-16. G. nephos, female genitalia. Figure 14. Complete, but corpus bursae deflated. Figure 15. Detail of lamella postvaginalis. Figure 16. Corpus bursae showing signum at left side. in one specimen from Dolores Co., Colorado in which the sacculus apex was armed with seven spines, three large outer spines and 4 smaller inner spines. Based on the male and female genitalia, G. incognitaria is closest to G. ig- navaria (Pearsall) [Arizona, Colorado, New Mexico] and G. foeminaria (Dyar) [Puebla, Mexico], and thus belongs in Group IV of Rindge (1959). In the male genitalia of G. ignavaria, the apex of the sacculus is armed with a cluster 5-8 elongate spines with an apical protuberance beyond the spines; in foeminaria the 2002 FERRIS & NORDIN: GLAUCINA FROM WYOMING AND COLORADO 253 length of the sacculus is considerably shorter than in G. incognitaria, and the apex is armed with 2 or 3 robust spines and 2 or 3 short spines. The overall aspect of the female genitalia in G. ignavaria and G. foeminaria is similar to that of G. incognitaria, with the major differences being the geometry of the ductus bursae and lamella postvaginalis. In G. ignavaria the lamella post- vaginalis is large and elliptical, while in G. foeminaria it is also elliptical, but smaller than in G. ignavaria. In the Rocky Mountain Region the most likely species with which G. incog¬ nitaria might be confused is G. interruptaria. Fresh specimens are easily separated by the lack of distinct dorsal forewing maculation in G. incognitaria, in contrast to the well-defined but light maculation in interruptaria. Genitalic dissection is required to separate worn specimens. In the male genitalia, the terminal portion of the sacculus arm in G. interruptaria is covered by numerous small short spines, while the terminal portion of the sacculus arm in G. incognitaria is equipped with two heavy outer spines and four smaller and less robust inner spines. The main characters in the female genitalia that separate the two species are: lamella post¬ vaginalis, essentially trapezoidal with the wider base rounded and convex in G. interruptaria, large asymmetrical oval (compressed at top) in G. incognitaria with convex top portion much wider than the nearly pointed rounded base; signum, sclerotization nearly symmetrical above and below inward-pointing median ridge in G. interruptaria, sclerotization asymmetric about inward-pointing median ridge in G. incognitaria with upper portion semicircular and uneven reduced lower portion. Glaucina nephos Rindge (female) (Figs. 5, 14-16) The female of this species was unknown to Rindge (1959). Males are relatively common in southeast Wyoming at black light, but females do not come readily to light. Over a number of years, the authors have managed between them to obtain 13 female specimens from several localities in Albany Co., Wyoming with collection dates from 18 May to 18 June at elevations from 2270 m to 2500 m. Similarity in wing pattern was used to associate the females with males of nephos. Two genitalic dissections were studied. Description of Female (Fig. 6 ).—Similar to the male as described by Rindge, except for filiform antennae and shorter and stouter abdomen. The dorsal dark wing markings are less distinct than in the males. Length of Forewing: 16-18 mm; average of 13 females = 17.0 mm. Female Genitalia (Fig. 14 ).—Sterigma with a large complex slightly sclerotized lamella postvagin¬ alis (Fig. 15), consisting of a smaller slightly distorted and displaced circle overlying a larger slightly distorted circle, bordered anteriorly and laterally by a few lightly sclerotized folds, with antevaginalis a small shallowly medially-indented sclerotized ridge; ductus bursae challis-like, sclerotized, flared laterally at top and tapering to the junction with the neck of the corpus bursae, slightly shorter than wide; corpus bursae (Fig. 16) with well-defined neck, in length at least half of length of corpus bursae, very weakly striated and dotted with small pits just below junction with ductus bursae, enlarging into an apically tapering terminal bulb; prominent signum, transverse with narrow inward-pointing median ridge, located at approximately mid-length of the tapered bulb; ovipositor lobes typical of the genus. Biology .—The biology and host plant of this species remain unknown. Observation .—Both G. incognitaria and G. nephos occupy the same habitats and may be taken at light on the same night, however the known geographic 254 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) range of G. nephos is much greater [Arizona, Colorado, Idaho, Wyoming] than that currently known for G. incognitaria. G. nephos is also a Group IV species. Acknowledgment The authors thank Dr. F. H. Rindge, Curator Emeritus, Lepidoptera, American Museum of Natural History for confirming the new Glaucina species, and for his encouragement to describe it. The authors acknowledge helpful changes to the manuscript suggested by Charles V. Covell Jr. and an anonymous reviewer. Literature Cited Rindge, F. H. 1959. A Revision of Glaucina, Synglochis, and Eubarnesia (Lepidoptera, Geometridae). Bulletin AMNH, 118 (6): 259-366. Received 3 April 2002; Accepted 4 Nov. 2002. PAN-PACIFIC ENTOMOLOGIST 78(4): 255-264, (2002) CHINESE SPECIES OF THE JUMPING SPIDER GENUS PORTIA KARSCH (ARANEAE: SALTICIDAE) Xianjin Peng and Shuqiang Li Institute of Zoology, Chinese Academy of Sciences, Beijing 100080, P. R. China Abstract .—The present paper presents a revision of Chinese Portia spiders. A total of six species, including one new species —Portia wui Peng & Li, are known from China. Descriptions of new species and diagnosis of known species are given. Distributional data, a key to Chinese species, and illustrations of body and genital organs are provided. Key Words. —Araneae, Salticidae, Portia, revision, new species, China. The spider genus Portia was erected by Karsch (1878: 774) to accommodate Portia schultzii Karsch. Most known species of Portia distributed in the Oriental region, few in the Ethiopia. To have a better understanding on Chinese represen¬ tatives of this jumping spider genus, we have examined the specimens of Portia deposited in the Institute of Zoology, Chinese Academy of Sciences (IZCAS), Hunan Normal University (HNU), and Lanzhou University (LZU). Results of this museum survey are reported in the present paper. Descriptions were made based on specimens fixed in 80% ethanol. Specimens were examined and figured under SZ40-Olympus stereomicroscope. Epigynum was figured before it was dissected from the spider abdomen, while vulva was figured after it was macerated in lactic acid. The sequence of leg segments in measurement data is as follows: Total (femur, patella + tibia, metatarsus, tarsus). Measurements are given in millimeter (mm). Terminology adopted is that used by Wanless (1978). Abbreviations used: AER-anterior eye row, AL-abdominal length, ALE-anterior lateral eye, AME-anterior median eye, AW-abdominal width, BTA-basal tibial apophyses, CD-conductor, CL-carapace length, CLYH-clypeus height, CW-cara- pace width, E-embolus, EFL-length of eye field, ITA- intermediate tibial apoph¬ yses, PER-posterior eye row, PLE-posterior lateral eye, RTA-retrolateral tibial apophyses, SD-sperm duct, T-tegulum, TA-tegular apophysis, TF-tegular furrow, TL-total length, VTA-ventral tibial apophysis. Portia Karsch, 1878 Sinis Thorell, 1878, Ann. Mus. civ. stor. nat. Genova 13: 269. Type species Sal- ticus fimbriatus Doleschall, 1859 by original designation. Portia Karsch, 1878, Zeitschr. ges. Naturw 51: 744. Type species Portia schultzii Karsch, 1878 by original designation. Boethoportia Hogg, 1915, Proc. Zool. Soc. Lond. 1915: 501. Type species Boeth- oportia ocellata Hogg, 1915. Neccocalus Roewer, 1965, Annls Mus. r. Afr. cent. ( Sci. Zool.) 139: 20. Type species Cocalus africanus Thorell, 1899 by original designation. Type Species.—Portia schultzii Karsch, 1878, by original designation. Diagnosis. —Medium to large spiders ranging from about 4.50 to 9.50 in length. Carapace high and elevated; usually with marked slope from PLE to posterior 256 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) margin of carapace; PME well developed, almost as big as ALE, about midway between ALE and PLE or closer to ALE; PER usually narrower than AER; EFL about 35-55 percent of carapace. Chelicera with 3 promarginal teeth and 3-6 retromarginal. Legs slender and long, with conspicuous fringes, spines numerous and strong. Abdomen usually ornate with tufts of hairs. Male palpal organ: bulb oval; embolus usually long and slender; tegulum with a deeply curved furrow, and sometimes with a small apophysis; tibia with numerous apophyses; cymbium usually with distinct flange. Epigynum weakly sclerotized, openings small and usually unclear; compulatory ducts short, wide and strongly sclerotized; spermathecae oval and big. Portia is represented by 14 species worldwide, including 1 new species de¬ scribed in this paper. These include —Portia albimana (Simon, 1900) (India to Vietnam), P. assamensis Wanless, 1978 (India to Malaysia), P. crassipalpis (Peckham & Peckham, 1907) (Singapore, Borneo), P. fimbriata (Doleschall, 1859) (Nepal, Sri Lanka to Australia), P. heteroidea Xie & Yin, 1991 (China), P. hoggi Zabka, 1985 (Vietnam), P. jianfeng Song & Zhu, 1998 (China), P. labiata (Thorell, 1887) (Sri Lanka to Philippines), P. orientalis Murphy & Murphy, 1983 (China), P. quei Zabka, 1985 (China, Vietnam), P. schultzi Karsch, 1878 (Central, East, Southern Africa, Madagascar), P. songi Tang & Yang, 1997 (China), P. strandi Caporiacco, 1941 (Ethiopia) and P. wui n. sp. (China). Up to now, a total of 6 Portia species including P. orientalis Murphy and Murphy, 1983 have been recorded from China. Key to Chinese Species of Portia 1. Male . 2 - Female .7 2. Tibia with more than 3 apophyses (Figs. 1C, 2B) . 3 — Tibia with 3 apophyses . 4 3. Embolus slender and long, encircled with conductor basely in ventral view . P- jianfeng (Fig. IB) — Embolus short with much larger base and sharp end, conductor invisible . P. songi (Fig. 2B) 4. Cymbium with a horn-shaped apophysis in addition to flange (Figs. 3C, 5. 6 . 7. 3D) . P. wui, NEW SPECIES Cymbium without apophysis, only with flange. 5 Tegulum furrow with deep curve . P.quei Tegulum furrow with shallower curve . 6 Chelicera with 4 retromarginal teeth, retrolateral tibial apophysis bar-like, with smooth end . P. heteroidea Chelicera with 3 retromarginal teeth, retrolateral tibial apophysis longer with sharp end. P. orientalis Spermathecae spherical . 8 Spermathecae (Fig. 2G) about cylindrical, its length twice its width. . P. songi 8. Epigynum with developed median septum, atrium circular ... P. heteroidea — Epigynum without septum, atrium transverse, slit-like . P. quei 2002 PENG & LI: CHINESE PORTIA 257 Portia heteroidea Xie & Yin, 1991 Portia heteroidea Xie & Yin, 1991: 31, figs. 5-13 (male & female); Peng et al., 1993: 187, figs. 653-659 (male & female); Song, Chen & Zhu, 1997: 1740, figs. 53a-c (male); Song, Zhu & Chen, 1999: 541, figs. 311J, 312E (male & female). Diagnosis. —Embolus of median length, its terminal end extended slightly be¬ yond the retrolateral margin of cymbium in ventral view. Three tibial apophyses, ventral apophyses short, conic; intermediate apophyses smallest and shortest; re¬ trolateral apophyses biggest and longest, bar-like, slightly swollen terminally. Te- gulum furrow procurved arc-like, shallow; no tegular apophysis. In dorsal view, cymbium flange long and robust, its upper base originated from the median por¬ tion of cymbium, its end extended to median front margin of tibial apophysis. Epigynum with large atria, almost circular; median septum developed, posterior margin wider with slight incision; spermathecae big spherical, compulatory duct invisible. Abdomen with 5 yellow-brown circles, posterior 3 circles covered by gray-white hairs. This species is closely related to P. quei Zabka, 1985, but differs in: 1) embolus shorter; 2) retrolateral tibial apophysis shorter with round end, that of P. quei with hook-like end; 3) atria larger and almost circular, that of the latter wide slit-like; 4) epigynum with median septum which is absent in that of the latter. Specimens Examined. —1 female, deposited inIZCAS, data: CHINA, SHAANXI PROVINCE, FUP- ING COUNTY Co.: 33.5° N, 108.0° E, 870-1000 m, 25 Jul 1998, by Chen Jun; 1 male, deposited in IZCAS, data: CHINA, GANSU PROVINCE, WENXIAN COUNTY Co.: 32.9° N, 104.7° E, 900- 1500 m, 25 Jun 1998, by Chen Jun. Distribution .—China (Gansu, Shaanxi, Hunnan, Hubei, Guizhou, Sichuan). Portia jianfeng Song & Zhu, 1998 (Fig. 1) Portia jianfeng Song & Zhu, 1998: 26, figs. 1-3 (male); Song, Zhu & Chen, 1999: 541, figs. 311 K-L (male). Diagnosis .—Embolus belt-like, tapering distally; conductor well developed, en¬ closing the base of embolus in ventral view; tegulum long and diagonal, lower end almost extended to the right bottom of the bulb; tegulum apophysis devel¬ oped, thin and triangular. 4 tibial apophyses: 3 in upper row, ventral one most stout; intermediate one short horn-like, bent ventrally in retrolateral view; retro¬ lateral apophysis finger-like in retrolateral and dorsal views; basal apophysis big¬ gest and very swollen, almost spherical in retrolateral view, and diagonal oblong in dorsal view. Abdomen with 2 gray longitudinal bands and pairs of gray patches. This species is allied to P. songi Tang & Yang, 1997, but can be distinguished from the latter by: 1) embolus much longer and thinner (Figs. IB, 1C, 2B, 2C); 2) conductor well developed (Figs. IB, 1C), enclosing the base of embolus in ventral view (Fig. IB), that of P. songi without conductor (Figs. 2B, 2C, 3) tegulum furrow almost longitudinal (Fig. IB) in ventral view, that of P. songi almost transverse (Fig. 2B); 4) cymbium thinner and longer (Figs. 1B-1D, 2B- 2D); 5) abdominal patterns also quite different (Figs. 1A, 2A). 258 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) Figure 1. Portia jianfeng Song & Zhu, 1998. Figure 1A. Body of male, dorsal view. Figure IB. Left palpal organ, ventral view. Figure 1C. Left palpal organ, retrolateral view. Figure ID. Tibial apophysis, dorsal view. Scale bar: Figure 1A = 1.00 mm, Figure lB-Figure ID = 0.10 mm. 2002 PENG & LI: CHINESE PORTIA 259 Specimens Examined. —2 males, deposited in IZCAS, data: CHINA, HAINAN PROVINCE, LE- DONG COUNTY, JIANFENGLING Co.: 18.7°N, 109.1°E, Apr 1994, by Liao Cong-Hui. Distribution. —China (Hainan). Portia orientalis Murphy & Murphy, 1983 Portia orientalis Murphy & Murphy, 1983: 40, figs. 6, 9, 12, 16, 20 (male). Diagnosis. —Embolus long and thin, narrowing gradually, terminal end extend¬ ed beyond the retrolateral margin of cymbium in ventral view. Tegulum furrow curve shallow and narrow, tegular apophysis indistinct. 3 tibial apophyses, ventral apophysis very short, hook-like in retrolateral view; intermediate apophysis thin, very pale, covered by a tuft of long white hairs; retrolateral apophysis longest, terminal end hook-like in retrolateral view. In dorsal view, cymbium flange long and large, overlapping dorsum of retrolateral tibial apophysis, terminal end be¬ yond the retromargin of tibial apophysis. This species resembles P. assamensis Wanless, 1978, but can be separated from the latter by: 1) retrolateral tibial apoph¬ ysis longer and thinner; 2) cymbial flange stouter and shorter; 3) embolus longer. Specimens Examined. —Type specimen was collected from Hong Kong and deposited in British Museum. No further specimens were collected from China. No specimens were examined in this study. The above information is after Murphy and Murphy (1983). Distribution. —China (Hong Kong). Portia quei Zabka, 1985 Portia quei Zabka, 1985: 438, figs. 497-501 (male); Song, Chen & Gong, 1990: 15, figs. 1—4 (male & female); Chen & Zhang, 1991: 314, figs. 334.1-6 (male & female); Peng et al., 1993: 188, figs. 660-666 (male & female); Song, Zhu & Chen, 1999: 541, figs. 311M-N, 312F-G (male & female). Diagnosis .—Embolus very long and thin, narrowing gradually, more than one third extended beyond the retrolateral margin of cymbium in ventral view; tegular furrow curve very deep, tegular apophysis indistinct. Three tibial apophyses, ven¬ tral apophysis thin and short, hook-like, bent retrolaterally; intermediate apophysis also very short, conical in retrolateral view; retrolateral apophysis long and thin, terminal portion hook-like. Cymbium flange thin, terminal portion overlapping on median portion of retrolateral tibial apophysis. Epigynum with wide slit-like atri¬ um near epigastric groove, no median septum. Spermathecae big and spherical, compulatory duct invisible. This species is closely allied to Portia heteroidea. Differences between them are discussed in the diagnosis of Portia heteroidea. Specimens Examined .—4 males, 6 immatures, deposited in HNU, data: CHINA, YUNNAN PROV¬ INCE, NUJIANG COUNTY, QIQI Co.: 27.'7° N, 98.7° E, 9-14 Jul 2000; 4 females, 3 immatures, deposited in HNU, data: CHINA, YUNNAN PROVINCE, GONGSHAN COUNTY Co.: 21.T N, 98.6° E, 29 Jun 2000; 1 male, deposited in IZCAS, data: CHINA, GUANGXI ZHUANG AUTON¬ OMOUS REGION, JINXIOU COUNTY CO.: 24.1° N, 110.1° E, 490 m, 1 Jul 2000, by Chen Jun; 1 female, deposited in IZCAS, data: CHINA, GUANGXI ZHUANG AUTONOMOUS REGION, JINX¬ IOU COUNTY CO.: 24.1° N, 110.1° E, 1050-1100 m, 2 Jul 2000, by Chen Jun. Distribution. —China (Hunan, Hubei, Guangxi, Sichuan, Guizhou, Yunnan), Viet Nam. 260 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) Portia songi Tang & Yang, 1997 (Fig. 2) Portia songi Tang & Yang, 1997: 353, figs. 1-8 (male & female); Song, Zhu & Chen, 1999: 541, figs. 3110-P, 312H, 313A, 328Q (male & female). Diagnosis. —Embolus short, basal portion large, terminal portion spine-like; tegular furrow slightly diagonal, tegular apophysis short and large; median apoph¬ ysis short, conic. 5 tibial apophyses: ventral apophysis biggest, conic; retrolateral apophysis longest, terminal portion hook-like in ventral and dorsal views; in ven¬ tral view, 3 intermediate apophyses arranged in a line, top one longest and finger¬ like, median one shortest and conic, bottom one biggest and conic. Cymbium flange big and short in retrolateral view. Epigynum longer than wide, transparent, 2 belts looped near epigastric furrow; spermathecae with 2 chambers, length twice its width; compulatory duct invisible. This species is allied to P. jianfeng Song & Zhu, 1998. Differences between them are discussed in the diagnosis of P. jianfeng. Specimens Examined. —1 male, 1 female, deposited in LZU, data: CHINA, GUNSU PROVINCE, WENXIAN COUNTY Co.: 32.9° N, 104.7° E, Jun 1992, by Tang Ying-Qiu. Distribution .—China (Gansu). Portia wui Peng & Li, NEW SPECIES (Fig. 3) Type .—Holotype, male, deposited in IZCAS, data: CHINA, GUANGXI ZHUANG AUTONOMOUS REGION, NAPO COUNTY, PINGMENG TOWN, BEIDOU TOWNSHIP, Co.: 23.4° N, 105.8° E, 500-550m, 10 Apr 1998, By WU Min (No. WM98GXsp.25). Measurements.— Male: TL 6.60, CL 3.00, CW 2.70, AL 3.60, AW 1.60; legs: I 12.50 (3.10, 4.50, 3.40, 1.50), II 10.00 (2.50, 4.00, 2.50, 1.00), III 9.40 (2.50, 3.60, 2.30, 1.00), IV 12.80 (3.20, 4.00, 4.40, 1.20), formula 4, 1, 2, 3. AER 2.10, PER 1.90, AME 0.75, ALE 0.35, PME 0.25, PLE0.30, EFL 1.40, CLYH 0.50. Description. —Male (holotype): Carapace (Fig. 3A) brown; ocular area light brown, base of AME brown, the other eyes surrounded with black bases; fovea black, longitudinal line-shaped; cervical and radial grooves black. Sternum yellow-brown, densely clothed in white and brown hair; margin dark brown with irregular black patches. Clypeus dark gray-brown, clothed in sparse hair; front margin gray-black. Chelicera dark gray-brown, anterior side darker, distal area and furrow margin clothed in gray-brown brush-like hair; furrow with 2 promarginal teeth and 3 retromarginal denticles (Fig. 3E). Endites and labium gray-black, distal area and inner sides clothed in gray-black long hair. Legs gray- brown with lighter annuli; ventral sides of tibiae and patellae clothed in dense brush-like long hair, which on tibia II is much denser and covers three fourth portion of tibia II; hair on the rest of segments very sparse; spines sparse and weak, 3 pairs on ventral sides of tibiae I and II, 2 pairs on ventral sides of metatarsi I and II. Abdomen cylindrical. Dorsum (Fig. 3A) gray-white with gray-black marks; cardiac pattern long bar-shaped, 2 muscular depressions darker and clear. Ventral side gray-black; each anterior side with a gray-white patch; 2 small gray-white circles on posterior median area. Spinnerets black brown. Palpal organ (Figs. 3B-D): embolus short and stout; seminal duct clear and S-shaped; 3 tibial apophyses, ventral one large and short, intermediate one smallest and finger-shaped, retrolateral one biggest and flag-shaped in dorsal view; cymbium flange slender and short; cymbium apophysis stout and horn-shaped. Female. —Unknown. Diagnosis. —The new species resembles Portia heteroidea Xie & Yin, 1991, but differs in: 1) embolus shorter and stouter; 2) retrolateral tibial apophysis much 2002 PENG & LI: CHINESE PORTIA 261 Figure 2. Portia songi Tang & Yang, 1997. Figure 2A. Body of male, dorsal view. Figure 2B. Left palpal organ, ventral view. Figure 2C. Left palpal organ, retrolateral view. Figure 2D. Left palpal organ, dorsal view. Figure 2E. Teeth of left male chelicera: upper—promargin, lower—retromargin. Figure 2F. Epigynum. Figure 2G. Vulva, dorsal view. Scale bar: Fig. 2A = 0.10 mm, Fig. 2B-Fig. 2D = 0.50 mm, Fig. 2F-Fig. 2G = 0.10 mm. 262 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) Figure 3. Portia wui Peng & Li, sp. nov. Figure 3A. Body of male. Figure 3B. Left palpal organ, ventral view. Figure 3C. Left palpal organ, retrolateral view. Figure 3D. Left palpal organ, dorsal view. Figure 3E. Teeth on left chelicera: upper—promargin, lower—retromargin. Scale bar: Fig. 3A = 1.00 mm, Fig. 3B-Fig. 3D = 0.5 mm. 2002 PENG & LI: CHINESE PORTIA 263 0115 120 125 130 135 Figure 4. Distribution of Chinese Species of Portia. A Portia heteroidea, K Portia jianfeng, Portia orientalis, $ Portia quei, ★ Portia songi, ■ Portia wui. bigger, flag-shaped in dorsal view (Fig. 3D), that of the latter bar-shaped; 3) cymbium flange (Fig. 3D) much shorter and more slender; 4) cymbium with a stout horn-shaped apophysis (Figs. 3C, 3D) nearing cymbium flange, which can¬ not be found in any other known species of the genus; 5) abdominal marks much more distinct. Etymology .—The new species is named in honor of Dr. WU Min, who collected the type specimen. Distribution .—China (Guangxi). Acknowledgment We are very grateful to Prof. YIN Changmin (HNU) for her continued support and encouragement during our study on Chinese jumping spiders. Our special thanks should be given to Dr. Wu Min (IZCAS) for his donation of the type of Portia wui Peng & Li, sp. nov., and to Prof. Tang Yingqiu (LZU) for supplying materials used in this study. The present study was supported by the National Natural Sciences Foundation of China to S. Li (Grant No. 39970102 and 30270183), and, in part, by the Special Support Project of the Department of Biology, Chinese Academy of Sciences (CAS) (STZ-00-19) and CAS Innovation Program. Literature Cited Chen, Z. F. & Z. H. Zhang. 1991. Fauna of Zhejiang: Araneida. Zhejiang Science and Technology Publishing House, Hangzhou, 356 pp. Karsch, E. 1878. Exotisch-araneologisches. Zeitschr. ges. Naturw, 51: 332-333, 771-826. 264 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) Hogg, H. R. 1915. On spiders of the family Salticidae collected by the British Ornitologists’ Union Expedition and the Wollaston Expedition in Dutch New Guinea. Proc. Zool. Soc. Lond, 1915: 501-528. Murphy, J. & F. Murphy. 1983. More about Portia (Araneae: Salticidae). Bull. Br. Arachnol. Soc., 6: 37-45. Peckham, G. W. & E. G. Peckham. 1885. Genera of the family Attidae: with a partial synonymy. Trans. Wis. Acad. Sci. Arts Lett., 6: 255-342. Peng, X. J., L. P. Xie, X. Q. Xiao & C. M. Yin. 1993. Salticids in China (Arachnida: Aranea). Hunan Normal University Press, Changsha, 270 pp. Roewer, C. F. 1965. Die Lyssomanidae und Salticidae-Pluridentati der Athiopischen Region (Araneae). Annls Mus. r. Afr. Cent. (Sci. Zool.), 139: 1-86. Song, D. X., Z. Q. Chen & L. S. Gong. 1990. Description of the female spider of the species Portia quei Zabka (Salticidae). Sichuan J. Zool., 9 (1): 15-16. Song, D. X., J. Chen & M. S. Zhu. 1997. Arachnida: Araneae. In Yang, X. K. (ed.). Insects of the Three Gorge Reservoir area of Yangtze River. Chongqing Publ. House, 2: 1704-1743. Song, D. X. & M. S. Zhu. 1998. Two new species of the family Salticidae (Araneae) from China. Acta Arachnol. Sin., 7: 26-29. Song, D. X., M. S. Zhu & J. Chen. 1999. The spiders of China. Hebei Sci. Technol. Publ. House, Shijiazhuang, 640 pp. Tang, Y. Q. & Y. T. Yang. 1997. A new species of the genus Portia from China (Araneae: Salticidae). Acta Zootaxon. Sin., 22: 353-355. Thorell, T. 1878. Studi sui ragni. Malesi e Papuanti. II. Ragni di Amboina raccolti Prof. O. Beccari. Ann. Mus. civ. stor. nat. Genova, 13: 1-317. Wanless, F. R., 1978. A revision of the spider genus Portia (Araneae: Salticidae) Bull. Br. Mus. Nat. Hist. (Zool.), 34 (3): 83-124. Xie, L. P. & C. M. Yin. 1991. Two new species of Salticidae from China (Arachnida: Araneae). Acta Zootaxon. Sin., 16: 30-34. Zabka, M. 1985. Systematic and zoogeographic study on the family Salticidae (Araneae) from Viet Nam. Ann. Zooll. Warsz., 39 (44): 1-465. Received 20 February 2002; Accepted 4 November 2002. PAN-PACIFIC ENTOMOLOGIST 78(4): 265-275, (2002) NEW GENERA AND NEW SPECIES OF NEOTROPICAL NEMATOPODINI (HEMIPTERA: HETEROPTERA: COREIDAE: COREINAE) Harry Brailovsky and Ernesto Barrera Departmento de Zoologfa, Instituto de Biologfa, Universidad Nacional Autonoma de Mexico, Apdo Postal 70153 Mexico D. F. 04510, Mexico e-mail: coreidae@ servidor.unam.mx Abstract .—Two new genera (Nectoquintius and Stenoquintius ) and three new species ( Necto- quintius alajuelensis, Stenoquintius matogrossensis, and Stenoquintius reclusa ) from Brasil, Cos¬ ta Rica, Ecuador and Venezuela are described in the tribe Nematopodini (Coreidae), and com¬ pared with the related genera Grammopoecilus Stal, Nematopus Berthold, Quintius Stal, and Saguntus Stal. Dorsal habitus illustrations and drawings of antennae, hind legs, and male genital capsule are provided. Key Words. —Insecta, Hemiptera, Heteroptera, Coreidae, Nematopodini, new genera, new spe¬ cies, neotropical region. The Nematopodini Amyot and Serville a New World tribe of the coreid sub¬ family Coreinae, is large and diverse. Members of this tribe are extremely abun¬ dant in the neotropics and despite the diversity of the fauna, many taxa remain undescribed. The twenty genera and one subgenus recognized in this tribe, have been revised recently by O’Shea (1980) and Brailovsky (1986, 1987, 1995). The Nematopodini are characterized by the hind femur ventrally armed, and usually strongly incrassate especially in males, tibiae sulcate, hind tibiae unarmed at apex; tylus projecting slightly beyond juga, antenniferous tubercles unarmed, occupying most of anterior head, ocellar tubercles small; metathoracic peritreme with two completely separated lobes and area between them depressed, and ab¬ dominal sterna unarmed in both sexes (O’Shea 1980, Packauskas 1994). In the present paper we describe two new genera and three new species from Brasil, Costa Rica, Ecuador and Venezuela. All measurements are in millimeters. Nectoquintius Brailovsky and Barrera, NEW GENUS Type species. — Nectoquintius alajuelensis Brailovsky and Barrera, NEW SPECIES. Description .—Body medium sized, relatively narrow and elongate. Head: Wider than long (across eyes), pentagonal, and declivant anteriorly; tylus unarmed, apically globose, raised, extending anteri¬ orly to and laterally higher than juga; juga unarmed, laterally expanded and thickened; antenniferous tubercle broad, widely separated, diverging anteriorly and unarmed; antennal segment I thicker than succeeding segments, and slightly curved outward; segments II and III, cylindrical and slender; seg¬ ment IV fusiform; antennal segment IV the longest, III the shortest, and I longer than II (Fig. 1); preocellar pit deep; ocellar tubercle small; eyes hemiespherical, prominent; postocular tubercle mod¬ erately protuberant; buccula rounded, short, raised, not projecting beyond antenniferous tubercle, with¬ out teeth, and closed posteriorly; tip of rostrum reaching middle third of mesosternum; genae and mandibular plate unarmed. Thorax. Pronotum: Wider than long, trapeziform, shallowly declivant; collar wide; frontal angles rounded; anterolateral borders obliquely straight, entire; humeral angles obtusely rounded; posterolateral borders sinuate; posterior border straight; triangular process narrow, apically subacute; calli transverse and conspicuously raised, and uniformly tuberculate. Anterior lobe 266 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) Figures 1-5. Antennae. Figure 1. Nectoquintius alajuelensis, new genus, new species. Figure 2. Quintius scenicum Brailovsky and Barrera. Figure 3. Stenoquintius reclusa new genus, new species. Figure 4. Quintius dentifer Stal. Figure 5. Saguntus pallens (Walker). Figures 6-9. Male genital cap¬ sule. Figure 6. Saguntus pallens (Walker). Figure 7. Stenoquintius reclusa new genus, new species. Figure 8. Nectoquintius alajuelensis new genus, new species. Figure 9. Quintius dentifer Stal. 2002 BRAILOVSKY & BARRERA: NEW NEOTROPICAL NEMATOPODINI 267 of metathoracic peritreme reniform, weakly elevated, posterior lobe sharp, small; mesosternum lacking longitudinal furrow. Legs: Femora not strongly incrassate, and ventrally armed with one subapical small teeth; tibiae unarmed, cylindrical, and sulcate; hind tibiae longer than hind femur; basal segment of hind tarsi longer than total length of middle and hind segment together (Fig. 10). Scutellum: Triangular, flat, longer than wide, with apex subacute. Hemelytra: Macropterous, reaching or extending beyond the apex of last abdominal segment; costal margin emarginate; apical margin almost obliquely straight. Abdomen: Lateral margins parallel; posterior angle of connexivum extending into short and acute spine; abdominal spiracle clearly elliptic, closed to anterior margin; abdominal sterna lacking medial furrow. Integument: Body surface shining; pronotum, scutellum, clavus, corium, propleura, posterior third of mesopleura and metapleura, acetabulae, and male genital capsule punctate; head, apex of scutellum, connexivum, prosternum, mesosternum and metasternum, anterior third of meso¬ pleura and metapleura, abdominal sterna, and female genital plates impunctate; scutellum transversely striate; dorsal surface glabrous; ventrally with few long bristle-like setae located into the sternal surface of thorax, and on the abdominal sterna; pubescence of antennal segments and tibiae short, mainly appressed, on rostral segments II to IV, femora, and tarsi longer, suberect to erect and rather dense; calli densely tuberculate. Male Genitalia .—Genital capsule broadly ovoid; posteroventral edge with broad tooth-like projec¬ tion at middle third, laterally deeply concave, and with lateral angles exposed, and subtruncated (Fig. 8). Female Genitalia .—Abdominal sternite VII with plica and fissura; the former curved, reduced, and transversely straight, the fissura with inner margin overlapping; gonocoxae I triangular, closed in caudal view, and with upper border rounded; paratergite VIII subtriangular with spiracle visible; paratergite IX squarish, and larged than paratergite VIII. Discussion .—This genus runs to O’Shea (1980) key at couplet 7 and its related particularly with Quintius Stal and Saguntus Stal. The relatively narrow and elongate body, the rounded humeral angles, the clear¬ ly elliptical abdominal spiracles close to the anterior margin, the ventrally armed femora, the cylindrical tibiae that are never dilated, the longer than wide scutel¬ lum, and the mesosternum lacking a longitudinal furrow suggest a relationship with Quintius. In Nectoquintius, the calli are conspicuous, transversely raised and uniformly tuberculate, the antennal segment III is slender (Fig. 1), cylindrical and longer than 1.70 mm, the posterocular tubercle is moderately protuberant, the hind femur is not strongly incrassate with only one subapical tooth, the hind tibia is longer than hind femur (Fig. 10), and the posterior angle of connexivum extends into a short, acute spine. In Quintius the calli are flat or barely convex and smooth; the antennal segment III is broad (Fig. 4) and shorter than 1.40 mm; the postocular tubercle is not visible, forming smooth curve with eye; the hind femur is incras¬ sate in both sexes (Figs. 10-12), especially in males, ventrally armed with two rows of spines running from middle third to subapical third, and is longer than hind tibia; and the posterior angles of each connexival segment unarmed. Saguntus is similar to Nectoquintius in having the body relatively narrow and elongate, antennal segment III slender, cylindrical and longer than 1.70 mm (Fig. 5), the humeral angles rounded, the posterior angle of each connexival segment extending into short and acute spine, the scutellum longer than wide, and the hind tibiae never dilated. In Nectoquintius the triangular processes of the posterior margin of pronotum are narrow and apically subacute, the postocular tubercle is moderately protuberant, the mesosternum lacks a longitudinal median furrow, the calli are transversely raised, the hind femur is not strongly incrassate and has only one subapical tooth, the hind tibiae is longer than hind femur, the male hind tibiae is cylindrical and unarmed (Fig. 10), and the abdominal spiracles are elliptic and 268 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) Figures 10-14. Hind leg. Figure 10. Nectoquintius alajuelensis new genus, new species (d). Figure 11. Quintius dentifer Stal (d). Figure 12. Quintius scenicum Brailovsky and Barrera (9). Figure 13. Saguntus pallens (Walker) (d). Figure 14. Stenoquintius reclusa new genus, new species (d). 2002 BRAILOVSKY & BARRERA: NEW NEOTROPICAL NEMATOPODINI 269 near to the anterior margin. In Saguntus the triangular processes are absent; the postocular tubercle is not visible; the mesosternum has a shallow, median longi¬ tudinal furrow; the calli are flat and smooth; the hind femur is incrassate in both sexes, especially in males, ventrally armed with two rows of spines, and longer than hind tibia; the male hind tibiae are curved and armed with one large, ventral spine at midpoint (Fig. 13); and the abdominal spiracles are circular and near to anterior margin. In caudal view the male genital capsule of Nectoquintius, Quintius, and Sa¬ guntus are remarkably different (Figs. 6, 8, 9). In Grammopoecilus Stal the abdomen is tapered inward from the base of pron- otum to the apex of abdomen, and the hind tibiae of male are armed distally with ventral and dorsal spines, which are absent in the new genus. In Nematopus Berthold, the lateral margins of the abdomen are more or less parallel, the humeral angles are sharp, and the hind femur of male is markedly incrassate and armed with a large curved spine at midpoint of ventral surface, which is absent in the new genus. Distribution. —Known from Costa Rica and Ecuador. Etymology. —Masculine: From the Latin “ necto ” (knit) plus the generic name Quintius, denoting the relationship between these genera. Nectoquintius alajuelensis Brailovsky and Barrera, NEW SPECIES (Figs. 1, 8, 10, 15) Types. —Holotype male: Costa Rica. Puntarenas Province, Peninsula de Osa, Rancho Quemado, 200 m, Die 1992, F. Quesada. Deposited in Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica. Paratypes: 1 female; data: same locality and date as holotype. Deposited in Instituto Nacional de Bio¬ diversidad, Santo Domingo de Heredia, Costa Rica. 1 female: Costa Rica. Alajuela Province, Sector San Ramon de Dos Rios, 620 m, 18 Mar 13 Apr 1995, F. A. Quesada. Deposited in Coleccion Entomologica del Instituto de Biologia, UNAM. 1 male: Costa Rica. Puntarenas Province, Peninsula de Osa, Fila Guerra, 1—100 m, Mar 1991, J. Quesada. Deposited in Coleccion Entomologica, del Instituto de Biologia, UNAM. 2 females: Costa Rica. Puntarenas Province, Parque Nacional Corcovado, Estacion Sirena, 0-100 m, Nov 1990 C. Saborio, and Die 1992, G. Fonseca. Deposited in Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica. 1 male: Costa Rica. Province Alajuela, 20 km S Upala, 10— 29 May 1991, F. D. Parker. Deposited in Department of Biology of the Utah State University. 1 female: Costa Rica. San Carlos, collection Schild-Burgdorf. Depos¬ ited in the Zoological Department of the Hungarian Natural History Museum. 1 male: Ecuador. Esmeralda Province, Zapallo Grande, 25-30 Oct 1987, M. Huy- bensz. Deposited in Museum of Comparative Zoology, Harvard University. Description.—Male (holotype). Dorsal coloration: Head yellow tinged with chestnut in front of ocelli; ocellar tubercle brownish; antennal segments I to III bright orange, and IV yellow; pronotal disc bright chestnut orange with collar, frontal angles, posterolateral borders, and middle third of posterior border yellow; calli with yellow and brown marks; anterolateral margins, posterolateral mar¬ gins (except the border), and posterior border (except middle third) black; scutellum yellow with lateral margins dark brown; clavus and corium dark brown to black with following areas yellow: claval vein, claval comissure, inner corial vein, costal margin and apical margin; hemelytral membrane dark am- barine, with basal angle darker; connexival segments I to V yellow, VI and VII black with anterior third yellow; dorsal abdominal segments dark brown to reddish brown with scars IV-V and V-VI 270 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) Figure 15. Dorsal view of Nectoquintius alajuelensis new genus, new species. yellow. Ventral coloration: Including rostral segments and legs yellow; apex of rostral segment IV, and caudal surface of genital capsule dark brown; mesopleura and metapleura with narrow and elongate creamy yellow hardened protuberance; metasternum yellow and tinged with orange. Female. —Coloration: Similar to male holotype. Connexival segments VIII and IX dark brown with anterior angle yellow; dorsal abdominal segments VIII and IX dark brown; genital plates yellow. Variation. —1, Anterolateral margins of pronotum bright chestnut orange. 2, calli almost entirely yellow. 3, Connexival segment VI with upper margin yellow and inner margin dark brown to reddish brown. 4, Metasternum orange. 5, Pleural abdominal sterna VI and VII yellow with posterior third dark brown. Measurements .—Male (female). Head length: 1.64 mm (1.84 mm); width across eyes: 2.28 mm (2.32 mm); interocular space: 1.08 mm (1.12 mm); preocular distance: 1.00 mm (1.02 mm); antennal segments lengths: I, 4.20 mm (3.80 mm); II, 3.96 mm (3.76 mm); III, 1.96 mm (1.84 mm); IV 4.76 mm (4.64 mm). Pronotal length: 3.00 mm (3.40 mm); width across frontal angles: 2.64 mm (2.88 mm); width across humeral angles: 4.16 mm (4.80 mm). Maximum length of hind femur: 6.30 mm 2002 BRAILOVSKY & BARRERA: NEW NEOTROPICAL NEMATOPODINI 271 (6.10 mm); maximum length of hind tibiae: 6.60 mm (6.60 mm). Scutellar length: 2.24 mm (2.48 mm); width: 1.84 mm (2.08 mm). Total body length: 15.77 mm (17.75 mm). Etymology .—The name refers to the Alajuela Province of Costa Rica. Stentoquintius Brailovsky and Barrera, NEW GENUS Type species. — Stenoquintius matogrossensis Brailovsky and Barrera, NEW SPECIES. Description .—Body medium sized, relatively narrow and elongate. Head: Wider than long, pentag¬ onal, and declivant anteriorly; tylus unarmed, apically globose, raised, extending anterior to and lat¬ erally higher than juga; juga unarmed, short, and thickened; antenniferous tubercles broad, widely separated, diverging anterirorly and unarmed; antennal segment I thicker than succeeding segments, and slightly curving; segments II and III cylindrical, and slender; segment IV fusiform; antennal segment IV the longest, III the shortest, and II longer than I (Fig. 3); preocellar pit deep; ocellar tubercle small; eyes hemispherical, prominent; postocular tubercle absent; buccula rounded, short, raised, not projecting beyond antenniferous tubercles, without teeth, and closed posteriorly; rostrum reaching anterior third of mesosternum; genae and mandibular plate unarmed. Thorax. Pronotum: Wider than long, trapeziform, shallowly declivant; collar wide; frontal angles rounded, not exposed; anterolateral borders obliquely straight, weakly nodulose; humeral angles produced laterally into short angulate spine; posterolateral borders sinuate, with outer third nodulose and inner third smooth; pos¬ terior border straight; triangular process absent; calli flat to weakly convex, separated along midline by two short longitudinal depressions. Anterior lobe of metathoracic peritreme elongate, reniform, posterior lobe rounded; mesosternum with median and deep sulcus in anterior and posterior third, and faint longitudinal furrow hard to see. Legs: Fore and middle femora not incrassate, ventrally with two rows of short and acute spines; hind femur slightly incrassate (much more in males), and ventrally with two rows of broad and acute spines; fore and middle tibiae unarmed, cylindrical and sulcate; hind tibiae of male longer than hind femur, weakly curved, flattened, armed with large ventral spine close to midpoint, and smaller spines along ventral surface; hind tibiae of female longer or shorter than hind femur, cylindrical, and unarmed (Fig. 14); basal segment of hind tarsi longer than total length of middle and hind segment together. Scutellum: Triangular, flat, longer than wide, with apex subacute. Hemelytra: Macropterous, reaching or extending beyond the apex of the last abdominal segment; costal margin emarginate; apical margin slightly sinuate. Abdomen: Lateral margins parallel; posterior angle of connexivum extending into short and acute pines; abdominal spiracle elliptic, closed to anterior margin; abdominal sterna lacking medial furrow. Integument: Body surface dull, and gla¬ brous; pronotum, clavus, corium, propleura, posterior margin of mesopleura and metapleura, aceta- bulae, and male genital capsule densely punctate; head, calli, apex of scutellum, connexivum, pro¬ sternum, mesosternum, and metasternum, anterior third of mesopleura and metapleura, abdominal sterna and female genital plates impunctate; pubescence of antennal segments, and legs, short, mainly appressed; scutellum transversely striate. Male Genitalia .—Genital capsule broadly ovoid; posteroventral edge transversely tuberculate or sinuate, with deep circular concavity at midpoint (Fig. 7). Female Genitalia .—Abdominal sternite VII with plica and fissura; the former curved, reduced, and transversely straight, the latter with inner margin overlapping; gonocoxae I triangular, closed in caudal view, and with upper border rounded; paratergite VIII subtriangular, with spiracle visible; paratergite IX squarish, and larged than paratergite VIII. Diagnosis .—The relatively narrow and elongate body, the abdominal spiracle elliptic and close to the anterior margin, the ventrally armed femora, the cylin¬ drical tibiae that are never dilated, the longer than wide scutellum, and the me¬ sosternum lacking a longitudinal medial furrow suggests a relationship with Nec- toquintius described in this paper, and Quintius Stal. In Stenoquintius, the antennal segment II is longer than I (Fig. 3), the postocular tubercle and the triangular processes of pronotum are absent, the calli are flat or barely convex, the mesosternum has a deep median, sulcus at anterior and pos¬ terior third, the femora are ventrally armed with two rows of spines, and hind 272 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) tibiae of male weakly and curved, flattened, and ventrally armed (Figs. 14, 16). In Nectoquintius, the antennal segment I is longer than II (Fig. 1), the postocular tubercle is moderately protuberant, the posterior margin of pronotum has the tri¬ angular processes narrow, and apically subacute, the are calli transverse and con¬ spicuously raised, the mesostemum lacks an anterior and posterior sulcus, each femur is armed with one subapical small tooth, and hind tibiae of male are cy¬ lindrical, slender, and unarmed (Fig. 10). In Quintius, like Stenoquintius, the post¬ ocular tubercle is not visible, the calli are flat or barely convex, and the male hind tibiae are ventrally armed (Figs. 11-12, 14), both the antennal segment III is broad, and shorter than 1.40 mm (Figs. 2, 4), the hind femora, particularly in males, are conspicuously incrassate, the posterior angle of connexival segments are unarmed, and the mesosternum lacks an anterior or posterior sulcus at middle third. Distribution .—Known from Venezuela and Brazil. Etymology. —Masculine: From the greek “ stenos ” (narrow), plus the generic name Quintius, denoting the relation between both genera. Stentoquintius matogrossensis Brailovsky and Barrera, NEW SPECIES (Fig. 16) Types. —Holotype male: Brasil. Mato Grosso, Sinop, Oct 1976, M. Alvarenga. Deposited in American Museum of Natural History, New York. Paratypes: 2 males, 1 female; data: same locality and date as holotype. Deposited in American Museum of Natural History, New York, and Coleccion Entomologica del Instituto de Biologia, UNAM. Description.—Male (holotype). Dorsal coloration: Head pale yellow; antennal segments dark yellow, tinged with green reflections; pronotum yellow, with green reflections, and dark brown punctures at humeral angles, posterolateral margins and posterior margin; scutellum yellow with lateral margins pale orange; clavus and corium yellow with punctures dark brown to chestnut orange; hemelytral membrane ambarine with basal angle darker; connexival segments yellow and VII with upper margin dark brown, basal and apical angle yellow, and inner margin reddish orange; dorsal abdominal seg¬ ments reddish orange with wide yellow longitudinal stripe running at middle third from I to VI segment. Ventral coloration: Head, prosternum, mesosternum, and metasternum, and abdominal sterna pale yellow; rostral segments (apex of IV dark brown), propleura, mesopleura, and metapleura, ace- tabulae, legs, pleural margin of abdominal sterna and genital capsule dark yellow, tinged with green reflections, and scattered with red to pink tiny spots; mesopleura and metapleura with wide and broad creamy yellow hardened protuberance. Genitalia .—Genital capsule. Posteroventral edge transversely sinuate, with deep circular concavity at midpoint. Female. —Coloration: Similar to the male holotype. Clavus and corium yellow, densely tinged with pink, and with punctures dark brown to chestnut orange; connexival segments I to VI yellow with upper margin tinged with pale brown marks, segment VII like male, and segments VIII and IX yellow with lateral margins brown; propleura, mesopleura, and metapleura with elongate and continuous creamy yellow hardened protuberance; abdominal sterna and genital plates yellow with pleural margins III to VII dirty chestnut brown, scattered with tiny red to pink spots. Measurements .—Male (female). Head length: 1.28 mm (1.38 mm); width across eyes: 1.96 mm (2.04 mm); interocular space: 0.94 mm (1.00 mm); preocular distance: 0.82 mm (0.96 mm); antennal segments lengths: I, 3.68 mm (3.24 mm); II, 3.84 mm (3.28 mm); III, 2.52 mm (2.24 mm); IV 4.48 mm (4.08 mm). Pronotal length: 3.00 mm (3.60 mm); width across frontal angles: 2.12 mm (2.28 mm); width across humeral angles: 4.00 mm (4.60 mm). Maximum length of hind femur: 5.90 mm (5.80 mm); maximum length of hind tibiae: 6.30 mm (6.00 mm). Scutellar length: 1.84 mm (2.16 mm); width: 1.60 mm (1.92 mm). Total body length: 14.57 mm (15.68 mm). 2002 BRAILOVSKY & BARRERA: NEW NEOTROPICAL NEMATOPODINI 273 Figure 16. Dorsal view of Stenoquintius matogrossensis new genus, new species. Etymology .—The name is a noun in apposition, referring to the State of Mato Grosso State in Brasil, source of the type series. Stentoquintius reclusa Brailovsky and Barrera, NEW SPECIES (Figs. 3, 7, 14) Types .—Holotype male: Venezuela. Territorio Federal Amazonas, San Carlos de Rio Negro, 10 Die 1984, R. Brown. Deposited in Universidad Central de Venezuela, Facultad de Agronomla, Maracay. Paratypes: 3 males; data: same lo¬ cality and date as holotype. Deposited in Universidad Central de Venezuela, Fa¬ cultad de Agronomla, Maracay, and Coleccion Entomologica, Instituto de Biol- ogla, UNAM. 1 male: Venezuela. Territorio Federal Amazonas, San Carlos de Rio Negro, 1 56' N-67 03' W, 6—12 Die 1984, R.L. Brown. Deposited in Mis¬ sissippi Entomological Museum, Mississippi State. 274 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) Description.—Male (holotype). Dorsal coloration: Head pale yellow; antennal segments yellow, tinged with green reflections; pronotum yellow with green reflections and scattered with reddish brown punctures at humeral angles, posterolateral margins and posterior margin; scutellum yellow with lateral margin orange; clavus yellow, tinged with brown, and with the punctures reddish brown; corium yellow with reddish brown punctures; hemelytral membrane ambarine with basal angle darker; con- nexivum yellow and segment VII with upper border yellow, and inner margin dark brown; dorsal abdominal segments dark brown with yellow longitudinal stripe running at midpoint from I to VI segment. Ventral coloration: Including rostral segments (apex of IV dark brown) and legs yellow, tinged with gree reflections; midpoint of mesosternum and metasternum pale yellow with orange reflections; mesopleura and metapleura with broad creamy yellow hardened protuberance; femora, abdominal sterna, and genital capsule pale yellow. Genitalia .—Genital capsule. Posteroventral edge transversely tuberculate, with deep circular con¬ cavity at midpoint (Fig. ??). Female. —Unknown. Measurements. — Male. Head length: 1.48 mm; width across eyes: 1.84 mm; interocular space: 0.94 mm; preocular distance: 0.94 mm; antennal segments lengths: I, 2.96 mm; II, 2.92 mm; III, 1.88 mm; IV 4.00 mm. Pronotal length: 3.24 mm; width across frontal angles: 1.64 mm; width across humeral angles: 4.08 mm. Maximum length of hind femur: 5.30 mm; maximum length of hind tibiae: 5.20 mm. Scutellar length: 1.88 mm; width: 1.60 mm. Total body length: 14.20 mm. Discussion.—Stenoquintius reclusa can be easily distinguished from S. mato- grossensis by the proportions of antennal segments I to IV which are conspicu¬ ously shorter (see measurements), the hind femur more incrassate, the hind tibia shorter than hind femur, and by the structure of the posteroventral edge of male genital capsule. Etymology .—The name “ reclusa ” refers to the secretive habits of this species, which is hard to found on the revised collections. Acknowledgments We thank the following colleagues and institutions for the loan of specimens and other assistance relevant to this study: Wilford J. Hanson (Utah State Uni¬ versity, Utah, U.S.A.), Terence Lee Schiefer (Mississippi Entomological Museum, Mississippi State University, State College, Mississippi), Eduardo Osuna (Univ- ersidad Central de Venezuela, Facultad de Agronomia, Maracay), Thomas Pape and Bert Viklund (Naturhistoriska Riksmuseet, Stockholm, Sweden), Stephen Pratt (Museum of Comparative Zoology, Harvard University, Cambridge, Mas¬ sachusetts); Randall T. Schuh (American Museum of Natural History, New York, New York), Jesus Ugalde (Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica), Tamas Vasarhelyi (Hungarian Natural History Museum, Budapest, Hungary), Mick Webb (The Natural History Museum, London, United Kingdom). Special thanks to Albino Luna and Cristina Urbina for the preparation of dorsal view illustrations. Literature Cited Brailovsky, H. 1987. Three new genera and six new species of Neotropical Coreidae (Heteroptera). J. New York Entomol. Soc., 95: 518-530. Brailovsky, H. 1995. New genera and new species of Neotropical Coreidae (Hemiptera: Heteroptera). Pan-Pacific Entomologist, 71: 217-226. Brailovsky, H. y E. Barrera. 1986 (1985). El genero Quintius Stal con descripcion de un subgenero nuevo y tres especies nuevas (Hemiptera-Heteroptera-Coreidae-Nematopodini). An. Inst. Biol. Univ. Nac. Auton. Mexico, Ser. Zool. 56: 437-452. 2002 BRAILOVSKY & BARRERA: NEW NEOTROPICAL NEMATOPODINI 275 O’Shea, R. 1980. A generic revision of the Nematopodini (Heteroptera: Coreidae: Coreinae). Stud. Neotropical Fauna Envir., 15: 197-225. Packauskas, R. J. 1994. Key to the subfamilies and tribes of the New World Coreidae (Hemiptera), with a checklist of published keys to genera and species. Proc. Entomol. Soc. Wash., 96: 44-53. Received 9 January 2002; Accepted 4 November 2002. PAN-PACIFIC ENTOMOLOGIST 78(4): 276-285, (2002) THREE NEW SPECIES OF SUNDARUS AMYOT & SERVILLE, AND KEY TO THE KNOWN SPECIES (HEMIPTERA: HETEROPTERA: COREIDAE: COREINAE: COREINI) Harry Brailovsky Departamento de Zoologfa, Instituto de Biologfa, Universidad Nacional Autonoma de Mexico, Apdo Postal 70153, Mexico D. F., 04510, Mexico Abstract .—Three new species of Sundarus Amyot and Serville from Brasil, Panama, and Peru are described and illustrated, and a key to the known species of the genus is included. Key Words. —Insecta, Hemiptera, Heteroptera, Coreidae, Coreini, Sundarus, new species, Brasil, Panama, Peru. Brailovsky (1988) revised the genus Sundarus Amyot and Serville, described 13 new species, and discussed the taxonomical importance of the humeral angles and sculpture of the pronotum, the shape of parameres and spermatheca and the general color of the body, including the distributional pattern of the metallic zone. In the same contribution he included a historical review of each species, added new records for the majority of the known species, and gave a key to the known taxa (except to S. muggei Schmidt). Previously, 28 species of Sundarus were known. In this contibution, three new species collected in Brasil, Panama, and Peru are described, and a revisioned key to the known species is included (except for S. muggei Schmidt). Sundarus occua Brailovsky, NEW SPECIES (Figs. 2 and 8) Types .—Holotype female: Panama. Canal Zone, Barro Colorado Isl., Jun 1939, J. Zetek. Deposited in the Coleccion Entomologica del Instituto de Biologfa, UNAM, Mexico. Description .—Female (holotype). Dorsal coloration: Head, pronotum, scutellum, clavus, and corium entirely bright orange; antennal segments I to III reddish brown with blue-green metallic reflections and segment IV reddish brown; hemelytral membrane black; connexivum black with upper margin yellow; dorsal abdominal segments black. Ventral coloration: Head bright orange; rostral segments reddish brown; prosternum, mesosternum, metasternum, anterior and posterior lobe of metathoracic peritreme and adjacent areas black; propleura metallic green with upper margin bright orange; me- sopleura and metapleura metallic green with upper border bright orange; legs reddish brown with blue- green metallic reflections; abdominal sterna and genital plates metallic green with pleural margin and posterior border of sterna V to VII yellow. Structure: Head: Rostrum reaching anterior border of metasternum. Pronotum: Anterolateral margins irregularly crenate; humeral angles broad, wider than long, exposed, raised, medium sized, hemispheric, directed upward, and each border crenate; postero¬ lateral and posterior border straigt and entire; calli transversely raised (Figs. 2 and 8). Male. —Unknown. Measurements. —Female. Head length: 1.56 mm; width across eyes: 2.40 mm; interocular space: 1.32 mm; interocellar space: 0.60 mm; antennal segments lengths: I, 4.08 mm; II, 3.36 mm; III, 3.68 mm; IV, 5.08 mm. Pronotal length: 3.80 mm; maximum width of anterior lobe: 3.20 mm; maximum width of posterior lobe: 6.60 mm; maximum length of humeral angle: 1.52 mm; maximum width of humeral angle: 2.56 mm. Scutellar length: 2.20 mm; width: 2.32 mm. Total body length: 19.28 mm. Discussion.—Sundarus occua appears to be closely related to S. ambarinus 2002 BRAILOVSKY: THREE NEW SUNDARUS SPECIES 277 Figures 1-6. Pronotum of Sundarus spp., in dorsal view. 1. S. ambarinus Brailovsky. 2. S. occua Brailovsky, NEW SPECIES. 3. S. rahmus Brailovsky, NEW SPECIES. 4. S. sheilae Brailovsky. 5. S. lugens Horvath. 6. S. xenia Brailovsky, NEW SPECIES. Brailovsky on the basis of the color of pronotum, clavus and corium which are entirely bright orange ( S. occua ) or entirely yellow ( S. ambarinus ). In the other known species of Sundarus each region displays extensive black areas. The most significant difference between the two species lies in the greater ex¬ pansion of the humeral angles in S. ambarinus, where that structure in lateral view almost covers the middle third of the head (Figs. 1 and 7), and by having the thorax entirely yellow. In S. occua the humeral expansions are shorter, in lateral view not extending beyond the anterior lobe of the pronotum (Figs. 2 and 8), and the thorax 278 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) Figures 7-12. Pronotum of Sundarus spp., in lateral view. 7. S. ambarinus Brailovsky. 8. S. occua Brailovsky, NEW SPECIES. 9. S. sheilae Brailovsky. 10. S. lugens Horvath. 11.5. rahmus Brailovsky, NEW SPECIES. 12. 5. xenia Brailovsky, NEW SPECIES. is metallic green with prosternum, mesosternum, metastemum and metathoracic peritreme black, with upper margin of pleural region bright orange. Etymology .—The species name is an arbitrary combination of letters and is to be treated as a noun. Sundarus rahmus Brailovsky, NEW SPECIES (Figs. 3, 11, 14) Types .—Holotype male: Peru. San Martin, 6 24' S—76 48' W, 4 Jul 1925, D. Melin. Deposited in Universitets Zoologiska Institut, Uppsala, Sweden. Paratypes: 2002 BRAILOVSKY: THREE NEW SUNDARUS SPECIES 279 1 male, 1 female; data: same as holotype. Deposited in Universitets Zoologiska Institut, Uppsala, Sweden and in Coleccion Entomologica del Instituto de Biol- ogia, UNAM, Mexico. Description .—Male (holotype). Dorsal coloration: Head black with purple metallic reflections; apex of tylus and ocellar tubercle bright to dark orange; inner angle of antenniferous tubercle yellow; antennal segment I red brick, segments II—III reddish brown with blue-green reflections, and segment IV reddish brown; pronotum black with anterolateral margins, anterior third of humeral angles, and short longitudinal stripe running between calli bright orange; collar and calli with blue-green metallic reflections; scutellum bright orange; clavus black with claval commissure creamy yellow; corium black with apical margin creamy yellow and costal border dark orange brown; hemelytral membrane black; connexivum black with upper margin yellow; dorsal abdominal segments black. Ventral coloration: Head reddish brown to black with buccula yellow with bright orange reflections; rostral segment I reddish brown with metallic green reflections, segments II to IV dark reddish brown; propleura black with purple and green metallic reflections, and with upper margin at anterior and middle third bright orange; mesopleura and metapleura black with purple and green metallic reflections; prosternum black; mesosternum and metasternum black with wide longitudinal stripe dark orange on midline; anterior lobe of metathoracic peritreme and adjacent areas black, posterior lobe bright orange; legs brick red; trochanters and femora with blue-green metallic reflections; middle third of abdomen black to dark reddish brown and laterally with three longitudinal rows of colors, each one clearly separated, the inner one metallic green, next metallic purple and the outer included pleural margin yellow; genital capsule black with green metallic reflections, and with posteroventral edge bright orange. Structure: Rostrum reaching posterior margin of metasternum. Pronotum: Anterolateral margins irregularly cre- nate; humeral angles broad, wider than long, conspicuously exposed, raised, large sized, hemispheric, directed upward, each border crenate, and in lateral view almost reaching the middle third of head (Figs. 3 and 11); posterolateral and posteror border straight, entire; calli transversely flat or slightly raised. Genital capsule: Posteroventral edge simple, with wide central concavity. Female. —Coloration: Similar to male. Antennal segments I to III red brick and IV reddish brown; collar and calli clearly with blue-green metallic reflections; connexival segments VIII and IX black with upper margin dark yellow; abdominal segments VIII and IX black; gonocoxae I black with blue- green reflections; paratergites VIII and IX black. Measurements .—Male (female). Head length: 1.72 mm (1.80 mm); width across eyes: 2.16 mm (2.32 mm); interocular space: 1.12 mm (1.22 mm); interocellar space: 0.42 mm (0.52 mm); antennal segments lengths: I, 4.24 mm (4.40 mm); II, 3.44 mm (3.16 mm); III, 3.52 mm (3.56 mm); IV, 5.40 mm (5.20 mm). Pronotal length: 3.44 mm (3.88 mm); maximum width of anterior lobe: 2.40 mm (2.64 mm); maximum width of posterior lobe: 7.30 mm (7.60 mm); maximum length of humeral angle: 2.76 mm (2.92 mm); maximum width of humeral angle: 3.08 mm (3.36 mm). Scutellar length: 2.00 mm (2.36 mm); width: 2.04 mm (2.52 mm). Total body length: 17.98 mm (20.15 mm). Discussion .—Like S. lugens Horvath, with humeral angles of pronotum pro¬ duced anteriorly into wing-like projections (Figs. 3, 5, 10-11), always bicolorous, with posterior third black, and with head in dorsal view black to reddish brown and always with metallic reflections. In S. rahmus, the anterior lobe of metathoracic peritreme and adjacent areas are black with posterior lobe bright orange, and abdominal sterna are black to reddish brown with three rows of colors clearly separated. In S. lugens, the an¬ terior and posterior lobe of metathoracic peritreme and adjacent areas are pale orange, and the color of the abdominal sterna exhibit a pink metallic reflection, with only the pleural margin yellow. Etymology .—The species name is an arbitrary combination of letters and is to be treated as a noun. SUNDARUS XENIA BRAILOVSKY, NEW SPECIES (Figs. 6, 12-13) Types .—Holotype male: Brasil. Rondonia, 62 km., SW Ariquemes, nr Fzda Rcho. Grande, 4—16 Nov 1997, J. E. Eger. Deposited in United States National Vol. 78(4) 280 THE PAN-PACIFIC ENTOMOLOGIST Figure 13. Dorsal view of Sundarus xenia Brailovsky, NEW SPECIES. 2002 BRAILOVSKY: THREE NEW SUNDARUS SPECIES 281 14 Figure 14. Dorsal view of Sundarus rahmus Brailovsky, NEW SPECIES. Museum of Natural History, Washington, D. C. (Drake Collection). Paratypes: 3 males, 4 females; data: same as holotype: 30 Mar-10 Apr 1992, J. E. Eger, and 5 Dec 1991, S. L. Heydon. Deposited in Coleccion Entomologica del Instituto de Biologfa, UN AM, Mexico; United States National Museum of Natural History, Washington, D. C. (Drake Collection); and Department of Entomology, University 282 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) of California, Davis, California. 1 male; data: Brasil. Rondonia, vie. Caucalandia, 0 32' S-62 48' W, 160-350 m, 14 Oct 1991, J. R. MacDonald. Deposited in Mississippi Entomological Museum, Mississippi State. 1 female; data: Brasil. Rondonia, 10 km S of Caucalandia (Linea C5) off B-65, 19-21 Mar 1991, B. Kondratieff. Deposited in Colorado State University, Department of Entomology, Fort Collins, Colorado. 1 female; data: Peru. Rio Taiche, 1923, H. Bassler. De¬ posited in American Museum of Natural History, New York. Description .—Male (holotype). Dorsal coloration: Head metallic green with apex of tylus, inner angle of antenniferous tubercle and ocellar tubercle dark yellow; antennal segments I to III black with blue-purple metallic reflections, and segment IV reddish brown; pronotum metallic green with wide bell-shaped orange spot at posterior lobe and below calli; scutellum orange; clavus and corium black with blue and purple metallic reflections and with claval commissure, apical margin and costal border creamy yellow; hemelytral membrane black; connexivm black with upper margin yellow; dorsal ab¬ dominal segments black, except segment VI with blue-purple metallic reflections. Ventral coloration: Head metallic green; buccula orange with green metallic reflections; rostral segments dark reddish brown with blue-green metallic reflections at segments I and II; thorax metallic green with anterior and posterior lobe of metathoracic peritreme and adjacent areas orange; prosternum and mesosternum reddish brown with median longitudinal stripe dark orange; metasternum reddish brown; abdominal sterna metallic green with pleural margin yellow; genital capsule metallic pink with green metallic reflections and with postero ventral edge yellow; coxae, trochanters and femora dark reddish brown with blue-green metallic reflections; tibiae dark brick red with blue-green metallic reflections and tarsi dark brick red. Structure: Rostrum reaching posterior margin of metasternum. Pronotum: Anterolateral margins obliquely straight, scarsely crenate; humeral angles not exposed, truncated with borders sin¬ uate (Figs. 6 and 12); posterolateral and posterior border straight and entire; calli transversely raised. Genital capsule. Posteroventral edge simple, broadly concave. Female. —Coloration: Similar to male. Connexival segments VIII and IX black with blue-green reflections; abdominal sterna metallic green with pink metallic reflections, and with pleural margin yellow; gonocoxae I metallic pink; paratergite VIII and IX black with green metallic reflections, and with upper margin yellow. Measurements .—Male (female). Head length: 1.46 mm (1.84 mm); width across eyes: 2.08 mm (2.36 mm); interocular space: 1.08 mm (1.34 mm); interocellar space: 0.47 mm (0.54 mm); antennal segments lengths: I, 3.28 mm (3.60 mm); II, 3.12 mm (3.40 mm); III, 3.40 mm (3.68 mm); IV, 4.64 mm (5.04 mm). Pronotal length: 2.80 mm (4.12 mm); maximum width of anterior lobe: 2.08 mm (3.20 mm); maximum width of posterior lobe: 3.84 mm (5.60 mm). Scutellar length: 1.76 mm (2.20 mm); width: 1.60 mm (2.32 mm). Total body length: 14.65 mm (20.97 mm). Discussion .—The pronotal shape, including the humeral angles not exposed and truncated (Figs. 6 and 12), the pronotum and thorax not entirely yellow, the pronotal disk not bulging outwards, and rostral segment I reddish brown and never yellow, somewhat similar to S. sheilae Brailovsky. Sundarus xenia described from Brazil is recognized by having an orange bell-shaped spot covering most of the posterior lobe of pronotal disk (Figs. 6, 12—13). In S. sheilae, only recorded from Bolivia, the posterior lobe of pronotal disk is metallic green with an orange lon¬ gitudinal and relatively narrow rectangular-shape median stripe (Figs. 4 and 9). Etymology .—The species name is an arbitrary combination of letters and is to be treated as a noun. Key to Sundarus Species** 1. Humeral angles of pronotum blunt, obtuse, not exposed or barely lami¬ nated (Figs. 4, 6, 9, 12) . 2 ** S. muggei Schmidt is excluded from the key. 2002 BRAILOVSKY: THREE NEW SUNDARUS SPECIES 283 1'. Humeral angles of pronotum produced into wing-like projections (Figs. 1, 3, 5, 7, 10-11) . 8 2. Pronotum and thorax entirely yellow . S. splendidus Distant 2'. Pronotum and thorax not entirely yellow . 3 3. Pronotum metallic purple; anterior third of pronotal disk strongly convex, with calli sunken; rostral segment I yellow . S. gibbus Brailovsky 3'. Pronotum not metallic purple; pronotal disk not remarkably convex; ros¬ tral segment I black to reddish brown . 4 4. Pronotum entirely metallic blue-green . S. castus Brailovsky 4'. Pronotum not entirely metallic blue-green . 5 5. Pronotum metallic green with an orange spot at posterior lobe. 6 5'. Pronotum not metallic green with an orange spot . 7 6. Posterior lobe of pronotum with an orange bell-shaped spot covering most of disk (Fig. 6) . S. xenia Brailovsky, NEW SPECIES 6'. Posterior lobe of pronotum with orange but narrow rectangular-shaped stripe on middle third (Fig. 4) . S. sheilae Brailovsky 7. Pronotum black, with anterolateral margins and collar yellow, and calli and adjacent region metallic green . S. ducalis (in part) 7'. Pronotum metallic green and posteriorly with two broad black spots lat¬ eral to middle line. S. rufoscutellatus (Gray) 8. Clavus and corium entirely yellow or bright orange . 9 8'. Clavus and corium always with black areas . 10 9. Pronotum, clavus and corium entirely yellow; humeral angles of pron¬ otum in lateral view almost covering the middle third of head (Fig. 7); thorax entirely yellow . S. ambarinus Brailovsky 9'. Pronotum, clavus and corium entirely bright orange; humeral angles of pronotum shorter, in lateral view not extending beyond anterior lobe (Fig. 8); thorax almost entirely metallic green and never yellow .... . S. occua Brailovsky, NEW SPECIES 10. Wing projections of pronotum bicolorous (Figs. 3 and 5) . 11 10'. Wing projections of pronotum unicolorous, usually yellow or orange and eventually with metallic green reflections at posterior margin. 17 11. Head dorsally yellow or orange without metallic reflections . . S. regalis (Westwood) 11'. Head dorsally with green, blue or purple metallic reflections. 12 12. Posterior lobe of pronotal disk entirely yellow or at least with broad or narrow longitudinal yellow stripe at middle third. 13 12'. Posterior lobe of pronotal disk black or metallic blue-green, and never with yellow or orange marks . 14 13. Posterior lobe of pronotal disk entirely yellow; wing-like projection re¬ markably expanded, in lateral view extending beyond the apex of ty- lus; calli yellow; black spot of wing-like projection surrounded by yellow . S. vulneratus Brailovsky 13'. Posterior lobe of pronotal disk black with yellow longitudinal stripe at mid point; wing-like projection shorter, in lateral view not extending beyond the anterior third of eyes; calli metallic green; posterior lobe of wing-like projection entirely black . S. collinus Brailovsky 284 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) 14. 14'. 15'. 15'. 16. 16'. 17. 17'. 18. 18'. 19. 19'. 20 . 20 '. 21 . 21 '. 22 . 22 '. 23. 23'. 24. 24'. 25. 25'. 26. 26'. 27. 27'. 28. 28'. 29. 29'. Pronotal disk behind calli with blue-green-purple metallic reflections . . . S. zonatus Brailovsky Pronotal disk behind calli black without metallic reflections. 15 Pronotal collar yellow; humeral angles of pronotum barely expanded . . . S. ducalis (Stal) (in part) Pronotal collar metallic green; wing-like projections conspicuously ex¬ panded (Figs. 3, 5, 10-11) . 16 Anterior lobe of metathoracic peritreme and adjacent areas black . . S. rahmus Brailovsky, NEW SPECIES Anterior lobe of metathoracic peritreme and adjacent areas pale orange . S. lugens Horvath Scutellum black. S. acutus Signoret Scutellum yellow or orange . 18 Anterior and posterior lobe of metathoracic peritreme and adjacent areas black to reddish brown. S. nigrosteolatus Brailovsky Anterior and posterior lobe of metathoracic peritreme yellow or orange without black areas . 19 Head dorsally yellow or orange without metallic reflections . 20 Head dorsally with green or pink metallic reflections . 25 Wing-like projections of pronotum longer than wide. 21 Wing-like projections of pronotum wider than long or subequal. 23 Wing-like projections rectangular, conspicuously expanded, in lateral view extending beyond the apex of tylus. S. selvaticus Brailovsky Wing-like projections medium sized, acute or rounded, in lateral view reaching the eyes. 22 Mesothorax and metathorax entirely orange; wing-like projections round¬ ed . S. flavicollis Signoret Mesothorax and metathorax metallic green with pink reflections; wing¬ like projections rectangular . S. tropicalis Brailovsky Prothorax metallic green with propleural expansions and posterior margin orange . S. perpictus Distant Prothorax yellow with acetabulae metallic green . 24 Wing-like projections, conspicuously rounded. S. paludum Brailovsky Wing-like projections shorter and slightly more elongate than rounded . S. horni Schmidt Thorax entirely yellow or orange. S. inca Breddin Thorax metallic green with or without pink reflections. 26 Posterior margin of prothorax entirely yellow or orange . 27 Posterior margin of prothorax metallic green or only with the border yellow but never covering entire margin . 28 Pronotum entirely yellow or orange . S. volutatorius Brailovsky Pronotum with collar and calli metallic green. S. bellus Brailovsky Calli yellow or orange . S. mucronatus Horvath Calli metallic green, with or without pink reflections . 29 Anterior lobe of pronotal disk entirely metallic green . . S. humeralis Horvath Anterior lobe of pronotal disk entirely yellow or orange . 30 2002 BRAILOVSKY: THREE NEW SUNDARUS SPECIES 285 30. Wing-like projections remarkably expanded, in lateral view extending beyond apex of tylus . S. palmatus Schmidt 30'. Wing-like projections shorter, in lateral view reaching the middle third of eyes. S. pontifex Buchana White Acknowledgments Thanks are due to Randall T. Schuh (American Museum of Natural History, New York), Boris Kondratieff (Colorado State University, Department of Ento¬ mology, Fort Collins, Colorado), R. L. Brown (Mississippi Entomological Mu¬ seum, Mississippi State), Mats Eriksson (Universitets Zoologiska Institut, Upp¬ sala, Sweden), the late R. Schuster (University of California, Department of En¬ tomology, Davis, California), and Thomas J. Henry (Systematic Entomology Lab¬ oratory, USDA, % United States National Museum of Natural History, Washington, D.C.) for support of this project. Special thanks are also given to Ernesto Barrera and Elvia Esparza, Instituto de Biologia UNAM, for the prepa¬ ration of illustrations. Literature Cited Brailovsky, H. 1988. Revision del genero Sundarus Amyot-Serville (Hemiptera-Coreidae-Coreini) para el Continente Americano. Anales Inst. Biol. UNAM 58 (1987), Ser. Zool., 2: 561-622. Received 23 June 2002, Accepted 4 November 2002. PAN-PACIFIC ENTOMOLOGIST 78(4): 286-288, (2002) Scientific Note DISCOVERY OF BRUCHIDIUS VILLOSUS F. (COLEOPTERA: BRUCHIDIDAE) ON SCOTCH BROOM IN CANADA Scotch broom ( Cytsius scoparius (L.) Link [Fabaceae]) is an invasive legu¬ minous shrub native to continental Europe (Waloff, N. 1966. J. Appl. Ecol. 3: 293-311). Scotch broom has now been disseminated to temperate regions of the world with movements of people. In North America it was introduced as a garden or ornamental hedge species and has since spread far beyond the bounds of cul¬ tivation in all locations (Zielke, K., J. O. Boateng, N. Caldicott & H. Williams. 1992. Broom and gorse in British Columbia: a forestry perspective problem anal¬ ysis. Ministry of Forests, Victoria, British Columbia) and is now distributed along both the Atlantic and Pacific Coasts. Scotch broom had become naturalized in British Columbia and in initial surveys was found on southern Vancouver Island, the Lower Mainland, and the Gulf Islands as far north as Cortes, Hernando, Savary and Texada Islands (Taylor, T. M. C. 1974. The pea family of British Columbia. British Columbia Provincial Museum Handbook No. 32, Victoria, Brit¬ ish Columbia). More recent spread has been reported to the northern limit of Vancouver Island and into the Queen Charlotte Islands (Zielke, K., J. O. Boateng, N. Caldicott & H. Williams. 1992. Broom and gorse in British Columbia: a for¬ estry perspective problem analysis. Ministry of Forests, Victoria, British Colum¬ bia). Scotch broom is also occurs on the Sunshine Coast to Powell River, through the Fraser and Chilliwack Valleys, into Hope, in the west Kootenay region in the British Columbia interior, and between Nelson and Castlegar (Zielke, K., J. O. Boateng, N. Caldicott & H. Williams. 1992. Broom and gorse in British Colum¬ bia: a forestry perspective problem analysis. Ministry of Forests, Victoria, British Columbia). Several European broom specialist insects are known from broom in western North America; Aceria genistae Nalepa (Acari: Eriphyidae), Agonopterix nervosa Haworth (Lepidoptera: Oecophoridae), Arytainilla spartiophila Forster (Hemiptera: Psyllidae), ( Ctenocallis setosa Kaltenbach (Homoptera: Aphididae), Gargara genistae F. (Hemiptera: Membracidae), Dictyonota fuliginosa (Costa) (Hemiptera: Tingidae), Leucoptera spartefolliela (Hiibner) (Lepidoptera: Lyone- tiidae), and three species of mirids (Chan, K. L. & C. E. Turner. 1998. Pan-Pacific. Entomol., 74(1): 55-57; Andres, L. A. & E. M. Coombs. 1995. Scotch broom. In Nechols, J. R.; L. A. Andres; J. W. Beardsly; R. D. Goeden; and C. G. Jackson (eds.). Biological control in the western United States: accomplishments and ben¬ efits of regional research project W-84. Berkeley, California, United States of America, University of California, Division of Agriculture and Natural Resources, Pub. No. 3361, pp. 303—305; Pfeiffer, D. G. 1986. J. Entomol. Sci., 21(3): 214— 218; Waloff, N. 1966. J. Appl. Ecol., 3: 293-311; Downes, W. 1957. Entomol. Soc. B. C., 54: 11-13; Footit, R. G. & W. R. Richards. 1993. The insects and arachnids of Canada. Part 22: The genera of aphids in Canada (Homoptera: Aphi- doidea and Phylloxeroidea), Agriculture Canada, Ottawa; Syrett, P, S. V. Fowler, E. M. Coombs, J. R. Hosking, G. P. Markin, Q. Paynter & A. W. Sheppard. 1999. 2002 SCIENTIFIC NOTE 287 Biocontrol News Inf., 20: 17N—34N). No biological control agents have been intentionally introduced as part of a biological control program for Scotch broom in Canada. We report here the discovery of another accidentally introduced insect specialist Bruchidius villosus F. (Coleoptera: Bruchididae) on Scotch broom in the vicinity of Victoria, British Columbia. The Scotch broom seed-feeding beetles were discovered in a seed collection conducted in mid-July 2000. Pods were collected from approximately 50 plants over a one-hectare area on a power line right-of-way that intersects Munn Road, Victoria (48°31'00" N, 123°26'00" W, elevation 81 m). The site is within the coastal Douglas-fir biogeoclimatic zone on a rock outcrop dominated by C. sco- parius, Quercus garryana, Holodiscus discolor, and grass species. After collec¬ tion, mature seeds were removed from pods and placed in vials for storage. In early August 2000, sixty-six B. villosus had emerged from the seeds through small holes. Examination of the remaining seeds revealed that the seeds infested with beetles had darkened noticeably since collection. Within 10 days the 416 beetles had emerged (51 failed to emerge) from a total of 1179 seeds (approximately 100 pods). The beetles were subsequently identified by J. M. Kingsolver of the Florida State Collections of Arthropods as Bruchidius villosus F. (Coleoptera: Bruchidi¬ dae), described from Scotch broom ( Sarothamnus scoparius Koch. [= Cytisus scoparius (L.) Link]) by B. J. Southgate (Southgate, B. J. 1963. Ann. Entomol. Soc. Am., 56: 795-797). It is unknown how or when these bruchids arrived in British Columbia. This is the first record of B. villosus in British Columbia. In Europe, Bruchidius villosus is restricted to Scotch broom and the adults only oviposit in the presence of a broom pod (Parnell, J. R. 1966. J. Anim. Ecol. 35: 157-188). Its native range includes the U.K., France, Portugal, Spain, Austria, Denmark, Germany, Hungary, Italy, and Switzerland (Frick, K. E. 1962 Unpub¬ lished file report, USDA—Agricultural Research Service; Syrett, P. & K. E. Em- berson. 1997. Biocontrol Sci. Technol., 7(3): 309—326). B. villosus has been in¬ tentionally introduced in Australia and New Zealand where broom has also be¬ come naturalized. Small numbers of B. villosus were released in the United States in 1998 in the foothills of the Cascades and on the Oregon coast. To date there have been no recoveries of B. villosus from the coastal 1998 release site, however, given the low number of adults (130) released, it is not surprising and is not considered a failed release. B. villosus was released at 18 sites in Oregon and Washington State in 1999. The beetle was recovered at two 1999 release sites inspected in 2000. There are now more than 30 releases on Scotch broom and one each on French and Portuguese broom (D. Isaacson, personal communica¬ tion). In 2001, an informal survey was conducted to determine the distribution of B. villosus on Scotch broom in British Columbia. B. villosus was found at 31 of 32 sites on Vancouver Island from 48°25' N to 50°01' N and at nine of 11 sites on the mainland from 49°02' N to 49°54' N. Seed damaged varied from 1% to 87% depending upon site. Based on my observations that B. villosus is common and widespread on Scotch broom in British Columbia, it seems unlikely that bruchids released in the pacific northwest of the United States are the source of the Canadian populations. B. villosus was first recorded on Scotch broom in Mas¬ sachusetts in 1918 after an accidental introduction (Bottimer, L. J. 1968. Can. Ent., 100: 139-145). One could postulate that B. villosus has spread from Mas- 288 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) sachusetts site to western Canada, utilizing other legumes as hosts in the absence of Scotch broom, however, given the host specificity of B. villosus, this is unlikely (Parnell, J. R. 1966. J. Anim. Ecol., 35: 157—188). Another hypothesis is that B. villosus was independently introduced into British Columbia after 1963 when Waloff (Waloff, N. 1966. J. Appl. Ecol., 3: 293—311) reported that there were no insects living inside the broom pods. Additional collections of Scotch broom pods will be made next season to de¬ termine the range and infestation levels of this species in British Columbia as well as its potential for control of the spread of Scotch broom. Records. —BRITISH COLUMBIA, VICTORIA: Munn Road, 18 July 2000, L. R. E. Hooper, Cytisus scoparius, seeds. Acknowledgement. —I am especially grateful to V. Nealis, R. Duncan, and L. Humble of the Pacific Forestry Centre in Victoria, BC and J. M. Kingsolver of Florida State Collections of Arthropods, Gainesville, Florida, USA who helped to identify specimens. The British Columbia Hydro Power Authority provided financial support for this work. Laura R. E. Hooper, Department of Zoology, University> of British Columbia, 6270 University Boulevard, Vancouver, British Columbia V6T 1Z4 Canada. Received 9 January 2002; Accepted 20 June 2002. PAN-PACIFIC ENTOMOLOGIST 78 ( 4 ): 289 - 306 , ( 2002 ) Scientific Note PRELIMINARY INSECT SURVEY ON WESTERN POISON OAK ( TOXICODENDRON DIVERSILOBUM (TORREY & GRAY)), COYOTE BRUSH {BACCHARIS PILULARIS DE CONDOLLE), AND TOYON ( HETEROMELES ARBUTIFOLIA (LINDLEY)) IN THE SANTA CRUZ MOUNTAINS, CALIFORNIA Chaparral represents one of the most widespread and unique vegetation types in California, covering roughly 9% of the state (Holland, V. L. & D. J. Keil, 1995, California vegetation. Kendall/Hunt Publishing, Dubuque, Iowa), yet little docu¬ mentation exists on the diversity of insects found within this community. Agri¬ cultural and urban expansion, grazing, fire suppression, and the introduction of exotic plants all threaten chaparral habitat. Over a one year period beginning June 1995, we examined the insect fauna present on three commonly encountered shrubs within a chaparral community: western poison oak ( Toxicodendron diver- silobum (Torrey & Gray)), coyote brush ( Baccharis pilularis De Condolle), and toyon ( Heteromeles arbutifolia (Lindley)). Our objective was to develop a prelim¬ inary inventory of the insect fauna for the chaparral community in the central region of California and forms the basis of this scientific note. Sampling was done in the Santa Cruz Mountains in portions of Santa Clara, Santa Cruz, and San Mateo counties in Northern California. Locations were ran¬ domly selected and used in this study if the three shrub species were present in quantities sufficient for sampling within a range of approximately 1 km. Nine new locations were sampled for each period. There were eight sampling periods throughout the year with five weeks between the start of periods except for two ten-week intervals during the winter. All sampling was done during the day in dry weather. At each location, beating samples were taken from each species of shrub, from ten healthy branches of each species. The ten branches were selected to give a fairly uniform distribution across the patch of shrubs within the sampling location; branches close to other plant species were avoided. Branches were beaten with a length of broomstick over a 0.5 m 2 linen beating tray-funnel and dislodged ma¬ terial collected in a large jar. Samples were kept in plastic bags and frozen before sorting arthropods from debris. Several resources were used for specimen identification, including specialists, identification keys, and by comparison with specimens in the collections at the California Academy of Sciences and San Jose State University. Due to lack of resources and time, identification efforts concentrated on the most common spe¬ cies, leaving many species undetermined. Undetermined species were separated by morphology. Voucher specimens have been deposited at the J. Gordon Edwards Museum at San Jose State University. A listing of the 826 insect species collected can be found in Appendix 1. Acknowledgments —We thank the following people for their invaluable help 290 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) with identifications: John Brown (Tortricidae), Douglass R. Miller (Psyllidae, Triozidae, Coccidae), Gary L. Miller (Aphididae), David A. Nickle (Orthoptera), and Thomas J. Henry (Miridae) (all with the Systematic Entomology Laboratory, Agricultural Research Service, U.S. Department of Agriculture); J. Gordon Ed¬ wards (San Jose State Univ.) (Chrysomelidae), Phil Ward (U.C. Davis) (Formi- cidae), Norman Penny (California Academy of Sciences) (adult Neuroptera), Richard Snider (Michigan State Univ.) ( Bourletiella ), and Catherine A. Tauber (Cornell Univ.) (immature Neuroptera). We also thank The Midpeninsula Re¬ gional Open Space Preserve and several private landowners for allowing us to sample on their property. Mark Isaak and Jeffrey Y. Honda, Department of Biological Sciences, San Jose State University. One Washington Square, San Jose, California 95192. Received 20 February 2002; Accepted 13 November 2002. Appendix 1. Number of insect specimens collected from coyote brush (C), poison oak (P), and toyon (T) over a 1 year period. Unless specified otherwise, Lepidoptera and Neuroptera are larvae and insects in other orders are adults. Order Family Species c p T Collembola Hypogastruidae undet. sp. 1 4 0 4 undet. sp. 2 2 0 0 Isotomidae undet. sp. 1 0 2 0 undet. sp. 2 0 0 1 undet. sp. 3 3 2 2 undet. sp. 4 0 0 1 undet. sp. 5 0 1 0 undet. sp. 6 0 0 1 Entomobryidae Entomobrya atrocincta Schott 16 2 6 Entomobrya multifasciata (Tullberg) 447 32 35 Entomobrya suzannae Schott 11 16 46 undet. sp. 1 4 0 8 undet. sp. 2 1 0 0 undet. sp. 3 0 0 4 undet. sp. 4 0 1 1 Sminthuridae Bourletiella n. sp. 1 61 153 9 Bourletiella n. sp. 2 489 364 277 Bourletiella sp. 3 0 62 0 Dicyrtoma beta Christiansen & Bellinger 179 68 192 Sminthurus sp. 1 0 1 Microcoryphia Machilidae undet. sp. 1 1 0 0 undet. sp. 2 0 1 0 Thysanura Lepismatidae undet. sp. 1 0 0 Ephemeroptera Undetermined undet. sp. 1 0 1 0 undet. sp. 2 0 0 1 2002 ISAAK & HONDA: INSECTS FOUND ON CHAPARRAL PLANTS 291 Appendix 1. Continued. Order Family Species c p T Phasmida Timemidae Timema califonica Scudder 20 1 115 Orthoptera Acrididae undet. sp. 1 1 0 0 undet. sp. 2 1 0 0 Tettigoniidae Microcentrum sp. 2 2 2 undet. sp. 2 0 1 0 undet. sp. 3 0 1 0 undet. sp. 4 0 0 1 Rhaphidophoridae Gammarotettix bilobatus Thomas 20 6 12 Gryllidae Hoplosphyrum sp. 2 0 1 Oecanthus californicus Saussure 1 9 8 undet. sp. 3 1 1 0 Dermaptera Forficulidae Forficula auricularia L. 5 2 8 Plecoptera Undetermined undet. sp. 1 1 0 4 undet. sp. 2 0 0 2 undet. sp. 3 0 0 1 Psocoptera Trogiidae Cerobasis guestfalica (Kolbe) 29 14 21 Liposcelididae Liposcelis sp. 2 0 1 Caeciliidae Caecilius burmeisteri Brauer 3 0 4 Caecilius maritimus Mockford 5 14 42 Caecilius sp. 3 1 1 1 Caecilius sp. 4 5 0 0 undet. sp. 5 0 0 1 Amphipsocidae Polypsocus corruptus (Hagen) 0 1 4 Dasydemellidae Teliapsocus conterminus (Walsh) 2 0 1 Stenopsocidae Graphopsocus cruciatus (L.) 30 52 167 Elipsocidae Elipsocus hyalinus (Stephens) 4 5 4 Philotarsidae Philotarsus picicornis (Fabricius) 1 6 2 Lachesillidae Lachesilla pacifica Chapman 5 2 17 Peripsocidae Peripsocus subfasciatus (Rambur) 1 0 0 Ectopsocidae Ectopsocus briggsi McLachlan 87 75 396 Ectopsocus californicus (Banks) 69 54 274 Trichopsocidae Trichopsocus acuminatus Badonnel 18 27 108 Psocidae Amphigerontia bifasciata (Latreille) 69 2 4 Indiopsocus sp. 0 5 1 Loensia maculosa (Banks) 23 1 1 undet. sp. 4 0 0 9 Undetermined undet. sp. 1 0 1 0 undet. sp. 2 1 0 0 Heteroptera Tingidae Corythucha incurvata Uhler 17 1 688 undet. sp. 2 2 0 0 undet. sp. 3 1 0 0 undet. sp. 4 0 0 1 Miridae Engytatus sp. 0 0 3 Irbisia sp. 1 1 17 0 292 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) Appendix 1. Continued. Order Family Species c p T Parthenicus brunneus Van Duzee 423 0 1 Plagiognathus diversus cruralis Van Duzee 0 58 0 undet. sp. 1 80 1 7 undet. sp. 2 0 0 23 undet. sp. 3 31 7 34 undet. sp. 4 227 0 1 undet. sp. 5 5 6 21 undet. sp. 6 1 6 13 undet. sp. 7 2 9 71 undet. sp. 8 3 5 68 undet. sp. 9 3 3 0 undet. sp. 11 0 0 4 undet. sp. 12 0 1 0 undet. sp. 13 17 0 0 undet. sp. 14 0 1 0 undet. sp. 15 5 0 0 undet. sp. 16 1 0 0 undet. sp. 17 1 0 0 undet. sp. 18 0 1 1 undet. sp. 19 1 0 0 Nabidae undet. sp. 1 0 0 Anthocoridae Orius insidiosus (Say) 22 0 3 undet. sp. 2 0 0 6 undet. sp. 3 1 0 0 Reduviidae Empicoris rubromaculatus (Blackburn) 0 0 7 undet. immature sp. 2 1 2 3 Berytidae undet. sp. 1 1 0 Lygaeidae Nysius tenellus Barber 85 0 2 undet. sp. 2 1 1 2 undet. sp. 3 2 0 1 undet. sp. 4 1 0 0 undet. sp. 5 1 0 0 Rhopalidae Leptocoris trivittatus (Say) 1 17 1 Pentatomidae undet. sp. 1 0 1 0 undet. sp. 2 0 0 2 undet. sp. 3 0 0 1 undet. sp. 4 4 0 0 undet. sp. 5 0 0 1 Homoptera Cicadidae undet. sp. 3 0 0 Membracidae Philya californiensis (Goding) 809 0 1 Stictocephala sp. 2 0 0 undet. sp. 3 0 0 3 undet. sp. 4 0 0 1 undet. sp. 5 2 0 3 Aphrophoridae undet. sp. 1 3 1 2 Cercopidae undet. sp. 1 15 2 11 undet. sp. 2 18 3 34 undet. sp. 3 0 1 0 undet. sp. 4 0 2 0 undet. sp. 5 1 0 0 undet. sp. 6 0 1 0 2002 ISAAK & HONDA: INSECTS FOUND ON CHAPARRAL PLANTS 293 Appendix 1. Continued. Order Family Species C p T Cicadellidae Acroneura sp. 0 13 0 Empoasca sp. 1 759 4 6 Empoasca sp. 2 134 1 3 Neocoelidiana obscura (Baker) 8 0 6 Pagaronia tredecimpunctata Ball 9 12 4 Stragania sp. 7 0 0 undet. sp. 7 8 0 0 undet. sp. 8 1 0 0 undet. sp. 9 1 0 0 undet. sp. 10 1 0 0 undet. sp. 11 3 0 0 undet. sp. 12 1 0 0 undet. sp. 13 1 3 0 undet. sp. 14 1 0 0 undet. sp. 15 1 0 0 undet. sp. 16 1 0 0 undet. sp. 17 0 1 0 undet. sp. 18 0 0 1 undet. sp. 19 1 0 0 undet. sp. 20 0 0 3 undet. sp. 21 1 0 1 undet. sp. 22 1 0 0 undet. sp. 23 1 0 0 undet. sp. 24 0 2 0 undet. sp. 25 0 1 0 undet. sp. 26 1 1 2 undet. sp. 27 1 0 0 undet. sp. 28 1 0 0 undet. sp. 29 2 1 0 undet. sp. 30 2 0 1 undet. sp. 31 2 0 0 undet. sp. 32 0 0 1 undet. sp. 33 0 1 0 undet. sp. 34 1 0 0 undet. sp. 35 0 0 2 undet. sp. 36 2 0 0 undet. sp. 37 0 2 1 undet. sp. 38 0 0 1 undet. sp. 39 0 0 1 undet. sp. 40 1 0 1 undet. sp. 41 3 3 2 undet. sp. 42 1 0 6 Delphacidae undet. sp. 3 0 0 Psyllidae Psylla sp. 3 1 2 undet. sp. 2 0 0 1 undet. sp. 3 0 1 0 undet. sp. 4 1 0 0 undet. sp. 5 0 0 1 undet. sp. 6 0 1 4 undet. sp. 7 1 0 0 undet. sp. 8 1 0 0 undet. sp. 9 0 0 2 294 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) Appendix 1. Continued. Order Family Species c p T Aleyrodidae undet. sp. 1 12 9 7 undet. sp. 2 6 9 0 undet. sp. 3 5 0 7 undet. sp. 4 0 0 1 undet. sp. 5 1 0 2 undet. sp. 6 0 0 2 undet. sp. 7 1 0 0 undet. sp. 8 2 1 1 Aphidae Aphis baccharicola Hille Ris Lambers 1383 10 1 Aphis sp. 3 5 1 5 Aphis sp. 4 15 2 0 Aphis spiraecola Patch 15 0 2 Brachycaudus helichrysi Kaltenbach 252 5 10 Dactynotus sp. 1 42 7 6 Dactynotus sp. 2 68 4 2 Hyadaphis foeniculi Passerini 31 7 46 Myzocallis agrifolicola Richards 7 4 15 Myzus persicae Sulzer 45 1 0 Ovatus sp. 14 5 5 Tuberculatus sp. 1 5 1 3 Tuberculatus sp. 2 0 4 5 undet. sp. 1 7 1 2 undet. sp. 2 14 3 0 undet. sp. 14 0 3 1 undet. sp. 15 3 0 0 undet. sp. 16 0 0 1 undet. sp. 17 7 0 0 undet. sp. 18 1 1 1 undet. sp. 19 0 2 2 undet. sp. 20 0 0 2 undet. sp. 21 2 0 1 undet. sp. 22 3 0 2 undet. sp. 23 0 3 0 undet. sp. 24 2 35 0 undet. sp. 25 0 4 0 undet. sp. 26 7 0 0 undet. sp. 27 2 0 0 undet. sp. 28 1 0 0 undet. sp. 29 1 0 0 undet. sp. 30 1 0 0 undet. sp. 31 0 0 1 undet. sp. 32 3 0 0 Coccidae Ceroplastes sp. 4 0 0 Triozidae Calinda longistylus Crawford 77 0 0 Undetermined male undet. scale sp. 1 1 0 0 male undet. scale sp. 2 0 0 1 male undet. scale sp. 3 1 0 0 male undet. scale sp. 4 0 2 0 male undet. scale sp. 5 1 0 0 undet. scale sp. 1 1 0 0 undet. scale sp. 2 1 0 0 undet. scale sp. 3 6 0 0 undet. scale sp. 4 1 1 0 2002 ISAAK & HONDA: INSECTS FOUND ON CHAPARRAL PLANTS 295 Appendix 1. Continued. Order Family Species c p T undet. scale sp. 5 0 0 10 undet. scale sp. 6 0 0 5 undet. scale sp. 7 0 0 2 undet. scale sp. 8 0 0 1 undet. sp. 1 0 0 1 undet. sp. 2 2 0 1 Thysanoptera Aeolothripidae Aeolothrips sp. 1 60 3 39 Aeolothrips sp. 2 1 0 0 Franklinothrips sp. 1 0 1 Melanothrips sp. 0 1 1 undet. sp. 2 3 2 5 undet. sp. 6 1 1 0 Thripidae Anaphothrips sp. 4 5 0 Chirothrips sp. 4 1 0 Frankliniella sp. 1 1060 102 18,397 Frankliniella sp. 2 15 137 17 Heliothrips haemorrhoidalis (Bouche) 0 11 98 undet. sp. 2 5 11 61 undet. sp. 3 73 1105 15 undet. sp. 4 3 26 7 undet. sp. 5 24 6 47 undet. sp. 6 3 3 11 undet. sp. 7 22 0 1 undet. sp. 8 0 1 0 undet. sp. 9 0 0 2 Phlaeothripidae Leptothrips sp. 362 71 184 Nesothrips sp. 1 0 0 undet. sp. 2 5 4 386 undet. sp. 3 1 3 0 undet. sp. 4 1 0 0 undet. sp. 5 0 1 1 Undetermined undet. sp. 1 0 0 1 undet. sp. 2 0 0 1 Neuroptera Raphidiidae Agulla sp. 1 1 1 Coniopterygidae Coniopteryx latipalpus Meinander 0 1 3 Coniopteryx pal pal pus Meinander 3 1 0 other adult Coniopteryx spp. 7 2 11 larva Coniopteryx spp. 28 1 13 Hemerobiidae Hemerobius sp. 1 17 5 12 Hemerobius sp. 2 1 0 2 Chrysopidae Chrysoperla carnea (Stephens) 26 17 20 Chrysoperla plorabunda (Fitch) 0 0 1 Pseudomallada sierra (Banks) 6 0 3 chrysopid sp. 4 0 0 1 chrysopid sp. 5 2 0 0 Coleopterera Leiodidae Agathidium rotundulum Mannerheim 3 0 38 Scydmaenidae undet. sp. 1 7 2 9 undet. sp. 2 6 0 4 undet. sp. 3 3 3 2 296 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) Appendix 1. Continued. Order Family Species c p T Staphylinidae Pseudolesteva sp. 0 28 0 undet. sp. 2 0 2 0 undet. sp. 3 0 0 2 undet. sp. 4 1 0 0 undet. sp. 5 0 0 1 undet. sp. 6 0 0 1 undet. sp. 7 1 0 0 Hydrophilidae undet. sp. 1 0 1 Buprestidae undet. sp. 1 2 0 0 Elateridae Limonius ornatulus (LeConte) 0 5 6 Sericus silaceus (Say) 1 10 0 undet. sp. 3 0 0 1 undet. sp. 4 0 10 0 undet. sp. 5 1 0 0 undet. sp. 6 1 0 0 undet. sp. 7 1 0 0 undet. sp. 8 0 0 1 undet. sp. 9 0 0 1 Cantharidae Cantharis sp. 1 0 10 2 Cantharis sp. 2 3 1 1 Malthinus sp. 0 2 0 Malthodes bicurvatus Fall 5 7 5 Podabrus pruniosus LeConte 4 0 1 Podabrus modulatus Fall 6 0 3 undet. sp. 7 0 0 1 undet. sp. 8 1 1 0 Dermestidae Cryptorhopalum apicale (Mannerheim) 3 16 6 undet. sp. 2 1 0 0 Anobiidae undet. sp. 1 0 0 1 undet. sp. 2 1 0 0 undet. sp. 3 5 0 0 undet. sp. 4 2 0 0 undet. sp. 5 0 1 0 undet. sp. 6 1 0 1 undet. sp. 7 1 0 0 undet. sp. 8 0 1 0 undet. sp. 9 0 0 1 Ptinidae Ptinus interruptus LeConte 8 0 5 undet. sp. 2 0 1 0 undet. sp. 3 3 0 1 Cleridae undet. sp. 1 0 2 1 undet. sp. 2 0 0 1 undet. sp. 3 1 1 0 undet. sp. 4 1 0 1 Melyridae Collops vitattus (Say) 0 2 2 Listrus canescens (Mannerheim) 1 11 6 undet. sp. 1 4 4 5 undet. sp. 2 1 1 9 Coccinellidae Cephaloscymnus occidentalis Horn 0 0 1 Chilocorus bipustulatus (L.) 0 0 1 Chilocorus nr. orbus 0 0 3 Coccinella californica Mannerheim 3 1 1 Coccinella septempunctata (L.) 1 0 1 2002 ISAAK & HONDA: INSECTS FOUND ON CHAPARRAL PLANTS 297 Appendix 1. Continued. Order Family Species c p T Coccinella trifasciata subversa LeConte 5 1 1 Cycloneda polita Casey 5 2 3 Hippodamia convergens Guerin 4 0 1 Hyperaspis quadrioculatus Motschulsky 4 0 3 Psyllobora vigintimaculata Say 80 7 21 Scymnus ( Pullus ) calaveras Casey 0 0 2 Scymnus ( Pullus ) pallens LeConte 0 1 5 Scymnus (Pullus ) sp. 1 10 1 1 Scymnus (Pullus ) sp. 2 15 0 0 Scymnus (Pullus ) sp. 3 1 0 1 Scymnus (Pullus ) sp. 4 1 0 0 Scymnus (Pullus ) sp. 5 0 1 1 Scymnus {Scymnus) nebulosus LeConte 1 0 0 Scymnus marginicollis Mannerheim 1 0 0 Stethorus punctum picipes Casey 1 2 0 Zagloba ornata (Horn) 1 0 10 undet. larva sp. 1 57 5 1 undet. larva sp. 2 1 0 2 undet. larva sp. 3 1 0 0 undet. larva sp. 4 2 0 0 undet. larva sp. 5 1 4 0 undet. larva sp. 6 0 0 2 undet. sp. 22 0 0 2 Corylophidae undet. sp. 1 1 0 0 undet. sp. 2 1 0 0 Byturidae Byturellus grisescens (Jayne) 4 3 6 Mordellidae undet. sp. 1 0 1 0 undet. sp. 2 0 0 2 undet. sp. 3 1 0 2 undet. sp. 4 0 0 1 undet. sp. 5 1 0 2 undet. sp. 6 0 0 12 undet. sp. 7 0 0 1 Meloidae undet. sp. 1 1 0 0 undet. sp. 2 0 1 0 undet. sp. 3 1 0 0 Anthicidae Anthicus nitidulus LeConte 14 4 13 undet. sp. 2 1 0 1 Cerambycidae undet. larva sp. 1 0 1 0 undet. sp. 1 0 0 1 undet. sp. 2 0 0 1 Bruchidae Stator limbatus (Horn) 10 3 1 undet. sp. 2 3 0 0 undet. sp. 3 1 0 0 undet. sp. 4 0 0 2 undet. sp. 5 0 0 1 Chrysomelidae Crepidodera sp. 1 8 0 1 Cryptocephalus confluentus White 3 0 0 Diabrotica undecimpunctata Mannerheim 7 0 1 Diachus auratus (Fabricius) 17 10 27 Epitrix hirtipennis (Melsheimer) 0 68 2 Trirhabda flavolimbada (Mannerheim) adult 14 0 0 Trirhabda flavolimbada (Mannerheim) larva 71 0 0 298 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) Appendix 1. Continued. Order Family Species c p T undet. larva sp. 2 1 9 3 undet. larva sp. 3 0 0 2 undet. larva sp. 4 0 0 2 undet. larva sp. 5 0 0 1 undet. sp. 7 0 0 3 undet. sp. 8 0 0 5 undet. sp. 9 0 0 1 undet. sp. 10 1 0 0 undet. sp. 11 0 1 0 undet. sp. 12 1 0 0 undet. sp. 13 0 0 1 Brentidae Apion cribricolle LeConte 11 0 9 Curculionidae Sibinia maculata (LeConte) 5 1 1 undet. sp. 2 0 0 7 undet. sp. 3 1 0 1 undet. sp. 4 0 0 1 undet. sp. 5 0 1 0 undet. sp. 6 0 0 1 undet. sp. 7 1 0 0 undet. sp. 8 0 0 1 undet. sp. 9 0 0 1 undet. sp. 10 0 0 3 Undetermined undet. sp. 1 0 0 4 undet. sp. 2 0 0 1 undet. sp. 3 1 0 2 undet. sp. 4 0 0 1 undet. sp. 5 0 0 2 undet. sp. 6 0 0 1 undet. sp. 7 1 0 0 undet. sp. 8 1 0 0 undet. larva sp. 1 6 1 0 undet. larva sp. 2 0 0 1 undet. larva sp. 3 0 1 0 undet. larva sp. 4 0 0 2 undet. larva sp. 5 0 1 0 undet. larva sp. 6 2 1 0 undet. larva sp. 7 0 1 0 undet. larva sp. 8 1 3 0 undet. larva sp. 9 0 1 0 undet. larva sp. 10 0 1 0 Diptera Tipulidae undet. sp. 1 0 0 1 undet. sp. 2 0 1 3 Mycetophilidae undet. sp. 0 0 1 Sciaridae Lycoriella sp. 2 0 2 undet. sp. 2 0 0 5 undet. sp. 3 3 0 1 undet. sp. 4 1 2 0 undet. sp. 5 0 2 0 undet. sp. 6 0 0 1 undet. sp. 7 1 0 0 undet. sp. 8 1 0 0 2002 ISAAK & HONDA: INSECTS FOUND ON CHAPARRAL PLANTS 299 Appendix 1. Continued. Order Family Species c p T Cecidomyiidae Contarinia sp. 39 3 0 Cordylomyia sp. 1 0 0 Lestremiinae sp. 0 0 1 Polystepha sp. 2 1 1 Porricondyla sp. 1 2 8 Rhopalomyia californica Felt 2 0 0 undet. sp. 5 0 0 1 undet. sp. 6 2 0 0 undet. sp. 8 0 1 0 undet. sp. 9 3 0 1 undet. sp. 10 0 1 5 undet. sp. 11 1 0 0 undet. sp. 12 3 0 0 undet. sp. 13 0 1 0 undet. sp. 14 0 0 1 Scatopsidae Psectrosciara sp. 0 0 2 Synneuridae undet. sp. 1 0 0 Simuliidae undet. sp. 1 0 0 1 undet. sp. 2 1 0 0 Ceratopogonidae undet. sp. 1 0 1 0 undet. sp. 2 0 0 1 Chironomidae Orthocladius sp. 28 3 6 undet. sp. 2 7 3 3 undet. sp. 3 17 1 34 undet. sp. 4 0 0 1 undet. sp. 5 0 0 1 undet. sp. 6 1 0 0 undet. sp. 7 1 2 0 undet. sp. 8 0 1 0 undet. sp. 9 1 0 0 undet. sp. 10 1 0 0 undet. sp. 11 2 0 1 undet. sp. 12 1 0 0 undet. sp. 13 1 0 0 Rhagionidae undet. sp. 0 0 2 Empididae undet. sp. 1 0 0 3 undet. sp. 2 0 1 0 undet. sp. 3 1 0 0 Dolichopodidae undet. sp. 1 1 0 0 undet. sp. 2 1 0 0 Phoridae undet. sp. 1 1 0 0 undet. sp. 2 0 0 1 undet. sp. 3 1 0 0 Tephritidae undet. sp. 1 1 0 0 undet. sp. 2 2 0 0 Agromyzidae undet. sp. 1 0 0 1 undet. sp. 2 1 0 0 undet. sp. 3 0 2 0 undet. sp. 4 0 1 0 Asteiidae undet. sp. 1 0 0 1 undet. sp. 2 1 0 0 Lauxaniidae undet. sp. 1 1 0 0 undet. sp. 2 1 0 0 300 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) Appendix 1. Continued. Order Family Species c p T Trixoscelididae undet. sp. 1 0 0 Chyromyidae undet. sp. 0 0 1 Drosophilidae undet. sp. 1 0 0 Chloropidae undet. sp. 1 0 1 0 undet. sp. 2 2 0 0 Undetermined undet. larva sp. 1 0 0 1 undet. larva sp. 2 0 1 0 Lepidoptera Lyonetiidae Bucculatrix variabilis Braun (adult) 87 0 3 Bucculatrix variabilis Braun (larva) 2265 1 4 Coleophoridae Coleophora lynosyridella Walsingham 59 0 0 Coleophora viscidiflorella Walsingham 313 1 0 Cossidae undet. sp. 1 0 3 1 Tortricidae undet. sp. 1 0 1 5 undet. sp. 2 0 0 3 Pterophoridae undet. sp. 1 0 1 0 undet. sp. 2 0 1 0 Pyralidae Phycitinae sp. 0 0 4 undet. sp. 1 4 1 0 undet. sp. 2 0 2 4 Hesperiidae undet. sp. 1 0 0 4 Lycaenidae undet. sp. 1 0 0 1 undet. sp. 2 0 0 1 Geometridae Dichorda illustraria (Hulst) 0 3 0 Elpiste marcescaria Guenee 314 1 2 Eupithecia rotundopuncta Packard 2 3 1 Nematocampa limbada (Haworth) 0 1 0 Nemoria leptalea Ferguson 0 0 3 Nemoria sp. 2 0 0 1 Neoalcis californica (Packard) 0 0 1 Prochoerodes truxaliata Guenee 8 0 0 Sabulodes aegrotata (Guenee) 7 9 11 undet. sp. 1 0 4 1 undet. sp. 2 1 0 6 undet. sp. 3 46 8 8 undet. sp. 4 4 0 0 undet. sp. 5 2 0 0 undet. sp. 6 3 0 0 undet. sp. 7 1 0 0 undet. sp. 8 3 0 1 undet. sp. 9 3 0 0 undet. sp. 10 1 0 0 undet. sp. 11 0 4 0 undet. sp. 12 0 1 0 undet. sp. 13 0 1 0 undet. sp. 14 0 1 0 undet. sp. 15 0 1 0 undet. sp. 16 0 1 0 undet. sp. 17 0 0 2 undet. sp. 18 0 0 2 undet. sp. 19 0 0 1 undet. sp. 20 0 0 1 2002 ISAAK & HONDA: INSECTS FOUND ON CHAPARRAL PLANTS 301 Table 1. Continued. Order Family Species c p T undet. sp. 21 0 0 2 undet. sp. 22 0 0 1 Lymantriidae Orgyia leucostigma (J. E. Smith) 0 0 1 undet. sp. 2 1 0 1 Noctuidae Cosmia canescens Behr. 0 0 1 Undetermined undet. larva sp. 1 49 0 0 undet. larva sp. 2 5 0 1 undet. larva sp. 3 2 0 0 undet. larva sp. 4 1 0 0 undet. larva sp. 5 1 0 0 undet. larva sp. 6 1 0 1 undet. larva sp. 7 1 0 0 undet. larva sp. 8 1 0 0 undet. larva sp. 9 0 1 0 undet. larva sp. 10 0 3 0 undet. larva sp. 11 0 2 1 undet. larva sp. 12 0 1 0 undet. larva sp. 13 0 1 0 undet. larva sp. 14 0 0 1 undet. larva sp. 15 0 0 1 undet. larva sp. 16 1 1 2 undet. larva sp. 17 2 1 3 undet. larva sp. 18 0 0 2 undet. larva sp. 19 0 0 1 undet. larva sp. 20 0 0 1 undet. larva sp. 21 0 0 2 undet. larva sp. 22 0 0 1 undet. larva sp. 23 1 0 1 undet. larva sp. 24 0 0 1 undet. larva sp. 25 1 0 1 undet. larva sp. 26 0 0 1 undet. larva sp. 27 0 0 1 undet. larva sp. 28 2 0 1 undet. larva sp. 29 0 0 2 undet. larva sp. 30 0 0 2 undet. larva sp. 31 0 1 0 undet. adult sp. 1 1 0 0 undet. adult sp. 2 9 0 0 undet. adult sp. 3 1 1 0 undet. adult sp. 4 1 0 0 Hymenoptera Ceraphronidae undet. sp. 1 1 0 0 undet. sp. 2 0 0 1 undet. sp. 3 0 0 1 undet. sp. 4 0 0 3 undet. sp. 5 0 1 0 undet. sp. 6 0 0 1 Braconidae undet. sp. 1 1 0 0 undet. sp. 2 2 0 0 undet. sp. 3 2 0 0 undet. sp. 4 1 0 1 undet. sp. 5 2 0 0 302 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) Appendix 1. Continued. Order Family Species c p T undet. sp. 6 2 0 0 undet. sp. 7 0 1 0 undet. sp. 8 0 0 1 undet. sp. 9 1 0 0 undet. sp. 10 0 0 1 undet. sp. 11 0 0 1 undet. sp. 12 0 0 1 undet. sp. 13 8 0 0 Ichneumonidae undet. sp. 1 0 0 3 undet. sp. 2 0 0 1 undet. sp. 3 0 1 0 undet. sp. 4 0 0 1 undet. sp. 5 0 0 1 undet. sp. 6 1 0 0 undet. sp. 7 0 1 0 undet. sp. 8 2 0 0 Mymaridae Gonatocerinae sp. 1 2 0 0 Gonatocerinae sp. 2 0 1 0 Gonatocerinae sp. 3 0 1 0 Gonatocerinae sp. 4 0 0 1 Gonatocerinae sp. 5 1 0 0 Gonatocerinae sp. 6 1 0 0 Mymarinae sp. 1 7 3 1 Mymarinae sp. 2 1 0 1 Mymarinae sp. 3 5 1 2 Mymarinae sp. 4 4 0 0 Mymarinae sp. 5 0 1 0 Mymarinae sp. 6 0 0 1 Mymarinae sp. 7 1 0 0 undet. sp. 3 4 0 1 Trichogrammatidae undet. sp. 1 1 0 0 undet. sp. 2 0 0 1 Eulophidae Tetrastichinae sp. 1 1 2 4 Tetrastichinae sp. 2 3 4 3 Tetrastichinae sp. 3 1 0 1 Tetrastichinae sp. 4 2 1 2 Tetrastichinae sp. 5 1 0 0 Tetrastichinae sp. 6 1 0 0 Tetrastichinae sp. 7 0 1 0 Tetrastichinae sp. 8 0 0 1 Tetrastichinae sp. 9 1 0 0 Tetrastichinae sp. 10 0 1 0 Tetrastichinae sp. 11 1 0 0 undet. sp. 1 3 0 0 undet. sp. 2 5 0 0 undet. sp. 3 8 0 0 undet. sp. 4 5 0 0 undet. sp. 5 0 1 1 undet. sp. 6 4 0 0 undet. sp. 7 0 2 0 undet. sp. 8 1 0 0 undet. sp. 9 0 2 5 undet. sp. 10 0 0 3 2002 ISAAK & HONDA: INSECTS FOUND ON CHAPARRAL PLANTS 303 Appendix 1. Continued. Order Family Species c p T undet. sp. 11 0 0 2 undet. sp. 12 0 0 1 undet. sp. 13 0 0 1 undet. sp. 14 0 1 0 undet. sp. 15 0 0 1 undet. sp. 16 1 0 0 undet. sp. 17 0 0 1 undet. sp. 18 3 0 0 Elasmidae undet. sp. 1 1 0 0 undet. sp. 2 1 0 0 Aphelinidae undet. sp. 1 2 4 1 undet. sp. 2 2 0 5 undet. sp. 3 0 0 1 undet. sp. 4 0 1 0 undet. sp. 5 0 0 1 undet. sp. 6 1 0 0 Signiphoridae undet. sp. 1 0 1 1 undet. sp. 2 1 0 0 undet. sp. 3 0 1 0 Encyrtidae undet. sp. 1 4 1 1 undet. sp. 2 0 0 4 undet. sp. 3 2 0 0 undet. sp. 4 2 0 3 undet. sp. 5 4 0 0 undet. sp. 6 2 0 2 undet. sp. 7 2 0 1 undet. sp. 8 2 0 2 undet. sp. 9 1 1 0 undet. sp. 10 1 0 0 undet. sp. 11 1 1 0 undet. sp. 12 1 1 0 undet. sp. 13 1 0 1 undet. sp. 14 1 0 0 undet. sp. 15 1 0 0 undet. sp. 16 1 0 0 undet. sp. 17 1 0 0 undet. sp. 18 1 0 0 undet. sp. 19 0 1 0 undet. sp. 20 2 0 0 undet. sp. 21 1 0 0 undet. sp. 22 1 0 0 undet. sp. 23 0 1 0 undet. sp. 24 1 0 0 undet. sp. 25 1 0 0 undet. sp. 26 0 0 1 undet. sp. 27 1 0 0 undet. sp. 28 0 1 0 undet. sp. 29 1 0 0 undet. sp. 30 1 0 0 undet. sp. 31 1 0 0 Eupelmidae Eupelminae sp. 0 0 0 1 Eupelminae sp. 1 0 1 0 Eupelminae sp. 2 4 0 0 304 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) Appendix 1. Continued. Order Family Species c p T Eupelminae sp. 3 2 0 0 Eupelminae sp. 4 3 0 0 Eupelminae sp. 5 7 0 0 Eupelminae sp. 6 3 0 0 Eupelminae sp. 7 1 0 0 Calosotinae sp. 1 2 0 0 Calosotinae sp. 2 1 0 0 Torymidae Toryminae sp. 1 3 0 0 Toryminae sp. 2 1 0 0 Toryminae sp. 3 1 0 0 undet. sp. 1 6 0 1 Pteromalidae Brachyscelidiphaginae sp. 43 0 0 Eunotinae sp. 3 0 0 undet. sp. 1 12 0 0 undet. sp. 3 7 0 0 undet. sp. 4 22 0 0 undet. sp. 5 8 0 0 undet. sp. 6 14 0 0 undet. sp. 7 6 0 0 undet. sp. 8 5 0 0 undet. sp. 9 3 0 0 undet. sp. 10 5 0 0 undet. sp. 11 1 0 1 undet. sp. 13 1 1 1 undet. sp. 14 2 0 0 undet. sp. 15 0 1 0 undet. sp. 16 0 1 0 undet. sp. 17 1 1 0 undet. sp. 18 1 0 1 undet. sp. 19 1 0 1 undet. sp. 20 0 1 1 undet. sp. 21 2 0 0 undet. sp. 22 1 0 0 undet. sp. 23 0 0 1 undet. sp. 24 0 0 1 undet. sp. 27 1 0 0 undet. sp. 28 1 0 0 undet. sp. 29 0 1 0 Eurytomidae Eurytominae sp. 1 1 0 0 Eurytominae sp. 2 0 1 0 Harmolitinae sp. 1 0 0 Chalcididae Brachymeriinae sp. 0 1 0 Cynipidae undet. sp. 1 2 7 1 undet. sp. 2 0 1 1 undet. sp. 3 1 0 2 undet. sp. 4 0 1 0 undet. sp. 5 0 0 1 undet. sp. 6 0 1 0 Diapriidae undet. sp. 1 1 0 1 undet. sp. 2 0 0 1 undet. sp. 3 0 0 1 Scelionidae undet. sp. 1 1 0 0 undet. sp. 2 0 0 1 2002 ISAAK & HONDA: INSECTS FOUND ON CHAPARRAL PLANTS 305 Appendix 1. Continued. Order Family Species c p T Platygastridae undet. sp. 1 197 2 1 undet. sp. 2 3 6 3 undet. sp. 3 0 1 4 undet. sp. 4 30 2 1 undet. sp. 5 1 0 1 undet. sp. 6 0 1 1 undet. sp. 7 0 0 1 undet. sp. 8 1 0 0 undet. sp. 9 0 1 0 undet. sp. 10 0 1 0 undet. sp. 11 0 0 4 Bethylidae undet. sp. 1 0 3 0 undet. sp. 2 1 3 0 undet. sp. 3 1 0 0 Halictidae undet. sp. 1 0 0 1 Tiphiidae undet. sp. 1 0 1 0 Formicidae Camponotus hyatti Emery 15 1 8 Camponotus vicinus Mayr 14 0 0 Camponotus sp. 3 0 1 0 Camponotus sp. 4 0 0 1 Camponotus sp. 5 0 0 1 Camponotus sp. 6 1 0 0 Camponotus sp. 7 1 0 0 Crematogaster coarctata Mayr 236 16 33 Formica moki Wheeler 46 2 2 Formica sp. 2 2 4 8 Formica sp. 3 112 0 0 Formica sp. 4 2 0 0 Formica sp. 5 1 0 0 Formica sp. 6 1 0 0 Formica sp. 7 0 0 1 Formica sp. 8 1 0 0 Formica sp. 9 0 3 0 Leptothorax gallae M. Smith 1 3 1 Leptothorax sp. 1 0 3 1 Leptothorax sp. 2 0 2 1 Leptothorax sp. 3 0 1 0 Leptothorax sp. 4 1 0 0 Leptothorax sp. 5 1 0 0 Line pit hema humile (Mayr) 1738 21 98 Monomorium ergatogyna Wheeler 77 6 0 Monomorium sp. 2 0 0 1 Prenolepis imparis (Say) 132 156 185 Tapinoma sessile (Say) 131 25 2 undet. sp. 29 0 0 2 undet. sp. 30 0 1 0 undet. sp. 31 0 0 1 undet. sp. 32 0 0 1 Undetermined undet. sawfly larva sp. 1 1 1 1 undet. sawfly larva sp. 2 0 1 0 undet. chalcidoid sp. 1 1 0 0 undet. chalcidoid sp. 2 0 0 1 306 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) Appendix 1. Continued. Order Family Species c p T undet. chalcidoid sp. 3 1 0 0 undet. chalcidoid sp. 4 0 0 3 undet. wasp sp. 1 1 0 0 undet. wasp sp. 2 1 0 0 PAN-PACIFIC ENTOMOLOGIST 78(4): 307, (2002) PROCEEDINGS OF THE PACIFIC COAST ENTOMOLOGICAL SOCIETY, 2001 Five Hundred Seventy-Second Meeting The 572nd meeting of the Pacific Coast Entomological Society was held at 8:00 PM on 14 Decem¬ ber 2001, in the Goethe Room of the California Academy of Sciences in San Francisco with Mr. Stanley E. Vaughn presiding. The following persons were introduced as guests: Mr. David Baumgardner by Dr. Norman D. Penny; Dr. Rob E. Roughley, a dytiscid specialist from the University of Manitoba, by Mr. Keve J. Ribardo; Dr. Catherine V. Milton of UC Berkeley by Dr. Paul H. Arnaud Jr.; and Dr. Elizabeth McGee of San Jose State University, David and Michele Vaughn, Lucille Mason, Lindsey Vaughn, and Jeffery Mills by Mr. Stanley E. Vaughn. Mr. Vaughn also recognized and welcomed Mike Solari, wishing him a happy 16th birthday. The following new slate of officers was voted on and approved by the society: Dr. Rolf L. Aalbu for President-elect, Dr. Robert L. Zuparko for Treasurer, Mr. Vincent F. Lee for Managing Secretary, and Dr. Katherine N. Schick for President. Mr. Vaughn then handed over the gavel to Dr. Schick, who presided over the remainder of the meeting. The membership committee announced that the 2001 membership was currently 329 members, including the 32 new members added this year. Mr. Gordon Nishida of Salinas, California was proposed and approved as a regular member of the society. The featured speaker, Mr. Stanley E. Vaughn, Curator of the Dr. J. Gordon Edwards Entomology Museum at San Jose State University, presented a blood-letting slide lecture entitled “Hippoboscid Flies of Madagascar Lemurs.” Aside from being another one of Mr. Vaughn’s legendary entomological fables of biting, stinging, hurting and bleeding, the talk updated the audience on his work with Eliz¬ abeth McGee in the Ranomafana National Park. In this ideal land of medical entomology, Mr. Vaughn surveyed three species of lemurs in primary and secondary rainforests, assessing the lemurs for ec¬ toparasite loads, with the long-term objective to assess the health risk associated with human and prosimian contact in disturbed ecosystems. During his work in the park, he attempted to determine how Allobosca crassipes (Speiser) find their hosts after pupating on the terrain. One possibility is they fly and then sheer off their wings once a host is acquired. Allobosca spp. are very host specific, with a life cycle that is still not fully understood, and no males have yet been described. Mr. Vaughn, now armed with 154 specimens of Allobosca spp., will continue research under better lab conditions. The meeting was adjourned at 9:17 PM, followed by a social hour held in the Department of Entomology conference room. The following 36 persons were present: (22 members) P. H. Arnaud Jr., R. M. Brown, H. K. Court, J. G. Edwards, J. J. Fairbanks, L. R. Faziola, E. M. Fisher, S. D. Gaimari, C. E. Griswold, D. K. James, V. F. Lee, T. C. Meikle, D. A. parks, N. D. Penny, W. W. Pitcher, A. E. Rackett, K. J. Ribardo, W. E. Savary, K. N. Schick, J. J. Schweikert, M. Solari, and S. E. Vaughn; (14 guests) B. Bianchini, D. Baumgardner, M. Delmas, L. J. Mason, E. McGee, J. Mills, K. Milton, J. Myatt, A. Myatt, R. Roughley, D. Silva, D. S. Vaughn, L. Vaughn, and M. Vaughn. PAN-PACIFIC ENTOMOLOGIST 78(4): 308, (2002) 2001 SPONSORING MEMBERS OF THE PACIFIC COAST ENTOMOLOGICAL SOCIETY Robert R Allen Ernest Anderson William F. Barr Paula & Robert Buickerood Helen K. Court Bryan K. Eya E. Eric Grissell Teresa C. Meikle & Charles E. Griswold John E. Hafernik Jr. Frank T. Hovore Alice S. Hunter Gordon A. Marsh Albert E. Rackett Norman E. Gershenz & Leslie S. Saul Warren E. Savary Evert I. & Marion B. Schlinger Harvey I. Scudder Frank E. Skinner Ryan S. Walters Thomas J. Zavortink PAN-PACIFIC ENTOMOLOGIST 78(4): 309-310, (2002) The Pan-Pacific Entomologist Index to Volume 78 (title and key words) Agrosteella biconvexa NEW SPECIES 80 Agrosteella cheni NEW SPECIES 80 Agrosteela fallaciosa 80 Agrosteella violaceilcollis NEW SPECIES 80 Agrosteella jinni NEW SPECIES 80 Agrosteella medvedevi 80 Agrosteella oligotricha NEW SPECIES 80 Agrosteella punctata NEW SPECIES 80 Allozyme phylogeny Of Lycaenidae 219 Amblyseius graminis 215 Anobiidae Parasitized by Heterosphilus 7 Anthocoridae Mating behavior of 43 Araneae Species found in woodrat middens 23 Predation in leaf rollers 140 Jumping spiders 255 Baccharis piluaris Insects found on 289 Baja California Sur New Dysphenges 88 Betelgeuse piceus NEW SPECIES 188 Betelgeuse variabilis NEW SPECIES 188 Biocontrol Potential of Diaeretus on pine aphids 56 And spider predation 140 Braconidae Parasitoid of Hemicoelus gibicollis 7 Parasitic on pine aphids 56 New Chinese Gnamptodon 184 Brazil Collembola 69 Buprestidae Abundance in pheromone traps 120 Bruchidius villosus on Scotch broom 286 Cerambycidae New host and distribution records for Eburia 66 Abundance in pheromone traps 120 Cheiracanthium mildei 140 China Gnamptodon 184 Revision of jumping spiders 255 Chryosmelidae Agrosteella NEW GENUS 80 Dysphenges from Mexico 88 Cicadellidae Egg parasitoids of 34 Copulation In Anthocoris 43 Coreidae New Mictis from Sulawesi 110 New genera and species from the neotropical region 265 New species of Sundarus 276 Cucujidae Abundance in pheromone traps 120 Dasymutilla sicheliana 230 Diaeretus essigellae NEW SPECIES 56 Douglas-fir Relative and seasonal abundance of beetles found in pheromone traps 120 Dysphenges eichlini NEW SPECIES 88 Dysphenges lagunae NEW SPECIES 88 Dysphenges rileyi NEW SPECIES 88 Embola powelli NEW SPECIES 132 Essigella californica 56 Euburia 66 Foil setae Found in Collembola 69 Eolsomia fimetaria NEW SPECIES 69 Geometridae New species from Wyoming and Colorado 247 Glaucina incognitaria NEW SPECIES 247 Gnamptodon chinensis NEW SPECIES 184 Gonatocerus atriclavus Hagen, Kenneth S. obituary 151 Hawaii Cydia parasitism rates found in 101 Heteromeles arbutifolia Insects found on 289 Heterospilus luridostigmus NEW SPECIES 7 Isotomidae Folsomia species 69 310 THE PAN-PACIFIC ENTOMOLOGIST Vol. 78(4) Lacanobia subjuncta Larval bionomics 1 Leptopodidae Species new to Oregon 62 Megachilidae High temperature responses of 235 Metopoplax ditomoides 63 Mexico Mymaridae 34 New Brachonidae species 188 Mictis riedeli NEW SPECIES 110 Mictis sulawesiana NEW SPECIES 110 Mirabilis 132 Monotomidae Review of Hesperobaenus 197 Nectoquinitus alajuelensis NEW SPECIES 265 Neotoma Spider fauna of 23 Nematopodini Nectoquintius NEW GENUS 265 Stenoquintius NEW GENUS 265 Nickel accumulation In arthropods 168 Noctuidae Lacanobia larval biology 1 North America New species of Oxycarenidae 63 Obituary Hagen, Kenneth S. 151 Oregon New species of leptopodid 62 Oviposition preference Of Scirtothrips perseae on avocado 177 Oxycarenidae New species in North America 63 Patapius spinosus 62 Per sea americana 111 Phytoseiulus persimilis 215 Portia wui NEW SPECIES 255 Salticidae 255 Serpentine arthropods 168 Sperm transfer In Anthocoris 43 Stenoquintius reclusa NEW SPECIES 265 Sulawesi New Coreidae species 110 Sundarus occua NEW SPECIES 276 Sundarus rahmus NEW SPECIES 276 Sundarus xenia NEW SPECIES 276 Taiwan New species of Trichoptera 74 Thailand Braconidae from 17 Thermotolerance Of leafcutting bees 235 Trichoptera New Uganatrichia species from Taiwan 74 Toxicodendron diversilobum Insects found on 289 Uganatrichia taiwanensis NEW SPECIES 74 Ulex europaeus 215 Wood rat middens Spider fauna of 23 Yelicones siamensis NEW SPECIES 17 PAN-PACIFIC ENTOMOLOGIST 78(4): 311, (2002) The Pan-Pacific Entomologist Reviewers Volume 78 Androw, R. A. Baranowski, R. M. Baumann, R. W. Bosch, J. Butler, L. Brown, J. J. Christinsen, K. Coveil, C. Coyle, F. Ferguson, D. Flowers, W. R. Furth, D. G. Gall, W. K. Hedin, M. Helmut, R. Henry, T. Heppner, J. B. Higbee, B. S. Hook, A. Hormiga, G. Kemp, W. Kerns, D. Knight, A. Krantz, G. Landry, B. Lattin, J. Luhman, J. C. MacRae, T. C. Martins, S. Mayer, D. F. McHugh, J. V. Metzger, M. Myatt, R. Nice, C. Opler, R Packauskus, R. Parajulee, M. N. Parker, K. Peck, T. Pfeiffer, J. Pieper, K. Pitts, J. Pogue, M. G. Polhemus, D. Pollard, J. Powell, J. Rulowski, R. Schuh, R. Schwartz, M. D. Seybold, S. Slater, J. A. Sharky, M. J. Shaw, S. Staines, C. L. Triapitsyn, S. V. Ubick, D. Vetter, R. Wharton, B. Wood, D. L. Young, D. K. Zack, R. S. PAN-PACIFIC ENTOMOLOGIST 78(4): 312-313, (2002) The Pan-Pacific Entomologist Contents for Volume 78 Areekul, B. & D. L. J. Quicke—A new species of Yelicones Cameron (Hymenoptera: Braconidae) from Thailand. 17 Barthell, J. F., J. M. Hranitz, R. W. Thorp, & M. K. Shue— High temperature responses in two exotic leaf cutting bee species: Megachile apicalis and M. rotundata (Hymenoptera: Megachilidae). 235 Bousquet, Y.—Review of the genus Hesperobaenus Leconte (Coleoptera: Monotomidae) of America, north of Mexico . 197 Brailovsky, H.—Two new species of Mictis Leach (Heteroptera: Coreidae: Mictini) . . . . 109 Brailovsky, H.—Three new species of Sundarus Amyot & Serville, and key to the known species (Hemiptera: Heteroptera: Coreidae: Coreinae: Coreini). 276 Brailovsky, H. & E. Barrera—N ew genera and new species of neotropical Nematopodini (Hemiptera: Heteroptera: Coreidae: Coreinae). 265 Brenner, G. J., P. T. Oboyski, & P. C. Banko— Parasitism of Cydia spp. (Lepidoptera: Tortricidae) on Sophora chrysophylla (Fabaceae) along an elevation gradient of dry subalpine forest on Mauna Kea, Hawaii . 100 Cabrera, B. J., P. M. Marsh, V. R. Lewis & S. J. Seybold—A new species of Heterosphilus (Hymenoptera: Braconidae) associated with the deathwatch beetle, Hemicoelus gibbicollis (Leconte) (Coleoptera: Anobiidae). 7 Chen, X., J. B. Whitfield, & J. He—T he discovery of the genus Gnamptodon Haliday (Hymenoptera: Braconidae) in China, with description of one new species. 184 Dodds, K. J. & D. W. Ross—Relative and seasonal abundance of wood borers (Buprestidae, Cerambycidae) and Cucujidae trapped in Douglas-fir beetle pheromone- baited traps in northern Idaho. 120 Ferris, C. D. & J. S. Nordin—A new species of Glaucina Hulst from Wyoming and Colorado, and description of the female of G. nephos Rindge (Lepidoptera: Geometridae) . 247 Ge, S. Q„ S. Y. Wang, X. K. Yang, & W. Z. Li—A revision of the genus Agrosteella Medvedev (Chrysomelidae: Chrysomelinae) . 80 Gilbert, A. J. & F. G. Andrews—S tudies on the Chrysomelidae (Coleoptera) of the Baja California Peninsula: the genus Dysphenges Horn (Galerucinae: Alticini). 88 Hoddle, M. S.—Oviposition preferences of Scirtothrips perseae Nakahara (Thysanoptera: Thripidae) in southern California avocado orchards. Ill Hooper, L. R. E.—Discovery of Bruchidius villosus F. (Coleoptera: Bruchididae) on scotch broom in Canada . 286 Horton, D. R., T. M. Lewis, & T. Hinojosa— Copulation duration in three species of Anthocoris (Heteroptera: Anthocoridae) at different temperatures and effects on insemination and ovarian development . . . . 43 Hsu, L. P. & C. H. Chen—A new species of Ugandatrichia (Trichoptera: Hydroptilidae) from Taiwan. 74 Hsu, Y. F.—Larval and pupal biology of a new sun moth in Southern California; novel host use strategy in the evolution of Heliodinidae (Lepidoptera: Yponomeutoidea) .... 128 Isaak, M. & J. Y. Honda—P reliminary insect survey on western poison oak (Toxicodendron diversilobum (Torrey & Gray)), coyote brush (Baccharis pilularis De Condolle), and toyon (Heteromeles arbutifolia (Lindley)) in the Santa Cruz Mountains, California . 289 Landolt, P. J.—Survival and development of Lacanobia subjuncta (Grote & Robinson) (Lepidoptera: Noctuidae) larvae on common weeds and crop plants of eastern Washington state . 1 Lattin, J. D.—The immigrant leptopodid, Patapius spinous (Rossi), in Oregon (Hemiptera: Heteroptera: Leptopodidae) . . . 62 Lattin, J. D. & K. Wetherill —Metopoplax ditomoides (Costa), a species of Oxycarenidae new to North America (Lygaeoidea: Hemiptera: Heteroptera). 63 Miliczky, E. R. & C. O. Calkins—S piders 2002 CONTENTS FOR VOLUME 78 313 (Araneae) as potential predators of leaf roller larvae and egg masses (Lepidoptera: Tortricidae) in central Washington apple and pear orchards . 140 The Pacific Coast Entomological Society: Proceedings 2001 . 307 The Pacific Coast Entomological Society: Sponsoring Members 2002 . 308 The Pan-Pacific Entomologist: Index for Volume 78 . 309 The Pan-Pacific Entomologist: Reviewers for Volume 78 . 311 The Pan-Pacific Entomologist: Table of Contents for Volume 78. 312 Potapov, M. & M. Culik—A new species of Folsomia (Collembola: Isotomidae) from Brazil, with notes on foil-setae in the Fimetaria group . 69 Peng, X. & X. Li—Chinese species of the jumping spider genus Portia Karsch (Araneae: Salticidae)) . 255 Pratt, P. D. & E. M. Coombs —Phytoseiid mite fauna on gorse, Ulex europaelus L., in western Oregon, USA with new records for Phytoseiulus persimilis athias Henriot and Amblyseius graminis (Chant) (Acari: Phytoseiidae) . 115 Shaw, S. R. — Two new species of Betelgeuse from Mexico (Hymenoptera: Braconidae: Euphorinae) . 118 Stary, P & R. L. Zuparko— Diaeretus essigellae (Hymenoptera: Braconidae), a new species parasitic on Essigella pine aphids (Homoptera: Aphididae) from California . . . 5 Szafranski, P — New host plant and distributional records for some Eburia Lepeletier & Audinet-Serville (Coleoptera: Cerambycidae) in North America including Mexico . 65 Triapitzyn, S. V., L. G. Bezark, & D. J. W. Morgan.— Redescription of Gonatocerus atriclavus Girault (Hymenoptera: Mymaridae), with notes on other egg parasitoids of sharpshooters (Homoptera: Cicadellidae: Proconiini) in northeastern Mexico . 34 Vetter, R. S. & T. R. Prentice —The spider fauna associated with litter under woodrat middens in southern California (Arachnida: Araneae) . 23 Wall, M. A. & R. S. Boyd — Nickel accumulation in serpentine arthropods from the Red Hills, California . 168 Xu, X., C. M. Yin, & C. E. Griswold— A new species of the spider genus Macrothele from the Gaoligong Mountains Yunnan, China (Araneae: Hexathelidae). 115 Zuparko, R. L. — Obituary and bibliography of Kenneth S. Hagen (1919-1997), dedicated entomologist and teacher. 151 PAN-PACIFIC ENTOMOLOGIST Information for Contributors See volume 74: 248-255, October 1997, for detailed general format information and the issues thereafter for examples; see below for discussion of this journal’s specific formats for taxonomic manuscripts and locality data for specimens. Manuscripts must be in English, but foreign language summaries are permitted. Manuscripts not meeting the format guidelines may be returned. Please maintain a copy of the article on a word-processor because revisions are usually necessary before acceptance, pending review and copy-editing. Format. —Type manuscripts in a legible serif font IN DOUBLE OR TRIPLE SPACE with 1.5 in margins on one side of 8.5 X II in, nonerasable, high quality paper. THREE (3) COPIES of each manuscript must be submitted, EACH INCLUDING REDUCTIONS OF ANY FIGURES TO THE 8.5 X 11 IN PAGE. Number pages as: title page (page 1 ), abstract and key words page (page 2), text pages (pages 3+), acknowledgment page, literature cited pages, footnote page, tables, figure caption page; place original figures last. List the corresponding author’s name, address including ZIP code, and phone number on the title page in the upper right comer. The title must include the taxon’s designation, where appropriate, as: (Order: Family). The ABSTRACT must not exceed 250 words; use five to seven words or concise phrases as KEYWORDS. Number FOOTNOTES sequentially and list on a separate page. Text. — Demarcate MAJOR HEADINGS as centered headings and MINOR HEADINGS as left indented paragraphs with lead phrases underlined and followed by a period and two hypens. CITATION FORMATS are: Coswell (1986), (Asher 1987a, Franks & Ebbet 1988, Dorly et al. 1989), (Burton in press) and (R. F. Tray, personal communication). For multiple papers by the same author use: (Weber 1932, 1936, 1941; Sebb 1950, 1952). For more detailed reference use: (Smith 1983: 149-153, Price 1985: fig. 7a, Nothwith 1987: table 3). Taxonomy. — Systematics manuscripts have special requirements outlined in volume 69(2): 194-198; if you do not have access to that volume, request a copy of the taxonomy/data format from the editor before submitting manuscripts for which these formats are applicable. These requirements include SEPARATE PARAGRAPHS FOR DIAGNOSES, TYPES AND MATERIAL EXAMINED (INCLUDING A SPECIFIC FORMAT), and a specific order for paragraphs in descriptions. List the unabbreviated taxonomic author of each species after its first mention. Data Formats. — All specimen data must be cited in the journal's locality data format. See volume 69(2), pages 196-198 for these format requirements; if you do not have access to that volume, request a copy of the taxonomy/data format from the editor before submitting manuscripts for which these formats are applicable. Literature Cited. —Format examples are: Anderson, T. W. 1984. An introduction to multivariate statistical analysis (2nd ed). John Wiley & Sons, New York. Blackman, R. L, P. A. Brown & V. F. Eastop. 1987. Problems in pest aphid taxonomy: can chromosomes plus morphometries provide some answers? pp. 233-238. In Holman, J., J. Pelikan, A. G. F. Dixon & L. Weismann (eds.). Population structure, genetics and taxonomy of aphids and Thysanoptera. Proc. international symposium held at Smolenice Czechoslovakia, Sept. 9-14, 1985. SPB Academic Publishing, The Hague, The Netherlands. Ferrari, J. A. & K. S. Rai. 1989. Phenotypic correlates of genome size variation in Aedes albopictus. Evolution, 42: 895-899. Sorensen, J. T. (in press). Three new species of Essigella (Homoptera: Aphididae). Pan-Pacif. Entomol. Illustrations. — Illustrations must be of high quality and large enough to ultimately reduce to 117 X 181 mm while maintaining label letter sizes of at least 1 mm; this reduction must also allow for space below the illustrations for the typeset figure captions. Authors are strongly encouraged to provide illustrations no larger than 8.5 X 11 in for easy handling. Number figures in the order presented. Mount all illustrations. Label illustrations on the back noting: (1) figure number, (2) direction of top, (3) author’s name, (4) title of the manuscript, and (5) journal. FIGURE CAPTIONS must be on a separate, numbered page; do not attach captions to the figures. Tables. — Keep tables to a minimum and do not reduce them. Table must be DOUBLE-SPACED THROUGHOUT and continued on additional sheets of paper as necessary. Designate footnotes within tables by alphabetic letter. Scientific Notes. — Notes use an abbreviated format and lack: an abstract, key words, footnotes, section headings and a Literature Cited section. Minimal references are listed in the text in the format: (Bohart, R. M. 1989. Pan-Pacific. Entomol., 65: 156-161.). A short acknowledgment is permitted as a minor headed paragraph. Authors and affiliations are listed in the last, left indented paragraph of the note with the affiliation underscored. Page Charges. — PCES members are charged $35.00 per page, for the first 20 (cumulative) pages per volume and full galley costs for pages thereafter. Nonmembers should contact the Treasurer for current nonmember page charge rates. Page charges do not include reprint costs, or charges for author changes to manuscripts after they are sent to the printer. Contributing authors will be sent a page charge fee notice with acknowledgment of initial receipt of manuscripts. Volume 78 THE PAN-PACIFIC ENTOMOLOGIST October 2002 Number 4 Contents PRATT, G. F. & D. N. WRIGHT.—Allozyme phylogeny of North American coppers (Lycaeninae: Lycaenidae) _ 219 MANLEY, D. G. & W. R. RADKE.—Synonymy of Dasymutilla sicheliana (Saussure) (Hymenoptera: Mutillidae) _ 230 BARTHELL, J. F„ J. M. HRANITZ, R. W. THORP, & M. K. SHUE.—-High temperature responses in two exotic leafcutting bee species: Megachile apicalis and M. rotundata (Hymenoptera: Megachilidae) _ 235 FERRIS, C. D. & J. S. NORDIN.—A new species of Glaucina Hulst from Wyoming and Colorado, and description of the female of G. nephos Rindge (Lepidoptera: Geometridae) _ 247 PENG, X. & S. LI.—Chinese species of the jumping spider genus Portia Karsch (Araneae: Salticidae) _ 255 BRAILOVSKY, H. & E. BARRERA.—New genera and new species of neotropical Nematopodini (Hemiptera: Heteroptera: Coreidae: Coreinae) _ 265 BRAILOVSKY, H.—Three new species of Sundarus Amyot & Serville, and key to the known species (Hemiptera: Heteroptera: Coreidae: Coreinae: Coreini) _ 276 SCIENTIFIC NOTES: HOOPER, L. R. E.—Discovery of Bruchidius villosus F. (Coleoptera: Bruchididae) on scotch broom in Canada _ 286 ISAAK, M. & J. Y. HONDA—Preliminary insect survey on western poison oak (Toxicodendron diversilobum (Torrey & Gray)), coyote brush (Baccharis pilularis De Condolle) and toyon (Heteromeles arbutifolia (Lindley)) in the Santa Cruz Mountains, California _ 289 THE PACIFIC COAST ENTOMOLOGICAL SOCIETY: Proceedings 2001 _ 307 THE PACIFIC COAST ENTOMOLOGICAL SOCIETY: Sponsoring Members 2001 _ 308 THE PAN-PACIFIC ENTOMOLOGIST: Index for Volume 78 _ 309 THE PAN-PACIFIC ENTOMOLOGIST: Reviewers for Volume78 _ 311 THE PAN-PACIFIC ENTOMOLOGIST: Table of Contents for Volume 7 8 _ 312