The PAN-PACIFIC ENTOMOLOGIST Volume 73 January 1997 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 R. V. Dowell, Editor R. M. Bohart R. L. Penrose, Associate Editor J. T. Doyen R. E. Somerby, Book Review Editor J. E. Hafernik, Jr. Julieta EK Parinas, Treasurer Warren E. Savary Published quarterly in January, April, July, and October with Society Proceed- ings usually appearing in the October issue. All communications regarding non- receipt of numbers should be addressed to: Vincent EK Lee, Managing Secretary; and financial communications should be addressed to: Julieta F Parinas, Treasurer; at: Pacific Coast Entomological Society, Dept. of Entomology, California Acad- emy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599. 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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. Pacific Coast Entomological Society OFFICERS FOR 1997 Warren E. Savary, President Vincent EK Lee, Managing Secretary Julieta FE Parinas, Treasurer Stanley E. Vaughn, Recording Secretary 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 14 February 1997 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 73(1): 1-3, (1997) A NEW POECILONOTA FROM SOUTHERN CALIFORNIA (COLEOPTERA: BUPRESTIDAE) G. H. NELSON College of Osteopathic Medicine of the Pacific Pomona, California 91766-1889 Abstract.—Poecilonota viridicyanea Nelson, NEW SPECIES, is described from southern Cali- fornia. It is compared to its closest relative, P. bridwelli Van Dyke. Key Words.—Insecta, Coleoptera, Buprestidae, Poecilonota, southern California A new species of Poecilonota, that is most similar to P. bridwelli Van Dyke, and is allopatric with it, has been discovered in desert areas of southern California. It is described here to make it available for a general work on Buprestidae of America north of Mexico. Abbreviations of collections are as indicated in Arnett et al. (1993). Measure- ments were made with the use of either a half millimeter scale ruler or an ocular net reticule. Poecilonota viridicyanea Nelson, NEW SPECIES (Figs. 1, 3, 5) Types.—Holotype, male; data: CALIFORNIA. SAN BERNARDINO Co.: Yer- mo, 4 May 1939, J. Helfer, flying near cottonwood; deposited: California Acad- emy of Sciences, San Francisco. Paratypes: data: IMPERIAL Co., Palo Verde, R. A. Flock, reared from ‘“‘desert willow’’, wood collected 2 May 1972, emerged 16 May 1972, 1 male; deposited: GHNC; Palo Verde, 6 Jun 1972, D. H. Harris, from willow, 1 male; deposited: GHNC. Description—Holoty pe, male (Fig. 1). Size, 15.0 X 5.6 mm. Green-blue above and below with brassy tints at elytral apices. Elongate-oval, moderately transversely convex. Head. Frontovertex flat- tened above, concave below; surface densely, rugosely punctate, punctate areas clothed by semire- cumbent white setae; frontovertex with fine midline sulcus on elongate raised carina that bifurcates at midpoint of eyes, bifurcations joining supraantennal ridges to surround concavity; clypeus impunctate, margin shallowly arcuately emarginate; antennae reaching middle of pronotum when laid alongside, serrate from antennomere 4, antennomere 11 elongate-oval and truncate apically. Pronotum. Width 1.5 X length; lateral margins expanding from posterior angles in nearly straight line to widest point at middle, then converging in straight line to narrowest point at anterior angles; anterior margin arcuately emarginate with faint median lobe; posterior margin subtruncate with weak median lobe; surface with median ¥; smooth and impunctate, Y; on either side smooth with some moderate. sized punctures, lateral /; except for margin densely punctate and clothed with short, semirecumbent, white setae with white pulverulentus. Scutellum transversely cordate, surface glabrous, impunctate. Elytra. Length 3.6 X pronotal length, slightly wider at base than pronotum; lateral margins weakly sinuately expanding to widest point at middle then arcuately converging to slightly prolonged obliquely truncate apices; sutural margin diverging near tips; disk with intervals variously raised, intervals 3 and 5 distinctly raised and uninterrupted from basal ¥% to apical ¥,, others interrupted by finely, densely punctate areas clothed by short, semirecumbent, white setae and pulverulentus. Ventral side. Proster- num with disk and process longitudinally concave, concavity finely, densely punctate and clothed with long, curved, slender, white setae, less densely punctate posteriorly, process with smooth, impunctate, raised lateral margins; thoracic sterna more densely punctate laterally, less densely medially, meta- sternum with midline sulcus, punctate areas clothed by semirecumbent white setae and with pulver- ulentus laterally; pro- and mesofemora slender fusiform, metafemora more parallel-sided; tibiae 2, THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(1) Figures 1-2. Male holotypes, dorsal views. Fig. 1. Poecilonota viridicyanea. Fig. 2. P. bridwelli. 5 6 Figures 3—4. Last visible abdominal sterna. Fig. 3. P. viridicyanea. Fig. 4. P. bridwelli (line = 1.0 mm). Figures 5—6. Male genitalia, dorsal views. Fig. 5. P. viridicyanea. Fig. 6. P. bridwelli (line = 1.0 mm). 1997 NELSON: A NEW POECILONOTA 3 straight;. abdominal sternum 1 impunctate and longitudinally concave medially; other sterna sparsely punctate medially, all more densely punctate laterally, punctate areas clothed by recumbent white setae; last visible sternum with apex broadly arcuately emarginate, lateral angles rounded (Fig. 3). Male genitalia (Fig. 5). Female— Unknown. Diagnosis.—Poecilonota viridicyanea is most similar to P. bridwelli Van Dyke and will key to that species in Evans (1957). It can, however, be distinguished as follows: it is vivid green-blue in color; the frontovertex of the head has a broad prominent carina; the pronotum has a median impunctate area that is Y; the total width and without a punctate channel on either side (Fig. 1); the prosternum and its process are deeply longitudinally concave; the apex of the last abdominal sternum has the posterolateral angles broad and blunt (Fig. 3); and the juncture of the parameres is rounded (Fig. 5). In P. bridwelli the color is dull green to purple; the carina on the frontovertex is narrow and not prominent; the median impunctate area of the pronotum is much less than its total width with a punctate channel on either side of it (Fig. 2); the prosternum and its process are weakly convex; the posterolateral angles of the last abdominal sternum are acute (Fig. 4); and the juncture of the parameres is acutely angulate (Fig. 6). Variation——The three males are closely similar. One paratype is the same size as the holotype, the other is 13:8 X 5.3 mm. The bifurcation of the frontal carina does not join the supraantennal ridges on one paratype. Distribution—Known only from Imperial and San Bernardino Counties. Hosts.—One specimen was reared from ‘“‘desert willow”, Chilopsis linearis (Cav.) Sweet (confirmed by the collector); one was collected on willow, Salix sp.; and one was found flying around cottonwood, Populus sp. The other species in this genus are associated either with Salix or Populus, so the rearing from Chil- opsis is remarkable. Etymology.—The specific name is derived from the green-blue color. ACKNOWLEDGMENT Sincere thanks is extended to the following for the loan of types and/or spec- imens in their care: California Academy of Sciences, R. Brett and D. Kavanaugh; University of Arizona, C. A. Olson, and to T. C. MacRae, Chesterfield, Missouri and R. L. Westcott, Salem, Oregon for helpful suggestions, R. A. Flock for spec- imens, Juliette Chitjian Wright for typing the manuscript, and Joe Marilo for the photographs. LITERATURE CITED Arnett, R. H., Jr, G. A. Samuelson, and G. M. Nishida. 1993. The insect and spider collections of the world (2nd ed.): Flora & Fauna Handbook No. 11, Sandhill Crane Press, Gainesville, Florida. Evans, D. 1957. A revision of the genus Poecilonota in America north of Mexico (Coleoptera: Buprestidae). Ann. Entomol. Soc. Amer., 50: 21-37. Received 26 Oct 1995; Accepted 22 Apr 1996. PAN-PACIFIC ENTOMOLOGIST 73(1): 4-8, (1997) EXPERIMENTAL ARENA FOR CONFINING THRIPS AND OTHER SMALL ARTHROPODS IN THE LABORATORY BROOK C. MurpHy, JANE ADAMS AND MICHAEL P. PARRELLA Department of Entomology, University of California, Davis, California 95616 Abstract—A new experimental arena to confine western flower thrips, Frankliniella occidentalis (Pergande) and other small arthropods in order to conduct laboratory bioassays and behavioral studies is described. Arenas are constructed from cardboard paper cartons using a clear plastic petri dish lid and weather stripping to form an insect tight seal. A water pic inserted into the side of the arena is used to maintain plant material within the arena. The configuration of the arenas allows for direct visual observation of arthropods on portions of whole plants using a dissection microscope while leaving the arena sealed. The arenas successfully confined western flower thrips adults for 14 d and survival rates in the arenas over 7 d were at levels acceptable for check treatments in laboratory bioassays. Key Words.—Insecta, experimental arena, bioassay, Frankliniella occidentalis Thrips are a major pest of ornamental and food crops and their pest status seems to be increasing on a wide range of crops (Mound & Teulon 1995). Plants are damaged by direct feeding on foliage, fruit and flowers and by thrips vectored disease. Management strategies have relied almost exclusively on repeated chemical applications. Intensive use of chemical controls has resulted in widespread devel- opment of resistance by economically important species such as the western flower thrips, Frankliniella occidentalis (Pergande) (Immaraju et al. 1992, Robb et al. 1995). As a result, there has been an increased interest in the development of alternative control agents such as predators, parasitoids and pathogens for thrips management in addition to evaluation of new biorational pesticides for thrips con- trol. The small size and intense activity of thrips, particularly adults, has made con- finement and treatment in experimental arenas for laboratory bioassays challeng- ing. Most researchers have relied on Munger cells (Munger 1942) or modifications thereof (Tashiro 1967, Morse et al. 1986) to conduct bioassays of chemical pes- ticides against thrips and other small arthropods. Typically, a known number of arthropods are confined on a leaf surface within the cells, exposed to a chemical pesticide and monitored for mortality over time. However, use of Munger cells has several disadvantages: their small size allows only a few insects to be assayed for extended periods of time; maintaining leaf vigor during the experiment can limit the length of bioassays; direct application of chemical treatments to thrips and other arthropods can be difficult; efficacy can only be determined in a sim- plified environment (a flat leaf surface); and realistic temperature and relative humidity conditions are difficult to manipulate within the cells. Nonetheless, use of Munger cells has proven to be a useful standard technique for evaluating pesticides against thrips and other small arthropods. The limitations of Munger cells reduces their usefulness for evaluations of alternative control agents such as microbial insecticides, natural enemies and some biorational insecticides which act more slowly than conventional pesticides and therefore require longer observational times. Maintaining plant vigor through this 1997 MURPHY ET AL.: THRIPS TEST ARENA 5 interval is critical. Thus, bioassays of alternative agents conducted on simple leaf surfaces in small enclosed cells may not simulate actual mortality in a field setting occurring Over a 2 week time period and within complex environments. Here we describe an alternative experimental arena used to evaluate the per- formance of entomopathogenic fungi against western flower thrips, Frankliniella occidentalis. Bioassays using entomopathogenic fungi can take 7 d or more and the degree of effectiveness can be influenced by temperature and relative humidity conditions. Furthermore, the behavior of thrips on plants may determine the de- gree of exposure to the pathogen and influence the extent of fungal infection and mortality. Therefore, we designed arenas with greater volume that can enclose larger amounts of plant material (whole leaves, terminal shoots, buds and flowers). This allowed assays of larger numbers of thrips in a situation similar to that in which thrips are found in the field or greenhouse while minimizing influence of the experiment on thrips behavior. Through ventilation of the containers we also achieved greater control over internal environmental conditions. As a result, ex- periments using these arenas are able to assess the efficacy of fungi on a wide range of different plant parts and under temperature and relative humidity regimes that more closely simulate field conditions. MATERIALS AND METHODS Construction.—Components of an experimental arena are shown in Fig. 1. Units are constructed using a pint (473 ml) paper can (Fonda Paper can, Fonda Group, P.O. Box 500953, St Louis, MO) cut to a height of 5.1 cm (Fig. la). Two 2.5 cm holes are cut in the sides of the paper can and covered with organdy polyester netting to allow for airflow. The top of the arena uses a 90 mm diameter plastic petri dish lid (100 X 15 mm petri dish, Fisher Scientific, Pittsburgh PA) lined with foam weather stripping (Fig, 1b). Two rubber bands compress the lid to the top of the paper can to form a seal preventing thrips from escaping. A 5 ml plastic specimen tube (Rohre tube, Sarstedt, Germany) is used as a water pic to support plant material (Fig. 1c). The cap of the specimen tube is mounted into the side of the paper can by cutting a hole just large enough so the cap can be inserted through the can and sealed using silicon glue. The tube can then be inserted into the cap mounted into the side of the can which operates as a water reservoir (Fig. 1d). A stem, branch or petiole can be inserted through a hole drilled in the cap on the inside of the arena. Replenishing water from the outside is inserted through a water hole drilled in the tube. Evaluation—The experimental arenas were evaluated for their ability to 1) successfully confine thrips for 14 d without escape or significant increases in adult mortality, 2) demonstrate and quantify variation in survival rates of adult thrips on rose foliage in the arenas over a 7 d period, and 3) monitor the internal temperature and relative humidity to determine the influence of arenas on envi- ronmental conditions. Adult thrips of various ages were collected from a thrips colony maintained in a greenhouse on potted chrysanthemums. For the confine- ment test 43 to 54 adults were placed in each of seven arenas, 4 arenas contained rose foliage and 3 contained chrysanthemum foliage. The arenas were maintained at 26° C and 60% RH in an environmental chamber during the course of the tests. For the survival tests 23 to 53 adults were placed in arenas containing rose foliage and held at 26° C and 75% RH for 7 d. The tests compared thrips survival rates 6 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(1) A. Arena Body 0.5 Pint Paper Container Air Hole With Organdy Polyester Netting Lid B Black Weather Stripping Plastic Lid |—___- 9.0 cm —— --| 5ml Rohre Specimen Tube Cs [76 cm Ul Tp) TubeCap Water Hole Assembled Arena Figure 1. Schematic of experimental arena components, a) arena body, b) lid, c) specimen tube and d) assembled arena. among five arenas within each experiment and 3 different experiments were per- formed on different dates. At the end of the experiments all thrips were removed from the arenas and the number of live and dead thrips recorded. Two other arenas containing plant material but no thrips were used to compare the influence of arenas on the temperature and relative humidity within the containers using a digital temperature and relative humidity meter (Fisher scientific, model 1055712). The expected background mortality of adult thrips over the 14d confinement trial and 7 d survival trial was estimated using longevity data for adult thrips reared 1997 MURPHY ET AL.: THRIPS TEST ARENA i Figure 2. Photograph of experimental arena test units showing rose host material for assays using entomopathogenic fungi against thrips. on chrysanthemum in environmental chambers from Robb (1989). At 26° C adult longevity is approximately 32 d. Assuming a normally distributed age distribution of adults was obtained from the colony, an expected mortality of approximately 44 percent at 14 d and 22 percent at 7 d was estimated and compared to actual mortality in the arenas. Statistical analyses were performed using f¢ tests. RESULTS AND DISCUSSION The experimental arenas successfully contained and maintained adult thrips during the 14 d period. For the containment test an average (+ SD) of 46.85 + 1.42 thrips were placed in the arenas and after 14d an average of 45.85 + 1.29 were recovered demonstrating a 97.9 + 1.16 percent recovery rate. Adult mor- tality during this period averaged (+ SD) 39.6 + 4.8 percent which was not significantly different from the expected 44 percent mortality (¢ = 0.92, df = 6, P < 0.05). Thus the results demonstrated thrips could be successfully confined within experimental arenas for extended periods of time without adversely af- fecting survival rates. The average (+ SD) percent adult thrips mortality for the 3 survival test experiments were 12.1 + 2.9, 11.3 + 0.75 and 19.2 + 4.3 percent after 7 d. Thrips mortality was significantly lower than expected for the first two trials (¢ = 3.41, df = 4, P < 0.05; t = 14.26, df = 4, P < 0.05) and not significantly different from expected for the third trial (¢ = 0.65, df = 4, P > 8 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(1) 0.05). Results demonstrated that efficacy trials could be conducted over a 7 d period without unexpected mortality and at survival rates that would be acceptable for check treatments in laboratory bioassays. Temperatures and relative humidity inside the arenas averaging 26.5° C, and 64 percent respectively, were found to be slightly higher than the environmental chamber of 26° C and 60 percent RH. Based on these results we have concluded that the arenas may be applicable for conducting bioassays using fungi and other microbial or biorational pesticides against thrips for at least 7 d in length. The depth of the container allows for a full range of observation of insect subjects using dissecting microscopes or through visual observation. We have found that dead insects tend to drop from the plant material and can be easily counted against the white background of the bottom of the container. Mortality as a function of time can therefore be deter- mined without opening the containers. Ventilation of the containers allows for similar internal environmental conditions to outside conditions and the containers are versatile enough to conduct bioassays on many different host plants and host plant parts. The arenas can be constructed rapidly and the materials are inexpen- sive and readily available. We believe the use of this design may provide a more realistic assessment of mortality under conditions that more closely resemble field conditions. In addition to thrips, we have successfully used these arenas for conducting efficacy trials against aphids and mites and have begun to evaluate the effects of entomopathogenic fungi on thrips natural enemies. This technique may also have additional experimental applications such as in behavioral studies, survival and longevity studies and experiments examining plant-insect and predator-prey in- teractions. ACKNOWLEDGMENT We thank Tunyalee Morisawa (Department of Entomology, University of Cal- ifornia, Davis) for assistance in construction of experimental arenas and con- ducting survival tests. This work was funded in part by the University of Cali- fornia Division of Agriculture and Natural Resources Special grant number 074, the Mycotech Corporation, Butte MT and the American Floral Endowment. LITERATURE CITED Immaraju, J. A., T. D. Paine, J. A. Bethke, K. L. Robb & J. PR Newman. 1992. Western flower thrips (Thysanoptera: Thripidae) resistance to insecticides in coastal California greenhouses. J. Econ. Entomol., 85: 9-14. Morse, J. G., T. S. Bellows & Y. Iwata. 1986. Technique for evaluating toxicity of pesticides to motile insects. J. Econ. Entomol., 79: 281-283. Mound L. A. & A. J. Teulon. 1995. Thysanoptera as phytophagous opportunists. pp. 3-19. In L. Parker, M. Skinner & T. Lewis (eds.). Thrips Biology and Management. Plenum Press, NY. Munger, EF 1942. A method for rearing citrus thrips in the laboratory. J. Econ. Entomol., 35: 373-375. Robb K. L., 1989. Analysis of Frankliniella occidentalis (Pergande) as a pest of floriculture crops in California greenhouses. Ph.D Dissertation, University of California, Riverside. Robb, K. L., J. Newman, J. K. Virzi & M. P. Parrella. 1995. Insecticide resistance in western flower thrips. pp. 341-346. Jn L. Parker, M. Skinner & T. Lewis (eds.). Thrips Biology and Manage- ment. Plenum Press, NY. Tashiro, H. 1967. Self-watering acrylic cages for confining insects and mites on detached leaves. J. Econ. Entomol., 60: 354—356. Received 16 May 1996; Accepted 20 Aug 1996. PAN-PACIFIC ENTOMOLOGIST 73(1): 9-15, (1997) AN EXAMINATION OF SPATIAL INPUT PARAMETERS IN ORDER TO IMPROVE CORN EARWORM (LEPIDOPTERA: NOCTUIDAE) DAMAGE PREDICTIONS FOR A PHEROMONE TRAP CATCH REGRESSION MODEL R. J. DRAPEK, B. A. CROFT, AND G. FISHER Department of Entomology, Oregon State University, Corvallis, OR 97331-2907 Abstract—A regression model relating cumulative pheromone trap catch and date of corn silking to subsequent damage in sweet corn by Helicoverpa zea (Boddie) was improved in one year by including additional spatial input parameters. Spatial inputs included information from timings and locations of corn plantings around the trap as well as from locations of wind blocking features. Wind blocking features were: tree rows, wooded areas, large buildings close to the trap, and abrupt hillsides. Pheromone traps were monitored at 28 and 30 sites in 1990 and 1991. Corn development through the year and damage levels at harvest (percent infestation) were also re- corded for these locations. Maps of all corn plantings and wind blocking features within 2.5 kilometers of the trap were created, digitized, and entered into a Geographical Information Sys- tem for each site. A stepwise regression analysis considering 18 spatial and two non-spatial variables resulted in a highly significant (P < 0.001) regression model with four variables (trap catch, silking date, the number of hectares of corn within 2.0 kilometers of the trap, and the average distance to wind blocking features on the north side of the trap) explaining 82% of the variability for the 1990 data. In 1991, of the 20 input parameters considered only one was significant, date of first silk. An extremely low corn earworm population was considered to have caused this inability to find correlations for the other parameters in 1991. Key Words.—Insecta, Helicoverpa zea, corn earworm, sweet corn, pheromone, trapping, moni- toring In the Willamette Valley of western Oregon, Helicoverpa zea (Boddie) popu- lations are low but periodically cause extensive damage to corn which is valued at $45 million per annum, statewide (Miles 1992). Earworm damage is usually limited to tips of ears, with older larvae tending to feed farther down the ear (Coop et al. 1992). This damage is intolerable on fresh market sweet corn where pesticides routinely are applied. Corn processors remove the tips of the ears whether or not earworm damage is present. Therefore earworm is only a problem in processed corn when populations are much larger than normal. In 1985, fol- lowing a year of severe earworm damage, a study was initiated to provide pest managers with the means to predict earworm damage on processed corn. Based on 5 years of pheromone trapping, a two variable regression model was developed that related cumulative pheromone trap catch from first tassel to first silk as well as the date of first silk to subsequent percent ears infested by H. zea (Coop et al. 1992). In this study, additional input variables were considered for inclusion into the regression model. The inclusion of these variable is based on two hypotheses: 1) that wind blocking features potentially interfere with normal wind flow around the trap or act to block pheromone plumes downwind of the trap and 2) that the amount, age, and position of corn plantings around the pheromone trap will affect 10 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(1) moth populations in the region around the trap as well as moth movement dy- namics. Therefore, positions and timings of corn plantings around the trap as well as positions of wind blocking features were considered as additional input vari- ables. MATERIALS AND METHODS Pheromone traps were placed in corn plantings prior to tasselling in 1990 and 1991. Traps were inspected every 3 d to record the number of moths trapped. Traps were a standard Texas wire cone trap (75—50 of Hartstack et al. 1979) with a Scentry pheromone cap containing Z-11-Hexadecenal, Z-9-Hexadecenal, Hex- ane, and Tenox 4. This trap/lure combination is one of the most cost effective monitoring tools available for H. zea (Drapek et al. 1990, Gauthier et al. 1991). In 1990 and 1991, 28 and 30 plantings were trapped respectively. Plantings cov- ered an area about 80 km. north to south by 40 km. east to west and consisted primarily of the variety ““Golden Jubilee’’. Traps were placed, when possible, 75 m within corn fields in accessible loca- tions. In some fields, irrigation equipment prevented placement of traps within the field. Instead, traps were placed as close to the edge of the field as possible. Witz et al. (1992) found that at low populations of H. zea (less than 50 moths per trap-night), traps placed within 5 m of the perimeter of cotton fields caught as many moths as interior traps. Willamette Valley trap catches seldom exceed 50 moths in a night. Traps were situated without regard to wind direction as previous tests had shown little effect of wind-side on trap catch (Drapek et al. 1990). When traps were placed in plantings prior to tasselling, they were set so that the bases were about 1.5 m from the ground. As the corn grew to the height of the trap base, the trap was raised so that the pheromone cap at the base of the trap was within 20 cm of the corn plants. With each checking of a pheromone trap, the stage of corn development was recorded. Corn plantings and wind blocking features within 2.5 km radius were mapped for every trap location. Wind blocking features included large buildings within 200 m of the trap, windrows, wooded areas, and large hills. For each mapped site, the mapping date and the stage of every corn planting were noted. Every feature was mapped using a compass to obtain directions and pacing to obtain distances (Figure 1). Maps were checked against 1:24,000 USGS topographic maps and were digitized to a geographical information system (GIS) for area calculations. Mature ears within study fields were collected as close to harvest date as pos- sible and inspected for earworms and damage. As plantings approached harvest, a 200 ear random sample was taken from every planting. If the corn field was not harvested within 6 d, a second sample was taken. A stepwise regression was used which considered 20 independent variables and ear infestation level at harvest as the dependent variable. Independent variables included: 1) cumulative trap catch from first tassel to first silk; 2) date of first silk measured as the number of days since 31 December; 3—5) average distance to wind blocking features north of the trap, south of the trap, and all around the trap; 6—10) hectares of corn older than the trapped planting within 0.5, 1.0, 1.5, 2.0, and 2.5 km of the trap; 11-15) hectares of corn the same age as the trapped planting within 0.5, 1.0, 1.5, 2.0, and 2.5 km. of the trap; and 16—20) hectares of 1997 DRAPEK ET AL.: IMPROVED CORN EARWORM MODEL 11 Etzel 2 | Coordinates of Trap (Center; in UTM) :\807910E 4951479N | Figure 1. Example map of one of the monitored plantings. Shaded polygons indicate locations of corn plantings. Numbers adjacent to the polygons indicate cumulative degree-day development as of August 1. The large unshaded polygon shows all locations within 2.5 kilometers of the trap that have no wind-blocking feature between them and the trap. all corn plantings within 0.5, 1.0, 1.5, 2.0, and 2.5 km. of the trap. The dependent variable was an arcsine-square-root transformation of the proportion of ears dam- aged. This transformation is suggested for variance stabilization by Weisberg (1985) for dependent variables that are binomial proportions and was used by Coop et al. (1992). A 0.5 km/radius around the trap was used to calculate average distances to wind blocking features because it was assumed that wind blocking features be- yond 0.5 km. would have minimal effect on the pheromone plume. The average was obtained by taking the mean distance to wind blocking features for radius lines drawn from the trap at 10° intervals. Average distances were calculated for each site for locations north, south, and all around the trap. A north-south distinction was made because early evening winds come predominantly from the north during summers in the Willamette Valley (Coop and Croft, 1995). Hectares of corn within the various radii away from the trap were calculated using the GIS. All plantings that at some point in the season were silking at that same time as the trapped field were called “‘same age’’ for the purposes of the model. ‘‘Silking”’ is defined as that period of time starting when silks could be observed on a substantial minority of corn stalks (~ 5%) and ending when a substantial minority (~ 5%) of stalks had silks with distinct browning (~ 50% of silks on an ear being brown). 12 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(1) 25 20 1S 10 Moths/Trap-Day - Teele tame ewee sentra ee" ) —EE = 5/8 17 26 6/4 13 22 7/1 10 19 28 8/6 15 24 9/2 11 20 Date Figure 2. Average moth catch per trap-day observed in 1990 and 1991. RESULTS Moth flight patterns as measured by pheromone trap catches did not display any unusual patterns in 1990 or 1991 (Fig. 2), compared to 1986-1988 (Coop et al. 1992, Drapek 1993). For both years a peak in catch started in early August, peaked in late August or early September, and dropped again towards the end of September. The first moths were captured on 8 May 1990 and 10 June 1991. The June 10 date was unusually late, reflecting a wet and cool spring. For both years the silking date proved to be a significant input variable (Table 1). Trap catch was a significant variable in 1990, but not in 1991. Significant Table 1. Models resulting from a stepwise regression considering 2 nonspatial and 18 spatial input parameters as predictors of percent ear infestation by Helicoverpa zea*. Year # Sites Variable Reg. coeff. SE F Prob > F 1990° 30 Intercept — 0.6026 0.2527 5.68 0.025 Ist Silk° 0.0039 0.0012 11.17 0.003 Corn Area 0.0004 0.0002 6.41 0.018 WBF* —0.1083 0.0587 3.40 0.077 Trap Catchf 0.0018 0.0003 39.96 <0.001 19918 27 Intercept — 0.9062 0.3226 —2.81 0.010 Ist Silk* 0.0047 0.0015 3.21 0.004 4 Dependent variable = Sin~'(Sqrt (proportion infested ears)). > R? = 0.82. Model P = 0.000. © 1st Silk is the date on which silking is first noted (measured as the number of days since Jan. 1). 4 Corn Area is the number of hectares of corn within 2.0 kilometers of the trap. © WBE is the average distance on the north side of the trap to a wind blocking feature. f Trap Catch is the cumulative trap catch from Ist tassel to 1st silk. 8 R? = 0.29. Model P = 0.004. 1997 DRAPEK ET AL.: IMPROVED CORN EARWORM MODEL 13 + 1990 x 1991 Percent Ears Infested e aa 0 20 40 60 80 100 120 140 160 180 200 Cumulative Trap Catch Figure 3. Trap catch and damage levels observed for individual sites in 1990 and 1991. Trap catch is the cumulative moth catch between lst tassel and Ist silk. Damage in measured as percent of ears infested with earworm at the time of harvest. input from spatial variables occurred in 1990, but not in 1991. The significant variables were corn area within 2.0 km of the trap and average distance on the north side of the trap to wind blocking features. Where trap catch and date of first silk were significant, the regression coefficients were positive. DISCUSSION The optimal multiple regression model obtained from 1990 data included four input variables: date of 1st silk, the number of hectares of corn within 2 km of the trap, the average distance north of the trap to wind blocking features, and cumulative trap catch. Date of Ist silk had a positive regression coefficient. This means that later silking corn plantings can be expected to have higher damage levels. This may result from timings of moth flight relative to corn development. Peak moth flights tended to be somewhat later than peak corn silking dates (Dra- pek et al. 1993). High trap catches imply large moth populations. Large moth populations imply that oviposition and subsequent damage should be high as well. Therefore, our expectation was that trap catch and damage would have a significant positive correlation. Moth flights and damage were so low both years, that the results did not meet our expectations (Figure 3). Coop et al. (1993) observed that corn ear- worm-caused damage did not equal control costs until infestation levels ap- proached 30% or higher. Over the 2 years we monitored earworms, infestation rates only reached that level for one planting, and all but 2 plantings had infes- tation levels less than 15%. Both plantings occurred in 1990. These 2 plantings were very influential in producing the significant regression coefficients between trap catch and damage in 1990. Because no plantings had such damage levels in 14 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(1) 1991, the relationship between trap catch and damage did not prove to be signif- icant for that year. The positive relationship between corn earworm trap catch, moth activity lev- els, and subsequent damage has been verified for other locations and years. Lath- eef et al. (1993) were able to demonstrate a significant correlation between daily pheromone trap catches and moth flights (both male and female) as measured by hand-net. Additionally, Latheef et al. (1991) observed a weak, but significant linear relationship between the number of eggs laid per day versus the daily pheromone trap catch. The positive trap catch versus subsequent damage rela- tionship concurs with observations made by Chowdhury et al. (1987) and Coop et al. (1992). In 1990, both spatial variables (corn area and average distance to north wind- blocking features) had negative regression coefficients. The negative coefficient for distance to wind-blocking features implies that the farther away from the trap north wind-blocking features occur, the lower should be the damage prediction. One explanation for this observation is that wind-blocking features interfere with the normal movement of the plume. In the Willamette Valley, evening winds come predominantly from the north during the summer (Coop & Croft, 1995). Therefore wind-blocking features on the north side of a field will probably have the most disruptive effect on the pheromone plume. In 1991 only the date of 1st silk proved to be significant as an input variable for prediction of percent earworm infestation. The regression coefficient for date of Ist silk was positive. The fact that trap catch and none of the potential spatial input parameters proved significant in 1991 does not necessarily invalidate them as useful model parameters. Earworm population and damage levels were unusu- ally low in 1991. Though the correlation between trap catch and damage levels for individual sites was low, the low trap catch levels observed for all sites did accurately reflect the low damage levels observed valley-wide. Anyone using pheromone traps in 1991 to monitor for earworms would have correctly deter- mined that earworms would not be a problem that year. When damage levels are so low, normal variation nullifies all but the strongest trends. Future observations from years with moderate to high earworm populations are required before a final evaluation can be made on the reliability of adjacent corn acreage and distance to wind blocking features as damage prediction modifying inputs. LITERATURE CITED Chowdhury, M. A., R. B. Chalfant & J. R. Young. 1987. Ear damage in sweet corn in relation to adult corn earworm populations. J. Econ. Entomol., 80: 867-869. Coop, L. B., R. J. Drapek, B. A. Croft & G. C. Fisher. 1992. Relationship of corn earworm pheromone catch and silking to infestation levels in Oregon sweet corn. J. Econ. Entomol., 85: 240-245. Coop, L. B., B. A. Croft & R. J. Drapek. 1993. Model of corn earworm development, damage, and crop loss in sweet corn. J. Econ. Entomol., 86: 906-916. Coop, L. B. & B. A. Croft. 1995. Neoseilus fallacis: dispersal and biological control of Tetranychus urticae following minimal inoculations into a strawberry field. Exper. & Appl. Acarol., 19: 31-43. Drapek, R. J., L. B. Coop, B. A. Croft & G. C Fisher. 1990. Heliothis zea Pheromone trapping: studies of trap and lure combinations and field placement in sweet corn. S. W. Entomol., 15: 63-70. Drapek, R. J. 1993. Use of a geographical information system to modify pheromone trap-based predictions of Helicoverpa zea (Boddie) damage. PhD. Thesis. Oregon State University. Gauthier, N. L., P. A. Logan, L. A. Tewksbury, C. E Hollingsworth, D. C. Weber & R. G. Adams. 1997 DRAPEK ET AL.: IMPROVED CORN EARWORM MODEL 15 1991. Field bioassay of pheromone lures and trap designs for monitoring adult corn earworm in sweet corn in southern New England. J. Econ. Entomol., 84: 1833-1836. Hartstack, A. W., J. A. Witz & D. R. Buck. 1979. Moth traps for the tobacco budworm. J. Econ. Entomol., 72: 519-522. Latheef, M. A., J. A. Witz & J. D. Lopez Jr. 1991. Relationships among pheromone trap catches of male corn earworm moths, egg numbers, and phenology in corn. Can. Entomol., 123: 271- 281. Latheef, M. A., J. D. Lopez Jr. & J. A. Witz. 1993. Capture of corn earworm in pheromone traps and hand nets: relationship to egg and adult densities in field corn, Texas Brazos River Valley. J. Econ. Entomol., 86: 407-415. Miles, S. D. 1992. 1991 Oregon County and State Agricultural Estimates. Oregon States University Extension Service. Special Report 790. Weisburg, S. 1985. Applied Linear Regression. John Wiley & Sons. New York. Witz, J. A., J. D. Lopez & J. L. Goodenough. 1992. Influence of sex pheromone trap placement relative to field edge on catch of bollworm males in cotton. S. W. Entomol., 17: 1-6. Received 6 Jun 1996; Accepted 7 Sep 1996. PAN-PACIFIC ENTOMOLOGIST 73(1): 16-20, (1997) CHRYSOMYA MEGACEPHALA (FABR.) IS MORE RESISTANT TO ATTACK BY CH. RUFIFACIES (MACQUART) IN A LABORATORY ARENA THAN IS COCHLIOMYIA MACELLARIA (FABR.) (DIPTERA: CALLIPHORIDAE) JEFFREY D. WELLS! AND HIROMU KURAHASHI Department of Medical Entomology, The National Institute of Health, Toyama 1-23-1, Shinjuku-ku, Tokyo 162, Japan Abstract——Chrysomya Robineau-Desvoidy blow flies recently introduced to the Americas in- clude two species, Ch. megacephala (Fabr.) and Ch. chloropyga Wiedemann (= Ch. putoria) with purely saprophagous larvae, and two, Ch. albiceps (Wiedemann) and Ch. rufifacies (Mac- quart), that are facultative predators on other maggots. Patterns of adult abundance suggest that the invading species suppress the saprophagous native Cochliomyia macellaria (Fabr.), and do so more effectively in combination than individually. We hypothesized that Ch. megacephala, historically sympatric with Ch. mufifacies, is relatively resistant to predation by Ch. rufifacies, which could provide it with a competitive advantage over a more vulnerable C. macellaria when larvae of all three occur together. To test this hypothesis, larvae of both prey species were individually paired with larvae of Ch. rufifacies in the laboratory. C. macellaria were consistently killed at a higher rate than were Ch. megace phala. Key Words.—Insecta, biological invasion, introduced species, competitive displacement, higher- order interaction, carrion, blow fly Old World blow flies in the genus Chrysomya Robineau-Desvoidy have been spectacularly successful following their recent invasion of the Western Hemi- sphere. Introduced at two locations in the mid 1970’s, Ch. albiceps (Wiedemann), Ch. chloropyga Wiedemann (= Ch. putoria) and Ch. megacephala (Fabr.) in Brazil (Guimaraes et al. 1979) and Ch. rufifacies (Macquart) in Costa Rica (Jir6n 1979), they quickly became widespread and abundant in Latin America (Baum- gartner & Greenberg 1984, Baumgartner 1988, Mariluis & Schnack 1989, Olsen et al. 1992., J. Mendez L., pers. comm., Kurahashi et al. 1994). Within two decades, Ch. rufifacies and Ch. megacephala had spread far enough to be firmly established at locations in the southern USA (Wells 1991, Baumgartner 1993), and Ch. chloropyga occurs as far north as Panama (J. Mendez L., personal comm.). Chrysomya chloropyga and Ch. megacephala are typical synanthopic pests, with saprophagous larvae usually found in carrion or feces (Greenberg 1971, Laurence 1986). Chrysomya albiceps and Ch. rufifacies, so similar to each other in form and natural history that their status as separate species has been debated (Tantawi & Greenberg 1993), have larvae that eat both carrion (and rarely live flesh) and other maggots (Fuller 1934, Ullyett 1950). The latter two species are also distinguished by the presence of prominent spiny tubercles (Fig. 1), which we believe serve to reduce cannibalism. As Chrysomya densities in the New World have increased, sympatric popula- ‘Current Address: Dept. ESPM, Div. Insect Biology, University of California, Berkeley, CA 94720. 1997 WELLS & KURAHASHI: RESISTANCE TO PREDATION BY CHRYSOMA_ 17 Figure 1. Larva of Chrysomya rufifacies (with tubercles) attacking a larva of Phormia regina (Meigen). The head of C. rufifacies, inserted into the other larva, is down in this view. 18 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(1) tions of the native calliphorid Cochliomyia macellaria Fabr. have decreased, sometimes precipitously, and this has been interpreted as competitive displace- ment (Guimaraes et al. 1979, Baumgartner & Greenberg 1984, Wells & Greenberg 1992, Paraluppi & Castellon 1994). A similar decline may be happening to Lucilia exima (Wiedemann) (Baumgartner 1993). Competition for food between carrion- fly larvae is often intense (Hanski 1987). Because the invaders don’t seem to be filling any previously unexploited niche, i.e., they are eating the same carrion and feces that are already exploited by native flies, the ability to out-compete native Species seems necessary for the success of Chrysomya in the Americas. Some evidence indeed suggests that the Chrysomya spp. have a stronger neg- ative effect on Co. macellaria in combination than individually. At a site in Peru where Ch. chloropyga became common but Ch. albiceps was rare, Co. macellaria dropped during a 4-year period from 46% to 11% of the adult population at baits, while at a site where both invaders were common, reduction of the native fly was from 89% to 0.2% (Baumgartner & Greenberg 1984). At Brazilian locations where all three Chrysomya are abundant, the previously common Co. macellaria has been described as rare or absent (Guimaraes et al. 1979, Paraluppi & Castellon 1994). Ch. rufifacies is the only member of the genus known to be established in Texas, and although it was experimentally shown to reduce the number of Co. macellaria bred from carrion, the native fly is still abundant (Wells & Greenberg 1992, 1994). The advance of these flies is in contrast to previous (and separate) introductions to Latin America of Ch. megacephala, Ch. chloropyga and Ch. rufifacies that failed (Baumgartner & Greenberg 1984, Baumgartner 1993). It is rather difficult to determine why an invasion did or did not succeed, but we hypothesize that the appearance of several Chrysomya spp. at the same time at least contributed to their successful establishment. Others have observed that coevolved sets of intro- duced species can be more able to invade because they have a greater impact on the invaded community than would be predicted from their individual interactions with native organisms (Simberloff 1991). In the case of Chrysomya, this would occur if the purely saprophagous larvae are adapted to resist or avoid the attack of their predaceous congeners with which they have long been sympatric. Chry- somya megacephala or Ch. chloropyga would then be more likely to successfully invade if Ch. albiceps or Ch. rufifacies were also present. Chrysomya megacephala is commonly found with Ch. rufifacies in the Oriental, Autralsasian and Oceanic regions (James 1977, Kurahashi 1989). In addition, its successional position within carrion is similar to Co. macellaria (making them almost certainly competitors for the same food), and both are species attacked by Ch. rufifacies in the field (Bohart & Gressitt 1951, Wells & Greenberg 1994). In this study, we measured the rate of predation by Ch. rufifacies larvae on Ch. megacephala and on Co. macellaria in a laboratory arena. METHODS AND MATERIALS All larvae used were third instars approximately one cm in length. During a trial, 20 Co. macellaria and 20 Ch. megacephala were individually paired with a single Ch. rufifacies (40 total) within a 60 X 15 mm plastic petri dish. Dishes were arranged in a 5 by 8 pattern on a laboratory shelf, with alternating prey Species in place. A trial began when a Ch. rufifacies larva was quickly dropped 1997 WELLS & KURAHASHI: RESISTANCE TO PREDATION BY CHRYSOMA_ 19 Table 1. Number of Cochliomyia macellaria and Chrysomya megacephala larvae, out of a total of 20 each, successfully attacked by Chrysomya rufifacies. Trial Co. macellaria Ch. megacephala 1 18 15 2 18 14 3 18 14 4 5 2 5 17 16 6 9 6 into each dish. After 20 min at 25 C, all larvae that, based on our experience, were damaged enough to be fatally wounded were counted. These included larvae that were shriveled, had body contents extruded through a hole in the cuticle, or were in the grip of a feeding Ch. rufifacies. Six trials were performed, each with a new generation of larvae. Larvae were obtained from two sets of fly colonies. For trials 1-4 these were: Co. macellaria from Kerr County, TX, USA, colonized for an unknown number of generations; Ch. megacephala and Ch. rufifacies from Yona, Okinawa, Japan, colonized for 3—10 generations. Colonies for trials 5 and 6 were: Co. macellaria from W. Lafayette, IN, USA, colonized for 4-5 generations; Ch. megacephala from Kimbe, New Britain, Papua New Guinea, colonized for 10—11 generations; Ch. rufifacies from Matsuda, Okinawa, Japan, colonized for 3—4 generations. RESULTS AND DISCUSSION In every trial, Ch. rufifacies killed or wounded a greater number of Co. ma- cellaria than Ch. megacephala (Table 1). Under these conditions, Ch. mega- cephala was more resistant to attack by its historically sympatric congener than was the previously allopatric Co. macellaria (sign test, p = 0.03). From casual observation, we believe that Ch. megacephala was more quick to struggle vig- orously and flee following contact with the mouthparts of Ch. rufifacies, although no effort was made to quantify such behavior. Although the total number killed per trial varied from in 7 to 33, the relative difference between the two prey species was similar for all trials. We suspect that each batch of Ch. rufifacies had a particular “hunger level’’ that influenced the overall probability and/or strength of attack, but that had no influence on the relative vulnerability of prey species. These results, albeit produced in a highly artificial setting, support the hypoth- esis that Ch. megacephala is a stronger competitor against Co. macellaria when Ch. rufifacies is present. To the extent that our observations apply to wild pop- ulations, it follows that the establishment and spread of Ch. megacephala within the range of Co. macellaria was aided by the presence of Ch. rufifacies or the nearly identical Ch. albiceps. Further experimental manipulations and field ob- servations are needed to confirm this complex interaction. ACKNOWLEDGMENT Ichiro Miyagi, University of the Ryukyus, Bernard Greenberg, University of Illinois at Chicago and S. Adam Shahid, Purdue University, kindly helped us to 20 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(1) obtain our fly colonies. JOW was supported by a fellowship from the National Science Foundation (INT-9311789) and the Japanese Science and Technology Agency (193040). LITERATURE CITED Baumgartner, D. L. 1988. Spread of introduced Chrysomya blowflies (Diptera, Calliphoridae) in the Neotropics with records new to Venezuela. Biotropica, 20: 167—168. Baumgartner, D. L. 1993. Review of Chrysomya rufifacies (Diptera: Calliphoridae). J. Med. Entomol., 30: 338-352. Baumgartner, D. L. & B. Greenberg. 1984. The genus Chrysomya (Diptera: Calliphoridae) in the New World. J. Med. Entomol., 21: 105-113. Bohart, G. E. & J. L. Gressitt. 1951. Filth-inhabiting flies of Guam. Bull. Bernice P. Bishop Mus., 204: 1-169. Fuller, M. E. 1934. The insect inhabitants of carrion: a study in animal ecology. Aust. Counc. Sci. Ind. Res. Bull., 82: 4-63. Greenberg, B. (ed.). 1971. Flies And disease. Vol. I. Ecology, classification and biotic associations. Princeton University Press, Princeton. Guimaraes, J. H., A. P do Prado & G. M. Buralli. 1979. Dispersal and distribution of three newly introduced species of Chrysomya Robineau-Desvoidy in Brazil (Diptera, Calliphoridae). Rev. Bras. Entomol., 23: 245-255. Hanski, I. 1987. Nutritional ecology of dung- and carrion-feeding insects. pp. 837-884. In Slansky, F & J. G. Rodriguez (eds.). Nutritional ecology of insects, mites, spiders and related inverte- brates. John-Wiley, New York. James, M. T. 1977. Family Calliphoridae. pp. 528-556. Jn. Delfinado, M. D. & D. E. Hardy (eds.). A catalog of the Diptera of the Oriental region. University Press of Hawaii, Honolulu. Jirén, L. EF 1979. Sobre moscas califoridas de Costa Rica (Diptera: Cyclorrhapha). Brenesia, 16: 221— 227; Kurahashi, H. 1989. Family Calliphoridae. pp. 702-717. In. Evenhuis, N. L. (ed.). Catalog of the Diptera of the Australasian and Oceanian regions. Bishop Museum Press, Honolulu. Kurahashi, H., J. D. Wells & K. Ogino. 1994. The Oriental latrine fly, Chrysomya megacephala (Fabricius) (Diptera), newly recorded from Honduras, Central America. Jpn. J. Entomol., 62: 860. Laurence, B. R. 1986. Old World blowflies in the New World. Parasit. Today, 2: 77-79. Mariluis, J. C. & J. A. Schnack. 1989. Ecology of the blow flies of an eusynanthropic habitat near Buenos Aires (Diptera, Calliphoridae). EOS, 65: 93-101. Olsen, A. R., S. C. Angold, D. E Gross & T. H. Sidebottom. 1992. New record of the blowfly, Chrysomya megace phala (Fabr.), from Ecuador. Pan-Pac. Entomol., 68:280—281. Paraluppi, N. D. & E. G. Castellon. 1994. Calliphoridae (Diptera) em Manaus: I. Levantamento taxonomica e sazonalidade. Rev. Bras. Entomol., 38: 661—668. Simberloff, D. 1991. Keystone species and community effects of biological introductions. pp. 1-19. In Ginzbeg, L. R. (ed.). Assessing ecological risks of biotechnology. Butterworths-Heinemann, Stoneham, MA. Tantawi, T. I. & B. Greenberg. 1993. Chrysomya albiceps and C. rufifacies (Diptera: Calliphoridae): contribution to an ongoing taxonomic problem. J. Med. Entomol., 30: 646-648. Ullyett, G. C. 1950. Competition for food and allied phenomena in sheep-blowfly populations. Phil. Trans. Roy. Soc. Ser. B., 234: 77-174. Wells, J. D. 1991. Chrysomya megacephala (Diptera: Calliphoridae) has reached the continental United States: review of its biology, pest status and spread around the world. J. Med. Entomol., 28: 471-472. Wells, J. D. & B. Greenberg. 1992. Interaction between Chrysomya rufifacies and Cochliomyia ma- cellaria (Diptera: Calliphoridae): the possible consequences of an invasion. Bull. Entomol. Res., 82: 133-137. Wells, J. D. & B. Greenberg. 1994. Resource use by an introduced and native carrion flies. Oecologia, 99: 181-187. Received 26 Apr 1996; Accepted 6 Aug 1996. PAN-PACIFIC ENTOMOLOGIST 73(1): 21-27, (1997) ECOLOGICAL STUDIES ON CARDIOCONDYLA ECTOPIA SNELLING (HYMENOPTERA: FORMICIDAE) IN SOUTHERN CALIFORNIA.! HANIF GULMAHAMAD 3547 Centurion Way, Ontario, California 91761 Abstract.—At this location, Cardiocondyla ectopia Snelling is a largely diurnal species which foraged throughout the year when ambient temperatures exceeded 18.9° C. Its major food source was nectar from sweet alyssum flowers, Lobularia maritima Desvaux. It preyed on insects, particularly small caterpillars, and it was also a scavenger. It nested in small cavities in mortar, cracks and expansion joints in concrete, and in soil. A colony was excavated from which 322 ants were retrieved. This represented the largest nest population ever recorded for a Cardiocon- dyla species. A number of behavioral strategies, in addition to a potent repellent chemical, probably allow C. ectopia to live in sympatry and synchrony with Linepithema humile (Mayr). Key Words.—Insecta, Formicidae, Cardiocondyla ectopia, foraging, food sources, nest sites, nest population, coexistence, Linepithema humile, Lobularia maritima Cardiocondyla ectopia Snelling was described in 1974 from specimens taken from Orange and Los Angeles counties, California (Snelling 1974). This species was also reported from Arizona (MacKay 1995). Although this species was re- corded from southern California, it has been rarely collected from this area. A survey of urban ants of California yielded 30 species but C. ectopia was not recorded (Knight & Rust 1990.) However, I collected this species on several occasions around structures in the cities of Downey and Long Beach (Los Angeles County), and Montclair and Ontario (San Bernardino County). This ant has been overlooked by structural pest control operators in southern California probably because of its small size, small colonies, cryptic nests, absence of trailing behav- ior, and its apparent inability to invade structures. All current biological information available on C. ectopia is contained in Creighton & Snelling (1974). Herein I provide additional information on C. ec- topia foraging behavior, food resource utilization, nest sites, nest population, and its coexistence with the Argentine ant, Line pithema humile (Mayr). MATERIALS AND METHODS I observed six colonies of C. ectopia located on my property in Ontario, San Bernardino County, California. Periodic observations, encompassing an hour or more at a time, began in January 1995 and continued through May 1996 as time permitted. On weekends, intermittent observations began as early as 08:00 h and continued for several hours at a time until 23:00 h. Most of the observations recorded here were made with an OptiVisor optical glass binocular magnifier (Donegan Optical Company, Kansas City, Missouri). From these observations, information was gathered on foraging behavior, food resource utilization, nest location, coexistence with the Argentine ant, Linepithema humile (Mayr), and ! Author page charges partially offset by a grant from the C. P. Alexander Fund, Pacific Coast Entomological Society. 22, THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(1) chemical defense strategy. On 25 Nov 1995, a colony of C. ectopia was excavated and the number of ants retrieved was recorded. RESULTS AND DISCUSSIONS Foraging Behavior.—Cardiocondyla ectopia is a largely diurnal species. Above ground activity of this ant is governed by temperature. Foraging activity in July 1972 was reported to begin when the ambient temperature ranges from 18.9 to 20 °C (Creighton & Snelling 1974). When asphalt temperature reached 26.1 °C foragers were no longer using the exposed surfaces (Creighton & Snelling 1974). I observed surface activity of this ant during the third and fourth weeks of De- cember 1995 when the concrete temperature was 15.6 °C (recorded with a surface temperature thermometer - Sybron Taylor Products, Arden, North Carolina). This activity was observed as early as 08:40 h when several workers and four alates were found around the nest entrance area. These ants were lethargic and a few appeared to be moribund. As the temperature increased ant activity increased. They may have been induced to emerge from this nest in an expansion joint in the slab because bright sunlight was shining directly on this area. Creighton & Snelling (1974) recorded some surface activity of this ant as late as 19:25 h in July 1972. During the last week of December 1995, foraging activity on sweet alyssum flowers, Lobularia maritima Desvaux was observed as late as 16:30 h. On 19 May 1996, foraging on these flowers was observed at 20:10 h. Some ant activity around the nest area was observed at 20:26 h on 19 May 1996 when observations had to be made with a flashlight as it was too dark to see these ants. Ant activities were observed as late as 23:05 hin July and August 1996 at unusual food sources such as dog food and soda. At this location, C. ectopia foraged throughout the year whenever temperatures were conducive to above ground ac- tivity. Cardiocondyla ectopia searched its foraging territory in a random manner. In- dividuals were found traversing areas as much as 6 m away from the nest. The colonies I observed foraged mostly on nectar from flowers of sweet alyssum. This landscape annual produces tiny white four-petaled flowers which are borne in clusters and the flowers emit a honeylike fragrance. Once a plant was located, C. ectopia foragers returned to this resource many times to exploit it. I observed numerous individuals from two colonies travelling directly to a clump of sweet alyssum located 3 m away from the nests. Tandem running and recruitment to a food source occurs in the genus Cardiocondyla (Wilson 1959a, Creighton & Snelling 1974). Cardiocondyla ectopia also exhibited this behavior, but it was not common. Associative learning appeared to be present in this species as these ants repeatedly returned to a productive floral food source by different routes over many days. I observed foragers returning to a location where I had removed a clump of flowers for up to two h after the plants were gone. Foragers of C. ectopia readily ascended various herbaceous plants and explored their surfaces. Many of these plants did not have flowers and it appeared that the ants were searching for other food sources, probably, extrafloral nectaries, glan- dular plant exudates, honeydew, live and dead insects, etc. On 26 Nov 1995, four foragers of C. ectopia were observed searching the surfaces of a small bean plant. The leaves of this plant exhibited typical caterpillar 1997 GULMAHAMAD: ECOLOGY OF CARDIOCONDYLA ECTOPIA 23 damage and it may have been the source of some of the first instar caterpillars retrieved from foragers returning to their nests. On 26 Nov 1995, five dead C. ectopia workers were found at different locations on the stem of a small tomato plant, Lycopersicon esculentum Miller. Microscopic examination of these ants revealed no obvious bodily injury. It is possible that these ants were killed by toxic glandular exudates from this plant. Glandular exudates, particularly sucrose and glucose esters from solanaceous plants such as tomato and tobacco, have been reported to immobilize small herbivores, inhibit and/or deter feeding, or to be toxic (Kennedy & Yamamoto 1979, Gregory et al. 1986, King et al. 1990). During the first three weeks of November 1995, several foragers of C. ectopia were observed ascending and descending a cardboard receptacle that was 0.6 m in height and was located about 0.6 m from their nest. This container held empty soda cans for recycling and C. ectopia workers were observed on the lids of several cans feeding on soda remnants. On 13 Aug 1996 as many as 75 ants were counted on this soda can. This is an example of an opportunistic ant species exploiting a man-made food resource. Food Resource Utilization.—Information on food resource utilization by mem- bers of the genus Cardiocondyla is scarce. Cardiocondyla venustula Wheeler is reported to be a scavenger (Wilson 1959a); C. emeryi Forel is thought to be omnivorous (Creighton & Snelling 1974); C. wroughtoni Forel is reported to be a predator (Lupo & Gerling 1984); and C. ectopia has been observed taking nectar from Chamaesyce serpens Small (Creighton & Snelling 1974). At this location, ants from all C. ectopia colonies fed primarily on nectar from flowers of sweet alyssum. Twelve foragers were recorded from one plant and as many as three workers were found on one tiny flower. Foragers often spent con- siderable time on flowers of this plant. Cardiocondyla ectopia also exhibited predatory behavior. The following live insects were retrieved from individual foragers returning to their nests: two first instar caterpillars, one second instar caterpillar, one first instar geometrid larva, an unknown small legless larva, and a small aphid. When freed from the man- dibles of the foragers, the caterpillars attempted to crawl away. On 25 Dec 1995, one forager was observed attempting to remove a second instar caterpillar from a sweet alyssum plant. It was experiencing great difficulty in doing so because the caterpillar was holding on to the plant. Finally, the ant and its quarry fell off the plant. Once on the ground, the ant was more successful in carrying the cat- erpillar especially when it grasped the caterpillar at about its midsection. Even when the prey was on the ground, the ant was not able to transport it more than a few centimeters at a time. Occasionally, it left the prey and ran about the area in a “‘frenzied’’ manner perhaps attempting to locate another nestmate which it could possibly recruit to this food resource. At this site, C. ectopia also functioned as a scavenger. The following dried, dead arthropods were retrieved from foragers returning to their nests: a small caterpillar, a cast larval skin, three small flies, a thrips, a drain fly (family Psy- chodidae) two collembolans, one earwig nymph, a chironomid midge, and a spi- derling. In August 1994, while evaluating commercial ant baits for Argentine ant con- trol, small amounts of Drax Ant Kill Gel—a sucrose/orthoboric acid ant gel bait, 24 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(1) (Waterbury Companies, Waterbury, Connecticut) were dispensed in short pieces of plastic straws and placed at various locations on the study site. Cardiocondyla ectopia foragers discovered this bait at one location and fed on it. The fact that this ant fed on soda and on a sugar-based ant bait indicates that, if accessible in nature, C. ectopia will probably feed at extrafloral nectaries. On 13 Jul 1996, C. ectopia were observed feeding on honeydew produced by aphids on cowpea plants at this location. Members of the genus Cardiocondyla have been reported to feed on honeydew (Smith 1944). Nest Location.—Around human habitation C. ectopia utilized various sites to establish its nests. At least six nests existed on this property during these inves- tigations. Two nests were located in cavities in mortar at the edge of bricks along the side of the concrete driveway. A third nest was located in soil on the south side of the house next to a concrete walkway. The fourth nest was found at the southeast corner of the building in an expansion joint where the concrete walkway met the foundation of the structure. The fifth nest was located in a crack in a concrete walkway at the southeast corner of the house. The sixth nest was located in soil at the edge of a concrete walkway leading to the front door of the structure. Only one of the nest entrances on this property was ever surrounded by a pile of debris as reported elsewhere (Creighton & Snelling 1974). However, nests of C. ectopia which were located in bare soil on a residential property in Montclair, California, were always surrounded by piles of miscellaneous materials. The pres- ence of debris around a nest entrance is probably influenced by soil type, nest location, and type of food utilized. On two occasions, when attempting to pinpoint nest entrances, the entry points were inadvertently enlarged by the author. The ants immediately closed off the entrances. They quickly gathered whatever materials were available from around the nest entrances and began piling them into the opening. Examples of materials used were small pieces of stucco, dirt, sand grains, small rocks, small pieces of mortar and concrete, flower petals of alyssum, and tiny pieces of grass clippings. Nest Population.—Reported colony size of some members of the genus Car- diocondyla are as follows: C. venustula—probably no more than 100 or 200 workers (Wilson 1959a); C. nuda—two dealated queens and 38 workers (Creigh- ton & Snelling 1974); C. paradoxa—S0O adults (Wilson 1959b); C. thoracica— 70 adults (Wilson 1959b); C. ectopia—eight dealate females, two alate females, 75 workers and 2 males. Immatures were not counted but they were estimated at 55 larvae and 15 pupae (Creighton & Snelling 1974). A total of 322 ants were recovered from an excavated colony of C. ectopia representing the largest number ever recorded for a Cardiocondyla species. Castes retrieved included 72 alate females, 13 dealate females, two males and 233 work- ers. Immatures were not counted. These numbers are considerably higher than the numbers previously recorded for a colony of C. ectopia (Creighton & Snelling 1974). Coexistence with the Argentine Ant—The Argentine ant Line pithema humile (Mayr), is a notoriously aggressive species which can negatively impact the bio- diversity of ecosystems (Smith 1936, Haskins 1939, Michener 1942, Haskins & Haskins 1965, Smith 1965, Wilson & Taylor 1967, Crowell 1968, Fluker & Beardsley 1970, Ebeling 1975, Erickson 1975, Lieberburg et al. 1975, Ward 1987, Gulmahamad 1995). However, C. ectopia was observed by the author coexisting 1997 GULMAHAMAD: ECOLOGY OF CARDIOCONDYLA ECTOPIA Pa) with the Argentine ant at four different geographical locations in southern Cali- fornia. At one site, it was surviving in a nest with the entrance located only 8 cm from a nest of the Argentine ant and only 3 cm from an active trail of this species. Cardiocondyla ectopia appears to employ a number of behavioral strategies to Survive in association with the pugnacious Argentine ant. Some examples of these behavioral strategies might be (1) it forages individually and seldom recruits to food sources by tandem running, thus maintaining a low profile presence in sym- patric and synchronous situations, (2) although it is omnivorous, its major food source is nectar from certain flowers, and its predaceous and scavenging activities do not appear to bring it in conflict with the Argentine ant as far as food resource utilization is concerned. No agonistic interactions were ever observed between the two species at any food source during these observations, (3) it maintains small colony populations and thus there are fewer opportunities for conflict among individuals of the two species, (4) it utilizes small, cryptic nest entrances, (5) only a small number of foragers of C. ectopia are usually above ground at a given time thus the potential for conflicts between the two species is much reduced, (6) foragers of C. ectopia enter and leave the nest singly, thus workers do not draw attention to the nest entrance (7) alates leave the nest singly and at intervals, thus there is no aggregation of swarmers at the nest entrance to draw unwanted atten- tion to the nest, and (8) during favorable temperature and daylight conditions, workers emerging from a nest crawl away from the nest entrance immediately thus there is no congregation of ants at or around the nest area. All of the above strategies probably enables C. ectopia to maintain a low profile existence in a hostile territory. Chemical Defense Strategy of C. ectopia.—Ants are known to use chemicals as defense weapons when threatened by other species (Blum 1981, Hermann & Blum 1981, Buschinger & Maschwitz 1984, Hermann 1984). Members of the genus Cardiocondyla are known to use chemicals to protect themselves from aggressive and carnivorous species (Creighton & Snelling 1974). These authors reported that C. emeryi often nest in close proximity to colonies of Solenopsis geminata (FE) and Pheidole dentata Mayr in Texas. They observed minor workers of P. dentata responding in a near panic manner in the presence of workers of C. emeryi. They postulated that the use of repellent chemicals may explain the nesting of C. emeryi close to other aggressive ants species. Creighton & Snelling (1974) also noted that Argentine ant workers often behaved in a very erratic fashion when they encountered workers of C. ectopia. I observed C. ectopia using a chemical to defend itself against the Argentine ant on many occasions. When this material was released, it elicited a dramatic frenzied and agitated running behavior in an affected Argentine ant. Shortly there- after, the recipient was often vigorously engaged in cleaning its eyes, antennae, and mouthparts with its forelegs. This material is of abdominal origin as crushed abdomens of C. ectopia elicited frenzied and agitated running behavior when touched on the heads of conspecifics as well as workers of L. humile. The repellent material elicited the most dramatic reaction when applied to the head of the ag- gressor. Chance encounters between individual ants of these species often did not result in the use of this repellent chemical. At least twelve interspecific contacts between these ants were observed which did not result in the use of the repellent 26 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(1) material. In these situations, the individuals involved briefly examined each other and then separated. This is a beneficial strategy because production of defensive chemicals represents a drain of resources which dictates prudent use. Possession of a potent repellent chemical does not always insure self preservation in a con- tinuous hostile territory. On two occasions, individual C. ectopia workers inad- vertently wandered into well established trails of the Argentine ant. In these sit- uations, L. humile workers attacked and dispatched these individuals with man- dibular cuts. Two encounters between individuals of C. ectopia and L. humile resulted in the loss of an antenna by C. ectopia. A combination of the various behavioral strategies described earlier in this paper and the prudent use of a repellent chemical probably enables C. ectopia to survive in disturbed urban environments around structures in southern California which are dominated by the Argentine ant. ACKNOWLEDGMENT I thank the following people who kindly read the manuscript and offered sug- gestions for its improvement: Roy Snelling, Los Angeles County Natural History Museum; Stoy Hedges, Terminix Int.; Rusty Bracho, ServiceMaster; Ken Hobbs, Entodendrex; William MacKay, University of Texas at El Paso, and William MacKay for providing the identification of C. ectopia. LITERATURE CITED Blum, M. S. 1981. Chemical defenses of Arthropods. Academic Press, New York. Bolton, B. 1982. Afrotropical species of the myrmicine ant genera Cardiocondyla, Leptothorax, Mel- issotarsus, Messor, and Cataulacus (Formicidae). Bull. Brit. Mus. Natur. History. (Entomol. Series), 45: 307-370. Buschinger, A. & U. Maschwitz. 1984. Defensive behavior and defensive mechanisms in ants. pp. 95-150. In Hermann, H.R. (ed.). Defensive mechanisms in social insects. Praeger, New York. Creighton, W. S. & R. R. Snelling. 1974. Notes on the behavior of three species of Cardiocondyla in the United States (Hymenoptera: Formicidae). J. N. Y. Entomol. Soc: 82: 82-92. Crowell, K. L. 1968. Rates of competitive exclusion by the Argentine ant in Bermuda. Ecology, 49: 531-555. Ebeling, W. 1975. Urban Entomology. Univ. Calif. Press, Berkeley, California. Erickson, J. M. 1971. The displacement of native ant species by the introduced Argentine ant, Jri- domyrmex humilis (Mayr). Psyche, 78: 257-266. Fluker, S. S. & J. W. Beardsley. 1970. Sympatric associations of three ants: Iridomyrmex humilis, Pheidole megacephala, and Anoplolepis longipes in Hawaii. Ann. Entomol. Soc. Amer., 63: 1290-1296. Gregory, P, D. A. Ave, P. Y. Bouthyette & W. M. Tingey. 1986. Insect defensive chemistry of potato glandular trichomes. Jn Juniper, B. E. & T. R. E. Southwood (eds.). Insects and the plant surface. Edward Arnolds, London. Gulmahamad, H. 1995. Argentine ant: the Genghis Khan of the ant world. Pest Mgmt, 14: 9-15. Haskins, C. P. 1939. Of ants and men. Prentice-Hall, New York. Haskins, C. P. & E. FE Haskins. 1965. Pheidole megacephala.and Iridomyrmex humilis in Bermuda- equilibrium or slow replacement? Ecology, 46: 736—740. Hermann, H. R. & M. S. Blum. 1981. Defensive mechanisms in the social Hymenoptera. pp 77-197. In Hermann, H. R. (ed.). Social insects, Vol. 2. Academic Press, New York. Kennedy, G. G. & R. T. Yamamoto. 1979. A toxic factor causing resistance in a wild tomato to the tomato hornworm and some other insects. Ent. Exp. Appli. 26: 121-126. King, R. R., L. A. Calhoun, R. P Singh & A. Boucher. 1990. Sucrose esters associated with glandular trichomes of wild Lycopersicon species. Phytochemistry, 29: 2115-2118. Knight, R. L. & M. K. Rust. 1990. The urban ants of California with distribution of imported species. Southwestern Entomol., 15: 167-178. 1997 GULMAHAMAD: ECOLOGY OF CARDIOCONDYLA ECTOPIA 27 Lieberburg, I., P. M. Kranz & A. Seip. 1975. Bermudian ants revisited: the status and interaction of Pheidole megacephala and Iridomyrmex humilis. Ecology, 56: 473-478. Lupo, A. & D. Gerling. 1984. Bionomics of the tamarix spindle-gall moth, Amblypalpis olivierella Rag. (Lepidoptera: Gelechiidae) and its natural enemies. Boll. Lab. Entomol. agr. Filippo Sil- vestri, 41: 71-90. MacKay, W. P. 1995. New distributional records for the ant genus Cardiocondyla in the New World. (Hymenoptera: Formicidae). Pan-Pacif. Entomol., 71: 169-172. Michener, C. D. 1942. The history and behavior of a colony of harvester ants. Sci. Mon., 55: 248-— 258. Smith, M. R. 1936. Distribution of the Argentine ant in the United States and suggestion for its control and eradication. U.S. Dept. Agric. Circ. No. 387. Smith, M. R. 1944. Ants of the genus Cardiocondyla Emery in the United States. Proc. Entomol. Soc. Wash., 46: 30-41. Smith, M. R. 1965. House-infesting ants of the eastern United States. Tech. Bull. Agr. Res. Svc., U.S. Dept. Agr. No. 1326. Snelling, R. R. 1974. Studies on California ants. 8. A new species of Cardiocondyla (Hymenoptera: Formicidae). J. N. Y. Entomol. Soc., 82: 76-81. Ward, P. S. 1987. Distribution of the introduced Argentine ant Uridomyrmex humilis) in natural habitats of the lower Sacramento Valley and its effects on the indigenous ant fauna. Hilgardia, 55: 1-16. Wilson, E. O. 1959a. Communication by tandem running in the ant genus Cardiocondyla. Psyche, 66: 29-34. Wilson, E. O. 1959b. Some ecological characteristics of ants in New Guinea rain forests. Ecology, 40: 437-447. Wilson, E. O. & R. W. Taylor. 1967. The ants of Polynesia (Hymenoptera: Formicidae). Pac. Ins. Monog. No. 14. Received 19 Jan 1996; Accepted I Jun 1996. PAN-PACIFIC ENTOMOLOGIST 73(1): 28-35, (1997) REPRODUCTIVE BEHAVIOR OF THE FEMALE CAROB MOTH, (LEPIDOPTERA: PYRALIDAE) RICHARD S. VETTER, STEVE TATEVOSSIAN,! AND THOMAS C. BAKER? Department of Entomology, University of California, Riverside, CA 92521 Abstract—Periodicities of the female reproductive behavior of the carob moth, Ectomyelois ceratoniae (Zeller), were investigated in regard to calling, mating, and oviposition. Under varying photoperiods (16:8, 14:10, 12:12 L:D h), female carob moths initiated calling about the mid- point of the scotophase to which they were entrained resulting in a shift to later mean initiation times as the nocturnal period lengthened. Matings were initiated during the fifth and sixth h of scotophase in a 16:8 L:D h light regime; this corresponded with the calling periodicity. Carob moth females laid significantly more eggs in the first hour of scotophase (16:8 L:D h) than in any other hour, after which oviposition declined significantly. Oviposition was greatest from the third through sixth scotophase after which it decreased. Oviposition periodicity was developed by the third scotophase, and peaked during the fourth. Key Words.—Insecta, Pheromone Behavior, Mating Periodicity, Ectomyelois ceratoniae The carob moth, Ectomyelois ceratoniae (Zeller), has occasionally been found in the southern United States. This species was most likely introduced from the Middle East where it is a pest of dates, almonds and pomegranates and was first noticed in California in 1982 (Eichlin 1982). It has since become a serious pest of dates in the Coachella Valley in southern California (Warner 1988, Warner et al. 1990a, b) and is of concern to growers as fewer insecticides are available for controlling this pest. In addition, there is concern that the carob moth may spread northward and threaten the almond and walnut industries in California’s Central Valley. Little information is available regarding behavior of the carob moth. Research has been performed on the effects of abiotic factors on development and diapause (Cox 1976, 1979), however, most research has focused on applied aspects in relation to agricultural crop damage (see Gothilf 1984 and references therein). Due to the recent immigration of the carob moth in the U.S., studies have been initiated to develop semiochemical control of this insect: assessment of male re- sponses to both female sex pheromone and a formate analog (Baker et al. 1991, Todd et al. 1992), and female responses to volatile date odors (Cossé et al. 1994). The goal of this study was to develop a knowledge of the reproductive behavior of the carob moth. MATERIALS AND METHODS Insects.—Moths were obtained from date (Phoenix dactylifera L.) gardens in the Coachella Valley, California (Lat. 30°30'N, Long. 116°W) in 1985 and main- tained year-round in the laboratory for >6 yr with no infusion of wild insects. Larvae were reared on a wheat bran-honey diet (Finney & Brinkman 1967) sup- Present Address: 'School of Dentistry, Loma Linda University, Loma Linda, CA 92350; ?Ento- mology Dept. Iowa State University Ames, IA 50011. 1997 VETTER ET AL.: CAROB MOTH REPRODUCTIVE BEHAVIOR 29 plemented with brewer’s yeast and maintained in clear, 4-liter, screened-lid glass jars at 28 + 2°C with a 16:8 L:D h photoperiod. Additional rearing methods were slightly modified from those of Strong et al. (1968). For the calling and mating studies, pupae were separated by sex and placed in moistened vermiculite-filled cups inside of screen cages (30 by 30 by 30 cm) at the light cycles under which the adult moths were eventually tested. Cups of pupae were removed daily to an empty cage, leaving behind moths of known age. When 3 differing light cycles were used in the calling experiment, pupae harvested each day were separated into three groups with one group set up in each light regime. In the mating experiment, male and female pupae were placed in separate environmental cham- bers and allowed to emerge. For the oviposition study, pupae were not separated by sex but were otherwise treated as above. All moths were maintained at 23 + 2° C throughout the course of the study and supplied with 8% sugar water solution ad libitum. Periodicity of Pheromone Calling.—Females were held under 16:8, 14:10 and 12:12 L:D h light cycles to investigate the periodicity of calling. Virgin females were individually placed into plastic, air-tight vials (70 mm by 33 mm) in the last hour of photophase; a 10 mm by 10 mm diam piece of moistened dental wick was added to each vial as a water source. Cohorts of females that had been adults for 1, 2, 3, 4 and 5 days were set up for each of the 16:8 and 14:10 L:D h cycles; a single cohort of Day 2 females was set up for the 12:12 L:D h cycle. In this experiment, the term “Day” is used to indicate a 24-h period starting with the first hr of scotophase; this is to avoid confusion between the ambiguous use of “day” for 24 h or for only its photoperiod. Insects labelled as “‘Day 2’ were entering their 2nd complete scotophase as moths and therefore were 25—48 h old. Each cohort consisted of 30 females except for Day 1 for the 16:8 (n = 24) and 14:10 (n = 10) light cycles. Females were moved into a bioassay room that was illuminated by dimmed white and red incandescent lighting (combined lighting level = 0.3 lux). Females were checked for calling (i.e., visible extrusion of the Ovipositor/sex pheromone gland and hence, presumed sex pheromone emission) every hr during the scotophase. Observations were made using a flashlight cov- ered by several pieces of red cellophane. This light did not appear to alter the females’ behavior. Observations in photophase were made at 2 h intervals until the next scotophase at which point the bioassay was terminated. Data were omitted for any moth which died in the course of the experiment. Periodicity of Mating Behavior——As a correlate of the calling periodicity, a mating study was performed. Virgin male-female pairs were placed together in screen mating cages (80 mm by 50 mm, 18 by 14 mesh), the ends of which were closed with plastic petri dishes. Moths were maintained on a 16:8 L:D h cycle and, in the last hour of photophase, a male-female pair (each of Day 3) was introduced into a mating cage; Day 3 moths were chosen because in many moth Species, males typically require several days to become sexually mature (Shorey et al. 1968). Pairs were then placed in a bioassay room with dimmed, white incandescent lighting (0.3 lux). Fan-forced air was circulated around the room and another fan continually exhausted the room air outside the building. Moths were observed every 30 min of scotophase until the first pair mated, whereafter they were observed every 15 min until the end of scotophase when the bioassay was terminated. Observations were aided with a red cellophane-covered flashlight, 30 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(1) although at 0.3 lux, there was sufficient light to see pairs coupled. Two replicates were run, 30 pairs per replicate. Data were excluded if either moth of a pairing was dead or moribund at the conclusion of the test. Periodicity of Oviposition—Female carob moths (16:8 L:D cycle) were re- moved from their emergence cage (which contained males of equal age as mating partners) and were set up in screen cages (80 mm by 50 mm) with an open end which was covered with a clear polyethylene sleeve. A cohort consisting of 10 females of known age (Day 2, 3, 4, 5, 6, 7 or 8) was placed in a cage; four cohorts were run for each age of female. The dates used as the oviposition sub- strate (variety Deglet Noor) have low water content and are preferred by the carob moth in the Coachella Valley over other, moister varieties (C. Kerby, pers. comm.). These dates were of commercial sale quality taken from a recent harvest, rinsed with water to remove elemental sulfur used for mite control, and air-dried. For the experiment, dates were impaled on a bent paper clip tied to a piece of string so they could be easily lowered into and pulled out of the cages. Decaying dates might prove more attractive as oviposition lures because female carob moths are attracted to odors of fermenting or fungus-infested host fruits (Gothilf et al. 1975, Warner 1988, Cossé et al. 1994). However, using them might introduce greater variation into the experiment; hence, non-decaying dates were used. One date was placed in each cage of 10 females and the plastic sleeve was folded over and clipped to minimize moth escape. Cages were transferred to a bioassay room described above. Dates were replaced hourly and deposited eggs were count- ed and totalled for each of the 8 h of scotophase. Because few eggs were laid on the cage that housed the moths, no attempt was made to count them. At initiation of photophase, females were given another date which was not removed until the end of the 16 h light period; we previously observed little oviposition occurs during photophase. The experiment was terminated at the start of the next sco- tophase. Females were transferred to vials containing 90% alcohol and later dis- sected to determine the number of spermatophores in the bursa copulatrix. Statistics—Calling periodicity was analyzed with one-way ANOVA with Tu- key-Compromise test or two-way ANOVA with Tukey’s studentized range tests separations. Mating periodicity was analyzed with a X? test for independence using Yate’s correction. Oviposition periodicity data were square-root transformed because there was a high variance within the data sets, and then analyzed using two-way ANOVA with Tukey’s studentized range test for separation (SAS Insti- tute, 1982). RESULTS Calling Periodicity.—Female carob moths initiated calling in the fourth through sixth h of scotophase (16:8) and fourth through seventh h (14:10) for all five age cohorts of females; Day 2 females on the 12:12 light cycle initiated calling in the fifth h (Fig. 1). ANOVA of the 14:10 and 16:8 cohorts revealed a significant difference among the groups for initial onset of calling (two-way ANOVA, F = 9.80; df = 9, 220; P < 0.0001). The 16:8 moths called significantly earlier than the 14:10 females (F = 61.97; df = 1; P < 0.0001), however, there was also a significant difference between cohorts within a light cycle (F = 3.51; df = 4; P < 0.01). The interaction of age X< light cycle was not significant (F = 2.08; df = 4; P > 0.05). In the 16:8 groups, as the age of the female cohort increased, 1997 VETTER ET AL.: CAROB MOTH REPRODUCTIVE BEHAVIOR 31 16:8 14:10 12:12 | =) kk) 7.06+0.73° 7.4340.79° DAY 1 18/24 6.52:0.747), 7.2810.84",, 7.4521.92 | DAY 2 22/30 BA tt tH HH 6.3620.99°° 7.8141.20" 0 12 24 DAY 3 hours 60 6.2911.06° 7.9211.41° ore comes DAY 4 21/30 20 ) 5.9641.179 7.0021.05° DAY 5 ce) 12 24 ce) 12 24 hours hours Figure 1. Percentage of calling female carob moths at 16:8, 14:10 and 12:12 L:D hr photoperiod regimes. Bar above symmetric histograms indicates dark:light cycle. Average initial onset of calling (x + SD) is indicated above the tick-marked axis; # females observed calling at least once/# females alive at experiment termination is indicated below. 16:8 versus 14:10 data were analyzed by two-way ANOVA. Means having none of the superscript letters (#®) in common indicate significant differences within a column (Tukey’s Studentized Range test). Day 2 females were compared with a one-way ANOVA. Means having none of the subscript letters (,,,,) in common indicate significant differences across the row (Tukey-Compromise test). their time of initial onset of calling occurred significantly earlier in the scotophase (Fig. 1). This behavior was not exhibited, however, with the 14:10 cohorts. Ad- ditionally, all calling terminated during the first hr of photophase and no calling was observed during the remainder of the photophase. When Day 2 females were analyzed across the three light cycles, the 16:8 females called significantly earlier than either of the other two groups (one-way ANOVA; F = 4.37; df = 2, 73; P < 0.01). The 14:10 and 12:12 groups were Statistically indistinguishable (Fig. 1). Mating Periodicity.—There was no difference between the 2 replicates for hour of mating initiation (X* = 3.13; df = 7; P > 0.05) or frequency of mating between pairs (X*? = 0.30; df = 1; P > 0.05) so the data were pooled. Mating occurred in 43 of 55 (76.4%) of the pairings; five pairs were excluded due to death or moribund condition of one of the partners. Mating times were not randomly dis- tributed throughout the scotophase (X? = 72.04; df = 7; P < 0.001) and carob 32 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(1) moths started mating in the fourth h of scotophase (N = 3). There was a signif- icant increase of initiated matings during the fifth and sixth h (N = 20 and 13 respectively; these values are statistically similar). After this, the number of mat- ings initiated within the seventh (N = 5), and eighth h (N = 2) of scotophase decreased significantly. The number of pairs observed in copula during each h was 3, 21, 35, 34 and 23 for the fourth, fifth, sixth, seventh and eighth h respec- tively. Pairs remained in the coupled position for 2.35 + 0.84 h. Oviposition Periodicity—There was a very pronounced nocturnal periodicity and age effect for carob moth female oviposition behavior (F = 11.03; df = 83, 168; P < 0.0001, Table 1). Two-way ANOVA revealed highly significant effects for hour of scotophase (F = 69.21; df = 8; P < 0.0001), age of female (F = 23.74; df = 6; P < 0.0001) and the interaction of these variables (F = 2.37; df = 48; P < 0.0001). Carob moths laid the greatest number of eggs during the first hour of scotophase (Table 1). There is a statistically significant decrease for each of the following 2 h periods followed by a diminishing of the behavior to near zero by the sixth h. Virtually no eggs were laid during the 16 h photophase. Considering the effect of age, egg deposition rose significantly by Day 3, peaked with Day 4 females whereafter egg production decreased to significantly lower levels by Day 7 (Table 1). Virtually all females were mated at the end of their second full scotophase (Day 3) as indicated by the presence of spermatophores, and the number of matings increased with age with a mode of one spermatophore for Day 3 thru Day 6 females, and two spermatophores beyond Day 6 (Table 1). This increase with age was significant across the group (F = 34.56; df = 6, 248; P < 0.0001) with Day 7 and 8 females having significantly more spermatophores than Day 3 through 6 females (which were all statistically similar). Day 2 had significantly fewer sper- matophores compared to every other group (Table 1). DISCUSSION Female carob moths exhibit periodicities in their reproductive behavior which may be useful in developing field control methods for the insect. The only pre- vious reference to female reproductive behavior of the carob moth was that of Cox (1976) in which he states that calling occurred “‘when it became darker’ and that oviposition occurred “‘during twilight and dark periods.’’ Carob moth calling was initiated near the mid-point of the scotophase over a range of ages and light regimes from short (16:8) to longer night (12:12) (Fig. 1). When ob- served as a group, carob moth females continued calling until the photophase, whereupon they abruptly ceased. When virgin carob moth females were placed with males under a 16:8 L:D cycle, matings were initiated at the same period (fifth to sixth h scotophase) as the initiation and rise in calling of 16:8 females (Fig. 1). Overall, most carob moth oviposition in this study occured during the first 3 h of scotophase with the highest number of eggs deposited during the first h (16:8 light regime) (Table 1). Oviposition decreased sharply by the mid-point of the scotophase and was virtually zero during the photophase. An ontogeny of ovi- position may occur where a periodicity develops by Day 3, production peaks at Day 4 and then drops off afterward. The lower performance of the Day 2 and Day 3 females may be partly due to some females still being virgins. This pattern Table 1. Hourly mean (SD) egg output and number of spermatophores detected for groups of 10 female carob moths (four replicates) at differing ages. Hour of scotophase 16h photophase Avr. for Avr. no. Ist 2nd 3rd 4th Sth 6th 7th 8th 9th—24th each day spermatophores n Day 2 6.3 7.0 LS ae 5.5 205 3.8 1.3 1.0 42.3 be 0.30 c (40) (8.5) (10.7) (9.7) (8.1) (1.9) (1.3) (2.6) (1.0) (0.8) (38.7) (0.46) Day 3 oles) 18.3 18.3 11.3 14.3 53 2 4.5 1.3 105.0 ab 1.10 b (38) (8.2) (10.3) (9.7) (7.4) (6.8) (1.9) (2.1) (2.6) (1.0) (32.0) (0.51) Day 4 43.5 41.0 31.3 16.0 8.5 7.3 6.3 8.3 25 164.8 a 1.26 b (35) (16.7) (40.2) (28.7) (11.7) (6.8) (3.9) (1.3) (4.7) (1.3) (88.7) (0.70) Day 5 44.5 15.6 10.8 9.0 8.0 4.3 2.0 4.3 0.5 99.0 abc 1.53 b (38) (21.0) (7.4) (9.2) (4.8) (1.2) (2.2) (1.4) (2.8) (1.0) (30.3) (0.60) Day 6 46.8 18.0 9.0 7.0 8.8 4.0 1.5 3.3 0.3 98.5 abc 1.54 b (35) (25.4) (11.2) (6.4) (5.6) (9.0) (3.6) (1.0) (1.7) (0.5) (56.4) (0.70) Day 7 31.0 13.0 5.3 33 2.3 2.0 2.8 1.0 0.3 60.8 be 2.32 a (37) (12.9) (7.8) (3.4) (1.0) (2.5) (1.8) (4.9) (1.4) (0.5) (24.8) (0.94) Day 8 32.5 11.8 2.8 3.0 0.8 1.5 0.3 1.0 0 5329-0 2.17 a (35) (17.2) (6.3) (1.0) (0.2) (1.0) (0.6) (0.5) (1.4) (0) (25.4) (1.07) Mean for 33.7 a 17.8 b 12.1 ¢ 8.lc¢ 6.9 cd 3.8 de 201 et 3.4e 0.9 f each h (19.7) (18.5) (14.6) (7.3) (6.2) (2.9) (2.7) (3.3) (1.1) Overall means having no letters in common are significantly different for hourly oviposition (row), daily egg output (column) and spermatophores (column). Oviposition data were analyzed with two-way ANOVA with Tukey’s studentized range test separation. Spermatophore data were analyzed with one-way ANOVA with Tukey’s compromise test separation. YOIAVHAd FAILONGOUdAY HLOW PONV) :"TV LA YALLYAA L66I te 34 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(1) of oviposition is found in at least five other species of pyralid moths (Bell 1981, Andrews et al. 1980). As the females age and egg production decreases, the number of spermatophores per female continues to increase suggesting that lack of sperm is probably not an explanation for decreased oviposition (Table 1). The information presented here may be of importance to date growers in their attempts to control the carob moth. Because females oviposit most heavily right after sunset, insecticide sprays during this time might increase the chances of killing egg-laden females. Similar studies with cotton pests documenting their nocturnal behavior patterns have resulted in changes in control strategies in that some crop-dusting is performed at night when the insects are more likely to contact pesticide (UC Press, 1984). In contrast, trying to control carob moths by targeting the mating behavior might be difficult because mating periodicity in moths is dependent on abiotic variables other than photoperiod (e.g., temperature, Baker & Cardé 1979, Kanno 1981). Also control methods may be ineffective for interrupting the mating behavior of the carob moth because the actual mating location in the date agrosystem is unknown. Therefore, targeting the ovipositing female may be the more effective control strategy because females are in the date canopy laying their eggs. ACKNOWLEDGMENT Thanks are extended to Neil Q. Vickers and P. K. Visscher for statistical help, Steve McElfresh for making copious improvements on an earlier draft of the manuscript, Judy Chari Kang and Dondi M. Flanagan for rearing the insects used in this study (all personnel from Univ. Calif., Riverside) and Cathy Kerby of Covalda Date Co., Coachella, CA for supplying the Deglet Noor dates. This research was funded in part by the Date Packer’s Council. LITERATURE CITED Andrews, K. L., M. M. Barnes & S. A. Josserand. 1980. Dispersal and oviposition by navel or- angeworm moths. Envir. Entomol., 9: 525-529. Baker, T. C. & R. T. Cardé. 1979. Endogenous and exogenous factors affecting periodicities of female calling and male sex pheromone response in Grapholitha molesta (Busck). J. Insect Physiol. 25: 943-950. Baker, T. C., W. Francke, J. G. Millar, C. Lofstedt, B. Hansson, J.-w. Du, P. L. Phelan, R. S. Vetter, R. Youngman & J. L. Todd. 1991. Identification and bioassay of sex pheromone components of carob moth, Ectomyelois ceratoniae (Zeller). J. Chem. Ecol., 17: 1973-1988. Bell, C. H. 1981. The influence of light cycle and circadian rhythm on oviposition in five Pyralid moth pests of stored products. Physiol. Entomol., 6: 231-239. Cossé, A., J. J. Endris, J. G. Millar & T. C. Baker. 1994. Identification of volatile components from fungus-infected date fruit that stimulate upwind flight in female Ectomyelois ceratoniae. En- tomol. exp. et appl., 72: 233-238. Cox, P. D. 1976. The influence of temperature and humidity on the life cycle of Ectomyelois cera- toniae (Zeller) (Lepidoptera: Phycitidae). J. stored Prod. Res., 12: 111-117. Cox, P. D. 1979. The influence of photoperiod on the life-cycle of Ectomyelois ceratoniae (Zeller) (Lepidoptera: Pyralidae). J. stored Prod. Res., 15: 111-115. Eichlin, T. D. 1982. Carob moth in California: New state record. Calif. Dept. Food Agric. Memo Nov. 26. Finney, G. L. & D. Brinkman. 1967. Rearing the navel orangeworm in the laboratory. J. Econ. Ent., 60: 1109-1111. Gothilf, S. 1984. Biology of Spectrobates on Almonds in Israel. Phytoparasitica, 12: 77-87. 1997 VETTER ET AL.: CAROB MOTH REPRODUCTIVE BEHAVIOR 39 Gothilf, S., E. C. Levy, R. Cooper & D. Lavie. 1975. Oviposition stimulants of the moth Ectomyelois ceratoniae: the effect of short-chain alcohols. J. Chem. Ecol., 1: 457-464. Kanno, H. 1981. Mating behaviour of the rice stem borer moth, Chilo suppressalis Walker (Lepi- doptera: Pyralidae). VI. Effects of photoperiod on the diel rhythms of mating behaviours. Appl. Entomol. Zool., 16: 406-411. SAS Institute. 1982. SAS user’s guide: statistics. SAS Institute, Cary, North Carolina, USA. Shorey, H. H, K. L. Morin, & L. K. Gaston. 1968. Sex pheromones of noctuid moths. XV. Timing of development of pheromone-responsiveness and other indicators of reproductive age in males of eight species. Ann. Entomol. Soc., Amer. 61: 857-861 Strong, R., G. J. Partida, & D. N. Warner. 1968. Rearing stored-products insects for laboratory studies: six species of moths. J. Econ. Entomol., 6: 1237-1249. Todd, J. L., J. G. Millar, R. S. Vetter, & T. C. Baker. 1992. Behavioral and electrophysiological activity of (Z,E)-7,9,11-dodecatrienyl formate, a mimic of the major sex pheromone component of carob moth, Ectomyelois ceratoniae. J. Chem. Ecol., 18: 2331-2351. Univ. Calif. Press 1984. Integrated Pest Management for Cotton in the Western Region of the United States. Univ. Calif. Div. Agric. Nat. Resources Pub. #3305. Warner, R. L. 1988. Contributions to the biology and the management of the carob moth, Ectomyelois ceratoniae (Zeller), in “‘Deglet Noor’ date gardens in the Coachella Valley of California. Ph.D dissert. University California, Riverside. Warner, R. L., M. M. Barnes, & E. FE Laird. 1990a. Reduction of insect infestation and fungal infection by cultural practice in date gardens. Envir. Entomol., 19: 1618-1623. Warner, R. L., M. M. Barnes, & E. F Laird. 1990b. Chemical control of a carob moth, Ectomyelois ceratoniae (Lepidoptera: Pyralidae) and various nitidulid beetles (Coleoptera) on ‘Deglet Noor’ dates in California. J. Econ. Entomol., 83: 2357-2361. Received 13 Jun 1996; Accepted 15 Aug 1996. PAN-PACIFIC ENTOMOLOGIST 73(1): 36-39, (1997) LILIOCERIS SP. (COLEOPTERA: CHRYSOMELIDAE) HERBIVORY ON CYCAS SIAMENSIS MIGUEL (TRACHEOPHYTA: CYCADALES) WILLIAM D. SHEPARD Department of Biological Sciences, California State University, Sacramento, Sacramento, California 95819 Abstract.—The chrysomelid Lilioceris sp. was found feeding on the leaflets of cycads in north- eastern Thailand. Many cycads were significantly damaged. Larval and adult Lilioceris sp. col- oration suggests aposematic coloration. Key Words.—Insecta, Chrysomelidae, Cricocerinae, Lilioceris, cycad, herbivory, aposematic col- oration Beetles are increasingly being associated with cycads either by entomologists (Crowson 1991) or by cycadologists (Norstog 1990). Most species either bore through the frond rachis or the trunk, or they attack the cones. Several species have been demonstrated to be pollinators. One species of chrysomelid beetle, Lilioceris clarki (Baly), has been reported to feed on the fronds of Cycas sp. in New Guinea (Szent-Ivany et al. 1956). Here I report Lilioceris sp. larvae feeding on the frond leaflets of Cycas siamensis Miquel in Thailand. The cycads were part of the understory vegetation of a “dry dipterocarpus”’ forest located along Highway 213, 2 km south of the headquarters of Phu Phan National Park, Sakhon Nakhon Province, in northeastern Thailand. The cycads were numerous and easily seen as a fire some months previously had reduced the understory cover. I estimated that about half of the cycads had either beetle larvae actively feeding or bore evidence of their feeding. Infested cycads usually sup- ported only a few larvae (> 10) on a few fronds, although some had heavy infestations (20+ larvae) and abundant frond damage (Fig. 1). In both cases there always were nearby cycads that had no larvae. Infested cycads could be distin- guished easily from a distance as the damaged leaflets were a pale brown color that contrasted sharply with the dark green of undamaged leaflets (Figs. 2—3, 5). Infested cycads were located both in and outside of the burn area. The fire may have made the cycads more visible to the beetles as the infestation was generally heavier in the burn area. Lilioceris sp. was present mainly as larvae. One adult was swept from the cycads. All larvae were located on the lower surfaces of the leaflets (Fig. 4). Feeding involved rasping away the lower epidermis and part of the mesophyll. Generally larvae only ate part of the tissue of any one leaflet before leaving to find another. However, when larval numbers were high the larvae remained on single leaflets until they were substantially consumed. On one heavily infested frond, the end of the rachis was chewed through. The ground under the cycads with heavy infestations was carpeted with feces (Fig. 6) which had a characteristic coiled shape both when being eliminated by the larva and when on the ground. One larva was dissected to examine the structure of the digestive tract. It was typical for herbivores, being voluminous in capacity, long and coiled. Attempts to rear larvae to adults in the lab failed due to lack of a proper pupation site, 1997 SHEPARD: LILIOCERIS LARVAE FEED ON CYCAS 37 Cycas siamensis frond heavily damaged by Lilioceris sp. larval herbivory. This frond Figure 1. has 20 larvae showing. Figure 2. Cycas siamensis with heavy damage caused by Lilioceris sp. larvae feeding on frond leaflets. Note end of rachis is chewed through. Figure 3. Cycas siamensis leaflets discolored by herbivory on the lower sides. Figure 4. Lilioceris sp. larva on underside of leaflet of Cycas siamensis. 38 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(1) Figure 5. Cycas siamensis fronds heavily damaged by insect herbivory. Figure 6. Accumulation of beetle larva feces under a heavily infested Cycas siamensis. which is thought to be the soil. Reared larvae that reached what appeared to be the last instar climbed down from the fronds and crawled around the laboratory. Larvae were bright red-orange in color and visible against the darker foliage, suggesting aposematic coloration warning of chemical defenses. Chemicals may be sequestered from the cycad tissue as they are known to have chemical defenses. None of the larvae evidenced signs of predation attempts or parasitism. When disturbed they remained immobile on the leaflets. The single adult Lilioceris sp. was also a bright red-orange in color. 1997 SHEPARD: LILIOCERIS LARVAE FEED ON CYCAS 39 Voucher specimens deposited at the Entomology Collection at California Acad- emy of Sciences (San Francisco) include one adult and five larval Lilioceris sp. plus several cycad leaflets fed upon by the larvae. ACKNOWLEDGMENT Special thanks go to Samang Homchuen who shared with me her wonderful country and its flora. Edward Riley provided the identification of the chrysomelid and a copy of Crowson’s paper. Travel to Thailand was provided by United States Information Agency Grant # IA-ASCS-G11902372. LITERATURE CITED Crowson, R. A. 1991. The relations of Coleoptera to Cycadales. pp. 13-28. Jn M. Zunino, X. Bellés & M. Blas (eds.). Advances in Coleopterology. AEC, Barcelona. Norstog, K. J. 1990. Studies of cycad reproduction at Fairchild Tropical Garden. pp. 63-81. Jn D. W. Stevenson (ed.). The biology, structure, and systematics of the Cycadales. Proceedings of the Symposium CYCAD 87, Beauliey-sur-Mer, France, April 17—22, 1987. Mem. N. Y. Bot. Garden, 57: 1-210. Szent-Ivany, J. J. H., J. S. Womersley & J. H. Ardley. 1956. Some insects of Cycas in New Guinea. Papua and New Guinea Agric. J., 11: 1-4. Received 21 Feb 1996; Accepted 16 Jun 1996. PAN-PACIFIC ENTOMOLOGIST 73(1): 40-45, (1997) ELECTROPHORETIC COMPARISON OF DENDROCTONUS PUNCTATUS LECONTE AND D. MICANS (KUGELANN) (COLEOPTERA: SCOLYTIDAE) SANDRA J. KEGLEY,! MALCOLM M. FuRNISS,? AND JEAN-CLAUDE GREGOIRE? Abstract—The taxonomic status of the American boreal spruce beetle, Dendroctonus punctatus LeConte, and its Eurasian sibling species, the European spruce beetle, D. micans (Kugelann), has been in doubt. The genetic relationship of adult D. punctatus from Montana, USA, and of D. micans from Belgium was examined by isozyme electrophoresis. Average heterozygosity and polymorphism was 0.075 and 37.5% for D. punctatus and 0.026 and 6.25% for D. micans, much lower than reported in other Dendroctonus species and which may result from the high degree of inbreeding that is characteristic of both species. Five of the 16 loci examined were fixed, or nearly fixed, for different alleles in the two species. Genetic identity between D. punctatus and D. micans was 0.693, lower than that reported between two other, uncontested, host-isolated sibling species, D. ponderosae Hopkins and D. jeffreyi Hopkins. Furthermore, the genetic dis- tance between D. punctatus and D. micans was 0.366, similar to another distinct pair of host- isolated sibling species, D. pseudotsugae Hopkins and D. simplex LeConte. These results support recent morphological evidence in favor of retaining D. punctatus and D. micans as separate species. Key Words.—Insecta, Scolytidae, Dendroctonus punctatus, Dendroctonus micans, isozyme elec- trophoresis The genus Dendroctonus Erichson is represented in North America by 17 spe- cies, including the boreal spruce beetle, D. punctatus LeConte, and in Eurasia by two species, including the European spruce beetle, D. micans (Kugelann) (Wood 1982, Wood & Bright 1992). Because the present center of diversity of Den- droctonus species is North America, the ancestor of D. micans is thought to have migrated from a spruce refugium in Alaska to Siberia via Beringia during the Wisconsinian glaciation and eventually reached Europe through intervening spruce forests. In recent times, D. micans has become of great economic impor- tance where it has invaded new territory in Europe, particularly in exotic spruce plantations in France and England (Bevan & King 1983, Grégoire 1988). On the other hand, D. punctatus has so far caused little economic damage, being appar- ently at a competitive disadvantage in Nearctic boreal spruces to the economically important spruce beetle, D. rufipennis (Kirby) (Furniss 1995). Taxonomists have noted the anatomical similarity of D. punctatus and D. micans and have expressed uncertainty about their status as separate species (Wood 1963, 1982). The biology of D. micans has been studied in Europe by Grégoire (1988); that of D. punctatus has been studied in western North America by Furniss (1995). Features that these two species share, differing from most others of the genus, are: a sex ratio strongly in favor of females; mating by siblings in the brood chamber prior to emergence (males never occur with females in egg galleries); ' USDA Forest Service, 3815 Schreiber Way, Coeur d’ Alene, Idaho 83814-8363. * Division of Entomology, University of Idaho, Moscow, Idaho 83844-2339. 3 Laboratoire de Biologie Animale et Cellulaire, Université Libre de Bruxelles, 1050 Bruxelles, Belgique. 1997 KEGLEY ET AL.: COMPARISON DENDROCTONUS SPP. 41 and aggregation in the larval stage rather than as attacking adults (Grégoire 1983, Furniss 1995). Their main biological difference appears to be that D. punctatus has four larval instars, whereas D. micans is reported to have five larval instars (Furniss 1995). Morphologically, D. punctatus and D. micans have been found to differ in 10 discrete characters (Furniss and Johnson 1989, Furniss 1996). This paper presents results of isozyme electrophoresis that further support the validity of these two species. MATERIALS AND METHODS Dendroctonus micans were collected as larvae at Wellin, Belgium, January 8, 1990 and sent to Moscow, Idaho, for rearing to the adult stage in 15 K 15 cm pieces of Norway spruce phloem (Picea abies (L.) Karst.) pressed between glass plates. Dendroctonus punctatus were F, progeny reared in Engelmann spruce logs (P. engelmannii Parry) (Furniss 1995). Their parents were collected from a white spruce hybrid (P. glauca (Moench.) Voss. X engelmannii) in Meagher Co., Mon- tana in June 1989. Representative voucher specimens are deposited in the W. FE Barr Entomological Museum, University of Idaho. Each adult was immersed in 1 cc of distilled water and macerated vigorously with pointed tweezers to develop a homogenate. Four filter paper wicks were soaked in each homogenate, wrapped individually in parafilm, and stored over- night at —18°C. Numbers of individuals tested were: D. micans = 58 female, 2 male; D. punctatus = 57 female, 2 male. Electrophoretic techniques used were those of Stock et al. (1987). Gels were made from a 13% solution of hydrolyzed potato starch and the appropriate buffer. Wicks containing beetle homogenate were inserted into slots in each gel and subjected to electrophoresis. Gels were then cut horizontally and stained for dif- ferent enzymes. Eleven enzyme systems were assayed (Table 1). Banding patterns were scored as homozygotes (appearing as a single band) or heterozygotes (ap- pearing as multiple bands) for each gene locus resolved. Genotype frequencies for each population were recorded. Genetic data were analyzed using BIOSYS-1, a computer program for analysis of allelic variation (Swofford and Selander 1989). Observed genotype frequencies were compared to values derived from random mating (Hardy-Weinberg expected proportions) using a chi-square test. Genetic diversity was estimated using Nei’s (1978) unbiased estimate of average hetero- zygosity and polymorphism (%). A locus was considered polymorphic when the frequency of the most common allele was less than or equal to 0.99. The rela- tionship between D. punctatus and D. micans was evaluated using Nei’s (1978) genetic identity value, Nei’s (1978) unbiased genetic distance value, and Rogers’ (1972) similarity index. RESULTS AND DISCUSSION Allele frequencies were calculated at 16 gene loci (Table 1). Major differences occurred between the two species at several loci. Six loci (Aat, Idh-2, Mdh-2, Me-1, Me-2, Mpi) were polymorphic in D. punctatus while only one (Me-2) was polymorphic in D. micans. As a result, average heterozygosity and polymorphism were much higher for D. punctatus (0.075 and 37.5%; respectively) than for D. micans (0.026 and 6.25%, respectively). 42 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(1) Table 1. Allele frequencies at 16 enzyme loci, percent polymorphism, average heterozygosity, and chi square comparisons of observed to expected (Hardy-Weinberg) numbers of each genotype for D. punctatus and D. micans. Enzyme abbrev. Allele D. punctatus D. micans Aspartate aminotransferase Aat A 0.064 - 1.0 B 0.936 0 x? 45 .8** Catalase Ck A 1.0 1.0 Esterase Est-1 A 1.0 1.0 Est-2 A 0 1.0 B 1.0 0 Est-3 A 1.0 1.0 Isocitrate dehydrogenase Idh-1 A 1.0 1.0 Idh-2 A 0.034 0 B 0.966. . 1.0 xX? 78.0** Malate dehydrogenase Mdh-1 A 1.0 1.0 Mdh-2 A 0.964 0 B 0.036 1.0 x? 74.0** Malic enzyme Me-1 A 0.034 BD 0.966 1.0 x? FOate Me-2 A 0.512 0.292 B 0.488 0.708 x? 0.7 6.3* Phosphomannose isomerase Mpi . A 0 1.0 B 0.750 0 G 0.250 0 x 20.8** Peptidase glycyl-leucine Pep-gl A 1.0 1.0 Peptidase leucyl-alanine Pep-la © A 1.0 1.0 Glucose phosphate isomerase Pgi A 1.0 0 B 0 1.0 Superoxide dismutase Sod A 1.0 1.0 % Polymorphism (0.99 criterion) 37.50 6.25 Av. heterozygosity 0.075 0.026 * = significant at 0.05 level, ** = 0.01 level. In an earlier study (Stock et al. 1987), average heterozygosity and polymor- phism of D. micans were 0.053 and 27%, respectively. However, the gene loci that were resolved and analyzed differed. somewhat between their study and ours, perhaps contributing to the difference in values for D. micans. For example, the latter authors reported that locus Est-2 had six alleles; we observed only two. An additional possible source of the difference is foreign protein such as from a parasitic nematode (Higby and Stock, 1982). With that in mind, we had examined prior to maceration each of our specimens for ecto- and endoparasitic nematodes; D. micans contained none. If, however, the D. micans that were analyzed by Stock et al. (1987) had nematodes, that might explain the difference in heterozygosity and polymorphism of D. micans in the two studies. In any case, we tested paired sets of D. micans and D. punctatus simultaneously in the same gels, and we are 1997 KEGLEY ET AL.: COMPARISON DENDROCTONUS SPP. 43 confident that the resultant values for heterozygosity and polymorphism truly reflect relative differences between the two species. Dendroctonus punctatus and D. micans were much less genetically diverse than 10 other North American Dendroctonus. species which have an average of 0.213 heterozygosity (range = 0.156—0.247) and 64% polymorphism (range = 50-72%) (Bentz and Stock 1986). Analysis of the deviations of alleles from Hardy-Wein- berg equilibrium showed significantly less heterozygosity in D. punctatus than would be expected of random mating in a population (Table 1). This may be explained by the high degree of inbreeding in this species (Furniss 1995). The one locus that was polymorphic for D. micans showed more heterozygosity than expected. Selection may be favoring heterozygotes at this locus. The fact that D. micans was less heterozygous and less polymorphic than D. punctatus may relate to the following. The immediate ancestor of D. micans must have migrated from an Alaskan glacial refugium. The particular genetic compo- sition of individuals of this isolated population may have been subjected to intense selective pressure in its various, entirely new, host species and the differing cli- mate as it extended thousands of miles eastward to Europe. On the other hand, during the time since D. micans migrated to Asia, its American ancestor has reunited with other population segments as its main host, Picea glauca, followed the retreating glaciers northward, eventually extending across the continent and throughout boreal North America. Five loci (Aat, Est-2, Mdh-2, Mpi, Pgi) (Table 1) were fixed, or nearly fixed, for different alleles in the two species. Fixation of different alleles at one or more loci is characteristic of separate species or geographically separated non-inter- breeding populations (Ayala and Powell 1972, Berlocher 1979, as cited by Higby and Stock 1982). Genetic similarity of D. punctatus and D. micans was 0.682. Conspecific populations of organisms commonly have similarity indices above 0.75 on a scale of 0-1 (Avise 1974, Ayala 1975). The genetic identity index of D. punctatus and D. micans was 0.693. In com- parison, the genetic identity index of two well-defined, host-isolated, sibling spe- cies, of Scolytidae, D. ponderosae Hopkins and D. jeffreyi Hopkins, was greater, being 0.83 (Higby and Stock 1982). That of conspecific populations of D. pon- derosae from Utah in two different species of host trees (Pinus contorta Douglas and P. ponderosa Lawson) was 0.992—0.993 (Stock and Amman 1980) and the genetic identity index of D. ponderosae populations in Alberta in three pine hosts were above 0.978 (Langor and Spence 1991). In further comparison, the genetic identity of humans and chimpanzees is 0.680 and that of humans and Borneo orangutans is 0.707 (Bruce & Ayala 1979). The genetic distance value for D. punctatus and D. micans was 0.366. This is somewhat similar to the genetic distance (0.305) of two uncontested sibling spe- cies, D. pseudotsugae Hopkins and D. simplex LeConte (Bentz and Stock 1986). No unique genetic difference was found between females and the few available males of D.. punctatus and D. micans. The genetic characters reported here supplement recent biological and morpho- logical evidence (Furniss and Johnson 1989, Furniss 1995, Furniss 1996) in sup- port of retaining D. punctatus and D. micans as separate species. 44 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(1) ACKNOWLEDGMENT Steven J. Brunsfeld, Department of Forest Resources, University of Idaho, pro- vided laboratory facilities and helped analyze data and interpret results. Molly W. Stock, Department of Forest Resources, University of Idaho, provided chemicals and laboratory materials and reviewed the manuscript. The manuscript was also reviewed by James B. Johnson, Division of Entomology, University of Idaho, and by David W. Langor, Northern Forestry Centre, Forestry Canada, Edmonton, AI]- berta. This is University of Idaho Agricultural Experiment Station Research Paper No. 95753. LITERATURE CITED Avise, J. C. 1974. Systematic value of electrophoretic data. Syst. Zool., 23: 465-481. Ayala, E J. 1975. Genetic differentiation during the speciation process. p. 1-78. Jn T. Dobzhansky, M. K. Hecht, and W. C. Steere, [eds.], Evolutionary biology. Vol. 8. Plenum Press, New York. Ayala, F J. & J. R. Powell. 1972. Allozymes as diagnostic characters of sibling species of Drosophila. Proc. Nat. Acad. Sci. USA, 69: 1094-1096. Bentz, B. J. & M. W. Stock. 1986. Genetic relationships among ten species of Dendroctonus bark beetles (Coleoptera: Scolytidae). Ann. Entomol. Soc. Amer., 79: 527-534. Berlocher, S. H. 1979. Biochemical approaches to strain, race, and species discriminations, pp. 137— 144. In M. A. Hoy and J. J. McKelvey, Jr. [eds.], Genetics in relation to insect management. Rockefeller Foundation. Bevan, D. & C. J. King. 1983. Dendroctonus micans Kug.—a new pest of spruce in U.K. Comm. For. Rev., 62: 41-51. Bruce, D.J. & EJ. Ayala. 1979. Phylogenetic relationships between man and the apes: electrophoretic evidence. Evolution, 33(4): 1040-1056. Furniss, M. M. & J. B. Johnson. 1989. Description of the gallery and larva of Dendroctonus punctatus LeConte (Coleoptera: Scolytidae). Can. Entomol., 121: 757-762. Furniss, M. M. 1995. Biology of Dendroctonus punctatus LeConte (Coleoptera: Scolytidae). Ann. Entomol. Soc. Amer., 88: 173-182. Furniss, M. M.| 1996. Taxonomic status of Dendroctonus punctatus and D. micans (Coleoptera: Scol- ytidae). Ann. Entomol. Soc. Amer., 89: 328-333. Grégoire, J.-C. 1983. Host colonization strategies in Dendroctonus: larval gregariousness or mass attack by adults? pp. 147-154. Proc. IUFRO Conf. Role of host in population dynamics of forest insects. Banff, Alta., Can. Sept. 4-7, 1983. Grégoire, J.-C. 1988. The greater European spruce beetle, pp. 455-478. In Berryman, A. A. [ed.], Dynamics of forest insect populations: patterns, causes, implications. Plenum, New York. Higby, P K. & M. W. Stock. 1982. Genetic relationships between two sibling species of bark beetle (Coleoptera: Scolytidae), Jeffrey pine beetle and mountain pine beetle, in northern California. Ann. Entomol. Soc. Amer., 75: 668—674. Langor, D. W. & J. R. Spence. 1991. Host effects on allozyme and morphological variation of the mountain pine beetle, Dendroctonus ponderosae Hopkins (Coleoptera: Scolytidae). Can. En- tomol., 123: 395-410. Nei, M. 1978. Estimation of average heterozygosity and genetic distance from a small number of individuals. Genetics, 89: 583-590. Rogers, J. S. 1972. Measures of genetic similarity and genetic distance. Studies in Genetics, Univ. Texas Publ 7213: 145-153. Stock, M. W. & G. D. Amman. 1980. Genetic differentiation among mountain pine beetle populations from lodgepole pine and ponderosa pine in northeast Utah. Ann. Entomol. Soc. Amer., 73: 472-478. Stock, M. W., J-C. Grégoire & M. M. Furniss. 1987. Electrophoretic comparison of European Den- droctonus micans and ten North American Dendroctonus species (Coleoptera: Scolytidae). Pan- Pac. Entomol., 63: 353-357. Swofford, D. L. & R. B. Selander. 1989. BIOSYS-1. A computer program for the analysis of allelic 1997 KEGLEY ET AL.: COMPARISON DENDROCTONUS SPP. 45 variation in population genetics and biochemical systematics. Illinois Natural History Survey. Univ. of Illinois, Urbana-Champaign. Wood, S. L. 1963. A revision of the bark beetle genus Dendroctonus Erichson (Coleoptera: Scoly- tidae). Grt. Bas. Nat., 23: 1-117. Wood, S. L. 1982. The bark and ambrosia beetles of North and Central America (Coleoptera: Scol- ytidae), a taxonomic monograph. Grt. Bas. Nat. Mem. 6. Wood, S. L. & D. E. Bright, Jr. 1992. A catalog of Scolytidae and Platypodidae (Coleoptera). Part 2: Taxonomic index. Grt. Bas. Nat. Mem. 13. Received 16 Jan 1996; Accepted 9 Oct 1996. PAN-PACIFIC ENTOMOLOGIST 73(1): 46, (1997) Scientific Note THE FIRST RECORD OF THE ANT PHEIDOLE MOERENS WHEELER FROM THE WESTERN UNITED STATES (HYMENOPTERA FORMICIDAE) Pheidole moerens (Wheeler) is native to the West Indies, originally described from Puerto Rico in 1908 by Wheeler in the U.S. It is known from Florida and Southern Alabama (Naves M.A. 1985. A monograph of the genus Pheidole in Florida (Hymenoptera Formicidae). Insecta Mundi, 1: 53—90). Pheidole moerens is easily confused with P. floridana (Emery) of the southeastern U.S. and which is not known from California. The light brown minor worker is minute, between 1.5 and 1.75 mm in total body length. The larger major worker is also small: about 2.5 to 2.75 mm in total body length and darker in color than the minor worker with a disproportionately larger and broader head that is characteristic of the genus. This ant is highly variable in color. On 8 Nov 1995, in a section of Shoreline Aquatic Park in the city of Long Beach called Palm Island, and in an adjacent area, I discovered P. moerens nesting at the base of and in the bark of several California fan palms, Washingtonia filifera (Lindley) Wendland. The trees are from 6 to 11 m high and 46 to 61 cm in diameter at the base. Workers, and one alate male, were collected on the above date and more workers and female alates were collected on 18 Nov and 9 Dec 1995. I also observed brood as I dug around the base of the trees and pulled apart some bark. The nests of P. moerens are small to moderate in size and were only found around the California fan palms. There were no nests along the edges or cracks in the sidewalk. In the immediate vicinity there are large aggressive nests of the southern fire ant, Solenopsis xyloni (McCook) which mostly nests on the edge of and cracks in the side walk; there are some nests around the Fan palms and at least one P. moerens nest was taken over by S. xyloni. Pheidole moerens appears to be a general scavenger ant with little or no economic importance. Material Examined.—CALIFORNIA LOS ANGELES Co.: Long Beach Shore- line Aquatic Park 200 West Shoreline Dr. Entrance on Shoreline Drive and Pine Ave. on Palm Island just southeast of the entrance across the lagoon on 8 Nov 1995. M.J. Martinez. Acknowled gments.—I thank my wife Charlean for her support; Dr. Rosser W. Garrison Los Angeles County entomologist and Robert J. Hamton, a fellow myr- mecologist, for confirming my identification of this ant to the Flavens group and for reviewing the manuscript; Roy R. Snelling Los Angeles County Natural His- tory Museum for determining this species as Pheidole moerens; and my nephew Eric Weis for typing the manuscript. Michael J. Martinez City of Long Beach, Dept. of Parks, Recreation and Ma- rine, 2760 Studebaker Road, Long Beach California 90815. Received 31 Jan 1996; Accepted I May 1996. PAN-PACIFIC ENTOMOLOGIST 73(1): 47-48, (1997) Scientific Note AGGREGATIONS OF THAUMATOMYIA GLABRA (MEIGEN) (DIPTERA: CHLOROPIDAE) ON WISTERIA FLOWERS (FABACEAE) During April, 1995, aggregations of Thaumatomyia glabra (Meigen) were no- ticed on the flower clusters of Wisteria sinensis (Sims) Sweet, at El Dorado Hills, in the Sierra Nevada foothills of California. Over a two week period, individuals were observed to each occupy an individual blossom, and flies were spread over all available flower clusters. All sampled flies were males. On each sequentially opening inflorescence, many blossoms, often adjacent, were occupied. The flies sat motionless, usually on one side of the front portion of the white banner of the blossom, but occasionally, if disturbed, they moved to the lavender Keel of the blossom. Movement of flies between blossoms was not observed, nor were en- counters among flies on blossoms. In one instance, two flies occupied the same blossom, with one on the front of the banner, and the other on the venter of the keel, so that they apparently could not see each other. The aggregations, which formed by mid-morning and dispersed at twilight, continued until all flower clus- ters had finished blooming. Mid-instar lepidopteran larvae, which had fallen or dropped from the overhanging canopy of Quercus douglasii W.J. Hooker & G.A. Walker-Arnott, and which occasionally crawled across occupied blossoms, did not elicit a reaction from the flies. Thaumatomyia glabra is considered an “‘almost cosmopolitan’’ chloropid spe- cies with a vast array of geographically variable phena (Sabrosky, C.W. 1943. Canad. Entomol., 75: 109-117). Therefore, we assume that T. glabra either rep- resents an adaptively polytypic species, or a group of sibling species. Sabrosky’s (1943: 114) description of a phena for “‘western and far western states,’ which has rather distinct mesonotal setae and yellow fore metatarsi, most closely matches the flies that we observed; his description of certain California populations that have reduced cheeks does not match. Because Wisteria sinensis is of oriental origin, and these 7. glabra have a western Nearctic phena, it seems implausible that this behavioral association could pre-date the introduction of Wisteria to the area. Thaumatomyia are predators of root aphids [Pemphigus sp.] (Alleyne, E. & F Morrison. 1977. Ann. Soc. Entomol. Que., 22 : 181-187; Roman, E. & C. Chauve. 1979. Bull. Mens Soc. Linn. Lyon, 48: 263-267), but there is a record of T. glabra from a spider egg sac (Sabrosky 1943). Because we did not observe interaction among the sitting flies on Wisteria, it is difficult to speculate about the evolutionary benefit of this behavior, except to mention that it seemed to resemble an apparent form of leking in the absence of observed interaction. It may involve an anticipation of mates drawn to the fragrant blossoms as a resource. Vouchers are deposited at the PPDC, CDFA, Sacramento. Acknowled gement.—We thank Eric M. Fisher (CDFA-PPDC) for the identifi- cations, information and discussions. 48 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(1) Material Examined—CALIFORNIA. EL DORADO Co.: El Dorado Hills, 4-18 Apr 1995, K.H. Sorensen, ex Wisteria sinensis blossoms. John T. Sorensen! and Kathleen H. Sorensen’; 'Plant Pest Diagnostics Center, California Dept. of Food & Agriculture, Sacramento, California 95832-1448; and 2EI Dorado Hills, California. Received 2 May 1995; Accepted I Aug 1996. PAN-PACIFIC ENTOMOLOGIST 73(1): 49-51, (1997) Scientific Note OCCURRENCE OF A NEOGREGARINE PROTOZOAN, OPHRYOCYSTIS ELEKTROSCIRRHA MCLAUGHLIN AND MYERS, IN POPULATIONS OF MONARCH AND QUEEN BUTTERFLIES The monarch butterfly, Danaus plexippus (L.), along with another New World species, the queen butterfly (Danaus gilippus berenice Cramer) are both suscep- tible to a neogregarine parasite, Ophryocystis elektroscirrha McLaughlin & My- ers, in Florida. The neogregarine parasite, described by McLaughin & Meyers (1970. J. Protozool., 17: 300—305), involves a life cycle strategy that incorporates the occurrence of spores on the scales and hairs of the infected butterflies. The neogregarine has been reported from monarch butterflies in California (Leong, K. et al. 1992. Ecol. Entomol., 17: 338-342), Hawaii and Mexico (Brower, L. et al. 1995. BioSci., 45: 540-544). However, except for California, the prevalence of infection and the occurrence of this neogregarine in other populations of the monarch butterfly and a closely related species, the queen butterfly, are unknown. Accordingly, we conducted a survey of monarch butterflies from various parts of the world and queen butterflies from Florida to gain a better insight into their distribution and prevalence of infection. Our survey confirms the presence of O. elektroscirrha on monarch butterfly adults collected from Hawaii, Mexico, Florida, Australia and New Zealand (Table 1). The survey data show that the percentage of butterflies with spores (based on a wash method by Leong et al. 1992) ranged from 3.8% (Mexico) to 100% (Maui, Hawaii). Among butterflies with spores, those collected from Oahu, Hawaii had the highest average number of spores per individual (197,480) and those from New Zealand, the least (10,500). Six of the 32 queen butterflies (D. gilippus berenice) examined had neogregarine spores (18.8%) with an average of 1500 spores per individual. The spore dimensions varied considerably depending upon the geographical regions and species (Table 2). The queen butterflies collected from Fort Lauder- dale, Florida had the smallest spores whereas monarch butterflies from California, Hawaii and the queen from Gainesville, Florida, had the largest. Notably, spores from the queen butterflies had both the smallest (Fort Lauderdale) and largest (Gainesville) measurements. Spores recovered from butterflies collected in Fort Lauderdale were significantly smaller than those recovered from queen butterflies collected in Gainesville or from monarch butterflies (P < 0.01, F = 12.8, df = 7,313; Table 2). The different sizes within the queen butterflies suggest the oc- currence of distinct strains or possibly another neogregarine species, perhaps one unique to the queen. The larger neogregarine spores isolated from D. gilippus berenice from Gaines- ville, Florida, were infectious to the monarch butterfly. When leaves of the blood flower milkweed, Asclepias curassavica L., were sprayed with 50,000 spores/ml until runoff and fed to first instars, 3 of 9 (33.3%) monarch larvae became in- fected. We were unable to test the pathogenicity of the smaller neogregarine spores 50 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(1) Table 1. Isolation of neogregarine spores from monarch butterflies (Danaus plexippus [L.]) from various geographical areas and from the queen butterfly (D. gilippus berenice Cramer) from Florida. . Species/Location n #S/n® (%) Mean? (Range)° M/F? Monarch butterfly United States of America Florida Broward County zi 6/7 (85.7%) 347,200 (200—232,000) 6/1 California San Luis Obispo County 160 92/160 (57.5%) 59,470 (200-565,000) 80/80 Santa Cruz County 130 = =86/130 = (66.2%) 76,000 (400—497,000) 70/60 Hawaii Oahu County 17 14/17 (82.3%) 197,480 (200—974,000) 11/6 Maui County 2 2/2 (100%) 17,900 (2000—33,800) 1/1 Kauai County 3 2/3 (66.7%) 47,600 (2000—94,800) 2/1 Hawaii County 7 5/7 (71.4%) 42,600 (400-—240,000) 6/1 Australia 20 12/20 (60%) 53,400 (200—436,000) 10/10 New Zealand f 1/7 (14.3%) 300 (300) 3/4 Mexico 26 1/26 (3.8%) 1400 (1400) 10/16 Queen butterfly Florida (combined) 32 6/32 (18.8%) 1500 (200-8600) 23/9 Gainesville 14 2/14 (14.3%) 600 (200-1000) 10/4 Fort Lauderdale 18 4/18 (22.2%) 2400 (200-8600) 13/5 @= no. of individuals with spores/total number of individuals. > = average spore level from abdomens of butterflies. ¢ = range of spore levels among individuals with spores. d = male/female. from Fort Lauderdale to monarchs because they were recovered earlier in our investigation (January 1993), and a colony of protozoan-free monarch butterflies was not available at that time. Neogregarine spores recovered from the monarch butterflies were not infectious to the silkworm, Bombyx mori (L.). When 30 first instar silkworm larvae were Table 2. The length, diameter and area of neogregarine spores recovered from monarch and queen butterflies from various geographical areas. Spore area with different superscript letters are significantly different (P < 0.01, F = 12.8; df = 7313). Species/Location n Length (um) Diameter (um) Area (um?) Queen, Fort Lauderdale, Florida, USA 37 1283 > OD S102 101.2 + 2.63 Monarch, Australia 50 13.9+ 0.1 7.8 + 0.3 114.0 + 1.7 Monarch, New Zealand 22 13.0 + 0.2 8.9 + 0.2 L722 Monarch, Fort Lauderdale, Florida, USA 50 14.1 + 0.1 8.4 + 0.1 118.2 + 1.2% Monarch, Mexican 42 13.9 + 0.1 8.6 + 0.2 119.6 + 2.9% Monarch, California, USA 50 13.7 + 0.1 8.8 + 0.1 120.6 + 2.2°¢ Monarch, Hawaii, USA 50 14.3 + 0.1 8.9 + 0.1 127.3-32 IR9S Queen, Gainesville, USA 30 13.2 4°02 96+ 0.1 IZT Ast 26° 1997 SCIENTIFIC NOTE 51 fed mulberry leaves sprayed with 100,000 spores/ml until run-off, none of the resulting adults had neogregarine spores. Thirty-five striated queen butterflies, D. g. strigosus (Bates), a subspecies found in the desert regions of Colorado and southern California, were examined for neogregarine spores. None of the adults surveyed had neogregarine spores. Their absence in the striated queen population may reflect an inadequate sample size, infection levels too low to be detected with our method, or the resistance of this subspecies to the protozoan. A more likely explanation is that the striated queen is susceptible to the protozoan, but are not exposed to the parasite because they do not share a common milkweed host plant with the monarch butterfly. The Striated queen larvae feed mainly on the rambling milkweed, Sarcostemma hir- tellum R. Holm, a plant not used for oviposition by monarch butterflies. The populations of queen and monarch butterflies of south central Florida are ecolog- ical competitors during early spring (March-May). The adults of the two species are found in the same habitat where they will feed on similar nectar sources and Oviposit on common Asclepias plants. The larvae of the two species have been reported on the same milkweed host (Brower, L. 1961. Ecol., 42: 76-83). Our previous observations indicated that the neogregarine is passed vertically from one generation to another by infected adults contaminating eggs or milkweed leaf surfaces with spores directly or with scales containing spores during ovipo- sition. The larvae become infected when they ingest the spores. Our survey data show that the infection level of butterflies overwintering in California ranges from 200 to > 900,000 spores per individual. Even at the higher levels of infection in the California monarch butterfly populations, the protozoan appeared to have little effect on the butterfly’s winter survival and mating successes (unpublished data). The low pathogenicity of this parasite on its host in nature has allowed it to persist widely within populations of monarch butterflies. Studies on the genus Danaus suggest that the monarch butterfly evolved in the New World, probably South or Central America (Kitching, I. et al. 1993 pp. 11-16. In Biology and conservation of the monarch butterfly, Natural History Museum of Los Angeles County, Los Angeles, CA). We hypothesize that the neogregarine co-evolved with its host and because of the high rate of infection, the parasite moved with its host into other geographic regions. We cannot discount the possibility that the queen butterfly or other related Danaid species were the original host for this neogreg- arine which subsequently became adapted to the monarch butterfly. Regardless, both the queen and monarch butterfly are hosts of O. elektroscirrha. Examination for neogregarine spores on other closely related species to the queen and monarch butterflies may provide further insights into the distribution of O. elektroscirrha and its host spectrum. Acknowled gment.—Authors thank J. Capinera, R. Giblin-Davis, T. A. Jackson, L. LeBeck, W. Sakai, L. Tsutsumi, and M. P. Zalucki for the monarch butterflies used in this survey. Authors page charges partially offset by a grant from the C. P. Alexander Fund, Pacific Coast Entomological Society. Kingston L. H. Leong,' Michael A. Yoshimura,' & Harry K. Kaya, ? 'Biological Sciences, California Polytechnic State University, San Luis Obispo, California, 93407; *Department of Nematology, University of California, Davis, California 95616. Received 15 May 1996; Accepted 15 Aug 1996. PAN-PACIFIC ENTOMOLOGIST 73(1): 52-54, (1997) Scientific Note FLOWER-VISITORS OF BACCHARIS PILULARIS DE CANDOLLE SUBSP. CONSANGUINEA (DE CANDOLLE) C.B. WOLF (ASTERACEAE) IN BERKELEY, CALIFORNIA Coyote brush (Baccharis pilularis De Candolle subspecies consanguinea (De Candolle) C.B. Wolf (Asteraceae)) is a dioecious evergreen perennial, native throughout cismontane California, Baja California, and as far north as Oregon (Wright, A.D. 1928. Ph.D. Thesis, University of California, Berkeley). It is a relatively common xerophyte ranging in altitude from sea-level to approximately 400 m, and although it does not compete well in areas where evaporation rates are particularly high, it can withstand seasonal drought stress (Wright 1928). Til- den’s (Tilden, J.W. 1951. Microentomology, 1: 149-188) study of B. pilularis catalogued the arthropod associates of coyote brush; this initial work, however, did not include the suite of insects visiting the flowers of coyote brush. As a supplementary study, I have provided a list of the insect flower-visitors collected in Strawberry Canyon in 1992. Large stands of coyote brush exist in the scrub oak communities of Strawberry Canyon (Berkeley, Alameda Co., CA). The two study sites of this project (both approximately 70 sq. meters) were within such plant communities. In addition to coyote brush, both sites had dense populations of Avena barbata Pott ex Link (Poaceae) (slender wild oat), Brassica nigra (L.) W.D.J. Koch (Brassicaceae) (black mustard), and Silybum marianum (L.) Gaertn. (Asteraceae) (milk thistle). The following plants occurred in much lower densities: Foeniculum vulgare P. Mill. (Apiaceae) (sweet fennel), Carduus sp. (L.) (Asteraceae), Bromus hordea- ceus L. (Poaceae) (soft chess), Cirsium arvense (L.) Scop. (Asteraceae) (Canada thistle), Genista monspessulana (L.) L. Johnson (Fabaceae) (French broom), Er- iogonum latifolium Sm. (Polygonaceae) (buckwheat), Phalaris aquatica L. (Po- aceae), Heteromeles arbutifolia (Lindl.) M. Roemer (Rosaceae), Quercus agrifolia Nee (Fagaceae) (live oak), and Nassella lepida (A.S. Hitchc.) Barkworth (Po- aceae) (needle grass). Soil moisture (% water of a 20cm soil-core) at both sites was approximately 7% during the sampling period. Daytime temperatures ranged from about 16° C to 37°. During the peak coyote brush flowering period in 1992 (mid-September through mid-October), insects visiting the inflorescences of gynoecious (female) and an- droecious (male) coyote brush plants were collected. The pistillate flower of coy- ote brush is a brush-type flower; the staminate flower is a disk-type. Collections were made on 20 and 26 Sep and 4 and 10 Oct 1992. They commenced at about 08:45 h, paused from noon to 12:45 h, and continued until about 15:45 h. Any insect seen on a flower or hovering directly above an inflorescence was collected using a small net and aspirator. Identifications were done by various specialists, as well as by the author. Representatives of at least 55 insect species were collected (five orders and 32 1997 SCIENTIFIC NOTE Table 1. List of insects visiting Baccharis flowers. Order/Family/(Subfamily) Hemiptera Cixiidae Lygaeidae undet. nymph undet. nymph Coleoptera Coccinellidae Chrysomelidae Staphylinidae Lepidoptera Nymphalidae Diptera Agromyzidae Anthomyiidae Anthomyiidae/Muscidae Bombyliidae Chamaemyiidae Muscidae Sarcophagidae Syrphidae (Syrphinae) (Microdontinae) Tachinidae Tephritidae Hymenoptera Apidae Braconidae (Agathidinae) (Braconinae). (Microgastrinae) Chalcididae Colletidae Eulophidae (Tetrastichinae) Eumenidae Eurytomidae Formicidae Genus/Species undetermined adult Nysius sp. Cryptolaemus montrouzieri Mulsant Psyllobora vigintimaculata Say Rhyzobius forestieri Mulsant Diabrotica undecimpunctata Mannerheim Diachus sp. undet. sp. Junonia coenia Hubner undet. sp. 1 sp. 2 undet. spp. undet. spp. Mythicomyia sp. 1 Mythicomyia sp. 2 Leucopsis sp. Coenosia sp. undet. spp. Allograpta sp. Paragus sp. Sphaerophoria sp. Syritta pipiens (L.) Chetogena parvipal pus Wulp Microchaetina sp. Tephritus sp. Trupanea sp. Apis mellifera L. Bombus sp. Agathis gibbosa (Say) Atanycolus sp. Apanteles sp. 1 (metacarpalis spp. group) Apanteles sp. 2 (ater spp. group) Apanteles sp. 3 (metacarpalis spp. group) Apanteles sp. (males) (metacarpalis spp. group) Dolichogenidea sp. (laevigatus spp. group) Spilochalcis sp. Hylaeus sp. Aprostocetus sp. undet. sp. Eurytoma sp. Linepithema humile (Mayr) KS WOW He — — — NON Pe FK OWN KF WN Nr RKP NR Rr KN ON — Re NWN KR OK CO — — —— O 80 53 54 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(1) Table 1. Continued. Order/Family/(Subfamily) Genus/Species No. Ichneumonidae (Cremastinae) undet. sp. i} Platygastridae Synopeas sp. 66 Pompilidae undet. sp. 1 1 sp. 2 1 sp. 3 1 Pteromalidae (Pteromalinae) undet. sp. 1 1 sp. 2 3 Sphecidae Sceliphron caementarium Drury 1 undet. sp. 1 Torymidae Megastigmus sp. 1 Unidentified Chalcidoidea undet. sp. 5 Vespidae Vespula pensvlvanica Saussure 2 families) including an undescribed Synopeas species near anomaliventre (Ash- mead) (Table 1.). Particularly well represented were Line pithema humile (Mayr) (Hymenoptera: Formicidae), Agathis gibbosa (Say) (Hymenoptera: Braconidae), Synopeas sp. (Hymenoptera: Platygastridae), Microgastrinae (Hymenoptera: Bra- conidae), and several chalcidoid species (Hymenoptera). Hymenoptera comprised approximately 81% of all insect specimens, Diptera accounted for 10%, and the remaining orders, 9%. It is worth noting that foraging A. gibbosa females fre- quently probed pistillate inflorescences with their ovipositors. The individual would repeatedly insert its ovipositor into the side of the flower and angle the thrusting motion downward. This behavior was restricted to A. gibbosa and usu- ally occurred whenever the wasp was present at a pistillate flower. Acknowled gment.—The author thanks Robert L. Bugg for support throughout the project (Information Group, SAREP, University of California, Davis). Leo- poldo Caltagirone (Laboratory of Biological Control, University of California, Berkeley) provided laboratory space and guidance in the identification of para- sitoids. Kenneth Hagen (Laboratory of Biological Control, University of Califor- nia, Berkeley) identified all Coleoptera as well as providing much support. Paul Amaud (Department of Entomology, California Academy of Sciences, San Fran- cisco) identified the Diptera. Robert Zuparko (Laboratory of Biological Control, University of California, Berkeley) and Howell Daly (Department of Environ- mental Science, Policy, and Management, University of California, Berkeley) helped with the identification of some Hymenoptera. Robert Wharton (Department of Entomology, Texas A&M University, College Station) provided the species name of the many Agathis specimens. James Whitfield (Department of Entomol- ogy, University of Arkansas, Fayatteville) provided the group names of the mi- crogastrines. Author page charges partially offset by a grant from the C.P. Alex- ander Fund, Pacific Coast Entomological Society. Shawn A. Steffan, Department of Entomology, University of Wisconsin, Mad- ison, Wisconsin 53706. Received 13 Feb 1996; Accepted 27 Sep 1996. PAN-PACIFIC ENTOMOLOGIST 73(1): 55-57, (1997) SCIENTIFIC NOTE THE IDENTITY OF CHELANOPS SERRATUS MOLES (PSEUDOSCORPIONIDA: CHERNETIDAE) Chelanops serratus was described on the basis of a single individual found ‘fon the window pane of the Pomona College greenhouse’”’ [Pomona, Los Angeles County, California] (Moles, M. M.[sic] 1914. J. Entomol. Zool., 6: 187-197). The specimen was mounted on a slide, which was said to be “‘so poorly made”’ [prob- ably by a student at Pomona College] that it was impossible to take measurements of any parts but the palps. The whereabouts of that holotype is now unknown, and it must be presumed lost. Nevertheless, the species was described in sufficient detail that many important characters are clear, especially of the palp, which was illustrated (1914: fig. 3). It is obvious that this is the same pseudoscorpion species as that later described by Chamberlin as Dinocheirus sicarius (1952. Bull. Amer. Mus. Nat. Hist., 99: 259-312), from the Frances Simes Hastings Natural History Reservation, Monterey County, California. This conclusion is supported by my own study of numerous representatives of the species from many locations in California, from San Diego County in the south to Modoc County in the north. Dinocheirus serratus (Moles) Chelanops serratus Moles, 1914: 193, fig. 3; Moles, M. & W. Moore. 1921. J. Entomol. Zool., 13: 7. Dinocheirus? serratus (Moles): Hoff, C. C. 1958. Amer. Mus. Novitates, 1875: 28. Dinocheirus serratus (Moles): Harvey, M. S. 1991. Catalogue of the Pseudo- scorpionida. Manchester Univ. Press: 573. Dinocheirus sicarius Chamberlin, 1952: 279-292, figs. 6-9; Hoff 1958: 28; Har- vey 1991: 573 (complete synonymy to 1989). NEW SYNONYMY Although Moles’s specimen of Chelanops serratus was apparently badly dam- aged, she was able to provide enough detail to allow some comparison with other chernetids in California. A glance at her illustration of the palp immediately sug- gests a close relationship to Dinocheirus sicarius Chamberlin. Though Moles did not mention the sex of her specimen, it is evident that it was a male, as the palp is like that of the holotype of D. sicarius (see Figs. 1, 2). The shapes and pro- portions of the segments are very similar—distinctive are the robust palpal seg- ments, especially the chela of the male, and the unusual configuration of the medial side of the femur. The latter was described by Moles (1914: 195), “‘femur - - - pedicellate, inner margin almost straight at base, then suddenly concave to tip,’’ and by Chamberlin (1952: 280), “unique angular protuberance on the inner or subdorsal face of the palpal femur of the male.’’ The form of the palpal femur is diagnostic for this species; I do not know of any other chernetid that looks like this. Actually, there is considerable variation in size and proportions of the palpal segments, especially in males, so that the chela may be more or less stout and the protuberance of the femur more or less pronounced (Chamberlin 1952: 282; personal observation). Sizes are comparable: Moles gave only the length of the pedipalp, 3 mm (1914: 56 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(1) Figure 1. Chelanops serratus, outline of palp of holotype (redrawn from Moles 1914: fig. 3). Figure 2. Dinocheirus sicarius, outline of palp of holotype male (redrawn from Chamberlin 1952: fig. 6B). 193); this compares well to 3.15 mm, the sum of the average lengths of the palpal segments reported by Chamberlin (1952: 284). Setae of the palps are similar: Chamberlin characterized them as “thickened and variously denticuloclavate’’ (p. 282); in profile, these appear “‘saw-like’’ (serrate), as described by Moles (p. 195). Moles described the cheliceral galea, or spinneret, as “‘small and transparent’ (p. 195), while Chamberlin showed that of the male to be smaller and more slender than that of the female (1952: fig. 6E, F). Moles and Chamberlin agreed that the carapace is granulate, but Moles (p. 193) apparently erred in stating that there are “no eye spots’’ (2 distinct eyespots are present) and only “one distinct median suture”’ (2 transverse furrows are present); it is understandable that she might not have seen these details on the damaged specimen she was studying. Altogether, it is clear that Moles and Chamberlin were dealing with the same species and that Dinocheirus sicarius Chamberlin (1952) is a junior synonym of Chelanops serratus Moles (1914). It is also quite clear, from the very detailed description by Chamberlin, that the species is a representative of the genus Di- nocheirus Chamberlin (see also Muchmore, W. B. 1974. J. Arachnol., 2: 34). The known range of Dinocheirus serratus now extends from San Diego County, California northward to Columbia County, Oregon (see below and Benedict, E. M. & D. R. Malcolm. 1982. J. Arachnol., 10: 100); there is also an unsubstantiated record from the Great Salt Lake Desert, Utah (Gering, R. L. 1956. p. 50 in: Woodbury, A. M., [ed.] Ecological Check Lists. The Great Salt Lake Desert Se- ries, Univ. of Utah, Dugway). I have made detailed studies on the following specimens. Material Examined.—CALIFORNIA. ALAMEDA Co.: Berkeley, nest of Neotoma fuscipes Baird, Nov 1926, 1 male, 3 females. COLUSA Co.: 4 km W of Stonyford, Neotoma nest at base of oak, 30 Apr 1980, E G. Andrews, 6 males, 9 females, 6 nymphs. MODOC Co.: Big Sage Reservoir, Neotoma nest at base of Juniperus occidentalis Hooker, 31 May 1978, E G. Andrews, 1 male. RIVERSIDE Co.: Mt. San Jacinto, 1250 m, litter from hollow of fallen oak, 28 Mar 1978, K. W. Cooper, 1 male, 2 females, 5 nymphs. SAN BERNARDINO Co.: E of Summit [only 40 km from Pomona], damp Neotoma nest, 5 Jun 1979, K. W. Cooper, 1 male, 3 females, 4 nymphs. SAN DIEGO Co.: 1.5 km E of Leucadia, 1997 SCIENTIFIC NOTE 57 Neotoma nest in chaparral, very dry, 8 Aug 1979, K. W. Cooper, 7 males, 5 females, 4 nymphs. (All deposited in Florida State Collection of Arthropods, Gainesville). Acknowled gment.—I am greatly indebted to EK G. Andrews and K. W. Cooper for sending me many pseudoscorpions for study. Author page charges partially offset by a grant from the C. P. Alexander Fund, Pacific Coast Entomological Society. William B. Muchmore, Department of Biology, University of Rochester, Roch- ester, New York 14627. Received 13 Feb 1996; Accepted 16 Oct 1996. PAN-PACIFIC ENTOMOLOGIST 73(1): 58-59, (1997) Scientific Note WEEVILS NEW TO THE STATE OF WASHINGTON (COLEOPTERA: CURCULIONIDAE) During 1994-95, a qualitative study to evaluate insect biodiversity was con- ducted at the Hanford Site which is located in southcentral Washington State. Situated in the semi-arid Columbia Plateau Basin, this 560 square mile site was closed to the general public in the early 1940s. Originally acquired by the United States federal government as a site for the production of plutonium to be used in weapons production, the site is currently administered by the Department of En- ergy for nuclear waste management, environmental restoration, and research and development. Our studies were confined primarily to the Fitzner-Eberhardt Arid Lands Ecol- ogy Reserve (ALE). An area of over 100 square miles, the ALE is located in the southwestern portion of the Hanford Site (latitude N 46°, longitude W 119°). Physiographically diverse, the site consists of a steeply rising, northeast facing slope (Rattlesnake Ridge 1150 m) and extensive flats that slope gently from 500 to 150 m. Vegetation consists primarily of a sagebrush-bitterbrush/Sandberg’s bluegrass-cheatgrass type, the general habitat is referred to as a shrub-steppe. Thirty-three species of Curculionidae were collected during this study. Twenty- six are species more common to the central basin of Washington State and are associated with primary vegetation including rabbitbrush, sagebrush, lupine, and balsamroot (Hatch, M. H. 1971. The Beetles of the Pacific Northwest., Univ. Washington Press). Seven of the species collected have previously not been re- corded for Washington. Distributional data primarily are from O’Brien and Wib- mer (1982. Mem. Amer. Entomol. Institute, 34: 1-382). The majority of speci- mens are in the M. T. James Entomological Collection, Washington State Uni- versity; voucher specimens are in the private collection of the senior author (CWOB). A list of species and exact locations for individual collections within the ALE are available from RSZ. First documented records for species from Washington State: Anthonomus cycliferus Fall: widespread but spotty distribution throughout the western states. Anthonomus sphaeralciae Fall: widespread throughout the southwest and cen- tral states, the species has also been found in Idaho. It is known to feed on several species of Sphaeralcea (Malvaceae) one of which, S. munroana (Dougl.) Spach, is widespread on the Hanford site. Ceutorhynchus erysimi (Fabr.): an introduced species, it is widespread through- out the eastern United States and Canada but previously recorded only from Or- egon in the western United States. The species feeds on various Cruciferae. Cleonidius erysimi (Fall): previously known only from scattered locations throughout the western United States and Canada. It has been predominantly taken in sand dune habitats, as were our specimens. Adults and immatures have been collected from a variety of plants, primarily Cruciferae and some Compositae (Anderson, R. S. 1988. Quaest. Entomol., 23: 431-709). 1997 SCIENTIFIC NOTE 59 Gymnetron pascuorum (Gyllenhal): an introduced species that is widespread throughout much of the eastern and western United States. It has been reared from Plantago lanceolata (L.) (English plantain) (Hatch, M. H. 1971. The Beetles of the Pacific Northwest. Univ. Washington Press) which is widespread and com- mon on the Hanford site. Lepesoma remota (Van Dyke): previously known only from Oregon. Mecinus pyraster (Herbst): an introduced species previously known only from Maryland, New Jersey, Florida, Virginia, and Oregon. The primary larval host is Plantago (Plantaginaceae) but Mecinus have also been found feeding in several genera of Scrophularaceae (Warner, R. E. 1955. Entomol. News, 66: 209-211). Acknowled gement.—This project was funded by The Nature Conservancy with awards from the U. S. Department of Energy, The Nature Conservancy of Wash- ington State, and The Bullitt Foundation. Charles W. O’Brien, Entomology-Biological Control, Florida A&M University, Tallahassee, Florida 32307-4100 and Richard S. Zack, James Entomological Col- lection, Department of Entomology, Washington State University, Pullman, Wash- ington 99164-6382. Received 27 Feb 1996; Accepted 16 Oct 1996. PAN-PACIFIC ENTOMOLOGIST 73(1): 60, (1997) Book Review Bleuzen, P. 1994. Les Coleopteres du Monde (The Beetles of the World). Prion- inae 1: Macrodontini: Macrodontia, Chalcoprionus, Ancistrotus, Acanthinod- era, Acalodegma: Prionini: Titanus, Braderochus. Volume 21. Sciences Nat. 92 pp., 16 plates. This is another volume in the marvelous series of beetle picture books. The text is done in French and English (Brian Morris). This volume contains keys for the determination of genera and species treated. There are distribution maps for the species although single locality records appear to be of little value in ascer- taining distributions. One new species of Macrodontia, M. jolyi is described from Venezuela. The genus Titanus Audinet-Serville is treated as monotypic and Braderochus Buquet is elevated to generic status with five species, three of these new (B. jolyi from Venezuela and Guyana; B. salcedoi from Venezuela, and B. shuteae from Hon- duras and Panama). One of these is unfortunately a synonym. The illustrations are of the usual superb quality of this series and are a great aid in species determinations. The bilingual text is very useful but the English version is often awkward and somewhat incomprehensible, probably due to a very literal translation from the French. This, however, does not appreciably detract from the overall usefullness of the book and the modest price of $150.00 makes this a very desirable item for stu- dents and collectors of large beetles, particularly Cerambycidae. The book is now available from Sciences Nat, 2, rue Andre Mellene, 60200 Venette, France. John A. Chemsak, Division of Insect Biology, University of California, Berke- ley, CA. 94720. PAN-PACIFIC ENTOMOLOGIST Information for Contributors See volume 66(1): 1-8, January 1990, 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 11 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 corner. 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 KEY WORDS. 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. FE Eastop. 1987. Problems in pest aphid taxonomy: can chromosomes plus morphometrics provide some answers? pp. 233-238. Jn Holman, J., J. Pelikan, A. G. EK 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. THE PAN-PACIFIC ENTOMOLOGIST Volume 73 January 1997 Number 1 Contents NELSON, G. H.—A new Poecilonota from southern California (Coleoptera: Buprestidae) MURPHY, B. C., J. ADAMS & M. P. PARRELLA—Experimental arena for confining thrips and other small arthropods in the laboratory DRAPEK, R. J., B. A. CROFT & G. FISHER—An examination of spatial input parameters in order to improve corn earworm (Lepidoptera: Noctuidae) damage predictions for a pher- omone trap catch regression WELLS, J. D. & H. KURAHASHI—Chrysomya megacephala (Fabr.) is more resistant to attack by Ch. rufifacies (MacQuart) in a laboratory arena that is Cochliomyia macellaria (Fabr.) (Diptera: Calliphoridae) GULMAHAMAD, H.—Ecological studies on Cardiocondyla ectopia Snelling (Hymenoptera: Formicidae) in southern California VETTER, R. S., S. TATEVOSSIAN & T. C. BAKER—Reproductive behavior of the female carob moth, (Lepidoptera: Pyralidae) SHEPARD, W. D.—Lilioceris sp. (Coleoptera: Chrysomelidae) herbivory on Cycas siamensis Miguel (Tracheophyta: Cycadales) KEGLEY, S. J..M. M. FURNISS & J. GREGOIRE—Electrophoretic comparison of Dendroc- tonus punctatus LeConte and D. micans (Kugelann) (Coleoptera: Scolytidae) -...--------- SCIENTIFIC NOTES MARTINEZ, M. J.—The first record of the ant Pheidole moerens Wheeler from the western United States (Hymenoptera: Formicidae) SORENSEN, J. T. & K. H. SORENSEN—Aggregations of Thaumatomyia glabra (Meigen) (Diptera: Chloropidae) on Wisteria flowers (Fabacae) LEONG, K. L. H., M. A. YOSHIMURA, & H. K. KAYA—Occurrence of a neogregarine protozoan, Ophryocystis elektroscirrha McLaughlin and Myers, in populations of mon- arch and queen butterflies STEFFAN, S. A.—Flower-visitors of Baccharis pilularis De Candolle subsp. consanguinea (De Candolle) C. B. Wolf (Asteraceae) in Berkeley, California MUCHMORE, W. B.—The identity of Chelanops serratus Moles (Pseudoscorpionida: Cher- netidae) O’BRIEN, C. W. & R. S. ZACK—Weevils new to the State of Washington (Coleoptera: Cur- culionidae) BOOK REVIEW CHEMSAK, J. A.—Bleuzen, P. 1994. Les Coleopteres du Monde (The Beetles of the World). Prioninae 1: Macrodontini: Macrodontia, Chalcoprionus, Ancistrotus, Acanthinodera, Acalodegma: Prionini: Titanus, Braderochus. Vol. 21. Sciences Nat. 92pp., 16 plates _ 21 28 36 40 46 47 49 52 oS) 58 60 The PAN-PACIFIC ENTOMOLOGIST Volume 73 April 1997 Number 2 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. V. Dowell, Editor R. M. Bohart R. L. Penrose, Associate Editor J. T. Doyen R. E. Somerby, Book Review Editor J. E. Hafernik, Jr. Julieta KF Parinas, Treasurer Warren E. Savary Published quarterly in January, April, July, and October with Society Proceed- ings usually appearing in the October issue. All communications regarding non- receipt of numbers should be addressed to: Vincent KF Lee, Managing Secretary; and financial communications should be addressed to: Julieta F Parinas, Treasurer; at: Pacific Coast Entomological Society, Dept. of Entomology, California Acad- emy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599. Application for membership in the Society and changes of address should be addressed to: William Hamersky, Membership Committee chair, Pacific Coast Entomological Society, Dept. of Entomology, California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599. Manuscripts, proofs, and all correspondence concerning editorial matters (but not aspects of publication charges or costs) should be sent to: Dr. Robert V. Dowell, Editor, Pan-Pacific Entomologist, California Dept. of Food & Agriculture, 1220 N St., Sacramento, CA 95814. See the back cover for Information-to-Con- tributors, and volume 66(1): 1—8, January 1990, for more detailed information. Information on format for taxonomic manuscripts 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. Pacific Coast Entomological Society OFFICERS FOR 1997 Warren E. Savary, President Vincent EK Lee, Managing Secretary Julieta EF Parinas, Treasurer Stanley E. Vaughn, Recording Secretary 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 22 April 1997 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 73(2): 61-69, (1997) FOUR NEW SPECIES OF COSTA RICAN CERAEOCHRYSA (NEUROPTERA: CHRYSOPIDAE) NORMAN D. PENNY Department of Entomology, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118 Abstract——Four new species of Costa Rican Ceraeochrysa are described, and compared to closely related congeneric species. Resumo.—Cuatro nuevas especiés de Ceraeochrysa costaricense son descrito, y comparadas a unas especiés cerca relativas congenericas. Key Words.—Insecta, Neuroptera, Chrysopidae, Ceraeochrysa, Costa Rica. The genus Ceraeochrysa was erected by Adams (1982) for 24 species of green lacewings distributed from southern Canada to Argentina. More recently, Brooks and Barnard (1990) included 40 valid species in this genus. They are primarily tropical, and some species may be valuable as biological control agents in plan- tation ecosystems (Adams & Penny 1987). Ceraeochrysa species are medium- sized (forewing length 9 to 15 mm) pale green lacewings that usually bear red or black markings on the pronotum and, less commonly, on the labial palps. An elon- gate gonapsis, the presence of entoprocesses, a decurved mediuncal apex, and the absence of a tignum are diagnostic traits of the male genitalia. In a recent study of the Neuroptera fauna of Costa Rica, 17 species of Ceraeo- chrysa were examined, five of which are undescribed. Four of the latter species are described here. The available material of the fifth species, a single female, is insufficient to warrant its description at the present time. MATERIALS AND METHODS The apical part of the abdomen of each specimen was broken off with fine for- ceps and macerated in 10% KOH, stained in Chlorazol Black E, and preserved in a glycerin-filled microvial pinned beneath the rest of the specimen. Wing tracings were made with a Ken-a-vision microprojector from temporary wing mounts on microscope slides. Following illustration, wings were glued to cards pinned be- neath the appropriate specimen. Body and genital drawings were made with the aid of a micrometer grid. Morphological terminology follows Adams & Penny (1987). CERAEOCHRYSA NIGRIPEDIS PENNY, NEW SPECIES Types.—Holotype, male: COSTA RICA. PUNTARENAS: Monteverde Biologi- cal Reserve, La Casona Station, UTM map coordinates L-N 253250, 449700, 1520 m, Nov 1991, N. Obando. Bar Code: INBIO, CR1000, 602561. Holotype depos- ited: Instituto de Biodiversidad INBIO), Santo Domingo de Heredia, Costa Rica. Allotype female, same data as holotype, except Bar Code: INBIO, CR1000, 602560; deposited California Academy of Sciences, San Francisco, California. Description.—Head (Fig. 3). Vertex, frons, clypeus, genae, labrum, and palpi pale yellow. Antennal scape, pedicel, and flagellum pale, without markings; flagellum slightly shorter than forewing length. 62 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(2) Thorax (Fig. 2). Pronotum wider than long, pale yellow, immaculate, with four small indentations. Meso- and metanota pale yellow, with a pair of round dark brown spots at the suture between pre- scutum and scutum on mesonotum and laterally on scutum of metanotum. Pleural and sternal areas pale yellow. Each leg pale yellow, except apical tarsomere contrastingly dark brown. Wings (Fig. 1). Forewing length—12.7 mm. Longitudinal veins pale green; crossveins and apical twiggings dark brown, except for apical costal crossveins. Gradate veins dark with strong infuscation of membrane along inner series. Seven inner and eight outer gradate veins. Very dark markings at apex of 1A, posterior cubitus, and cua-cup crossvein, forming a point of visual attraction on the wing. Hindwing length—11.5 mm. All veins pale green, except for most gradate veins dark. Six inner and seven outer gradate veins. Abdomen. Pale yellow. Female subgenitale (Fig. 4) broadly heart-shaped, with deep central cleft. Spermatheca strongly arched anterior to spermathecal ducts. Male ectoproct + tergite 9 (Fig. 5) short and broad, with scattered setae with thickened bases; dorsal apodeme simple, reaching callo cerci, without ventral lobe. Sternite 9 with broad, latero-dorsal subapical projection; numerous setae with thickened bases. Gonarcus (Figs. 7-8) medially narrow, with broad lateral plates. Ventral arms of gonarcus laterally embedded in simple, membranous gonosaccus. Entoprocesses of gonarcus- absent. Mediuncus very long, narrow, straight, with decurved apical point. Gonapsis (Fig. 6) elongate, with slightly upturned and expanded, smoothly rounded apex. Diagnosis.—No other species of Ceraeochrysa has this spotting pattern on the thorax, nor the darkened apical tarsal segment. Ceraeochrysa nigripedis appears to be most closely related to C. tauberae, with which it shares an elongate, straight arcessus and dark area of visual attraction on the forewing at 1A. They differ by C. nigripedis having the aforementioned characteristics, in addition to a much longer ventral projection of the gonarcus and no strong gonosetae on the gonosaccus. Etymology.—This species is named for the distinctively dark apical tarsal seg- ments. Material Examined.—In addition to holotype and allotype, one additional female from INBIO: same data as holotype, except collected Jul 1992. CERAEOCHRYSA INBIO PENNY, NEW SPECIES Types.—Holotype, male: from COSTA RICA. CARTAGO: La Amistad Bio- sphere Reserve, Guayabo National Monument, UTM grid coordinates L-N 217400, 570000, 1100 m, Jul 1994, G. Fonseca. Bar code: INBIO, CR1001, 887909. Holotype deposited: Instituto de Biodiversidad (INBIO), Santo Domingo de Heredia, Costa Rica. Description.— Head (Fig. 10). Vertex, frons, clypeus, genae, labrum, and palpi bright yellow. An- tennal scape and pedicel bright yellow, with scape bearing longitudinal, mid-dorsal brown stripe. An- tennal flagella missing. Thorax (Fig. 11). Pronotum wider than long; pale green with dark brown spots in anterio-lateral and posterio-lateral corners of sclerite. Meso- and metanota pale yellow, immacu- late, except for a pair of small, elongate, brown spots on mesonotum along suture between pre- scutum and scutum. Pleural and sternal areas pale green, immaculate. Legs pale green, becoming gradu- > Figures 1-8. Ceraeochrysa nigripedis, NEW SPECIES. Figure 1. Right wings. Figure 2. Head and thorax, dorsal view. Figure 3. Head, frontal view. Figure 4. Female genitalia, ventral view. Figure 5. Apex of male abdomen, lateral view. Figure 6. Gonapsis, dorsal view. Figure 7. Male genitalia, lateral view. Figure 8. Male genitalia, dorsal view. Abbreviations: arc = arcessus, cl = clypeus, epr = tergite 9 + ectoproct, fr = frons, gps = gonapsis, gs = gonarcus, ig = inner gradate veins, lbr = labrum, msn = mesonotum, mtn = metanotum, og = outer gradate veins, prn = pronotum, Pscu = pseudocubitus, Psm = Pseudomedia, Rs = radial sector, sg = subgenitale, sp = spermatheca, vtx = vertex, 1A = third anal vein, 2A = second anal vein, 3A = third anal vein, 8T = tergite 8. 1997 PENNY: NEW COSTA RICAN LACEWINGS 63 64 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(2) ally dark yellow on tarsi. Wings (Fig. 9). Forewing length—17.3 mm. Longitudinal veins pale green with dark brown crossveins, except apical costal and radial crossveins pale. Gradate crossveins dark, with some membrane infuscation along inner gradates; nine inner and ten outer gradate veins. Costa slightly swollen and darkened basad of apex of 3A. Hindwing length—15.0 mm. Longitudinal and crossveins completely pale green, immaculate. Eight inner gradate and nine outer gradate veins. Ab- domen. Tergites dark green, with pair of small dark spots at posterior margin of tergites 2 and 3. Sternites pale yellow. Male terminalia: ectoproct + tergite 9 elongate, without a ventral lobe (Fig. 12); dorsal apodeme forked, with straight, posterodorsally branch apically forked around callo cerci; ventral fork posteroventrally directed, elongate, straight, heavily-sclerotized projection terminating in a small, acute, ventral point. Sternite 9 apically tapering sharply to ventral point; bearing numerous setae with expanded (stalked) bases. Gonarcus (Figs. 14-15) narrow medially, tapering to relatively elongate, narrow, lateral plates. Entoprocesses elongate, evenly tapering points extending about 0.67 length of arcessus. Arcessus a broad, flat plate, terminating in a short, decurved median hook, and a pair of smaller bilaterally symmetrical lateral points. Ventral arms of gonarcus embedded in and sup- porting lateral margins of gonosaccus. Gonosaccus lacking gonosetae and gonocristae; membrane be- tween gonosaccus and sternite 9 bearing a broad field of small gonocristae, giving membrane a rugose appearance. Gonapsis elongate, upturned for posterior fifth, terminating in unexpanded, smoothly rounded apex (Fig. 13). Diagnosis.—Ceraeochrysa inbio is a part of the cincta species group, all of which have a caudally recurved ventral fork of the dorsal apodeme and a field of gonocristae between the gonosaccus and ninth sternite in males. Ceraeochrysa inbio differs from C. claveri (Navas) in the latter’s distinctive gonapsis, which terminates in a broad, U-shaped bifurcation, and in the lack of pronounced ento- processes. Ceraeochrysa cincta (Schneider) also has much shorter entoprocesses and a shorter, more highly curved dorsal apodeme. Ceraeochrysa arioles (Banks) has a much larger, more recurved apex of the ninth sternite. Ceraeochrysa inbio is very similar to C. caligata (Banks), except for the former’s uniquely spotted pronotum and small apical point on the male ninth sternite. Etymology.—This species is named after the Instituto de Biodiversidad, com- monly referred to as “INBIO”,, to recognize the hard work and pioneering meth- ods that its staff has instigated to increase knowledge about the biodiversity of Costa Rica. Material Examined.—See Type. CERAEOCHRYSA COSTARICENSIS PENNY, NEW SPECIES Type.—Holotype, male: COSTA RICA. PUNTARENAS: Los Alturas, 1360 m, 27 Feb 1991, Helen Sparrow. Holotype deposited: Instituto de Biodiversidad (INBIO), Santo Domingo de Heredia, Costa Rica. Description—Head (Fig. 18). Vertex, frons, clypeus, genae, labrum, and palps bright yellow. An- tenna slightly shorter than forewing length. Antennal scape suffuse orange dorsally, pale yellow ven- trally; pedicel and flagellum pale yellow. Thorax (Fig. 17). Pronotum wider than long, pale green with pair of large reddish brown spots laterally. Smaller red brown spot on membrane under anterio- lateral margins on either side of pronotum. Meso- and metanota pale green, immaculate. Pleural and sternal areas pale green to pale yellow. Legs entirely pale green. Wings (Fig. 16). Forewing length— 13.5 mm. Longitudinal veins pale green; crossveins dark, except for apical costal crossveins. Four —= Figures 9-15. Ceraeochrysa inbio, NEW SPECIES. Figure 9. Right wings. Figure 10. Head, frontal view. Figure 11. Head and thorax, dorsal view. Figure 12. Apex of male abdomen, lateral view. Figure 13. Gonapsis, dorsal view. Figure 14. Male genitalia, lateral view. Figure 15. Male genitalia, dorsal view. 1997 PENNY: NEW COSTA RICAN LACEWINGS 65 a Uiifiideg iYflitastill LO EE MT << PS ES IL PRE Rae em ees T 66 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(2) inner and seven outer gradate veins in parallel series forming cells approximately twice as long as wide. Hindwing length—12.0 mm. Longitudinal and crossveins uniformly pale green. Three inner and seven outer gradate veins. Apex angulate. Abdomen. Pale green. Male ectoproct + tergite 9 (Fig. 19) elongate, with scattered chalazate setae; an acute posteroventral lobe supported by ventral lobe of dorsal apodeme. Sternite 9 tapering to apex bearing chalazate setae. Gonarcus (Figs. 21-22) narrow medially, and narrow, caudally-projecting flat plates laterally. Broad dorsal plate at medial part of - gonarcus. Mediuncus laterally forming pair of elongate, upturned plates, which end in dorso-caudal acute point. Mediuncus with pair of small projections basally and an evenly decurved medial point apico-ventrally. Convoluted field of small gonocristae on gonosaccus. Gonapsis (Fig. 20) relatively short, with subapical lateral arms and acute, upturned apical point. Diagnosis.—This species is very closely related to C. everes (Banks), with which it shares a very distinctive dorsal hood medially on the gonarcus, upturned plate of the arcessus, and thickened setal bases at the apex of the ninth sternite. They differ in that C. costaricensis has two pairs of pronotal spots (not stripes), antennal bases pale (not infused with red), and apex of gonapsis simply upturned (not toothed dorsally with a ventral medial lobe). Etymology.—This species is named after Costa Rica, in homage to all of the efforts that have been made to make known and conserve its fauna. Material Examined.—See Type. CERAEOCHRYSA TAUBERAE PENNY, NEW SPECIES Type.—Holotype, male: COSTA RICA. CARTAGO: near Eslabén, 21 Dec 1994, C.A., M.J. and PJ. Tauber. Holotype deposited: Instituto de Buiodiversidad (INBIO), Santo Domingo de Heredia, Costa Rica. Description—Head (Fig. 25). Vertex, frons, clypeus, genae, labrum, and palps bright yellow. An- tennal scape and pedicel bright yellow. Antennal flagella missing. Thorax (Fig. 24). Pronotum as wide as long, pale yellow, with faint indications of darker markings at anterio-lateral and posterio-lateral margins. Meso- and metanota entirely pale green. Pleural and ventral surfaces of pterothorax pale yellow, immaculate. Legs pale yellow, changing to pale brown on tarsi. Wings (Fig. 23). Forewing length—10.0 mm. Longitudinal veins pale green, crossveins and apical twiggings brown. Three inner and four outer gradate veins present, dark brown with extensive infuscation of membrane adjacent to veins. Two additional areas of forewing heavily infuscated: apex of CuP and adjacent crossveins, and 2A-3A crossvein. Hindwing length—8.8 mm. Apex angulate. All longitudinal and crossveins entirely pale green. Two inner and four outer gradate veins. Abdomen. Entirely green. Male ectoproct + ter- gite 9 (Fig. 26) short, broadly ovate, only slightly extended ventrally, bearing scattered chalazate se- tae. Dorsal apodeme apically forked around callo cerci, but without ventral projecting lobe. Sternite 9 elongate, notably narrowed at mid-length, bearing scattered stalked setae. Gonarcus (Figs. 28, 29) formed by two very broad lateral plates which almost fuse along midline. Entoprocesses poorly de- veloped as short, pointed lobes caudally and tiny points anteriorly. Ventral lobes of gonarcus absent. Mediuncus elongate, straight, tapering to apical decurved point. Gonosaccus bearing numerous gono- setae. Gonapsis (Fig. 27) elongate, apically forked, with both forks apically rounded. Diagnosis.—This species appears most closely related to C. nigripedis, in part due to the long, straight arcessus and shape of the male ectoproct and ninth ster- nite. There is also a similar area visual attraction on the forewing at the apex of —_ Figures 16-22. Ceraeochrysa costaricensis, NEW SPECIES. Figure 16. Right wings. Figure 17. Head and thorax, dorsal view. Figure 18. Head, frontal view. Figure 19. Apex of male abdomen, lat- eral view. Figure 20. Gonapsis, dorsal view. Figure 21. Male genitalia, lateral view. Figure 22. Male genitalia, caudal view. 1997 PENNY: NEW COSTA RICAN LACEWINGS 67 Lf ie Z PEELE = 19 - ALL PR a ' an 68 THE PAN-PACIFIC ENTOMOLOGIST y Mi tllbi Z c IE WA 823 woe Vol. 73(2) 1997 PENNY: NEW COSTA RICAN LACEWINGS 69 CuP. Interestingly, there is a second illusory maculate area; Ceraeochrysa nigripe- dis has a pair of dark spots laterally on the metanotum, while in C. tauberae the forewing 2A—3A crossvein spot reposes over much the same metanotal area when the wings are at rest—forming the illusion of a metanotal spot in this species as well. These species also differ in that C. tauberae has a very wide gonarcal arch (not narrow), and numerous gonosetae on the gonosaccus (not smooth membrane). Etymology.—This species is named after Catherine A. Tauber, who is a noted authority on chrysopid biology and the taxonomy of their immature stages. Material Examined.—See Type. ACKNOWLEDGMENT The author thanks Connie Yan, Victoria Saxe and Diane Wong for drawings of the head, thorax and wings of the four species. Kady and Maurice Tauber gra- ciously donated the type specimen of C. tauberae to INBIO. Manuel Zumbado and the staff of INBIO provided many of the specimens studied, and Angel Solis provided access and equipment during visits to INBIO. I wish to thank George Gorman for logistical support within Costa Rica, as well as Helen Sparrow and Tom Sisk for providing assistance and logistical support at the Los Alturas Field Station during a collecting trip in 1991. Phillip A. Adams generously provided illustrations and background information about the genus Ceraeochrysa. The former chairman of the California Academy of Science’s Entomology Department, Charles Griswold, provided funding and encouragement for the ongoing study of Costa Rican neuropterans, as did the California Academy of Sciences In-House Research Fund. LITERATURE CITED Adams, P. A. 1982. Ceraeochrysa, a new genus of Chrysopinae (Neuroptera) (Studies in New World Chrysopidae, Part II). Neuroptera International, 2: 69-75. 12 figures. Adams, P. A. & N. D. Penny. 1987. Neuroptera of the Amazon Basin. Part lla. Introduction and Chrysopini. Acta Amazénica, 15(1985): 413-479. 213 + 29 figures. 1 table. Brooks, S. J. & P. C. Bamard. 1990. The green lacewings of the world: a generic review (Neurop- tera: Chrysopidae). Bulletin of the British Museum of Natural History, Entomology, 59: 117— 286. 578 figures. 1 table. Received 26 Mar 1996; Accepted 4 Nov 1996. > ‘4 Figure 1. Head of Wesmaelia lizanoi, anterior view, antennae removed. Figure 2. Posterior half of mesosoma and metasoma of Wesmaelia lizanoi, lateral view, right wing removed. 106 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(2) Figure 3. Wings of Wesmaelia lizanoi. by trace rugulose sculpture; face 1.9 wider than tall; clypeus with lower margin rounded; malar space very narrow, 0.9 basal width of mandible; mandibles long and slender, nearly completely over- lapping when closed. Mesosoma 1.4 longer than tall in lateral view; pronotum and prosternum mostly areolate-rugose; mesonotum and scutellar disc mostly smooth, except notauli foveate; mesopleuron mostly smooth and devoid of setae on medial disc, except antero-ventral sternaulus foveate; pro- podeum deeply excavated medially; propodeal sculpture finely areolate, except smooth area dorsad metasomal insertion delimited dorsally by a transverse carina; hind tibia with apical fringe of flat setae along inner margin; wings as in Figs. 2—3; pterostigma 2.7 longer than tall; vein Rs sharply curved, meeting wing margin well before apex; length of marginal cell along wing margin 0.75 pterostigma length. Metasoma as in Fig. 2, smooth, highly polished, and mostly devoid of setae ex- cept along posterior margins of posterior segments; petiolate first metasomal segment exceptionally long and slender, 8.4 longer than midpoint width in lateral view, longer than entire mesosoma, and strongly curved in lateral view (Fig. 2); metasoma beyond petiole with 6 visible segments, 2.7 < longer than tall in lateral view, terga weakly sclerotized and somewhat indented posteromedially; hypopyg- ium truncated, finely carinate medially; ovipositor sheath 4.4 longer than tall in lateral view; needle- like tip of ovipositor just visible beyond sheath apex. Color: Ocellar triangle, vertex posteriorly, me- sosoma, petiolate first metasomal segment, metasomal terga 2 + 3, and ovipositor sheath dark brown to black; eyes silver; antenna, remainder of head, legs, and remainder of metasoma mostly light yellow- brown; membranous parts of metasoma white; wing membrane transparent; pterostigma and wing ve- nation pale brown. Diagnosis.—Wesmaelia lizanoi can be distinguished from related species by its unique combination of short antennae with 17—18 flagellomeres in the females, 21-22 flagellomeres in the males, eyes barely converging ventrally (Fig. 1), the exceptionally long and slender first metasomal segment (Fig. 2), and extensive black coloration on the top of the head, the entire mesosoma, and much of the metasoma. Variation.—All variation noted is based on study of the paratypes. Females with body length 3.3-4.1 mm; forewing length 2.3—2.8 mm; antenna with 17-18 flag- ellomeres; light-colored parts of body varying from yellow-brown to pale brown- white or white; metasoma posteriorly and ventrally infused with varying amounts of dark brown pigment. Color of the face is more pure white in specimens from Vara Blanca, as compared with those from other sites. There is substantial varia- tion in the apparent size of the metasoma beyond the petiolate first segment (vary- ing from 2.7-4.1X longer than tall in lateral view), depending on the degree to which segments are telescoped outwards. The great length of the first metasomal segment, along with the telescoping capability of the remaining segments, allow the metasoma to swing ventrally and anteriorly, presumably allowing oviposition 1997 SHAW: COSTA RICAN WESMAELIA TS ek LEED OIE cer ar A LE = AGLrish Figure 4. Lateral habitus of Wesmaelia lizanoi female, with metasoma advanced in ovipositional posture. in front of the body of the wasp (Fig. 4). Several specimens died with the meta- soma extended in this forward position. Males with body length 2.9—3.7 mm; forewing length 2.4—3.0 mm; antenna with 21-22 flagellomeres; flagellum and metasoma much darker than in female, mostly dark brown to black; propodeal sculpture entirely finely areolate, lacking a smooth area above the metasomal insertion. Distribution.—Known only from three Costa Rican sites in Heredia, San Jose, and Puntarenas Provinces, at elevations ranging from 1200 to 2100 meters. The site at Zurqui de Moravia is located on the main highway east from San Jose to Limon, just before the entrance to Braulio Carillo National Park, behind the La Fonda restaurant. The Zurqui Malaise trap has been situated for several years just over the crest of a grassy hill on the edge of (and overlooking) a moist primary cloud forest, rich in epiphytes. The site at Vara Blanca is located at Finca Geor- gina, a farm belonging to the president of the Dos Pinos dairy company. The Finca Georgina Malaise trap was situated on the edge of a strip of primary forest re- 107 108 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(2) maining in a ravine. The site at San Vito is located at the Jardin Botanica Las Cruces, also known as the Wilson Botanic Garden. Biology.—The hosts of W. lizanoi are unknown, but if its habits are similar to W. pendula, then this new species also parasitizes late instar nymphs and adults of nabid bugs. Adults of W. lizanoi have been collected during almost every month of the year, indicating that this species is probably multivoltine. Etymology.—This species is named in honor of Sr. Jorge Arturo Lizano, owner of the La Fonda restaurant, behind which the “Zurqui de Moravia” Malaise trap was situated. Through his courtesy many new insects have become known to sci- ence. Material Examined.—See Types. ACKNOWLEDGMENT Special thanks to Paul Hanson for providing specimens and detailed informa- tion on the sites of the Malaise traps. Scanning electron microscopy for Figs. 1—2 was done by Paul Marsh. The habitus drawing (Fig. 4) was done by Ms. Amy Irish. This research was supported by grant DEB-930-0517 from the National Sci- ence Foundation. LITERATURE CITED Belokobylskij, S. A. 1992. Wesmaelia and Syrrhizus species (Hymenoptera, Braconidae, Euphori- nae) in the Far East. Vest. Zool., 1992: 8—16 (in Russian). Foerster, A. 1862. Synopsis der Familien und Gattuungen der Braconen. Verh. Naturh. Ver. Preuss. Rheinl. 19: 224-288. Hanson, P. E. & I. D. Gauld. 1995. The Hymenoptera of Costa Rica. Oxford University Press, Oxford. Harris, R. A. 1979. A glossary of surface sculpturing. Occas. Pap. Entomol. 28: 1-31. Huber, J. T. & M.J. Sharkey. 1993. Structure. pp. 13-59. Jn: Goulet, H. & J. T. Huber (eds.), Hy- menoptera of the World: An identificaiton guide to families. Centre for Land and Biological Resources Research, Ottawa, Ontario. Research Branch, Agriculture Canada, Publication 1894/E. Marsh, P. M. 1979. Family Braconidae. pp. 144-313. In: Krombein, K. V., P. D. Hurd Jr., D. R. Smith, & B. D. Burks (eds.), Catalog of Hymenoptera in America North of Mexico. Smithso- nian Institution Press, Washington, D.C. Marsh, P. M., S. R. Shaw & R. A. Wharton. 1987. An identification manual for the North American genera of the Family Braconidae (Hymenoptera). Mem. Entomol. Soc. Wash., 13: 1-98. Muesebeck, C. FW. 1963. Host relationships of the Euphorinae (Hymenoptera: Braconidae). Proc. Entomol. Soc. Wash., 65: 306. Papp, J. 1990. New braconid wasps (Hymenoptera: Braconidae) in the Hungarian Natural History Museum, 1. Ann. hist.-nat. Mus. natn. Hung., 82: 175-190. Papp, J., & L-y. Chou. 1995. The genus Wesmaelia Foerster of Taiwan (Hymenoptera: Braconidae: Euphorinae). Chin. J. Entomol., 15: 345-354. Sharkey, M. J. 1993. Family Braconidae. pp. 362-394. Jn: Goulet, H. and J. T. Huber (eds.). Hy- menoptera of the World: An identification guide to families. Centre for Land and Biological Resources Research, Ottawa, Ontario. Research Branch, Agriculture Canada, Publication 1894/E. Shaw, M. R. & T. Huddleston. 1991. Classification and biology of braconid wasps (Hymenoptera: Braconidae). Handbooks for the Identification of British Insects, Volume 7, Part 11. Royal En- tomological Society of London. Shaw, S. R. 1985. A phylogenetic study of the Subfamilies Meteorinae and Euphorinae (Hymenoptera: Braconidae). Entomography, 3: 277-370. Shaw, S. R. 1987. Orionis, anew genus from Central America, with an analysis of its phylogenetic placement in the Tribe Euphorini (Hymenoptera: Braconidae). Syst. Entomol., 12: 103-109. 1997 SHAW: COSTA RICAN WESMAELIA 109 Shaw, S. R. 1988. Euphorine phylogeny: the evolution of diversity in host-utilization by parasitoid wasps (Hymenoptera: Braconidae). Ecol. Entomol., 13: 323-335. Shaw, S. R. 1995. Braconidae. pp. 431-463. Jn: Hanson, P. E. and I D. Gauld (eds.). The Hy- menoptera of Costa Rica. Oxford University Press, Oxford. Received 23 May 1996; Accepted 4 Nov 1996 PAN-PACIFIC ENTOMOLOGIST 73(2): 110-121, (1997) POPULATION DENSITY AND DISPERSAL ABILITY IN DARWIN’S DARKLINGS: FLIGHTLESS BEETLES OF THE GALAPAGOS ISLANDS! T. L. FINsTon?, S. B. PECK, AND R. B. PERRY? Department of Biology, Carleton University Ottawa, Ontario K1S 5B6 Canada Abstract—This study is the first to combine both field and genetic data to examine population structure in flightless beetles from the Galapagos Islands. Field studies were conducted on four species of tenebrionid beetles belonging to three genera, Ammophorus Giierin-Méneville, Blap- stinus Latreille, and Stomion Waterhouse. The dynamics of the beetle community at the study site, Tortuga Bay, Santa Cruz Island, were analyzed in an attempt to examine patterns of activity and to quantify species abundances, population sizes, densities and levels of individual vagility. Beetle activity was found to vary with temperature, precipitation and number of sunlight hours. Although the number of recaptures was low, densities in the quadrats ranged from eight B. lugubris Boheman per hectare to 1238 S. laevigatum Waterhouse per hectare. Individual vagility is shown to be low among S. laevigatum, the most abundant species at the study site, as the dispersion index (DI) showed that captures were aggregated in three of the four quadrats, sug- gesting little movement. In addition, beetle captures occurred more frequently than expected in internal traps, again revealing limited movement into or out of the quadrats. These results were confirmed by a separate analysis of genetic differentiation among demes of S. laevigatum which showed the number of migrants to be less than one per generation. Key Words.—Insecta, Coleoptera, ecology, mark and recapture, population structure, tenebrionid beetles The Galapagos Islands have provided striking examples of species radiations under conditions of allopatry. The role of geographic isolation in island archi- pelagoes has long been recognized in limiting gene flow and promoting repro- ductive isolation (Mayr 1963). Geographic isolation may also be imposed by life- history parameters such as low dispersal ability or flightlessness. Isolation may also result from habitat restriction, whereby populations are spatially separated by regions of unsuitable habitat (King 1987, Crouau-Roy 1989), or by host plant specificity (McCauley & Eanes 1987). Genetic studies on flightless insects (Zera 1981, Finston & Peck 1995) and birds (Baker et al. 1995) have revealed high levels of genetic differentiation among populations and low genetic variability. The migration of individuals can be measured using both direct and indirect methods. Mark and recapture studies can be used to directly measure individual vagility and provide estimates of the number of migrants between demes. In particular, pitfall traps have commonly been used in population studies of surface dwelling insects, although their use is highly dependent upon a number of factors, including vagility of the organism under study, and ecological factors such as the influence of substrate type, vegetation and weather patterns on activity levels (Ahearn 1971, Thomas & Sleeper 1977). Alternately, population genetic structure 1 Authors page charges partially offset by a grant from the C. P. Alexander Fund, Pacific Coast Entomological Society. Current Address: Department of Zoology, University of Guelph, Guelph, Ontario, Canada N1G 2W1. 1997 FINSTON ET AL.: DISPERSAL ABILITY OF DARWIN’S DARKLINGS | I11 can be measured, and the amount of gene flow and migration inferred. Investi- gations into the role of limited vagility may provide an insight into patterns of microevolutionary change. The extent of isolation and gene flow among spatially separated populations determines the potential for phenotypic and genetic differ- entiation. Indeed, speciation is most active where geographic barriers are greatest- among insular habitats (Mayr 1942, 1966). Prior work on species radiations in the Galapagos Islands has focused on the vertebrate fauna (Van Denbergh 1914, Hendrickson 1966, Lack 1968, Grant 1986), however, the invertebrate fauna also provides striking examples of species radiations (Coppois 1984, Peck & Kukalova-Peck 1990). In particular, the Gala- pagos supports a rich assemblage of tenebrionid (darkling) beetles. Darkling bee- tles are often abundant members of surface dwelling communities in arid and semi-arid environments (Crawford 1990). Tenebrionid species occur sympatrically in arid regions all over the world (Thomas 1983), and most are flightless, night active members of these communities (Doyen & Tschinkel 1974). A total of 51 Species are known in 14 genera from the Galapagos Islands. Most notable are Darwin’s darklings, Ammo phorus Gterin-Méneville, Blapstinus Latreille, and Sto- mion Waterhouse, three genera containing 38 described species of flightless, ground-dwelling beetles (Peck & Kukalova-Peck 1990). Collectively, this group represents the largest radiation of beetle species in the archipelago. Four of these species, belonging to all three genera, were found sympatrically at a single site, and were the focus of this study. The present study represents the first attempt to quantify population sizes, densities, species abundances and beetle activity in a tenebrionid community in the Galapagos Islands, and provides the first exami- nation of dispersal ability in a Galapagos insect, employing both field and pop- ulation genetic data. MATERIALS AND METHODS The present study involved two field components: a mark and recapture study, and a weekly transect study. The field site was the dune area located at Tortuga Bay on the island of Santa Cruz, Galapagos. Beetles were identified using the key of Van Dyke (1953). Field work took place March to July, 1992. Mark and Recapture Study.—In order to assess species assemblages, population sizes and densities, and dispersal ability, four separate quadrats were arranged, each consisting of five rows of five traps. Each trap was 2 m from any other trap in a quadrat, and consisted of small, unbaited cylindrical plastic containers 10 cm in diameter and 6 cm deep. Each trap was equipped with a cover which was used to close the traps and a coarse wire screen (used when the traps were open) to prevent beetle predation by birds and lizards. Each quadrat covered 64 m?. The quadrats were arranged as in Fig. 1, such that each quadrat was situated in the corner of a 32 m X 102 m rectangle. The vegetation was not homogeneous over the entire study area; the quadrats differed both in plant species present and in the proportion of ground covered by vegetation. The plant composition in order of abundance for each quadrat is as follows, with the estimated percent of ground cover noted in brackets: I. Scaevola plumieri, Ipomoea pes-caprae, Sesuvium portulacastrum, Heliotropium sp., some grasses (80%); II. Scaevola plumieri, Ipomoea pes-caprae, Sesuvium portulacas- trum, Prosopis juliflora (90%); Il. Scaevola plumieri, Sesuvium portulacastrum, 112 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(2) 102 m Figure 1. Arrangement of traps and quadrats used in the mark and recapture study. Ipomoea pes-caprae, Heliotropium sp. (60%); IV. Heliotropium sp., Tiquilia dar- winil, Cryptocarpus pyriformis, Prosopis juliflora (15%). The traps were opened for a twelve hour period, from dusk to dawn, at five day intervals from 15 Jun to 8 Jul 1992. Individuals caught in the traps were recorded by quadrat and trap number, and marked with a dot of white typing correction fluid in the corner of the elytron which corresponded to the quadrat in which it was caught. Thus, recaptured individuals could be identified with respect to their original quadrat of capture. The beetle was then released at the site of capture. Estimates of population size (N) were calculated for those quadrats in which marked beetles were recaptured using the geometric model described by Thomas & Sleeper (1977): N = M/[I1 - (M/t)]; s? = M?’t(t - n)/r? where M = number of individuals marked; r = number of individuals recaptured, and t = total number of captures (= total number marked + recaptures). This model was chosen because it does not assume equal catchability rates among individuals, and minimizes stochastic events by permitting the use of cumulative results. Population density was calculated by dividing N by the unit area in hec- tares. The dispersion index (DI) was calculated for S. laevigatum Waterhouse only, the most abundant beetle in the quadrat area. The DI (variance/mean) approxi- mates unity when individuals are randomly distributed; a higher DI suggests ag- gregations of individuals (Fowler & Cohen 1990). A chi-square analysis was used to test the random distribution of individuals in each of the four quadrats. In addition, a chi-square test of the observed and expected number of captures (as- suming a uniform distribution) from peripheral and internal traps was performed as a further inspection of the dispersal habits of S. laevigatum. Transect Census.—As an independent assessment of community assemblages and beetle activity, a 100 m transect was marked along a one m wide foot trail which crossed the littoral and arid vegetation zones. The substrate changed from a fine white marine sand to volcanic rock approximately 20 m from the start of the transect. Sampling took place every five days and continued for eight weeks, 1997 FINSTON ET AL.: DISPERSAL ABILITY OF DARWIN’S DARKLINGS — 113 from 8 May to 7 Jul 1992. Baits of 5 gm of dry oatmeal were laid out at 2 m intervals at dusk, such that each bait site covered approximately 2 m *. The tran- sect was inspected one hour later, beginning at the site closest to the beach. The numbers of individuals of each species belonging to the three tenebrionid genera were recorded for each bait site. In order to examine the effects of local climatic conditions on species abundance and activity, weather data collected at the Charles Darwin Research Station was used for analysis. The research station lies approx- imately 2 km from the study site at Tortuga Bay, and is thus representative of the climatic conditions at the study site. Data included in this analysis were temper- ature and humidity at 18:00 h, total precipitation, number of hours of sunlight, and minimum and maximum temperatures for the day in which the count was made. In addition, the total precipitation for the seven day period preceding each sampling date was used. Pearson correlations and their corresponding probabilities were calculated between these ecological data, and weekly species richness (num- ber of different species found in the evening’s collection) and individual species abundances using the CORR module in SYSTAT (Wilkinson 1988). Genetic Analysis—Stomion laevigatum were collected from the following 5 sites on 4 different islands: Caamafio-sea lion beach (CAAM); Isabela-Alcedo Rim (ARIM), Tagus Cove (TAGU); Santa Cruz-Tortuga Bay (BTOR) Santiago- Playa Espumilla (SANT). Where possible, 44 individuals were analyzed for vari- ation at eight polymorphic enzyme loci using cellulose acetate electrophoresis (Hebert & Beaton 1993). The loci analyzed were as follows: hexokinase (Hk-2), mannose-6-phosphate isomerase (Mpi), phosphoglucomutase (Pgm-/), peptidase-A (Pep, utilizing phe-pro), supernatant and mitochondrial glutamate ox- aloacetate transamerase (Got-s, Got-m), 6-phosphogluconate dehydrogenase (6pgdh), and phosphoglucose isomerase (Pgi). Genetic analyses were carried out using BIOSYS-1 (Swofford & Selander 1991) unless otherwise indicated. Allele frequencies were calculated for each site, and Wright’s F-statistics were analyzed hierarchically such that gene frequency divergence could be measured among demes on the same island as well as among demes on different islands. (Wright 1940). Two models of gene flow were employed to obtain estimates of the number of migrants between demes. Wright’s F-statistics were used to calculate the num- ber of migrants using the following formula: Foy = 1/(4N,, + 1) where N,, = the effective number of migrants per generation. Slatkin’s rare alleles model (Slatkin 1985, 1987) was used to directly estimate the number of migrants between demes using the frequency of unique alleles as indicators of gene flow: In [p(1)] = aln(N,,) + 5B, where p(1) is the average frequency of alleles found only in single populations and a and b are constants that depend on the population size. In order to use Slatkin’s constants which were based on a sample size of 25, N,, was multiplied by the ratio 25/n where n is the mean sample size of those populations possessing unique alleles (Slatkin 1985, 1987). RESULTS Mark and Recapture Study.—Four species, Ammophorus bifoveatus Water- house, Blapstinus lugubris Boheman, B . spatulatus Van Dyke, and Stomion lae- 114 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(2) Table 1. The total number of individuals marked/recaptured for each species for all four quadrats. Week A. bifoveatus B. lugubris B. spatulatus S. laevigatum 1 7/0 0/0 22/0 36/0 2 20/0 0/0 30/1 42/0 3 5/0 0/0 28/1 44/0 4 2/0 0/0 19/1 25/0 5 3/1 1/0 10/0 24/0 Total 37/1 1/0 119/3 171/0 vigatum were found at the study site. Of the 328 beetles captured, marked, and released, only four were recaptured (Table 1). A single individual of A. bifoveatus, three individuals of B. spatulatus, and no individuals of S. laevigatum or B. lu- gubris were recaptured. All four individuals were recaptured in their original quadrat of capture. Population sizes obtained from the model presented by Thom- as & Sleeper (1977) are not reported here, as the estimates resulted in standard deviations which approached or exceeded the estimates themselves. Because of the poor recapture rate, population density was instead calculated each week by dividing the total number of individuals of each species in a quadrat by the unit area (64 m7’). The arithmetic mean over all four quadrats was then calculated over all weeks. Density estimates ranged from 8 B. lugubris per hectare to 1340 S. laevigatum per hectare (Table 2). The DI showed that S. laevigatum captures were not random in three of the four quadrats, as this ratio approximated unity only in quadrat I (Table 3). Some traps caught large numbers of beetles, while others caught no beetles over the course of the study. Tests of distribution models showed that aggregation of bee- tles occurred in quadrats II, III and IV. Quadrat I fitted neither a Poisson nor a binomial distribution, although its chi-square value (8.00) was very close to the critical value (7.81) for acceptance of the random (Poisson) distribution model. Furthermore, a chi-square test showed significant deviations between the number of observed and expected captures in both peripheral and internal traps. Beetle catches occurred more frequently than expected in the internal traps and less frequently in the peripheral traps in three of the four quadrats (Table 4). The number of captures per quadrat was not found to be significantly related to: the estimated amount of vegetation cover (Pearson correlation = 0.765, P = 0.235), but the greatest number of captures did come from quadrats I and II, Table 2. Weekly density estimates (per hectare) for four tenebrionid species averaged over each of four quadrats. Week S. laevigatum B. lugubris B. spatulatus A. bifoveatus 1: 1410 0 860 270 we 1640 0 1170 780 a 1220 0 1090 190 4 980 0 740 80 5 940 40 390 120 Mean 1340 8 850 288 (+SD) (294.5) (17.9) (309.8) (284.4) 1997 FINSTON ET AL.: DISPERSAL ABILITY OF DARWIN’S DARKLINGS © 115 Table 3. Mean number of catches per trap (x), standard deviation (SD), dispersion index (DI) and chi-square values (x?) for random (Poisson) and aggregated (binomial) distribution model tests for each of four quadrats in the mark and recapture study. * = P < 0.05. Poisson Binomial Quadrat ‘t SD DI x? (df) x? (df) I 1.80 1.83 1.02 8.00* (3) 8.93* (2) II 3.44 5.24 1.52 25.98* (9) 8.72 (8) Ill 0.60 0.92 1.53 6.32* (2) 2.04 (1) IV i Bs 1.69 1.51 14.10* (4) 7.67 (3) which were estimated to be 80% and 90% vegetated, respectively. The fewest captures came from quadrats III and IV, which were an estimated 60% and 15% vegetated, respectively. Stomion laevigatum was the most abundant species in all four quadrats. Ammophorus bifoveatus and B. spatulatus were particularly rare in the less vegetated quadrats. No A. bifoveatus were captured in quadrat III and S. laevigatum was more than four times more abundant than B. spatulatus. In quadrat IV, S. laevigatum was twenty-seven times more abundant than B. spatulatus and fourteen times more abundant than A. bifoveatus. Blapstinus lugubris was cap- tured only once, in quadrat II. Transect Census.—The four species found in the quadrat area, A. bifoveatus, B. lugubris, B. spatulatus, and S. laevigatum were also found along the transect. A total of 1199 beetles were encountered, with B. spatulatus and A. bifoveatus being most common (Table 5). In contrast to the mark and recapture site, S. laevigatum was the least abundant species encountered. Evening beetle counts ranged from two to 208 individuals, and the number of species observed ranged from one to four. Correlation of species abundance and richness with seven environmental con- ditions showed significant but negative correlations for several variables (Table 6). For example, the number of individuals of B. spatulatus and A. bifoveatus were negatively correlated with maximum daily temperature, temperature at 18:00 h, the days’ precipitation, hours of sunlight during the day, and the total precipitation for the week preceding the sampling date. In contrast, the number of different species observed in a sample was shown to be positively correlated with percent relative humidity. The days’ minimum temperature showed no effect on species abundance or diversity. The numbers of S. laevigatum and B. lugubris were not correlated with any measured environmental conditions. Table 4. Chi-square test of number of expected versus number of observed captures in internal and peripheral traps for four quadrats. * = P < 0.05. df = 1. Internal Peripheral Quadrat ~ Obs. Exp | Obs. Exp. x? I PS 16.2 0.632 32 28.8 0.356 II 43 31.2 5.38 4] 53.8 3.03 Il a 5.4 0.474 8 9.6 0.267 IV 12 10.1 0.366 16 17.9 0.206 116 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(2) Table 5. The number of individuals of each of four species of tenebrionids counted at the Tortuga Bay transect site on Santa Cruz Island, Galapagos. Week S. laevigatum B. lugubris B. spatulatus A. bifoveatus 1 i, 0 0 0 2 9 4 37 37 3 4 44 58 96 4 1 0 53 74 5 5 19 65 73 6 1 14 80 112 ei 1 8 89 110 8 2 19 69 113 Totals bs 108 451 615 Genetic Analysis.—Although there were no allelic substitutions among sites at any of the eight loci surveyed for S. laevigatum, allele frequencies did differ among sites (Table 7). Genetic differentiation was substantial among demes at both hierarchical levels, where on average, nearly 40% of the total observed variation in allele frequencies was due to variation among demes, whether on the same island or on different islands (Table 8). Approximately 60% of the variation resulted from differences within demes (1-F,;). Both models of gene flow pro- duced migration estimates of considerably less than one migrant per generation among demes (Table 8). Wright’s model produced an estimate of 0.429 migrants per generation among demes on the same island and 0.383 migrants per generation among demes on different islands. Similarly, Slatkin’s model produced an estimate of 0.417 migrants per generation among demes on different islands. DISCUSSION The utility of pit-fall traps in mark and recapture studies for the estimation of population sizes has fallen under scrutiny (e.g. Southwood 1966, Thomas & Sleeper 1977). Many non-random effects such as changes in activity patterns or abundance of populations, and biases introduced by the trapping methodology, may affect catchability and population size estimates. However, these effects can be overcome when the proper precautionary measures are observed, such as main- Table 6. Pearson correlation coefficients for individual species’ abundances and species richness and seven environmental variables. T = temperature at 18:00, PP = daily precipitation, H = relative humidity at 18:00, SOL = number of sunlight hours, TMAX = maximum daily temperature, TMIN = minimum daily temperature, TPP = total precipitation in the week preceding the collection, TI = total number of individuals in the collection, TS = total number of species in the collection. * = P < 0.05. Factor T PP H SOL TMAX TMIN TPP Taxon S. laevigatum 0.38 —0.16 0.39 0.38 0.48 0.25 —0.07 B. lugubris —0.23 —0.38 0.07 —0.35 —0.35 0.11 —0.36 B. spatulatus —0.93* —0.82* 0.52 —0.73* —0.96* —0.68 —0.86* A. bifoveatus —0.87* —0.77* 0.37 —0.76* —0.96* —0.52 —0.81* TI —0.85* —0.80* 0.42 —0.74* —0.94* —0.51 —0.84* TS —0.70 —0.94* 0.71* —0.51 —0.70 —0.46 =().93* 1997 FINSTON ET AL.: DISPERSAL ABILITY OF DARWIN’S DARKLINGS — 117 Table 7. Allele frequencies for five populations of S. laevigatum. n = sample size. Pop. SANT BTOR CAAM TAGU ARIM n 44 44 42 43 44 HK-2 3 0.989 0.977 1.000 1.000 1.000 4 0.011 = —4 =a an 5 ee 0.023 = = oa MPI 2 0.034 = = 0.012 = 3 0.375 = 0.012 sd 0.080 4 0.580 0.372 0.666 0.928 0.920 5 0.011 0.581 0.012 0.060 as 6 ea 0.047 0.310 cs = PGM-1 1 0.011 0.011 = = = 2 0.011 0.034 a 0.058 sa 3 0.967 0.921 0.857 0.907 0.989 4 0.011 0.034 0.143 0.035 0.011 PEP 2 zt 0.284 0.119 0.081 0.080 3 0.989 0.693 0.881 0.919 0.920 4 0.011 = £5 = = 5 = 0.023 = ze = GOT-M a) 0.625 = 0.488 0.907 0.909 3 0.375 0.405 0.512 0.093 0.091 4 2s, 0.595 = = = GOT-S 1 0.012 a = = _ 2 0.035 = me = a 3 0.720 0.667 a = = 4 0.233 0.333 0.561 ies 0.966 5 = = 0.439 1.000 0.034 6PGDH 2 = 0.035 0.012 0.012 = 3 1.000 0.930 0.988 0.163 0.068 4 = 0.035 a2, 0.826 0.898 5 2 we 2 = 0.034 PGI 1 Zs 0.023 = = a 2 0.125 0.080 = 0.186 0.011 3 0.546 0.874 1.000 0.791 0.989 4 0.295 0.023 zs = =e 5 0.034 5 = 0.023 ais taining uniform trap size and distance between traps, as well as using unbaited traps (Thomas & Sleeper 1977). Moreover, some surface-dwelling arthropods, such as tenebrionids, may be particularly amenable to pit-fall trapping techniques. This method was shown to be most effective in evaluating changes in population 118 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(2) Table 8. Estimates of the number of migrants per generation using two models of gene flow. Fy; = Wright’s F-statistic, N,, = number of migrants. Subgroup (S) Total group (T) For N,, (Wright) N,, (Slatkin) Site island 0.368 0.429 — Site total 0.395 0.383 0.417 behaviour, density, and dispersal, but least effective at obtaining estimates of population size (Aheam 1971). The number of recaptures in the present study was too few to allow a reliable estimation of population size for any of the species, therefore, distribution, population density and vagility of the beetles were all ex- amined in the quadrats using the total number of captures. Capture success may have been a function of both trapping methodology and patterns of beetles activity and abundance. For example, mortality rates may have been higher in marked beetles, that is, the correction fluid may have been toxic (although no dead, marked beetles were found in the study area), or they may have been more visible to predators. Alternately, the mark may have rubbed off during the course of the study as the beetle brushed against vegetation or sand while scavenging, although marked beetles held in captivity during the course of the study did not lose their markings (pers. obs.). Individual ranges may have been larger than the area covered by the four quadrats. Beetles, once marked, may have wandered out of the area. Finally, weekly patterns of beetle activity may also have been a contributing factor. Beetle activity on any given day may be affected by temperature, relative humidity, moon phase, recent rainfall, or amount of cloud cover (Thomas 1979). Indeed, the results of the transect study suggested that for at least two species, sand temperature and humidity may affect beetle activity and therefore recapture success. The highest number of evening sightings occurred following days with the lowest number of sunlight hours (less than five hours), perhaps as a result of lowered sand surface temperatures. In addition, the greatest species richness was noted on evenings with the greatest degree of daytime relative humidity, suggesting that cooler temperatures and some moisture favoured beetle activity. However, beetle activity was greatly reduced during heavy rains, such as on the first evening of collection. Density estimates made from the total capture data were generally lower but in accordance with estimates for tenebrionids from other mark and recapture stud- ies, ranging from 8—1340 beetles/hectare. Aheam (1971) found the average den- sity for five tenebrionid species to be about 1700 beetles/hectare at a site in South Mountain Desert Park in Phoenix, Arizona. Thomas & Sleeper (1977) found a range of 96—2755 beetles/hectare for six species in Rock Valley, Nevada. Simi- larly, Thomas (1979) found approximately 1000 beetles/hectare for the two most abundant species over a two year period in the Mojave Desert. The results of this study further suggested that Darwin’s darklings may have limited vagility. If individuals had large ranges, or good dispersal tendency, we would expect to capture some beetles in a quadrat different from that in which they were marked, or outside the quadrats. However, of the four beetles which were recaptured, each was found in the same quadrat in which it was marked. In addition, active searches in the area surrounding the quadrats produced no marked 1997 FINSTON ET AL.: DISPERSAL ABILITY OF DARWIN’S DARKLINGS _ 119 beetles. Because each beetle was recaptured in its original quadrat of capture, no direct estimates of displacement distances could be made. However, the DI for S. laevigatum showed that beetles tended to be aggregated in three of the four quad- rats, suggesting little movement, although the use of pheromones as attracting agents cannot be discounted in explaining the observed aggregations. In addition, an analysis of the number of beetles trapped in peripheral and internal traps showed that more beetles were trapped in the internal traps than the peripheral traps. If populations were vagile, we would expect to trap more beetles in the peripheral traps, as beetles move in from the surrounding area. Doyen & Tschinkel (1974) found vagility to be low among some tenebrionid species in their study in the Chiricahua Mountains; beetles were frequently recaptured in the same or a nearby quadrat as that in which they were marked, and taxa which showed the greatest levels of aggregation were those which were least vagile. The compara- tively low number of S. laevigatum at the near-by transect site also suggests that this taxon is limited in its range. These results were confirmed by an analysis of genetic differentiation among demes. Estimates of the number of migrants among demes using both Wright’s F-statistics and Slatkin’s rare alleles model for S. lae- vigatum showed dispersal to be very low, revealing less than one migrant per generation for both models (Finston & Peck 1995). The dispersal ability of individuals of a species is largely responsible for the establishment and maintenance of geographical isolates (Mayr 1966:565). Dar- win’s darklings show narrow distributions; each species is found on only one or a few geographically close islands (Van Dyke 1953). The flightless condition of the beetles limits their dispersal ability, particularly across water gaps. Peck (1994a,b) showed that various Coleoptera are occasionally present as aerial plank- ton or as pleuston between islands in the archipelago. Darwin’s darklings, how- ever, range from 5-10 mm and are larger than the small-bodied Coleoptera typ- ically found as aerial plankton. It is likely that only single or few founding in- dividuals, carried as sea surface pleuston, were responsible for the colonization of new islands. Although the Galapagos climate is semi-arid, it is seasonal, having both a wet and a dry season. Nevertheless, as with other, more stable tropical habitats, there may be little pressure for seasonal movement of beetles in search of favourable habitats. Furthermore, the highly stratified vegetation zones of the islands (Wiggins & Porter 1971) may pose geographical barriers for tenebrionid beetles, which are largely restricted to the littoral and arid zones (Van Dyke 1953, Finston 1993). This, and a previous study (Finston & Peck 1995) showed that dispersal ability and gene flow are limited even on a local scale; the number of migrants between populations was less than one per generation between popula- tions on the same island. The potential for few founders and limited subsequent movement of flightless beetles provides a model for the establishment and main- tenance of geographical isolation of populations. Such isolation is necessary for Microevolutionary change- localized differentiation through both adaptation and stochastic events such as founder effects and genetic drift- and ultimately, spe- ciation. ACKNOWLEDGMENT E Cepeda and A. Izurieta, Superintendents, Galapagos National Park (Depart- ment of Forestry, Ministry of Agriculture, Republic of Ecuador) issued scientific 120 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(2) research permits. Field logistical support and meteorological data were provided by the Charles Darwin Research Station, Santa Cruz Island, D. Evans, and C. Blanton, Directors. Field work was partially supported by a research grant to SBP from the Natural Sciences and Engineering Research Council of Canada and an OGS award to TLE Field sampling was aided by Sandra Abedrabbo, Joyce Cook, Moraima Inca, Bernard Landry, Ricardo Palma, and Eduardo Villema. The manu- script was improved by comments from Margaret Beaton, Konjev Desender, Chris Wilson, and several anonymous reviewers. LITERATURE CITED Ahearn, G. A. 1971. Ecological factors affecting population sampling of desert tenebrionid beetles. Amer. Midl. Nature., 86: 385—406. Baker, A. J., C. H. Daugherty, R. Colbourne, & J. L. McLennan. 1995. 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Sc. thesis, Carleton University. Finston, T. L. & S. B. Peck. 1995. Population structure and gene flow in Stomion: a species swarm of flightless beetles of the Galapagos Islands. Heredity, 75: 390-397. Fowler, J. & L. Cohen. 1990. Practical Statistics for Field Biology. Open University Press, Milton Keynes, Philadelphia. Grant, P. R. 1986. Ecology and evolution of Darwin’s Finches. Princeton University Press, Princeton, NJ. Herbert, P D. N., & M. J. Beaton. 1993. Methodologies for Allozyme Analysis using Cellulose Acetate Electrophoresis, 2nd edn. Helena Laboratories, Beaumont, TX. Hendrickson, J. R. 1966. The Galapagos Tortoises, Geochelone Fitzinger 1835 (Testudo Linnaeus 1758 in part). In Bowman, R. I. (ed.), Proceedings of the Symposium of the Galapagos Inter- national Scientific Project. University of California Press, Berkeley and Los Angeles. King, P. S. 1987. Macro- and microgeographic structure of a spatially isolated beetle species in nature. Evolution, 41: 401-416. Lack, D. 1968. Darwin’s Finches- an essay on the general biological theory of evolution. Peter Smith Publishers, Glouster, Mass. Mayr, E. 1942. Systematics and the origin of species. Columbia University Press, New York. Mayr, E. 1963. Animal Species and Evolution. The Belknap Press of Harvard University Press. Cambridge, Mass. McCauley, D. E. & W. E Eanes. 1987. Hierarchical population structure analysis of the milkweed beetle, Tetraopes tetraophthalmus (Forster). Heredity, 58: 193-201. Peck, S. B. 1994a. Aerial dispersal of insects between and to islands in the Galapagos archipelago, Ecuador. Ann. Entomol. Soc. Amer., 87: 218—224. Peck, S. B. 1994b. Sea-surface (pleuston) transport of insects between islands in the Galapagos archipelago, Ecuador. Ann. Entomol. Soc. Amer., 87: 576-582. Peck, S. B. & J. Kukalova-Peck. 1990. Origin and biogeography of the beetles (Coleoptera) of the Galapagos Archipelago, Ecuador. Can. J. Zool., 68: 1617-1638. Slatkin, M. J. 1985. 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Amer., 70: 242-248. Van Denbergh, J. 1914. The gigantic land tortoises of the Galapagos archipelago. Proc. Calif. Acad. Sci., 1: 7-288. Van Dyke, E. C. 1953. The Coleoptera of the Galapagos Islands. Occas. Papers Calif. Acad. Sci., 22: 1-181. Wiggins, I. L. & D. M. Porter. 1971. Flora of the Galapagos Islands. Stanford University Press. Stanford, California. Wilkenson, L. 1988. SYSTAT: The System for Statistics. SYSTAT, Evanston, IL. Wright, S. 1940. Isolation by distance. Am. Nat., 74: 232-248. Zera, A. J. 1981. Genetic structure of two species of waterstriders (Gerridae: Hemiptera) with differing degrees of winglessness. Evolution, 35: 218-225. Received 30 Apr 1996; Accepted I Sep 1996. PAN-PACIFIC ENTOMOLOGIST 73(2): 122-126, (1997) A NEW SPECIES OF LITOPROSOPUS (LEPIDOPTERA: NOCTUIDAE) FROM BAJA CALIFORNIA, MEXICO JOHN W. Brown! AND DaAvip K. FAULKNER Entomology Department, San Diego Natural History Museum, P.O. Box 1390, San Diego, California 92112 Abstract.—Litoprosopus bajaensis Brown and Faulkner, NEW SPECIES, a taxon endemic to the northern portion of the Vizcaino Desert, Baja California, Mexico, is described and illus- trated. The description is based on six specimens (two males and four females) collected be- tween 31 March and 1 May (1975-1985). The combination of the long curved ampulla, short third segment of the labial palpus, and pale fawn-gray forewing color separate L. bajaensis from all other species of Litoprosopus. Key Words.—Insecta, Lepidoptera, Noctuidae, Litoprosopus bajaensis, Baja California. Although our knowledge of the lepidopterous fauna of the peninsula of Baja California, Mexico, has increased dramatically over the last ten years (e.g., Brown & Donahue 1989, Brown, Real & Faulkner 1992, Eichlin 1993), considerable por- tions of the fauna remain poorly studied. Numerous endemic taxa are undescribed, and published information is lacking for several major groups (e.g., microlepi- doptera, Arctiidae, Noctuidae). This paper describes a new species of Litoproso- pus (Noctuidae) from the northern portion of the Vizcaino Desert. The new spe- cies is geographically isolated and morphologically distinct from its congeners. Dissection methodology followed that presented in Clarke (1941). Terminology for structures of the genitalia follows Eichlin (1975). Depositories and Abbreviations.—Specimens of Litoprosopus were borrowed from the following collections: Natural History Museum of Los Angeles County (LACM); University of California, Berkeley (UCB); and San Diego Natural His- tory Museum (SDNHM). Comparative material was borrowed from The Natural History Museum, London, England; National Museum of Natural History, Wash- ington, D.C.; American Museum of Natural History, New York, New York; and Camegie Museum of Natural History, Pittsburgh, Pennsylvania. Litoprosopus bajaensis Brown & Faulkner, NEW SPECIES (Figs. 1-3) Type Material.—Holotype: male: MEXICO: BAJA CALIFORNIA: 1.6 km (1 mi) E of Santa Inez, 6 Apr 1985, D. K. Faulkner, UV light (SDNHM). Five para- types as follows: MEXICO: BAJA CALIFORNIA: 5 km N of Catavina, 2 fe- males, 1 May 1975, E. M. Fisher (LACM); 16 km (10 mi) SE of El Rosario, 1 male, 31 Mar 1976, black light trap (UCB); 1.6 km (1 mi) E of Santa Inez, 2 females, 6 Apr 1985, D. K. Faulkner (SDNHM). Description.—Adult Male. Head: Frons sparsely scaled below mid-eye; vertex long-scaled, pale fawn-gray. Ocelli small. Labial palpus strongly upturned, pale fawn-gray; segment III short and blunt, approximately 0.25 times length of segment II. Thorax: Smooth-scaled, densely clothed in fawn-gray 1 Current Address: USDA, Systematic Entomology Laboratory, c/o National Museum of Natural His- tory, Washington, D.C. 20560 1997 BROWN & FAULKNER: NEW BAJA CALIFORNIA LITOPROSOPUS 123 Figure 1. Adult female of Litoprosopus bajaensis. scales. Forewing: Length 20-22 mm (x = 21; n = 2) (Fig. 1). Uniform pale fawn-gray above, with few small brown specks in basal half of discal cell. Underside glossy pale tan. Hindwing: White above, with fawn-brown border and faint mottling; “eye-spot” at margin comprised of a pair of small, dark brown, subrectangular spots between veins 1A + 2A — CuP and CuP — CuA,. Underside glossy white. Genitalia: As in Fig. 2 (drawn from JWB slide no. 136, Baja California, Mexico; n = 1). Uncus long, slender, nearly uniform in width, but attenuate distally. Tegumen narrow. Juxta trapezoi- dal. Valva long, broadest distally, narrowest subbasally. Sacculus with broadly hook-shaped base, sub- tended distally by long, narrow digitate process; broad basal portion of sacculus attenuate at approxi- mately 0.4 distance from base to apex; long, narrow, curved ampulla from sacculus approximately 0.5 distance from base to apex, arising from narrow, strongly sclerotized portion of sacculus; sacculus ending near apex of valva with several small warts bearing setae. Aedeagus moderately long, broad; two large patches of scobination in basal portion of vesica. Adult Female.—Essentially as described for male. Forewing length 19-24 mm (x = 21; n = 4). Genitalia: As in Fig. 3 (drawn from JWB slide no. 137, Baja California, Mexico; n = 2). Papillae anales simple, unmodified. Sterigma a sclerotized band; ostium bursae represented by a strong cup- shaped process. Ductus bursae with dense patches of scobination posteriorly, lines of sclerotization anteriorly. Corpus bursae long, moderately narrow, unsclerotized, with large region caudal to attach- ment of ductus bursae (“apex” sensu Eichlin 1975). Ductus seminalis from posterior portion of apex. Diagnosis.—Litoprosopus bajaensis can be distinguished superficially from all other species of Litoprosopus by its nearly uniform pale fawn-gray forewing color. Most species in the genus are characterized by varying shades of brown on the forewing, usually with some lighter or darker mottling. The closest species geo- graphically, L. coachella Hill, has a pale tan-ocherous forewing, with traces of two, narrow, brown, curved lateral bands. The two apparently are allopatric, with L. coachella restricted to southern California and L. bajaensis restricted to the Vizcaino Desert of Baja California, Mexico. The male genitalia of L. bajaensis can be distinguished from L. futilis (Grote & Robinson), L. confligens (Walker), and L. coachella by its longer and more strongly curved process from the ampulla, which is reminiscent of L. puncticosta Hampson and L. bahamensis Hampson, both from the Caribbean. Litoprosopus bajaensis can be distinguished from the latter two by its longer region of sub- basal constriction of the valva and the warty bases of setae at the distal end of the sacculus. 124 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(2) Figures 2-3. Genitalia of Litoprosopus bajaensis. Figure 2a. Male, valvae spread. Figure 2b. Aedeagus removed. Figure 3. Female. The third segment of the labial palpus is short (approximately 0.25 the length of the second segment) and blunt in L. bajaensis. This is similar to L. confligens but differs from all other species examined in which the third segment is 0.6—0.8 times as long as the second. Distribution and Life History—Litoprosopus bajaensis is known only from the northern portion of the Vizcaino Desert in Baja California, Mexico. Capture records extend from 10 km SE of El Rosario to just north of Catavina (Fig. 4); all specimens (n = 6) were collected between 31 March and 1 May (1975-1985). Since adults of L. coachella have been collected from April through October, it is likely that L. bajaensis has a considerably longer flight period than the available records suggest. Based on life history information of L. coachella (e.g., Com- stock 1933, 1956; Flock 1951), it is suspected that larvae of L. bajaensis feed on the flowers, buds, and possibly fruits of fan palms in the genus Washingtonia (Are- caceae), two species of which occur within its range—W. /filifera (Linden) Wendl. and W. robusta Wendl. (Wiggins 1980). Discussion.—Poole (1989) included seven species in Litoprosopus: bahamen- sis, coachella, confligens, futilis, haitiensis, hatuey (Poey), and punticosta. We have examined the types of all but L hatuey; it is likely that all are distinct spe- cies. The type of hatuey may be deposited in a collection in Cuba, but we were unable to locate it. This is particularly troubling because it is the type species of the genus. Species of Litoprosopus are distributed from the southern United States (i.e., Florida, Georgia, Texas, Arizona, California) south through Central America, with a few records from northern South America. Although most exhibit a compara- 1997 BROWN & FAULKNER: NEW BAJA CALIFORNIA LITOPROSOPUS 125 100 200 KILOMETERS Figure 4. Geographical distribution of Litoprosopus bajaensis. tively limited distribution, L. confligens ranges from southern Florida to South America. The phylogenetic position of the genus within the Noctuidae is unresolved, and species level problems may be present in the Caribbean region (including Florida). The paucity of material from this region inhibits our ability to discriminate be- tween intraspecific variation and interspecific differences. Material Examined.—See Types. ACKNOWLEDGMENT We thank the following for allowing us to examine specimens in their care: Robert W. Poole, United States National Museum of Natural History, Washing- ton, D.C.; John E. Rawlins, Carnegie Museum of Natural History, Pittsburgh, 126 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(2) Pennsylvania; Jerry A. Powell, Essig Museum of Entomology, University of Cali- fornia Berkeley; Julian P. Donahue, Natural History Museum of Los Angeles County, Los Angeles, California; James Miller, American Museum of Natural His- tory, New York, New York; and Ian Kitching, The Natural History Museum, Lon- don, England. We thank R. Poole and J. Rawlins for encouragement, assistance, and discussions. LITERATURE CITED Brown, J. W. & J. P. Donahue. 1989. The Sphingidae (Lepidoptera) of Baja California, Mexico. J. Lepid. Soc., 43: 184-209. Brown, J. W., H. G. Real & D. K. Faulkner. 1992. Butterflies of Baja California, Mexico. Lepidopt- era Research Foundation, Beverly Hills, California. Clarke, J. FG. 1941. The preparation of the genitalia of Lepidoptera. Bull. Brooklyn Entomol. Soc., 36: 149-161. Comstock, J. A. 1933. Studies in Pacific Coast Lepidoptera. Bull. South. Calif. Acad. Sci., 32: 113- 120. Comstock, J. A. 1956. Is this a new, and giant clothes moth? Bull. South. Calif. Acad. Sci., 55: 51-53. Eichlin, T. D. 1975. Guide to the adult and larval Plusiinae of California (Lepidoptera: Noctuidae). Occas. Pap. Entomol., Calif. Dept. Food & Agri. No. 21. Eichlin, T. D. 1993. Clearwing moths of Baja California. Trop. Lepid., 3: 135-150. Flock, R. A. 1951. Damage to household goods by the fan palm caterpillar. J. Econ. Entomol., 44: 260-261. Poole, R. 1989. Lepidopterorum Catalogus (new series), fascicle 118: Noctuidae. E. J. Brill/Flora and Fauna Publ., Leiden, The Netherlands. Three volumes, xii + 1341 pp. Wiggins, I. L. 1980. Flora of Baja California. Stanford University Press, Stanford. Received 22 Apr 1996; Accepted 16 Dec 1996 PAN-PACIFIC ENTOMOLOGIST 73(2): 127-134, (1997) THE NAUCORIDAE (HETEROPTERA) OF SOUTHERN THAILAND ROBERT W. SITES,! BECKy J. NICHOLS,! AND SURAKRAI PERMKAM? 'Wilbur R. Enns Entomology Museum, Department of Entomology, University of Missouri, Columbia, Missouri 65211; *Faculty of Natural Resources, Prince of Songkla University, Hat Yai, Thailand Abstract—In January 1995, the southernmost seven provinces of Thailand were surveyed for their naucorid fauna. Eight species representing five genera and four subfamilies were collected from waterfalls, streams, and ponds. An annotated list of taxa and illustrated taxonomic key are presented. uNndagea - OUUNSIAU weod IdtiinisdisoDUoUAzWWIN ludUN a dondamald noudwuovUszinalne wuvoudenard ¢ din dodnodlu ¢ ana uaz ¢ doddou law lAUMos woINUSIOMUthNaN 419615 Nazindotiadodu 4 fnosuiwsiwaxl5yaov lwWavlisiazviia tazsUde1u Tatitauelund Key Words.—Insecta, Naucoridae, Thailand, fauna, aquatic Tropical peninsular Thailand is topographically diverse with many mountain ranges and associated waterfalls and streams. These numerous aquatic systems as well as vegetated ponds harbor a diverse aquatic insect fauna. The composition of lotic insect communities in southern Thailand is shaped in part by natural biogeographic distributions as well as by disturbances from a variety of natural and anthropogenic origins. Scouring monsoons occur primarily from October through December; however, rainfall occurs throughout the year with an average of approximately 432 cm per year (Nuttonson 1963), and a recorded high of 660 cm (Pendleton 1962). Most streams of peninsular Thailand have their origins in the forested mountains. Although extensive deforestation occurred in the 1970’s and 1980’s, resulting in the removal of riparian vegetation, the government banned commercial logging in 1989, affording greater protection for these aquatic systems and associated organisms. In addition, use of streams for personal hygiene and for the disposal of acids during the commercial production of rubber contributes to the presence or absence of particular members of the aquatic insect community. Naucoridae constitutes a family of predacious aquatic Heteroptera which is known to inhabit a wide variety of lotic and lentic situations. This family is most speciose in both the New and Old World tropics, although representatives also occur in temperate regions. Naucorids are considered keystone consumers (Sites & Willig 1991); thus, they constitute an important component of the trophic web of aquatic systems, particularly of tropical streams (Sites in press). Riparian de- forestation has been reported to potentially have a substantial effect on populations of Naucoridae by resulting in increased densities (Polhemus & Polhemus 1988, Sites in press). Although most treatments of Naucoridae are of taxonomic focus, several faunal lists are available for regions of Southeast Asia, including Sri Lanka (Mendis & Fernando 1962), India (Tonapi 1959, and references therein), Indonesia (Nieser 128 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(2) & Chen 1991, 1992), peninsular Malaysia (Fernando & Cheng 1963), the Phil- ippine Islands (Usinger 1937), India, Sri Lanka, and Burma (Distant 1911), and Sumatra, Java, and Bali (Lundblad 1933). No naucorid species are given in the list of insects of southern Thailand (Chinajariyawong et al. 1986). Herein, we present an annotated list of naucorid species collected in southern Thailand and an illustrated taxonomic key. FIELD COLLECTIONS Thirty collections were made in the southernmost seven provinces of Thailand (Narathiwat, Pattani, Phattalung, Satun, Songkhla, Trang, Yala), including, in some cases, in national parks (with permission). Because of political instability, collecting was not conducted in extreme southeastern Narathiwat Province. This area is mountainous with waterfalls and streams, and holds promise as harboring Species not included herein. Collecting was performed with an aquatic D-net. In streams, the substrate was kick-sampled, allowing the current to carry organic debris, including insects, into the net. Along stream margins and in ponds, veg- etation was swept with the D-net. All insects were placed into 80% ethyl alcohol. It is common for more than one species of naucorid (even congeners) to be present in a particular body of water. For some species, both sexes are required for accurate identification. Further, other taxa such as the common lotic genus, Aphelocheirus, are polymorphic with respect to wing development. Therefore, when possible, series of specimens should be collected to sufficiently characterize the species and maximize the likelihood of obtaining accurate specific determi- nations. Voucher specimens have been deposited in the museum of the Department of Pest Management, Faculty of Natural Resources, Prince of Songkla University (PSU), Hat Yai, Thailand; and the Wilbur R. Enns Entomology Museum, Uni- versity of Missouri, Columbia, Missouri. Although we collected more species than expected from the region based on the literature, the possibility exists that addi- tional species may be found in southern Thailand. Therefore, this key may require modification in the future if additional species are discovered. KEY TO THE ADULTS OF NAUCORIDAE OF SOUTHERN THAILAND Identification of species of Aphelocheirus requires the distinction between male and female individuals. Males are asymmetrical in the terminal abdominal sterna, whereas females are symmetrical. la Labium reaching posteriorly at most to prothoracic coxae; antennae short, not reaching lateral margins of head ..................... 2. 1b Labium extending posteriorly to near mesothoracic coxae; antennae long, extending past lateral margins of head .................... 5 2a(la) Posterior margin of pronotum with lateral %4 widely separated from anterior margin of mesothorax and hemelytra; = 13 mm in length se Ne ce, hes Rie islet setae Gestroiella limnocoroides Montandon 2b Posterior margin of pronotum closely appressed to anterior margin of mesothorax and hemelytra; < 13 mm in length ................. 3 3a(2b) Prothoracic leg with pretarsus with single minute claw; protarsus one- SQ CMC tS sh ye seer ci, ss ORE ace aE Naucoris scutellaris Stal 1997 SITES ET AL.: NAUCORIDAE OF THAILAND 129 Figure 1. Distal end of metatibia of Heleocoris. Figure 2. Distal end of metatibia of Ctenipocoris asiaticus. Figure 3. Subgenital plate of female Aphelocheirus grik. Figure 4. Abdominal sternite V of male Aphelocheirus grik. Figure 5. Subgenital plate of female Aphelocheirus femoratus. Figure 6. Subgenital plate of female Aphelocheirus malayanus. 3b Prothoracic leg with pretarsus with two conspicuous claws; protarsus two-segmented (one-segmented in Heleocoris females) .......... 4 4a(3b) Metatibia ventrally with subapical, stout, parallel spines arranged in two or more rows (Fig. 1); fringe of natatorial hairs on mesal surface of metatibia dense, without gaps between hairs ..... Heleocoris spp. 4b Metatibia with circlet of stout, divergent, subapical spines, not ar- ranged in rows (Fig. 2); fringe of hairs on mesal surface with gaps between haits =. 4.4 6.x 8 ct eye See Ctenipocoris asiaticus Montandon 5a(1b) Female with subgenital plate truncate (Fig. 3); male with tab projecting from posterior margin of abdominal sternite V (Fig. 4); never with raised, brown patch on hind trochanter and base of femur ......... Ree tA Ae. tn, Aan re Aphelocheirus grik Polhemus & Polhemus Sb Female with subgenital plate triangular (Figs. 5, 6); male with or with- out weakly developed tab on abdominal sternite V; if tab is present, then with raised, brown patch on hind trochanter and base of femur 6 130 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(2) 6a(5b) Male with hind trochanter and base of femur each bearing well-de- fined, raised, brown patch; female with posterolateral angle of ter- gite VII 90° (Fig. 5), subgenital plate without peg-like setae, al- though hair-like setae present, peg-like setae near posterior margin at midline of sterna IV-V and occasionally VI .................... et tsa cal. ee Aphelocheirus femoratus Polhemus & Polhemus 6b Male with hind trochanter and base of femur devoid of well-defined brown patch, although poorly-defined dark streak may occur over much of femur; female with posterolateral angle of tergite VII ca. 130°, 4-6 peg like setae near apex of subgenital plate (may be dif- ficult to see) (Fig. 6), peg-like setae also near posterior margin at inidlinie cof steria TV > VT get eke ra ayers ey ah edn tre Pa cn Bese opie ANNOTATED LIST OF TAXA SUBFAMILY APHELOCHEIRINAE GENUS APHELOCHEIRUS Three species of Aphelocheirus were collected from southern Thailand. Five additional species have been recorded from northern Thailand (Polhemus & Pol- hemus 1988) but have not been recorded from the peninsular part of the country. Species in this genus are represented by brachypterous and macropterous forms. The subfamily is represented by only the genus Aphelocheirus and is considered by some to represent a distinct family level taxon (Stys & Jansson 1988, and citations therein). Aphelocheirus femoratus Polhemus & Polhemus Aphelocheirus femoratus Polhemus & Polhemus 1988: Raffles Bull. Zool. 36: 214-216. Diagnosis.—The male is distinctive in having well-defined brown patches on the hind trochanters and bases of the hind femora. The seventh abdominal terga of the female have posterolateral angles of 90° and the subgenital plate is trian- gular. Brachypterous and macropterous forms are consistent in expression of the diagnostic characters. Discussion.—This species occurs in gravel and rocky substrates of streams. This is the most common Aphelocheirus species in southern Thailand, and has been recorded from peninsular Malaysia north to Chiang Mai Province in northern Thailand (Polhemus & Polhemus 1988). This species occurred syntopically with A. grik at Banglang National Park and 9 km N Than To, Heleocoris sp. nr Sri- sakhon, and Gestroiella limnocoroides at Ton Nga Chang. Material Examined ——NARATHIWAT: stream 14 km W of Srisakhon, 15 Jan 1995, L-77 (11 bra- chypterous males, 1 macropterous male, 4 brachypterous females, 7 macropterous females, 46 nymphs); stream below Bacho Waterfall, 15 Jan 1995, L-78 (5 brachypterous males, 4 brachypterous females, 57 nymphs). SONGKHLA: stream from Ton Plieuw, 7 Jan 1995, L-62 (17 brachypterous males, 9 brachypterous females, 13 nymphs); same locality, 8 Jan 1995, L-64 (2 brachypterous males, 2 brachypterous females, 2 nymphs); Ton Nga Chang National Park, stream at Buddhist temple, 6 Jan 1995, L-59 (1 macropterous male, 7 nymphs); same locality, 7 Jan 1995, L-60 (14 brachypterous males, 14 brachypterous females, 13 nymphs); same locality, 8 Jan 1995, L-65 (15 brachypterous 1997 SITES ET AL.: NAUCORIDAE OF THAILAND 131 males, 10 brachypterous females, 33 nymphs); same locality, 30 Jan 1995, L-81 (13 brachypterous males, 11 brachypterous females, 6 nymphs); Ton Nga Chang National Park, waterfall levels 2 and 3, 6 Jan 1995, L-66 (1 macropterous male, 1 brachypterous female, 1 nymph). YALA: Banglang National Park, Than To, 14 Jan 1995, L-73 (7 brachypterous males, 4 brachypterous females); 9 km N of Than To, 15 Jan 1995, L-76 (1 macropterous male). Aphelocheirus grik Polhemus & Polhemus Aphelocheirus grik Polhemus & Polhemus 1988: Raffles Bull. Zool. 36: 218-220. Diagnosis.—The male has a tab extending from the posterior margin of abdom- inal sternite V and lacks the raised brown patches characteristic of A. femoratus. The female is easily recognized by the truncate subgenital plate. Brachypterous and macropterous forms are consistent in expression of these diagnostic charac- ters. Discussion.—This species occurs in gravel and rocky substrates of streams and has been recorded from peninsular Malaysia north to Chiang Mai Province in northern Thailand (Polhemus & Polhemus 1988). This species occurred syntopi- cally with A. femoratus at Banglang National Park and near Than To, and A. malayanus near Khao Ka Chong. Material Examined——TRANG: ca. 10 km E of Khao Ka Chong National Park on hwy 4, 12 Jan 1995, L-69 (1 macropterous male). YALA: Banglang National Park, Than To, 14 Jan 1995, L-73 (1 macropterous male, 1 brachypterous female, 4 macropterous females); 9 km N of Than To, 15 Jan 1995, L-76 (14 brachypterous males, 13 brachypterous females). Aphelocheirus malayanus Polhemus & Polhemus Aphelocheirus malayanus Polhemus & Polhemus 1988: Raffles Bull. Zool. 36: 216-218. Diagnosis.—This species is similar to A. femoratus although it is slightly larger. The male lacks the raised brown patches characteristic of A. femoratus and the tab on abdominal sternite V characteristic of A. grik. The female has a triangular subgenital plate with 4-6 obscure peg-like setae near the apex, and the postero- lateral angle of tergum VII is ca. 130°. Brachypterous and macropterous forms are consistent in expression of these diagnostic characters. Discussion.—This species occurs in the gravel and rocky substrates of streams and previously was known only from peninsular Malaysia. This is the first record of A. malayanus from Thailand and it occurred syntopically with A. grik. Material Examined ——TRANG: ca. 10 km E of Khao Ka Chong National Park on hwy 4, 12 Jan 1995, L-69 (4 brachypterous males, 6 brachypterous females, 4 macropterous females, 16 nymphs). SUBFAMILY CHEIROCHELINAE GENUS GESTROIELLA Gestroiella limnocoroides Montandon Gestroiella limnocoroides Montandon 1897: Ann. Mus. Civ. Storia Nat. Genova 17s 374372; Diagnosis.—This species is the largest known naucorid in southern Thailand. Length ranges from 13—17 mm, although specimens from other regions of South- east Asia are considerably larger. The posterior margin of the pronotum is widely separated from the anterior margin of the mesonotum and embolia of the hemel- 132 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(2) ytra. The lateral margins of the head and pronotum are markedly straight and convergent. Discussion.—This species occurs in the gravel and rocky substrate of streams. We also collected specimens from rock pools of the waterfall which were dis- continuous from the main body of water. This record represents the southernmost known extent of the range of G. limnocoroides and the nearest known population is at Chiang Mai in northern Thailand. This species occurred syntopically with A. femoratus at Ton Nga Chang. The specific status of this population is equivocal because this genus is in need of revision. Material Examined—SONGKHLA: Ton Nga Chang National Park, stream at Buddhist temple, 6 Jan 1995, L-59 (9 nymphs, 1 exuviae); same locality, 7 Jan 1995, L-60 (12 nymphs); same locality, 8 Jan 1995, L-65 (1 male, 10 nymphs); same locality, 30 Jan 1995, L-81 (1 male, 26 nymphs); same locality, waterfall levels 2 and 3, 6 Jan 1995, L-66 (1 female, 14 nymphs). SUBFAMILY LACCOCORINAE GENUS CTENIPOCORIS Ctenipocoris asiaticus Montandon Ctenipocoris asiaticus Montandon 1897: Ann. Mus. Civ. Storia Nat. Genova 17: 374-376. Diagnosis.—In most somatic characters, this genus resembles Heleocoris, al- though it is smaller and the degree of spination of the hind legs is characteristic of Ctenipocoris. Specifically, a whorl of spines encircles the distal end of the hind femur, whereas in Heleocoris several rows of parallel spines are located ventrally at the distal end. Also, the spines of Ctenipocoris are stouter than in Heleocoris. Discussion.—This species is rare, seldom collected in series, and generally occurs in ponds or in vegetation along the quiet margins of streams. The only specimen collected occurred syntopically with Naucoris scutellaris in vegetated ponds on the campus at Prince of Songkla University in Hat Yai. Material Examined.—SONGKHLA: Hat Yai, PSU campus, 5 Jan 1995, vegetated ponds, L-56 (1 male). Genus Heleocoris Diagnosis.—Members of the subfamily Laccocorinae all have the prothoracic pretarsus with two claws (Usinger 1941). The prothoracic tarsal segmentation is sexually dimorphic in Heleocoris: Males have two segments whereas females have only one. In Ctenipocoris, both sexes have two segments. Spination of the hind femur will distinguish species in this genus from Ctenipocoris. Specifically, two or more rows of parallel spines are located ventrally at the distal end of the hind femur in Heleocoris, whereas a whorl of spines encircles the distal end of the hind femur in Ctenipocoris. Also, the spines of Heleocoris are not as stout as in Ctenipocoris. Discussion.—Two species of Heleocoris were collected in southern Thailand. Because of the lack of availability of authoritatively identified comparative ma- terial in major collections, these two species are not reliably identifiable. In ad- dition, because of the dire need for taxonomic revision (Nieser & Chen 1992), these species possibly will be removed from Heleocoris because the type species of this genus is African, and differences exist between the African and Asian 1997 SITES ET AL.: NAUCORIDAE OF THAILAND 133 species (J. T. Polhemus, personal communication). One species (sp. A, which may be H. ovatus Montandon) was common when present, occurring in vegetated stream margins and very shallow, slow riffles (<< 5 cm depth). Species B was collected on only one occasion and was in a leaf pack in the plunge pool of a small waterfall. Material Examined —SPECIES A: NARATHIWAT: stream 14 km W of Srisakhon, 15 Jan 1995, L-77 (29 males, 33 females, 5 nymphs); stream below Bacho Waterfall, 15 Jan 1995, L-78 (1 female, 3 nymphs). YALA: Banglang National Park, Than To, 14 Jan 1995, L-73 (4 males, 5 females, 2 nymphs); 9 km N of Than To, 15 Jan 1995, L-76 (19 males, 14 females). SPECIES B: SONGKHLA: Hat Yai, stream on PSU campus, leaf pack of plunge pool at base of waterfall, 5 Jan 1995, L-57 (1 female, 1 nymph). SUBFAMILY NAUCORINAE GENUS NAUCORIS Naucoris scutellaris Stal Naucoris scutellaris Stal 1860: Kongl. Sv. Freg. Eugenies Jord. 266. Diagnosis.—This is the smallest of the naucorids in southern Thailand (length, 6.5—7.5 mm). The underside, particularly that of the legs, is conspicuously speck- led with dark brown spots. The forelegs have the tarsus one-segmented and pre- tarsus with one claw. The labium is short, barely reaching the fore coxae. Discussion.—This species is a common inhabitant of ponds and vegetated stream margins throughout southern Thailand. The known range of this species extends from Java to India (La Rivers 1971) and includes Thailand (Nieser & Chen 1992). Material Examined.—PHATTALU NG: Khao Chai Son, pond nr hot springs, 12 Jan 1995, L-71 (8 nymphs). SONGKHLA: reservoir at end of stream from Ton Plieuw, 7 Jan 1995, L-61 (1 female, 2 nymphs); Hat Yai, PSU campus, vegetated ponds, 4 Jan 1995, L-55 (1 male); same locality, 8 Jan 1995, L-67 (1 male, 1 female, 2 nymphs); same locality, 30 Jan 1995, L-83 (1 male, 2 females, 2 nymphs); Ton Nga Chang National Park, stream at Buddhist temple, 7 Jan 1995, L-60 (1 male). YALA: Banglang National Park, Than To, 14 Jan 1995, L-73 (2 males, 6 females); 9 km N of Than To, 15 Jan 1995, L-76 (2 males, 1 female). ACKNOWLEDGMENT We are very grateful to Prasert Chitapong and Soontorn Pipithsangchan (Prince of Songkla University) for logistical support in Thailand. We thank Wasant Atis- abda for assistance with the Thai abstract, and J. T. Polhemus for assistance with identifications and for reviewing this manuscript. We also thank Michael Nolan, University of Missouri, for providing partial support for this research, including the International Agriculture Programs in the College of Agriculture, Food, and Natural Resources for RWS and the Brown Fellowship for BJN. Partial funding for RWS also was provided by MU project PSSLO232. This is Missouri Agri- cultural Experiment Station journal series paper 12,425. LITERATURE CITED Chinajariyawong, A., J. Pecharat, S. Kritsaneepaiboon & S. Permkhamm. 1986. Scientific and com- mon names of insects in southern Thailand. Thai-Australian Prince of Songkla University Proj- ect, Faculty of Natural Resources, 39 p. Distant, W. L. 1911. The fauna of British India, including Ceylon and Burma (Rhynchota). 5: i-xii, 1-362. Taylor & Francis, London. ; 134 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(2) Fernando, C. H. & L. Cheng. 1963. A guide to Malayan water bugs (Hemiptera-Heteroptera) with keys to the genera. Dept. Zoology, Univ. Singapore, 31 pp. unpublished document. La Rivers, I. 1971. Studies of Naucoridae (Hemiptera). Biol. Soc. Nevada Mem. 2. 111 + 120 p. Lundblad, O. 1933. Zur Kenntnis der aquatilen und semi-aquatilen Hemipteren von Sumatra, Java und Bali. Arch. Hydrobiol. suppl. 12, 1-195; 263-489, 21 Taf. Mendis, A. S. & C. H. Fernando. 1962. A guide to the freshwater fauna of Ceylon. Bull. Fish. Res. Stn. Ceylon 12, 160 p. Montandon, A. L. 1897. Viaggio di Leonardo Fea in Birmania e regioni Vicine LXXV. Hemiptera Cryptocerata. Annali del Museo Civico di Storia Naturale di Genova, 17(2): 365-377. Nieser, N. & P. Chen. 1991. Naucoridae, Nepidae and Notonectidae, mainly from Sulawesi and Pulau Buton (Indonesia). Tijdschr. Entomol., 134: 47-67. Nieser, N. & P Chen. 1992. Notes on Indonesian waterbugs (Nepomorpha & Gerromorpha). Storkia, 1: 30-40. Nuttonson, M. Y. 1963. The physical environment and agriculture of Thailand. A study based on field survey data and on pertinent records, material, and reports. Am. Inst. Crop Ecol., Wash- ington D.C., 256 p. Pendleton, R. L. 1962. Thailand. Aspects of landscape and life. Am. Geograph. Soc. Handbook, Duell, Sloan and Pearce, New York. Polhemus, D. A. & J. T. Polhemus. 1988. The Aphelocheirinae of tropical Asia (Heteroptera: Nau- coridae). Raffles Bull. Zool. (1989), 36: 167-300. Sites, R. W. (in press). Economic importance of Naucoridae. In: C. W. Schaefer & A. R. Panizzi [eds.], Heteropterans of economic importance. Sites, R. W. & M. R. Willig. 1991. Microhabitat associations of three sympatric species of Naucoridae. Environ. Entomol., 20: 127-134. Stal, C. 1860. Zoologi. I. Insecta. Hemiptera. Kongl. Sv. Freg. Eugenies Jord., etc. Vettens. Lakt. IT. 2(1): 219-299, Norstedt & Sodner, Stockholm. Stys, PR & A. Jansson. 1988. Check-list of recent family-group and genus-group names of Nepomorpha (Heteroptera) of the world. Acta Entomol. Fennica, 50: 1-44. Tonapi, G. T. 1959. Studies on the aquatic insect fauna of Poona (aquatic Heteroptera). Proc. Nat. Inst. Sci. India (B), 25: 321-332. Usinger, R. L. 1937. The Naucoridae of the Philippine Islands (Hemiptera). Philip. J. Sci. (1938), 64: 299-311. Usinger, R. L. 1941. Key to the subfamilies of Naucoridae with a generic synopsis of the new subfamily Ambrysinae (Hemiptera). Ann. Entomol. Soc. Am., 34: 5-16. Received 3 Jan 1996; Accepted 26 Sep 1996. PAN-PACIFIC ENTOMOLOGIST 73(2): 135-136, (1997) Scientific Note APHODIUS ALTERNATUS HORN (APHODIINAE: SCARABAEIDAE), FIRST RECORD OF A SEMIAQUATIC SCARAB BEETLE Aphodius alternatus Horn belongs to a species group that is associated with moist habitats, generally along lake or stream margins (Gordon, R. 1977. Proc. Entomol. Soc. Wash; 79: 157—167) and are characterized by males that exhibit a large, down-curved laterally flattened anterior tibial spur (Horn, G. H. 1887. Trans. Amer. Entomol. Soc; (Philadelphia), 14: 1-110). Aphodius alternatus has been reported from moist habitats along stream, pond, slough, and prairie pothole margins (Gordon 1977) and is widespread in distribu- tion. It is recorded from Alberta, British Colombia, and Manitoba in Canada, and is reported from California, Colorado, Idaho, Iowa, Michigan, Montana, Nevada, North Dakota, Oregon, South Dakota, Utah, Washington, Wisconsin, and Wyo- ming in the United States. While studying vernal pool invertebrates, I occasionally collected A. alternatus floating on the surface or clinging to floating vegetation. These beetles were never common, and I initially believed them to have fallen into the pools inadvertently. When sampling over a larger geographic range, pools were located in which A. alternatus occurred in larger numbers (12 to 14 per square meter). Aphodius sp. collected from droppings of nearby range cattle and the droppings chambers of the local ground squirrels and gophers and were found to be differ- ent species. Aphodius alternatus were observed and collected from California grassland ver- nal pools from Shasta County south to Tulare and Monterey Counties. The biol- ogy of A. alternatus was observed in Shasta, Tehama, Placer, Sacramento, and Monterey Counties. Field observations were made from first appearance in Janu- ary until the beetles died at the end of the vernal pool season. Adults first appeared in vernal pools near the end of January. They were found clinging to floating vegetation, with one side of the abdomen exposed to air. When dislodged from the floating substrate, a beetle would reorient itself, dorsum up, flail its legs in a similar, although slower fashion as is observed in beetles of the family Hydrophilidae, and propel itself to another holdfast. Adults often let go of the vegetation to “swim” to another floating plant. Aphodius alternatus were observed feeding on insects that had fallen into the pools and drowned; however, the beetles were often forced away from their food by the large numbers of aquatic gastropods (Physatella sp. and Lymnaea sp.), and numerous turbellarians also feeding on dead insects. On warm, calm days at the end of March and into April, adults climbed as high as possible onto emergent vegetation, and then flew away. The beetles flew up to a height of 1.5—2 m in a slow zig-zag pattern. They then flew straight to another pool, paused about 0.5 m above the water surface, closed their wings and dropped into the water. During the zig-zag searching flight, the beetles flew back and forth over a 3 m wide area for a distance of up to 30 meters to find a pool. Some 136 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(2) beetles were observed flying across approximately 144 m? in five minutes. When the beetles located a pool, their flight speed increased to almost 1 m per second. Flight was observed only on still sunny days, with air temperatures at or above i. Mating occurred on the surface of the pools, when air temperatures rose above 19.5° C and the water temperature reached 10° C. Pairs in copula were only ob- served clinging to floating vegetation. Egg deposition was not observed. Toward the end of April, the vernal pools begin evaporating. As the pools dried, invertebrates and amphibian larvae were concentrated into the deeper parts of the pool where they eventually died. Adult A. alternatus were observed to feed ac- tively in these deposits of carrion which varied in weight from 2 to 26 g. Apho- dius alternatus densities varied greatly between deposits, with no apparent rela- tionship between the numbers of beetles and the weight or size of carrion. No movement of A. alternatus between carrion deposits was observed. At various times after the drying of the pools, both carrion and soil samples were taken and sieved in an attempt to find A. alternatus larvae. Samples were collected from ten pools in which A. alternatus had previously been collected in large numbers. Soil samples were taken from beneath the carrion, various sites within the pools, from pool margins, and from areas between pools. Fifteen soil samples were taken per pool, along with a variable number of carrion samples depending on the number of carrion deposition sites within the pool. In only one sample, from Shasta County, was there an aphodine larva, which died before pu- pating. Gordon (1977) reported that extensive variation occurs in A. alternatus through- out its range and that the species is made up of numerous disjunct populations. This is probable in California as populations seem to be restricted to specific ver- nal pool complexes. Acknowled gment.—A. alternatus were identified by Robert Gordon, Systematic Entomology Laboratory, Smithsonian Institute. D. Christopher Rogers, Jones & Stokes Associates, Inc. 2600 V Street, Sacra- mento CA, 95818-1914. (916)737-3000. Received 15 May 1996; Accepted 24 Dec 1996. PAN-PACIFIC ENTOMOLOGIST 73(2): 137-140, (1997) Scientific Note DISTRIBUTIONAL LIMITS OF EUGLOSSINE AND MELIPONINE BEES (HYMENOPTERA: APIDAE) IN NORTHWESTERN MEXICO Euglossine and meliponine bees are predominantly distributed within the Amer- ican tropics (Dressler, R.L. 1982. Ann. Rev. Ecol. Syst., 13: 373-384; Roubik, D.W. 1989. Ecology and Natural History of Tropical Bees. Cambridge, Univ. Press). Despite the extensive collections of euglossine bees with chemical baits, few extra-tropical records have been reported (Moure, J.S. 1967. At. Simp. Biota Amaz6on., 5: 395-415; Kimsey, L.S. & Dressler, R.L. 1986. Pan-Pacific Entomol., 62: 229-236; Kimsey, L.S. 1987. Syst. Entomol., 12: 63—72). Nevertheless, some species occur outside the geographic tropics. In South America there are reports of euglossine and meliponine bees as far south as 32° S (Moure 1967. Wittmann, D., Hoffmann, M. & Scholz, E. 1988. Entomol. Generalis, 14: 53-60), but in the Northern Hemisphere it has been thought, until recently, that their distribution was restricted to about 25° N (Roubik 1989), in the Sierra Madre Oriental and central México. During the summers of 1991 and 1993 to 1995 bee collections have been made in several localities in southern Sonora, México (Table 1). These include Nannotrigona perilampoides (Cresson), and Euglossa viridissima Friese. The former has been collected in wild nests in trunks of Jpomoea arborescens (Humb. & Bonpl. ex Willd.) G. Don in the Sierra de Alamos, and from domestic hives kept by Rafael Figueroa at his carpentry in Alamos, Sonora. Male and female specimens of Eg. viridissima have been collected visiting flowers of Te- coma stans (L.) Juss ex H.B.K. and Thevetia peruviana (Pers.) Scum ex Engler & Prantl. However, other plants known to be visited and pollinated almost strictly by euglossine bees are present in the area. These include a large complement of orchids, tropical trees, and vines like Dalechampia scandens L. (Armbuster, W.S. & Webster, G.L. 1979. Biotropica, 11: 278-283) that are known to be used by euglossine and meliponine bees as food, and also as fragrance and resin sources for attraction and nest building. Males of euglossine bees were also lured with fragrances of eugenol, methyl salycilate, vanillin, and eucalyptol. However, in this study they were only attracted to eugenol. The area where the bees have been collected is abrupt, with a broken topog- raphy dissected by numerous streams, some of them in deeply incised canyons. The vegetation consists of tropical deciduous forest and ecotones to Foothills Thornscrub and Sonoran Desert in the lowlands, and oak woodlands and pine- oak forests on the upper elevations (Gentry, H.S. 1942. Rio Mayo Plants. Camegie Inst. Washington. Publ. 527; Burquez, A., Martinez-Yrizar, A. & Felger, R.S. 1996. In: Biodiversity and Conservation in the Sonoran Desert. Robichaux, R.H. [ed.] Univ. Arizona Press). The capture of specimens of Eg. viridissima in south- ern Sonora, and the occasional reports of other Euglossini in NW Mexico, places the northern Sierra Madre Occidental in the states of Sonora and Chihuahua, as the present absolute northern limit for viable populations of this neotropical group of bees. The repeated collection of male and female individuals of Eg. viridissima, 138 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(2) Table 1. Tropical taxa collected near Alamos, Sonora, México or farther north (see Fig. 1). Taxa are ordered by collection date and plant visited. All specimens deposited at the reference collection of Estacién Regional Noroeste, Centro de Ecologia, UNAM. Elev = elevation in meters. n = number of individuals, M = male, F = female. TDF = Tropical deciduous forest. Taxon Date Collected on Locality Elev n Habitat Euglossa viridissima 03 Sep 1991 Eugenol bait Sierra de Alamos (E side) 400 1M TDF 04 Sep 1991 Thevetia peruviana Alamos 400 1F town garden 17 May 1991 Eugenol bait Sierra de Alamos (E side) 750 6M TDF 26 Oct 1994 Thevetia peruviana Alamos 400 1M town garden 26 Oct 1994 Thevetia peruviana Yocojihua 350 2M 1F TDF/thorn- scrub 30 Oct 1995 Tecoma stans Piedras Verdes 190 1M 2F town garden 30 Oct 1995 Eugenol bait Sierra Alamos (N side) 500 1M TDF 30 Oct 1995 Thevetia peruviana Yocojihua 350 2M 1F town garden 30 Oct 1995 Eugenol bait Sierra de Alamos (E side) 750 2M TDF Eulaema polychroma 04 Sep 1991 Martynia annua Arroyo Alamos (2 km E) 450 1 TDF 30 Oct 1995 Tecoma stans Alamos 400 2 town garden Nannotrigona perilampoides 30 May 1992 domestic hive Alamos 400 12 TDF/thorn- scrub 30 May 1992 nest Sierra de Alamos (E side) 750 60 TDF 30 Oct 1995 nest Sierra de Alamos (E side) 800 5 TDF Mesocheira bicolor 30 Oct 1995 Tecoma stans Piedras Verdes 190 1 town garden Mesoplia sp. 23 Abr 1990 ‘Vitex mollis Tonichi 250 1 thornscrub 13 Ago 1991 Antigonon leptopus El Gavilan (E Hermosillo) 325 1 thornscrub Xylocopa guatemalensis 17 May 1991 Martynia annua Sierra de Alamos (E side) 750 1 TDF Xylocopa muscaria 17 May 1991 Martynia annua Sierra de Alamos (E side) 750 = 1 TDF the lack of wear on their wings, and the presence of extensive tropical vegetation, along the sierran foothills indicate that these are members of persistent bee pop- ulations, rather than long-distance transient vagrants. The collection of N. peri- lampoides at their nests and domestic hives, confirms their presence and use farther north than previously reported (Schwarz, H. FE 1949. An. Inst. Biol. Mex., XX: 357-370). Other bee species found in the region near their extreme northern distribution are: Partamona bilineata (Say) [see Rozen, J. 1992. Melissa, 5: 1— 2], Eufriesea caerulescens (Lepeletier) [reported in Kimsey. & Dressler 1986. at Maguarichic, Chihuahua, but close by is Maguarichi, Sonora, both in the Rio Fuerte drainage], Eulaema polychroma (Mocsary), Mesoplia sp., Mesocheira bi- color Fabricius, Xylocopa muscaria Fabricius and X. guatemalensis Cockerell (this report, Table 1). The occurrence of these strictly tropical bee species, add support to the remarkable deep intrusion of tropical elements along the Pacific 1997 SCIENTIFIC NOTE 139 CHIHUAHUAN DESERT/ GRASSLANDS SONORAN DESERT | THORNSCRUB TROPICAL DECIDUOUS FOREST | MADREAN FORESTS AND WOODLANDS GUAYMAS 27° GULF | OF | CALIFORNIA L Figure 1. Diagram showing the vegetation of NW México-SW USA. Triangles show the collection sites mentioned in this report. coast of Mexico far north of the tropic line. This phenomenon is also evident by the numerous strictly tropical plant taxa extending their ranges northwards along the same region (Gentry 1942, P. Jenkins pers. com.). Euglossine and meliponine bees, as happen with other tropical bees, probably follow the tropical vegetation corridors along the Sierra Madre Oriental and Sierra Madre Occidental. The most northerly tropical communities (tropical deciduous 140 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(2) forest), are present on the western face of the Sierra Madre Occidental (Gentry 1942; Rzedowsky, J. 1978. Vegetaciédn de México. Limusa, México). These reach their northernmost occurrence in eastern Sonora, at the Pacific side of the Sierra, developing as an extensive vegetation belt between the Sonoran Desert and the oak Madrean Woodlands, or as isolated patches along the deep sierran canyons (Figure 1; Burquez et al. 1996). An extreme example of dispersal along the Sierra Madre biological corridors was the recent report of a single male of E. polychroma in the Sonoran Desert captured near Tucson, Arizona that probably strayed from populations from the Sierra Madre Occidental (Minckley, R. L. & Reyes, S. G. 1995. J. Kansas Entomol. Soc., 69: 102-104). Some euglossine bee species, although almost strictly tropical follow these tropical corridors in both hemispheres. Species at their extreme distribution range include Eufriesea chalybaea (Friese) that reaches 32° S, near Cérdoba, Argentina (Moure 1967), Ef. violacea (Blanchard), Eu. nigrita Lepeletier, Eg. cordata (L.), Eg. sp. indet., and Ef. sp. indet., near Rio Grande do Sul, Brazil (ca. 30° S; Wittmann et al. 1988.), while in the Northern Hemisphere, Ef. mexicana (Moc- sary) has been collected at Presidio, Durango, México (25° N), and Ef. caerules- cens (but perhaps the reputed synonym, Ef. simillima [Moure & Michener in Moure] D. Yanega pers. com.) has been collected near General Trias, México at 29° N (Rozen 1992, Minckley & Reyes 1995., Kimsey & Dressler 1986). The Sonoran collections of Eg. viridissima and N. perilampoides (this report) sets the distribution of the genera up to 27° N. However, it is probable that these species extend along the tropical deciduous forests in areas northward up to 29° N. Eu. polychroma collected at 32° N (Minckley & Reyes 1995.) is the absolute north- ermnmost range of any euglossine bee, but as Minckley & Reyes (1995) have noted, persistent populations might be farther south, in the Pacific slope of the Sierra Madre Occidental in Southern Sonora, where their reproductive populations may live sympatrically along with other euglossine and meliponine species. Acknowled gment.—Bob Minckley provided encouragement and revised previ- ous versions of the manuscript. Two anonymous reviewers gave useful criticisms. G. Eickwort+, B. Alexander+, R. Ayala, T. Griswold, R. McGinley, D. Michener, W. LaBerge, J. Rozen, D. Roubik and other PCAM members introduced me to bees. S. Meyer called my attention on the presence of domestic meliponine hives in the Alamos region. To all of them I am grateful. Alberto Burquez, Centro de Ecologia, Universidad Nacional Auténoma de Mé- xico, Apartado Postal 1354, Hermosillo, Sonora 83000, México. Received 20 Jun 1996; Accepted I Sep 1996. PAN-PACIFIC ENTOMOLOGIST Information for Contributors See volume 66(1): 1-8, January 1990, 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 INDOUBLE OR TRIPLE SPACE with 1.5 in margins on one side of 8.5 X 11 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. 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D.—Four new species of Costa Rican Ceraeochrysa (Neuroptera: Chrysopidae) -___- BRAILOVSKY, H.—Sibuyanhygia, a new genus of Colpurini from the Philippine Republic, with descriptions of three new species (Heteroptera: Coridae) AHN, K-J—A review of Liparocephalus Maklin (Coleoptera: Staphylinidae: Aleocharinae) with descriptions of larvae HYNES, C. D.—The immature stages and biology of the craneflies Toxorhina caledonica and Elephantomyia garrigouana (Diptera: Limoniidae) AREFINA, T. I—A new species of the genus Ceraclea Stephens (Trichoptera: Leptoceridae) from Zelyonyi Island (South Kuril Islands) SHAW, S. R.—The Costa Rican species of Wesmaelia Foerster with description of a new species (Hymenoptera: Braconidae: Euphorinae) FINSTON, T. L., S. B. PECK & R. B. PERRY—Population density and dispersal ability in Darwin’s darklings: Flightless beetles of the Galapagos Islands BROWN, J. W. & D. K. FAULKNER—A new species of Litoprosopus (Lepidoptera: Noctui- dae) from Baja California, Mexico SITES, R. W., B. J. NICHOLS & S. PERMKAM—tThe Naucoridae (Heteroptera) of southern Sihavlandaec some» Worn cise a a WA Te Pea a ey ye Oak Ts NL SCIENTIFIC NOTES ROGERS, D. C.—Aphodius alternatus Horn (Aphodiinae: Scarabaeidae), first record of a semi- aquatic scarab beetle BURQUEZ, A.—Distributional limits of Euglossine and Meliponine bees (Hymenoptera: Ap- idae) in northwestern Mexico 70 phe. 3 fo. 100 103 110 122 127 The PAN-PACIFIC ENTOMOLOGIST Volume 73 July 1997 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. V. Dowell, Editor R. M. Bohart R. L. Penrose, Associate Editor J. T. Doyen R. E. Somerby, Book Review Editor J. E. Hafernik, Jr. Julieta F Parinas, Treasurer Warren E. Savary Published quarterly in January, April, July, and October with Society Proceed- ings usually appearing in the October issue. All communications regarding non- receipt of numbers should be addressed to: Vincent EK Lee, Managing Secretary; and financial communications should be addressed to: Julieta F Parinas, Treasurer; at: Pacific Coast Entomological Society, Dept. of Entomology, California Acad- emy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599. Application for membership in the Society and changes of address should be addressed to: William Hamersky, Membership Committee chair, Pacific Coast Entomological Society, Dept. of Entomology, California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599. Manuscripts, proofs, and all correspondence concerning editorial matters (but not aspects of publication charges or costs) should be sent to: Dr. Robert V. Dowell, Editor, Pan-Pacific Entomologist, California Dept. of Food & Agriculture, 1220 N St., Sacramento, CA 95814. See the back cover for Information-to-Con- tributors, and volume 66(1): 1—8, January 1990, for more detailed information. Information on format for taxonomic manuscripts 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. Pacific Coast Entomological Society OFFICERS FOR 1997 Warren E. Savary, President Vincent EK Lee, Managing Secretary Julieta F Parinas, Treasurer Stanley E. Vaughn, Recording Secretary 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 31 July 1997 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 73(3): 141-151, (1997) OSMIA (HYMENOPTERA: MEGACHILIDAE) DIVERSITY AT A SITE IN CENTRAL COASTAL CALIFORNIA JOHN FE BARTHELL!”, TERRY L. GRISWOLD?, GORDON W. FRANKIE!, & ROBBIN W. THORP* ! Division of Entomology, University of California, Berkeley, California 94720 3 USDA-ARS Bee Biology & Systematics Lab, Utah State University, Logan, Utah 84322 4 Department of Entomology, University of California, Davis, California 95616 Abstract.—Thirty species of the megachilid bee genus Osmia were recorded at a research res- ervation in central coastal California during two survey periods: 1937—43 and 1987-92. Diversity remained constant at 23 species between surveys. However, cumulative diversity increased from 23 to 30 species. The total number of species at this geographic locale is relatively high when compared with nine other surveys but is most typical of diversities found at other montane, mid- elevation latitudes. Differences between study periods suggest that long-term surveys are required to accurately assess species diversity. Key Words.—Insecta, Hymenoptera, Megachilidae, Osmia, species diversity. Studies of biodiversity often rely upon knowing numbers of species, information that is critical to testing some of the most frequently discussed hypotheses in basic and applied ecology, including island biogeography and species/area relationships (reviewed in Williamson 1981, MacArthur & Wilson 1967). Within the Apoidea, taxonomic summaries for broad geographic regions have been assembled (Michener et al. 1994, Ayala et al. 1993, Roubik 1989, Westrich 1989a & b, Rust et al. 1983, Tepedino 1982, Michener 1979, Moldenke 1976, Stephen et al. 1969), but fewer studies seek to determine local species diversity in preserved, natural settings (e.g. Thorp et al. 1994, Thorp & Gordon 1992, Ayala 1988, Rust et al. 1985). We surveyed such a region for the solitary bee genus Osmia. This group rep- resents the fifth largest genus of bees in North America, containing approximately 130 species (Rust 1974). Like other genera in the family Megachilidae, many spe- cies use pre-existing cavities in their environment for nesting. This trait makes them amenable to studies that use artificial nesting sites, including trap-nests (Krombein 1967). The present study concerns collections made by hand during two historical periods and at a single locale, the Hastings Natural History Reservation (Hastings). Our objectives were two-fold. First, we wished to gain a relatively complete record of Osmia species at a preserved site in Califomia. Secondly, we wanted to compare collection results from two survey periods (1937—43 and 1987-92) for differences. MATERIALS AND METHODS Study Site —The Hastings Natural History Reservation (Hastings) is located 42 km SE of Carmel in the Santa Lucia foothills of Monterey County, California (36°23' N, 121°33' W). Hastings is part of the University of California Natural Reserve System and is managed through the Museum of Vertebrate Zoology (U.C. Berkeley). It originally encompassed 664 ha. of land when established in 1937 Current Address: Department of Biology, University of Central Oklahoma, Edmond, Oklahoma 73034. 142 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(3) and currently covers 914 ha., ranging in elevation from 467—953 m (Griffin 1988). Griffin (1974, 1990) describes six major plant communities at Hastings with 90 of the 576 plant species (15%) being introduced (Knops et al. 1995). Early research at Hastings was directed by the resident zoologist Jean M. Lins- dale who helped to accumulate a wealth of information on the natural history and species composition of the reserve (Linsdale 1947). Several taxonomic surveys were produced, including an extensive insect collection made by C. D. Michener in the late 1930s. The rediscovery of a portion of this collection in metal storage cabinets during the late 1980s coincided with our efforts to survey bee species at Hastings and other California sites (Thorp et al. 1992). Here we compare Osmia diversity at Hastings between the two survey periods as well as its overall diver- sity to other North American surveys. Collections were made primarily with a hand-held insect net with the date recorded for each specimen. Some specimens collected during the early survey period (1937—43) were stored in gelatin capsules and held in glassine envelopes with labelled, 7.6 cm X 21.7 cm index cards. These latter specimens were later mounted on insect pins to facilitate species determination. Other specimens from that period were deposited in the Snow Entomology Museum (SEM) at the Uni- versity of Kansas. These specimens were re-examined for this study and later redeposited at SEM. Specimens from the most recent survey will be deposited at University of California (Berkeley and Davis campuses) entomological museums. Host plant associations were recorded for specimens during both survey peri- ods, but most often during the second one. When possible, pollen sources were determined by observation of foraging female bees and microscopic comparison of pollen taken from flower and bee specimens. RESULTS AND DISCUSSION A total of 377 Osmia specimens were collected during the first survey period (1937-1943). Most (360) were collected during 1938 with only four specimens from 1939, 12 from 1940 and one from 1943 (none in 1937, 1941 and 1942). During the 1987—92 period, 602 Osmia specimens were collected, most of these during the last two years of the study, 1991 (227) and 1992 (222); fifteen specimens were collected during 1987, none during 1988, 52 during 1989 and 86 during 1990. During both survey periods, 23 Osmia species were documented. Seven species were unique to each survey, with a total of 30 species overall (Table 1). Geographic Comparisons of Diversity—Osmia species diversity at Hastings is relatively high. The 30 species collected there represent over 20% of the 130 Species estimated for North America by Rust (1974) and 10 of the 12 subgenera (excepting Diceratosmia and Nothosmia). Despite its relatively small size (914 hectares), Hastings has over one-third (27 of 75) of the Osmia species estimated in California by Moldenke & Neff (1974). A comparison of our results with nine other North American surveys that in- clude Osmia (Table 2) confirms the generalization that Osmia represent “‘boreal’’ species (Linsley 1958). The five most diverse Osmia locales (including Hastings) were at higher elevations (> 450 m) with substantial grassland/meadow compo- nents. The combined studies of Thorp et al. (1994) and Rust et al. (1985) yielded one-third as many Osmia species on Santa Cruz Island (off the coast of southern California) as in this survey, even though Hastings represents < 1% of the area 1997 BARTHELL ET AL.: OSMIA DIVERSITY 143 Table 1. A comparison of Osmia species collected during two study periods at the Hastings Natural History Reservation: 1937—43 and 1987-92. Collection period 1937-43 1987-92 Species Pres* (No, %) ~Pres.*¥ (No, %) Osmia (Acanthosmioides) nifoata Cockerell @ (7, 1.9) @ (16, 2.7) Osmia (Centrosmia) bakeri Sandhouse O (0, 0.0) ® (16, 2.7) Osmia (Cephalosmia) californica Cresson O (0, 0.0) @ (4, 0.7) Osmia (Cephalosmia) montana Cresson @ (15, 4.0) O (0, 0.0) Osmia (Chalcosmia) coloradensis Cresson @ (2, 0.5) O (0, 0.0) Osmia (Chalcosmia) texana Cresson @ (1, 0.3) & (30, 5.0) Osmia (Chenosmia) aglaia Sandhouse @ (19, 5.0) @ (65, 10.8) Osmia (Chenosmia) calla Cockerell @ (2, 0.5) O (0, 0.0) Osmia (Chenosmia) clarescens Cockerell © (2, 0.5) O (0, 0.0) Osmia (Chenosmia) cyanopoda Cockerell O (0, 0.0) ® (2, 0.3) Osmia (Chenosmia) granulosa Cockerell @ (10, 2.7) @ (9, 1.5) Osmia (Chenosmia) kincaidii Cockerell @ (4, 1.1) @ (5, 0.8) Osmia (Chenosmia) laeta Sandhouse @ C1282) e@ (28, 4.7) Osmia (Chenosmia) pusilla Cresson @ (1, 0.3) O (0, 0.0) Osmia (Chenosmia) regulina Cockerell © (29, 7.7) oO (17, 2.8) Osmia (Chenosmia) trevoris Cockerell O (0, 0.0) @ (5, 0.8) Osmia (Chenosmia) tristella Cockerell e@ (2, 0.5) @ (6, 1.0) Osmia (Chenosmia) zephyros Sandhouse O (0, 0.0) @ (7, 1.2) Osmia (Euthosmia) glauca (Fowler) @ (193, 51.2) ® (55, 9.1) Osmia (Monilosmia) albolateralis Sandhouse @ (5, 1.3) @ (2, 0.3) Osmia (Monilosmia) atrocyanea (Cockerell) @ (8, 2.1) @ (101, 16.8) Osmia (Monilosmia) brevis Cresson @ (4, 1.1) oO (8, 1.3) Osmia (Monilosmia) cara Cockerell td) (1, 0.3) O (0, 0.0) Osmia (Monilosmia) cyanella Cockerell @ (21, 5.6) @ (41, 6.8) Osmia (Monilosmia) gabrielis Cockerell @ (8, 2.1) e@ (135 22) Osmia (Mystacosmia) nemoris Sandhouse @ (21, 5.6) ) (25, 4.2) Osmia (Osmia) lignaria Cresson e@ (8, 2.1) @ (145, 24.1) Osmia (Osmia) ribifloris Michener O (0, 0.0) @ (1, 0.2) Osmia (Trichinosmia) latisulcata Michener @ (1, 0.3) O (0, 0.0) Osmia (Unassigned) claremontensis Michener O (0, 0.0) @ (1, 0.2) Osmia undetermined specimens — (1, 0.3) — (0, 0.0) Totals 23 (377, 100.2) 23 (602, 100.2) * Species presence (@) or absence (©). of the island. This result probably reflects the fact that Santa Cruz Island is spe- cies-poor relative to the mainland bee fauna (Thorp et al. 1994). The latitudinal extremes of Alaska (Armbruster & Gunn 1989) and Mexico (Ayala 1988) show little Osmia diversity as do the surveys conducted at Nevada and Utah sand dune habitats (Rust et al. 1983, Griswold, unpublished data). Differences Between Surveys at Hastings.—Osmia lignaria Cresson was the most frequently collected species during the 1987—92 period (24% of the collec- tion) while Osmia glauca was most common during the 1937—43 survey (193 of 377 specimens). All but one of the latter species were collected on the same day, 1 Jun, in 1938. Relatively few (eight) O. lignaria were collected during the early survey period, an artifact of a delayed initiation of the survey. The least common Species in the current survey were. Osmia ribifloris and Osmia claremontensis Table 2. A comparison of selected North American faunal surveys that include Osmia species. Survey citation Armbruster & Gunn (1989) Griswold (unpub.) Tepedino (1982) Griswold (unpub.) Griswold (unpub.) Rust et al. (1983) Griswold (unpub.) Current study Thorp et al. (1994)! Ayala (1988) Locale (disjunct sites) Habitat types USA—Alaska (32) variable—interior arctic USA—Idaho (1) montane meadow USA—Wyoming (2) shortgrass prairie USA—California (4) montane meadow USA—Nevada (5) montane meadow USA—Nevada (2) sand dunes USA—Utah (8) sand dunes USA—California (1) coastal foothill USA—California (1) variable—island MEX—Jalisco (1) seasonal lowland Years Latitude 64.0-70.0° N 42.0—43.0° N 41.0—42.0° N 40.0—41.0° N 40.0—41.0° N 39.0—40.0° N 38.0—39.0° N 36.0—37.0° N 33.5-34.5° N 19.0—20.0° N 1 Based upon the work of Rust et al. (1985) which includes additional years of collections. + Number shared with current study. Elevation (m) unreported 1768 2250-2425 1342-2286 2316-3109 1250-1400 1372-1585 467-953 0-753 unreported Method Net Malaise Net Net Net Net Net Net Net Net No. subgenera Total (sharedT) 2 (2) 7 (7) 7 (7) 8 (7) 5 (5) 1 (1) 5 (4) 10 (-) 4 () pare) No. spp. Total (sharedt) 4 (0) 28 (13) 20 (8) 34 (17) 20 (7) 1 (0) 7 (1) 30 (—) 10 (9) 0(-) vl LSIDO'IOWO.LNA OMIOVd-NVd AHL (Q)EL TOA 1997 BARTHELL ET AL.: OSMIA DIVERSITY 145 whereas O. cara, O. pusilla, O. latisulcata and O. texana were the rarest of the original survey (all species represented by a single specimen). Fifteen species from the current survey were represented by = 20 specimens, including all 7 Species not recorded during the original collection period. Although most species undetected between surveys were relatively rare or in- conspicuous, the absence of O. coloradensis and O. montana in the recent survey is enigmatic. Both are well documented in California (Rust 1974) and are rela- tively large in size with a dark metallic appearance that makes them especially conspicuous to collectors. Both species prefer asteraceous pollen (Hurd 1979) and we have collected the closely related Osmia californica and Osmia texana from the thistle species Cirsium occidentale. Nectar sources such as Salvia mellifera and Vicia villosa were also monitored without encountering either Osmia (see Appendix). If either of these species were at Hastings during the current survey, they were in such low numbers as to be undetectable. There may be several explanations for the apparent absence of species between survey periods. First, naturally occurring factors such as annual host plant vari- ation could produce temporary declines in bee population levels. Bloom intensity of Lupinus nanus can vary 11-fold between years at Hastings, for example, po- tentially influencing those bee species that require its pollen (Knops & Barthell 1996). Population fluctuations may also reflect parsivoltinism, a condition in some Osmia species that delays emergence (Torchio & Tepedino 1982). The current survey was conducted during consecutive years, however, and we are likely to have encountered such species if they inhabited Hastings in high numbers. Most plant species introductions occurred well before the first complete floral survey conducted at Hastings (Linsdale 1955). Knops et al. (1995), indicate that very early agricultural practices (~ 1800s) have profoundly altered plant com- munities at Hastings. Over 28% of the grassland and 19% of the herbaceous communities, both important sources of host plants, are now alien species. How these long-term introductions have influenced Osmia is difficult to ascertain. Al- though bee species were collected from introduced host plants, most of these Species were already at Hastings when the original survey was conducted. Ten fires are known to have at least minimally affected Hastings in this century, seven since the reserve was established (Griffin 1988). In total, they affected ~ 206 hectares of the current reserve property and 88 ha (~ 13%) of the original property. The extensive “‘Marble Cone”’ fire of 1977 did not reach Hastings but did alter nearby plant communities by producing an abundance of herbaceous species (Talley & Griffin 1980). This may have influenced pollen and nectar availability to bees, but there is no direct evidence of this occurring. Such fires also consume dead trees and limbs which might otherwise be used for nesting sites. There is little evidence that burns have had a serious effect on the avail- ability of downed wood, however. Species diversity has remained constant at Hastings between surveys (23 spe- cies) although cumulative species diversity has increased. Each survey had seven unique species, suggesting that species may become alternately present (detect- able) and absent (undetectable) in the environment over time. The absence of conspicuous species such as Osmia montana and Osmia coloradensis in the most recent survey indicates that these absences are not necessarily an artifact of dif- 146 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(3) ferent collectors, underscoring the need for long-term surveys to better estimate Species diversity. ACKNOWLEDGMENT We thank A. Barthell, K. Delate, D. Fruitt, J. Knops, M. Prentice and R. Stone for field assistance and H. Daly, E Pitelka, M. Stromberg and D. Wood for their logistical support. Charles Michener collected most of the material in the early survey and loaned us that portion of the material housed in the Snow Entomo- logical Museum, University of Kansas. The staff of Hastings Natural History Reservation (University of California, Berkeley) provided advice and an enthu- Siastic atmosphere for our research efforts. Earlier drafts of the manuscript were evaluated by H. Daly, T. Duncan, C. Michener, E Parker and N. Williams. Valu- able botanical discussions were provided by J. Griffin and J. Knops. The research was supported by the U.C. Agricultural Experiment Station and through disser- tation support to JFB by a U.C. Natural Reserve System Grant and the U.C. Entomology Department’s Walker Fund. Posthumously, we thank J. Linsdale who, with help from numerous assistants, oversaw collection and curation of specimens at Hastings. This paper was presented as part of a memorial symposium in honor of George Eickwort held at Cornell University in April of 1995. LITERATURE CITED Armbruster, W. S. & D. A. Gunn. 1989. The solitary bee fauna (Hymenoptera: Apoidea) of interior and arctic Alaska: flower associations, habitat use, and phenology. J. Kansas Entomol. Soc., 62: 468-483. Ayala, R. 1988. Abejas silvestres de Chamela, Jalisco, Mexico. Fol. Entomol. Mex., 77: 395-493. Ayala, R., T: Griswold & S. Bullock. 1993. pp. 179-227. The native bees of Mexico. Jn Ramamoorthy, T. PB, R. Bye & A. Lot (eds.). Biological diversity of Mexico: Origins and distribution. Oxford Univ. Press, New York. Griffin, J. R. 1974. Botanical resources of the Hastings reservation, Monterey county, California. Madrofio, 22: 329-333. Griffin, J. R. 1988. A natural history of Hastings reservation. Report Number 1. Hastings Natural History Reservation, Museum of Vertebrate Zoology, Natural Reserve System, University of California. Griffin, J. R. 1990. Flora of Hastings Reservation Carmel Valley, California. Report No. 2. Hastings Natural History Reservation, Museum of Vertebrate Zoology, Natural Reserve System, Uni- versity of California. Hurd, P. D., Jr. 1979. Superfamily Apoidea. pp. 1741-2209. In K. V. Krombein, P. D. Hurd, Jr., D. R. Smith & B. D. Burks, (eds.). Catalog of Hymenoptera in América North of Mexico, Volume 2, Apocrita. Smithsonian Institution Press, Washington, D. C. Knops, J. M. H. & J. E Barthell. 1996. Flower abundance in a population of sky lupine (Lupinus nanus) over three years in central coastal California. Madrofio, 43: 85-92. Knops, J. M. H., J. R. Griffin & A. C. Royalty. 1995. Introduced and native plants at the Hastings Reservation, central coastal California: a comparison. Biol. Conservation, 71: 115-123. Krombein, K. V. 1967. Trap-nesting wasps and bees: life histories, nests, and associates. Smithsonian Press, Washington, D.C. Linsdale, J. M. 1947. The Francis Simes Hastings Natural History Reservation 1937—47. University of California, Berkeley. Linsdale, J. M. 1955. Check list of ferns and seed plants of the Frances Simes Hastings Natural History Reservation. Leaf. West. Bot., 7: 201-218. Linsley, E. G. 1958. The ecology of solitary bees. Hilgardia, 27: 543-599. MacArthur, R. H. & E. O. Wilson. 1967. The theory of island biogeography. Princeton University Press, Princeton. Michener, 1979. Biogeography of the bees. Ann. Missouri Bot. Gard. 66: 277-347. 1997 BARTHELL ET AL.: OSMIA DIVERSITY 147 Michener, C. D., R. J. McGinley & B. N. Danforth. 1994. The bee genera of North and Central America (Hymenoptera: Apoidea). Smithsonian Institution Press, Washington, D. C. Moldenke, A. R. 1976. California pollination ecology and vegetation types. Phytologia, 34: 147-178. Moldenke, A. R. & J. L. Neff. 1974. The bees of California: a catalogue with special relevance to pollination and ecological research. Technical Report 74-3 (Part III of 3). Board of Studies in Biology, University of California, Santa Cruz. Roubik, D. W. 1989. Ecology and natural history of tropical bees. Cambridge University Press, Cam- bridge, Massachusetts. Rust, R. W. 1974. The systematics and biology of the genus Osmia, Subgenera Osmia, Chalcosmia, and Cephalosmia (Hymenoptera: Megachilidae). Wasmann J. Biol., 32: 1-93. Rust, R. W., L. M. Hanks & R. C. Bechtel. 1983. Aculeata Hymenoptera of Sand Mountain and Blow Sand Mountains, Nevada. Great Basin Naturalist., 43: 403-408. Rust, R., A. Menke & D. Miller. 1985. A biogeographic comparison of the bees, sphecid wasps, and mealybugs of the California channel islands (Hymenoptera, Homoptera). pp. 29-58. In Menke, A. S. & D. R. Miller (eds.). Entomology of the California Channel Islands: Proceedings of the First Symposium, Santa Barbara Museum of Natural History. Stephen, W. P., G. E. Bohart & P. E Torchio. 1969. The biology and external morphology of bees. Oregon State University Press, Corvallis, Oregon. Talley, S. N. & J. R. Griffin. 1980. Fire ecology of a montane pine forest, Junipero Serra Peak, California. Madrofio, 27: 49-60. Tepedino, V. J. 1982. Flower visitation and pollen collection records for bees of high altitude shortgrass prairie in southeastern Wyoming. Southwestern Entomol., 7: 16-25. Thorp, R. W. & D. M. Gordon. 1992. Biodiversity and pollination biology of bees in coastal nature preserves. pp. 105-111. Jn Turner, H. M. (ed.), Proc. Symp. Biodiversity of NW Calif. (28-30 Oct. 1991, Santa Rosa). Univ. of Calif. DANR Publ. Wildland Research Center, Berkeley, CA Rept. 29. Thorp, R. W., G. W. Frankie, J. Barthell, D. Gordon, L. Newstrom, T. Griswold, J. Schmidt & S. Thoenes. 1992. Ecological Research. Long-term studies to gauge effects of invading bees. Cal. Ag., 46: 20-23. Thorp, R. W., A. M. Wenner & J. EK Barthell. 1994. Flowers visited by honey bees and native bees on Santa Cruz Island. In Halverson, W. L. & G. J. Maender (eds.). Fourth California Islands Symposium: Update on the Status of Resources. Santa Barbara Museum of Natural History, Santa Barbara, California. Torchio, P. FE & V. J. Tepedino. 1982. Parsivoltinism in three species of Osmia bees. Psyche, 89: 221- 238. Westrich, P. 1989a. Die Wild-Bienen-Wiirttembergs. Allgemiener Teil: Lebensraiime, Okologie und Schutz. Stuttgart, pp. 5-431. Westrich, P. 1989b. Die Wild-Bienen-Wiirttembergs. Spezieller Teil: die Gattungen und Arten. Stutt- gart, pp. 437-972. Williamson, M. H. 1981. Island populations. Oxford University Press, Oxford. Received 28 Aug 1996; Accepted 6 Jan 1997. Appendix. Flower visitation records of Osmia species for which records are available. Amaryllidaceae Triteleia ixoides (S. Watson) E. Greene Asteraceae Centaurea melitensis L.* Cirsium occidentale (Nutt.) Jepson Boraginaceae Plagiobothrys nothofulvus (A. Gray) A. Gray Brassicaceae Cardamine californica (Torrey & A. Gray) E. Greene Caprifoliaceae Lonicera interrupta Benth. Ericaceae Arbutus menziesii Pursh Oy +0 Oy +0 Oy +0 Oy +0 O. (Acanth.) nifoata O. (Centr.) bakeri O. (Cephal.) californica O. (Chalc.) texana O. (Chen.) aglaia O. (Chen.) cyanopoda O. (Chen.) granulosa O. (Chen.) kincaidii O. (Chen.) laeta O. (Chen.) regulina O. (Chen.) trevoris O. (Chen.) tristella O. (Chen.) zephyros O. (Euth.) glauca O. (Monil.) albolateralis O. (Monil.) atrocyanea O. (Monil.) brevis — O. (Monil.) cyanella O. (Monil.) gabrielis O. (Mystac.) nemoris O. (Osmia) lignaria O. (Osmia) ribifloris O. (None) claremontensis 81 LSIDO'IONOLNY OWIOVd-NVd dH (CNEL ‘TOA 4 AS SNULIDUSOY avieiqe’] om RS 2 2G ¢ 3 P FF Se > 2s 29 § SF F EF EB = = $2 £ F F FE gs z. y S 5 w g = 5 e = s sg 5 > & & E 3 a @ : a Se er 5 a SS Qe. — a a 2. eS oS = a = S&S S g Se — 5 pe QW+O B10 8 10 O 10 OJ 10 GB 10 G 10 G4 40 Ee le” pila lll4 lel, Hee ee O. (Acanth.) nifoata ite sired aibes al a (ea Ais neem tA ed O. (Cephal.) californica CoGy GAM IMAL leery emer aN ee O- (Chen) aslaia | | | | fac ean en (es Eel US a O. (Chen.) cyanopoda Ce it plociotler) la ene) th O. (Chen.) granulosa | J ee sar ae | bas Pash as acai eg O. (Chen.) kincaidii lee ammiie Chil tne ibrar bone ee Peep hk mm Retest cca) S031; HI MIRO ose O. (Chen.) regulina 1 ae Sa] lotsa Shh ae righ iiss) ol O. (Chen.) trevoris me leh} Mintle Ya lie tee O. (Chen.) tristella [tty | (ee te ile ele rare O. (Chen.) zephyros en ere Sa hol i) see Pt On ceney plane ler all wi ees lL Noy eth leee O. (Monil.) albolateralis & — ll de eS Ada Jor 2° ls O. (Monil.) atrocyanea Point aps Pa hl eg Fel la O. (Monil.) brevis bag lt Pelt lett tl | - O. (Monil.) cyanella hie 4 BB [fe alll se [eS] me 5 O. (Monil.) gabrielis Fm eo reed fee © a a jo te ee fe, O. (Mystac.) nemoris ile s? RIT eile aliearet Ml iter ee oy oI O. (Osmia) lignaria latte yg teeORT SSS peste taeda tel O. (Osmia) ribifloris alee a LE ate Pe i) O. (None) claremontensis ALISHYHAIC VIWNSO -"TV LY TISHLAVa L66l (CNEL TOA ‘YOsy VIIUOfIVI snuumDpyy avoovUUIeyYy ‘Yo wnjoynoiospf wnuosougq oevaovuosA[og x] avadpo-sad sijvxQ QvIOvTeXO Ag[pury vyojnoinsun viy4v]D aed0vISVUO yds snudy JT e £ QS = al > 8 = cf a a s 3 , S @ ~~ g 5 (a) * +O Oy 40 OY He Le | | | || Heke eae | Pees cep dl ply | | a [ball ae’ LF lbeessal | | Pelee Pee Fale ih | ee al | | | ales P| | | a | YE | | | | eee x] PADSINA WINIGnAAD IY dvooRIUIB'T o.oo. & 6 (Acanth.) nifoata (Centr.) bakeri (Cephal.) californica (Chalc.) texana (Chen.) aglaia (Chen.) cyanopoda . (Chen.) granulosa . (Chen.) kincaidii . (Chen.) laeta (Chen.) regulina . (Chen.) trevoris . (Chen.) tristella . (Chen.) zephyros (Euth.) glauca . (Monil.) albolateralis . (Monil.) atrocyanea . (Monil.) brevis . (Monil.) cyanella . (Monil.) gabrielis . (Mystac.) nemoris . (Osmia) lignaria . (Osmia) ribifloris . (None) claremontensis LSIDO'IOWO.LNA OWIOVd-NVd FHL ‘ponuyuoy “xipueddy OST ~| vs) > Whee tis = s = § ey Q 8 3 |or|>S er FF ©CS BB é “Se (o88 FB FE 8B B reajog@ FR EE Ef 5 : a = > o os & 8 0 Psp 5 =. x © “* fe) S30 S 3 S S > io) ny» = i) ie @ n' Do fo} io) ~ : oO g = 5 Cf Ge. o b. x 6 2 = e g. : & ees Os +0 Oy 40 Os 4 Oy +0 eal | | laa | | QO. (Acanth.) nifoata || || ic? S | | O. (Centr.) bakeri | | | | | | Laat O. (Cephal.) californica || - | || | | O. (Chalc.) texana | | | stata | | O. (Chen.) aglaia | | | | | | a8 O. (Chen.) cyanopoda | | | | me | O. (Chen.) granulosa Meet [ J al O. (Chen.) kincaidii || | | on ae | | O. (Chen.) laeta | | | | ae | | O. (Chen.) regulina | | | | | | FE "| O. (Chen.) trevoris | | | | ae | | O. (Chen.) tristella \ te Pe al | | O. (Chen.) zephyros || }- aa | | O. (Euth.) glauca hit | | Ea ia O. (Monil.) albolateralis a || oe | | O. (Monil.) atrocyanea | | | | N & | | O. (Monil.) brevis || | i S | | O. (Monil.) cyanella et | | | | | | O. (Monil.) gabrielis | | ors lies iy, O. (Mystac.) nemoris || ll la ol O. (Osmia) lignaria | | | | | | | | O. (Osmia) ribifloris | | | | = | | | O. (None) claremontensis IST ALISYHAIC VINSO "TV LH TIHHLAVa L66l PAN-PACIFIC ENTOMOLOGIST 73(3): 152-155, (1997) A NEW SPECIES OF ENICOSCOLUS (DIPTERA: BIBIONIDAE) FROM BRAZIL, WITH ADDITIONAL DISTRIBUTION RECORDS FOR THE GENUS SCOTT FITZGERALD Colorado State University, Department of Entomology, Fort Collins, CO 80523 Abstract—A new species of the rare bibionid genus Enicoscolus is described from Brazil and a holotype female is designated. Characters separating Enicoscolus hardyi new species from the three other known Enicoscolus species are summarized. Illustrations to distinguish the three New World species and a key to the world species are provided. New range extentions and seasonal distribution are also given for the New World species. Key Words.—Insecta, Diptera, Bibionidae, Enicoscolus, new species, Mexico, Brazil. The bibionid genus Enicoscolus Hardy was erected for two species, E. bra- chycephalus Hardy and E. dolichocephalus Hardy, represented by five females collected in the state of Morelos, Mexico (Hardy 1961). An additional species, E. collessi Hardy, was added to the genus, represented by one female, from Queensland, Australia (Hardy 1962). Hardy (1982) reports three additional fe- males of E. collessi from the island of New Guinea, which brings the number of known specimens of the genus to a total of nine females. No fossils of Enicoscolus are known. An examination of neotropical material from various institutions pro- duced an additional seventeen females, one representing a new species from Bra- zil. Because the genus was previously known from the Australian region and Mex- ico, the discovery of a new species of Enicoscolus from South America was expected (Hardy 1962) and provides additional support for an ancient Antarctic land connection between the Australian and South American land masses. How- ever, without a phylogeny only speculation can be made as to the origin of the genus, dispersal events, and the order of vicariance events that may have led to its present-day distribution. A Gondwanian distribution makes it possible that the genus may be found in Africa, and due to its scarcity and small size, additional unknown species seem likely. Males of Enicoscolus are unknown. With the exception of brachypterous Pen- thetria funebris Meigen, no other bibionids are known to be apterous, brachyp- terous, or parthenogenetic. There is a possibility that Enicoscolus may exhibit one of these derived states, making males either difficult to collect or absent. Depositories—Califomia Academy of Sciences, San Francisco (CASC); Ca- nadian National Collection, Ottawa (CNCI); Essig Museum of Entomology, Uni- versity of California, Berkeley (EMEC); Utah State University, Logan (EMUS); Snow Entomological Museum, University of Kansas, Lawrence (SEMC); The Bohart Museum of Entomology, University of California, Davis (UCDC); Uni- versity of California, Riverside (UCRC); Coleccion Entomologica, Instituto de Biologia, Universidad Nacional Autonoma de Mexico (UNAM); United States National Museum of Natural History, Washington, D.C. (USNM). Terminology of morphology follows McAlpine (1981). 1997 FITZGERALD: A NEW BRAZILIAN ENICOSCOLUS 153 KEY TO THE SPECIES OF ENICOSCOLUS la Rostrum developed (Fig. 3); Mexico 1b Rostrum undeveloped (Figs. 1 & 2) 2a(1b) Apical segment of palpus short (see Hardy 1962: 784, fig. a); Austra- TECROUN Ce ON ST Me SOO, See ere Faas ee ee ee collessi 2b Apical segment of palpus long (Fig. 1 & see Hardy 1961: 83, fig. 1); PSK COMMMC! TREAZIT 5 se cctse eee Se ets Hace etre aticecaee Wa se oo ee eds oe 3 3a(2b) Dorsum of thorax orange; shape of head as Fig. 1; hind basitarsus slender, elongate, relative to tarsomeres 2 and 3 (Fig. 4); Brazil ........ hardyi 3b Dorsum of thorax black; shape of head as Fig. 2; hind basitarsus not so slender, elongate, relative to tarsomeres 2 and 3 (Fig. 5); Mexico RS Re rn kd Sn RI oc! Se, Sot 0 brachyce phalus Enicoscolus hardyi Fitzgerald, NEW SPECIES (Figs. 1, 4) Type.—Holotype female. BRAZIL. West border, Mato Grosso, May 1931, R.C. Shannon; deposited: Snow Entomological Museum, University of Kansas (SEMC). Flagellum of one antenna, one hind leg, and apical two tarsal segments of the remaining hind leg missing. Female.—Entirely orange-yellow with the exception of brown-orange abdomen and black occiput. Head: Antennae capitate. Antenna with six or seven flagellomeres (apical segments difficult to distin- guish); first two flagellomeres easily distinguishable, remaining flagellomeres composing a capitate knob. Palpus four- or five- segmented (basal segments difficult to distinguish). Apical palpal segment elongate, 1.5 X length of preceding segment (Fig. 1). Rostrum undeveloped (Fig. 1). Thorax: Meso- notum anteriorly (presuturally) setose, posteriorly (postsuturally) with setae in dorsocentral rows and laterally. Wing: About 2.5 mm long, anterior veins orange to light brown, posterior veins and stigma concolorous with membrane. Membrane with microtrichia. Basal portion of Rs subequal to r-m cross vein. Legs: Fore tibia with anterior spur rudimentary, posterior spine strongly developed. Hind basi- tarsus slender, elongate (~ 0.5 mm), as long as next two segments combined (Fig. 4). Hind tibial spurs short, slender, flat, apically rounded. Male.—Unknown. Diagnosis——Enicoscolus hardyi can be separated from other Enicoscolus species by the undeveloped rostrum (Fig. 1), elongate apical segment of the palpus (Fig. 1), orange thorax, hind basitarsus more slender, elongate relative to tarsomeres two and three (Fig. 4), and geographic occurrence (distribution; Brazil). Enicoscolus dolicho- cephalus differs by the developed rostrum (Fig. 3), apical segment of palpus short, hind basitarsus not so slender, elongate, relative to tarsomeres two and three (Fig. 6), and known only from Mexico. Enicoscolus collessi differs by the apical segment of palpus short (see Hardy 1962: 784, fig. a), dorsum of the thorax black, and known only from the Australian region. Enicoscolus hardyi is most similar to E. brachy- cephalus, but the latter species differs by general shape of the head (compare Figs. 1 and 2), dorsum of the thorax black, hind basitarsus not so slender, elongate, relative to tarsomeres two and three (Fig. 5), and known only from Mexico. Etymology.—The specific name honors Elmo Hardy, University of Hawaii, whose comprehensive works on world Bibionidae have made further studies of the family possible. Material Examined.—See Type. 154 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(3) ho 5 f Figures 1-3. Enicoscolus spp., female head, lateral view. Figure 1. EF. hardyi. Figure 2. E. bra- chycephalus. Figure 3. E. dolichocephalus. Figures 4—6. Enicoscolus spp., hind leg (arrow indicates basitarsus). Figure 4. E. hardyi. Figure 5. E. brachycephalus. Figure 6. E. dolichocephalus. Enicoscolus brachyce phalus Hardy (Figs. 2, 5) Enicoscolus brachycephalus Hardy 1961: 82. Type Material Examined.—Holotype female; MEXICO. MORELOS: Yautepec, 29 Oct 1956, R. & K. Dreisbach; deposited USNM. This species was previously represented by two females collected from Mo- relos, Mexico, in September and October (Hardy 1961). The following additional 1997 FITZGERALD: A NEW BRAZILIAN ENICOSCOLUS 155 records expand the geographic distribution of the species about 1,140 km north- westward to southern Chihuahua, Mexico, and expand the seasonal distribution of the species from July-November. Other Specimens Examined—MEXICO. CHIHUAHUA: 4.8 km W of Santa Barbara, 22 Jul 1967, 1 female (UCDC). JALISCO: Guadalajara, 2 Oct 1966, G.E. & A.S. Bohart, 1 female (EMUS). MORELOS: Cuernavaca, Nov 1944, N.H.L. Krauss, 2 females (USNM). PUEBLA: river E of Tepexco, 1250 m, Highway 160, 24 Aug 1977, E.I. Schlinger, 1 female (EMEC); narrow canyon 8 km S of Tecamachalco, 2103 m, flight trap, 10 Aug 1967, M.E. Irwin, 3 females (UCRC). VERACRUZ: Nov 1963, N.H.L. Krauss, 1 female (USNM). Enicoscolus dolichocephalus Hardy (Figs. 3, 6) Enicoscolus dolichocephalus Hardy 1961: 82. Type Material Examined.—Holotype female; MEXICO. MORELOS: Tepoztlan, 20 Oct 1957, R. & K. Dreisbach; deposited USNM. This species was previously represented by three females collected from Mo- relos, Mexico, in October (Hardy 1961). The following additional records of this Species extend the geographic distribution 1,425 km northwestward to Sonora, Mexico, and expand the seasonal distribution from September—October. Other Material Examined—MEXICO. MORELOS: 19.2 km E of Cuernavaca, 1310 m, 14 Aug 1954, J.G. Chillcott, 1 female (CNCI); Canon de Lobos, 6 Sep 1976, J.M. Pino, 1 female (UNAM); Canon de Lobos, 6-11 Sep 1976, J. Butze, 1 female (UNAM); (MORELOS?), Highway 95D, km 62, 3.2 km SE of LaPera, (lava beds), 30 Oct 1973, C.W. O’Brien, 1 female (CASC). NAYARIT: Tepic, 15—17 Sep 1953, B. Malkin, 1 female (CASC). SONORA: Alamos, 7 Sep 1970, G.E. & R.M. Bohart, 2 females (EMUS). ACKNOWLEDGMENT I sincerely thank Boris C. Kondratieff, Colorado State University, for support of this research and a critical review of the manuscript. I thank Paul Amaud, Jr., Cal- ifomia Academy of Sciences; Cheryl B. Barr, University of Califormia, Berkeley; Robert W. Brooks, University of Kansas, Atilano Contreras-Ramos, Coleccion En- tomologica, Instituto de Biologia, Universidad Nacional Autonoma de Mexico; Jeff Cumming, Canadian National Collection; Saul I. Frommer, University of California, Riverside; Wilford J. Hanson, Utah State University; S. L. Heydon, University of Califomia, Davis; and FE Christian Thompson, Systematic Entomology Laboratory, USDA, for the loan of material. My visit to the United States National Museum of Natural History was supported by the Samuel Wendell Williston Diptera Research fund and my visit to the Coleccion Entomologica, Instituto de Biologia, Universidad Nacional Autonoma de Mexico was supported by the Dipterology Fund. I thank Atilano Contreras-Ramos for his hospitality during my visit to the Instituto de Biol- ogia, Universidad Nacional Autonoma de Mexico. LITERATURE CITED Hardy, D. E. 1961. Notes and descriptions of exotic Bibionidae. Proc. Entomol. Soc. Wash. 63: 81-99. Hardy, D. E. 1962. A remarkable new bibionid fly from Australia. Pac. Insects 4: 783-785. Hardy, D. E. 1982. The Bibionidae (Diptera) of Australia. Aust. J. Zool. 30: 805-855. McAlpine, J. EK 1981. Morphology and terminology—adults. [Chapter] 2. pp. 9-63. In McAlpine J. F, B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, and D. M. Wood. Manual of Nearctic Diptera. Vol. 1. Agriculture Canada Monograph 27. Received 24 Jul 1996; Accepted 20 Mar 1997. PAN-PACIFIC ENTOMOLOGIST 73(3): 156-167, (1997) NEW SPECIES OF DIOXYPTERUS FAIRMAIRE FROM TONGA AND FIJI, WITH NEW DISTRIBUTION RECORDS, A TRIBAL REASSIGNMENT, AND KEY TO THE SPECIES OF THE REGION (COLEOPTERA: ELATERIDAE) PAUL J. JOHNSON Insect Research Collection, South Dakota State University, Brookings, South Dakota, 57007 Abstract—Dioxypterus tonga NEW SPECIES and D. eua NEW SPECIES are described from Tonga and are the first species of Dioxypterus reported from east of Fiji. Dioxypterus beaveri NEW SPECIES is described and D. ovalauensis Van Zwaluwenburg newly recorded from Viti Levu, Fiji. Dioxypterus is removed from the Hemirhipini (Agrypninae) and reassigned to the Dicrepidiini (Elaterinae). A key to the species of Fiji and Tonga is provided. Key Words.—Insecta, Elateridae, Dioxypterus, taxonomy, Tonga, Fiji. Dioxypterus Fairmaire is a moderately diverse genus of 31 species, including those herein described. Species of this genus are restricted to the south-central Pacific, from northeastern Papua New Guinea, through the Bismarck and Solomon archipelagos, south to Tanna I., Vanuatu, and east to Fiji. Here, two species are reported from Tonga for the first time, both previously undescribed. In addition, a new species is also described from Viti Levu, Fiji, and D. ovalauensis Van Zwaluwenburg is newly reported from Viti Levu. Adult Dioxypterus are readily recognized by their fusiform body that usually appears humped in profile at the elytral base, attenuate and acuminate elytra, antennae short and serrate reaching only to pronotal posterior margin, a frontal margin that is often incompletely carinate medially, closed pronotosternal sutures, and a prosternal process that is greatly expanded and bidentate posteriorly. This form of prosternal process is known only from the species of Dioxypterus, the Melanesian Symphostethus Schwarz, and the neotropical Ypsilosthetus semiotulus Candéze. Among the South Pacific elaterids, Dioxypterus species are distinctive and are not usually confused with other click beetles. The genus was diagnosed by Fairmaire (1881) with four Fijian species origi- nally attributed. Hyslop (1921) designated D. nigrotransversus Fairmaire as type species. Neither Schwarz (1902) nor Van Zwaluwenburg (1933, 1940) provided revised generic diagnoses of the genus while describing many of the valid species. Because of this lack of discreet characterization a redescription including the salient traits of generic value is presented below. In the following descriptions, mensural traits given are length and width. Body length is measured from the frontal margin to elytral apex, and width measured at the elytral humeri. Antennal and tarsal segment length ratios are measured along the dorsum of each segment. The ocular index is also used (Campbell & Marshall 1964). Terminology for genital structures follows Lawrence & Britton (1991), and wing venation terminology follows Kukalova-Peck & Lawrence (1993). Geo- graphic names follow Motteler (1986). 1997 JOHNSON: DIOXYPTERUS FROM TONGA & FIJI 157 Depositories—Holotypes are deposited at the Bernice P. Bishop Museum, Ho- nolulu (BPBM). Additional specimens are deposited at BPBM or in the author’s personal collection (PJJC). DIOXYPTERUS FAIRMAIRE Fairmaire’s (1881) original description lacked nearly all characteristics of value for modern interpretation of click beetle relationships. Since no subsequent author has provided a revised generic characterization one is presented here to facilitate future studies. Redescription.—Body fusiform; dorsum shallowly convex, venter strongly convex. Head convex on frons; supra-antennal ridges obtuse to subcarinate, anteromedially directed, conjoined medially to form a complete and usually subcarinate fronto-clypeal margin; antenna 11-segmented, serrate from segment 4; mandible strongly arcuate ectally; maxillary and labial palps with ultimate segment elon- gate, narrowly subsecuriform. Prothorax with pronotum trapezoidal, basal incisures absent, hind angles bicarinate dorsally; pronotosternal sutures closed anteriorly, mesal margin of hypomeron with a narrow polished and flattened bead; prosternal intercoxal process broadly, dorsoventrally arcuate posteriorly. Mesosternum with sides of median fossa subvertically declivous; mesepimeron and mesepisternum reaching coxal cavity. Meso-metasternal suture connate, with or without surface trace. Elytra attenuate, apices mucronate as an extension of strial interval 3, intervals flat to shallowly convex, striae shallowly impressed and with small punctures. Metathoracic wings with RP,, RP, and RP; apical sclerotizations, CuA,-CuA,,, crossvein present, CuA,-MP,,, juncture proximal of MP;-MP, fork. Legs proportionately long, slender, femur and tibia subequal in length; tarsus with segment 1 long, = 2X length of segment 2, segments 2—4 or 3—4 with densely setose membranous pads ventrally, segment 4 with setose pad extended anteroapically and briefly divided; pretarsal claw simple, asetose. Male with abdominal ventrite 5 broadly rounded or subtruncately lobed. Female with abdominal ventrite 5 broadly, shallowly emarginate, with or without a median subtruncate lobe. Aedeagus with lateral lobe hooked apically, and with a single or only a few apical setae. Gonocoxites slender, lightly sclerotized; styli elongate; bursa copulatrix trisaccate with a heavily sclerotized and spinose collar. Dioxypterus eua Johnson, NEW SPECIES (Figs. 1, 8-9, 11-13, 15, 17, 19, 21) Types.—Holotype, male; data: TONGA. EUA I.: Hafu, 100—200 m, Feb 1972, N.L.H. Krauss; deposited: Bernice P. Bishop Museum, Honolulu. Paratypes: Hafu, 150—200 m, Feb 1969, N.L.H. Krauss, 1 female; Pangai, 0-100 m, Jan 1979, N.L.H. Krauss, 1 male; hills above Pangai, 100—300 m, Jan 1979, N.L.H. Krauss, 1 male; Parker’s Hill area, 200—300 m, Mar 1969, N.L.H. Krauss, 1 female; Ohonua, Feb 1956, N.L.H. Krauss, 1 female. Paratypes deposited: Bernice P. Bishop Museum, Honolulu. Description Length 9.1-11.6 mm, width 2.8-3.4 mm; body flavotestaceous, with castaneous high- lights, the following structures and patterns piceous: a pair of bifurcate vittae on frons and median spot on frontal margin, anterolateral portions and hind angles of pronotum, anterior third of hypom- eron, prosternum, elevated rim of mesosternal fossa, mediolateral portions of metasternum, medial margin of metacoxa, sutural margins of ventral sclerites, and humeral region and strial punctures of elytra. Antennae and legs brunneous. Pubescence aurantaceous, moderately-dense, forming a pair of whorls on pronotal disc. Head evenly, moderately-sparsely, finely punctured; ocular index = 66; supra- antennal ridges subcarinate, shallowly arcuate over antennal fossa, then transverse, evanescently con- joined medially; clypeal region obsolescent, coarsely punctured. Antenna (Fig. 1) short, reaching to apex of pronotal hind angle; segment 2 short, segment 3 subcylindrical; segments 4—10 short, serrate; segment 2-11 length ratio = 1.0:1.2:2.0:1.8:1.8:1.8:1.9:1.8:1.8:2.2. Labrum broadly rounded anteri- orly, slightly transverse, coarsely punctured. Pronotum with medial length 0.83X width across hind angles at posterior margin, moderately-sparsely and finely punctured on disc, becoming denser and coarser anterolaterally; lateral margin carinate, remaining distinct to anterior margin; hind angles nar- 158 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(3) Figures 1-3. Dioxypterus species, antenna. Figure 1. D. eua. Figure 2. D. tonga. Figure 3. D. beaveri. Figures 4-8. Dioxypterus species. Figure 4. D. beaveri, prosternal process, lateral aspect; Figure 5. D. tonga. mesosternal profile, posterior portion. Figure 6. D. tonga, pronotum. Figure 7. D. tonga, mesosternum, ventral aspect. Figure 8. D. eua, left hypomeron. 1997 JOHNSON: DIOXYPTERUS FROM TONGA & FIJI 159 14 Figures 9-14. Dioxypterus species. Figure 9. D. eua, metathoracic wing. Figure 10. D. tonga, elytral apex. Figure 11. D. eua, elytral apex. Figure 12. D. eua, metatarsus. Figure 13. D. eua, me- tacoxal lamina. Figure 14. D. tonga, metacoxal lamina. rowly rounded at apex, dorsal carinae subequal in length, anterior ends terminating abruptly. Hypom- eron (Fig. 8) moderately-sparsely, shallowly punctured; mesal margin with narrow, flattened, polished, sparsely punctured bead widening posteriorly; posterior margin with median, subquadrate lobe. Pro- sternum with punctures fine, shallow; anterior lobe broadly arcuate; intercoxal process arcuate, com- pressed, acute at apex, expanded dorsally and subvertical posteriorly. Mesosternum with sides of median fossa slightly elevated, subvertically declivous; fossa narrowly U-shaped in ventral aspect; mesepisternum narrowly rounded at mesocoxal cavity. Elytral apex (Fig. 11) acute, shallowly arcuate mesally. Metathoracic wing (Fig. 9) with radial cell large; RP,;, RP, and RP, not conjoined, RP, obsolescent. Metasternum finely, shallowly, moderately-densely punctured; connate with mesosternum, sutural trace absent; midline shallowly engraved throughout; coxal lamina (Fig. 13) shallowly sinuate posteriorly; tarsus (Fig. 12) with segment length ratio = 1.0:0.4:0.3:0.2:0.5, segment 4 obliquely extended ventroapically. Male—Abdominal ventrite 5 (Fig. 19) with median subrectangular lobe at apex. Aedeagus (Fig. 15) with median lobe subparallel apically, apex obtuse; lateral lobe strongly narrowing at midlength, narrow and hooked at apex, with single seta at apex. Female.—Abdominal ventrite 5 (Fig. 17) emarginate and shallowly impressed at apex. Bursa cop- ulatrix as in Fig. 21. Diagnosis.—This species is similar in size and coloration to D. ovalauensis Van Zwaluwenburg, from Fiji. These two can be separated by color pattern, dis- tribution, and genital morphology, as given in the key, below. 160 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(3) 19 20 Figures 15-20. Dioxypterus species. Figure 15. Aedeagus of D. eua, dorsal aspect. Figure 16. Aedeagus of D. tonga, dorsal aspect. Figure 17. D. eua, female ventrite 5, outline of apex. Figure 18. D. beaveri, female ventrite 5, outline of apex. Figure 19. D. eua, male ventrite 5, outline of apex. Figure 20. D. tonga, male ventrite 5, outline of apex. Etymology.—Named after the island of provenance, Eua, and treated as a noun in apposition. Distribution.—Known only from Eua Island. Material Examined.—See Types. Dioxypterus tonga Johnson, NEW SPECIES (Figs. 2, 5-7, 14, 16, 20) Type.—Holotype, male; data) TONGA. EUA I, Hafu, 100—200 m, Feb 1972, N.L.H. Krauss; deposited: Bernice P. Bishop Museum, Honolulu. Description—Length 18.2 mm, width 5.9 mm; Body brunneotestaceous, with infuscate highlights; frons with a pair of narrow infuscate maculae between eyes, pronotum with large infuscate lateral maculae extending along margins and hind angles and narrowly conjoined along anterior margin, midline and posterior third brunneotestaceous; scutellum brunneotestaceous; elytra brunneotestaceous, except narrowly infuscate striae. Pubescence aurantaceous, moderately-dense, directed posteriorly ex- 1997 JOHNSON: DIOXYPTERUS FROM TONGA & FIJI 161 21 O93 Figures 21-22. Dioxypterus species, bursa copulatrix and accessory gland duct. Figure 21. D. eua. Figure 22. D. beaveri. cept as follows: directed anteriorly on head, pronotum with two discal whirls and setae directed anteriorly on anterior half and laterally on sides. Head evenly, moderately-sparsely, finely punctured; ocular index = 59; frontal margin obtuse, not cariniform, supra-antennal portion shallowly arcuate over each antennal fossa, median portion transverse; clypeal region obsolescent, coarsely punctured. Antenna (Fig. 2) short, reaching apex of pronotal hind angle; segment 2 short, segment 3 subcylindr- ical, segments 4—10 serrate; segments 2—11 length ratio = 1.0:1.8:2.5:2.4:2.4:2.0:2.0:2.0:2.0:2.9. La- brum broadly rounded anteriorly, slightly transverse, finely punctured. Pronotum (Fig. 6) with medial length 0.76X width across hind angles at posterior margin, moderately-sparsely and finely punctured on disc, denser and coarser anterolaterally; lateral margin carinate, evanescent anteriorly; hind angles narrowly rounded at apex, lateral dorsal carina 1.8X length of mesal carina, anterior ends of carinae obsolescent. Hypomeron coarsely, umbilicately punctured; mesal margin with moderately broad, flat- tened, polished, sparsely punctured anteriorly; posterior margin with median, subquadrate lobe. Pro- sternum with punctures shallowly umbilicate; anterior lobe broadly, shallowly arcuate; intercoxal pro- cess shallowly arcuate, compressed, obtuse at apex, expanded dorsally and broadly concave posteriorly. Mesosternum (Figs. 5, 7) with sides of median fossa strongly elevated, subvertically declivous; fossa narrowly V-shaped in ventral aspect; mesepisternum truncate at mesocoxal cavity. Elytral apex evenly attenuate, oblique mesally. Metathoracic wing similar to preceding species. Metasternum finely, shal- lowly, moderately-densely punctured; connate with mesosternum with partial sutural trace present laterally; midline shallowly engraved throughout; coxal lamina (Fig. 14) arcuate posteriorly; tarsus with segment 4 extended ventroapically, segment length ratio = 1.0:0.6:0.4:0.3:0.8. Male—Abdominal ventrite 5 evenly rounded apically (Fig. 20). Aedeagus (Fig. 16) with median lobe narrow, attenuate at apex; lateral lobes strongly narrowing, arcuate laterally, and with 4 subapical setae. Female—Unknown. Diagnosis.—This specimen is distinct from all known species of Dioxypterus in the combination of its size, coloration, pubescence, aedeagal morphology, and distribution. Only D. gressitti Van Zwaluwenburg is similar in size and general coloration, but differs considerably in most other traits and is only known from Guadalcanal, Solomon Islands. Dioxypterus tonga lacks immediate known affin- 162 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(3) ities and possesses certain characteristics which are tentatively regarded as an- cestral in expression, such as the ecarinate frontal margin, integument and pu- bescence lacking discrete patterns and contrasting coloration, meso-metasternal suture with surface trace restricted to lateral portions, and male abdominal ventrite 5 evenly rounded at apex. Etymology.—Named after the country of origin, Tonga, and is treated as a noun in apposition. Distribution.—Known only from Eua Island. Material Examined.—See Type. Dioxypterus beaveri Johnson, NEW SPECIES (Figs. 3-4, 11, 18, 22) Type.—Holotype, female; data: FIJI. VIT7 LEVU, Savura Creek, 1-7 Sep 1981, malaise trap, luminous, 39 09, R.A. Beaver; deposited: Bernice P. Bishop Mu- seum, Honolulu. Description—Length 11.0 mm, width 3.1 mm; integument of head and antennal segments 4-11 piceous, and pronotum, elytra and most of venter brunneopiceous, with antennal segments 1—3, labrum, anterior lobe of prosternum, hypomeron, mesosternum, mesepisternum, epipleuron, and medio-basal portion of elytra testaceous, legs infuscate. Sculpture of small, simple, moderately-dense punctures, sparse on hypomeron. Pubescence long, directed laterally from median line on pronotum, otherwise longitudinally arranged; color is generally pale testaceous, but matches ground color to form a trans- verse, midlength elytral fascia and apico-lateral inverted “‘L” pattern. Head with supra-antennal ridges obtuse, shallowly arched, obsolete medially; frontal margin coarsely punctate medially; clypeal region narrow, coplanar with frons, not separated by ridge or carina medially; ocular index = 62. Antenna (Fig. 3) short, reaching posterior margin of pronotum; segment 2 subquadrate, segment 3 subcylindr- ical, segments 4—10 serrate; segments 2-11 length ratio = 1.0:1.4:2.1:1.9:1.9:1.9:1.9:1.9:1.9:2.4. La- brum broadly rounded anteriorly. Pronotum with medial length 0.84X width across hind angles at posterior margin, moderately-sparsely and finely punctured on disc, becoming denser and coarser laterally; lateral margin carinate; hind angles narrowly rounded at apex, lateral dorsal carina 1.4X length of mesal carina, strongly elevated and slightly reflexed laterally. Hypomeron sparsely set with shallow and small to moderate sized punctures; mesal margin with moderately broad, flattened and polished, slightly elevated bead and a narrow sulcus adjacent to bead; posterior margin with median, subquadrate lobe. Prosternum punctured as hypomeron; anterior lobe broadly, evenly arcuate; inter- coxal process (Fig. 4) strongly arcuate, compressed, acute at apex, expanded dorsally and subvertical posteriorly. Mesosternum with sides of median fossa strongly elevated, subvertically declivous; fossa narrowly V-shaped in ventral aspect; mesepisternum narrowly adjacent to mesocoxal cavity. Elytral apex similar to Fig. 11. Metathoracic wing similar to that of preceding species. Metasternum finely and sparsely punctured; connate with mesosternum, without trace of suture; midline shallowly en- graved throughout; coxal lamina similar to Fig. 14, shallowly sinuate posteriorly; tarsus with segment 4 extended ventroapically, segment length ratio = 1.0:0.4:0.3: 0.2:0.5. Female—Abdominal ventrite 5 (Fig. 18) with transverse, rectangular, median projection at apex. Bursa copulatrix as in Fig. 22. Male.—Unknown. Diagnosis.—This species differs from other described Fijian Dioxypterus by its relatively small size, dark dorsal integument, and the rectangular apical projection on ventrite 5 of the female. This species is most similar to D. vagepictus Fairmaire in general coloration and patterns of pubescence on the elytra, but is readily distinguished by the dark and unicolorous pronotum of D. beaveri, versus a tes- taceous pronotum with 2 longitudinal piceous vittae on the disc of D. vagepictus. Etymology.—Named in honor of Roger A. Beaver, Chiang Mai, Thailand, in 1997 JOHNSON: DIOXYPTERUS FROM TONGA & FIJI 163 gratitude for collecting this interesting specimen and his contributions to coleop- terology. Material Examined.—See Type. Dioxypterus ovalauensis Van Zwaluwenburg Dioxypterus ovalauensis was originally described (Van Zwaluwenburg 1933) from Ovalau, Fiji. There are no subsequent island records published for this spe- cies. Specimens were examined from two localities on Viti Levu and these rep- resent a new island record for this species. Material Examined.—FIJ1. VITI LEVU: Navai, 700-800 m, 29 Sep 1970, N.L.H. Krause, 1 male (deposited BPBM); Colo-i-suva, 3-6 Mar 1963, C.M. Yoshimoto, 1 female (deposited BPBM); Savura Creek, 3-9 Apr 83, malaise trap, 58 40, R.A. Beaver; 1 female (deposited PJJC). KEY TO THE SPECIES OF DIOXYPTERUS FROM TONGA AND FIJI la. 1b. 2a (la). 2b. 3a (2a). 3b. 4a (3a). Ab. 5a (4a). Sb. 6a (5a). 6b. Ta (2b). Small, =15.0 mm in length; elytra bicolored, with maculae and TARTS eh Bete cat lh Soe RN Ae Nee lalate ade ee ed, eo eMC erem ete eae 2 Size large, ca. 18.0 mm in length; elytra bruneotestaceous; pubes- cence golden; Tonga: Eua I tonga NEW SPECIES Body flavous to orange; tarsi, pronotal hind angles, elytral maculae piceous to black Body testaceous, brunneous or brunneopiceous; elytra with flavous or. brunneous: maculac*or bands 4.00.45 «dasa os bas 7 Pronotal disc unicolorous, hind angles piceous; elytral ground color same as on venter and pronotum ..................0-e ee eee 4 Pronotum flavous, disc with pair of longitudinal infuscate vittae; elytral ground color flavobrunneous, with 3 oblique infuscate bands; Fijae Viti Mevauk cre ote). e eles ee vagepictus Fairmaire Elytral basal half concolorous with pronotum, piceous to black in apical half, apical quarter with triangular patch of testaceous pu- ESC OMICS a coca asc scealrs, ate Moaale arorelen ian sc oemagiele F wiapatalch stots Belcan ate rogs neal 5 Elytra orange in basal half, brunneous in apical third with testa- ceous pubescence, a transverse black band at midlength, and an oblique black band from humerus to suture and along suture to transverse band; Fiji: Viti Levu ...... nigrotransversus Fairmaire Elytra lacking transverse band and postscutellar patch, anterior margin of apical black area emarginate Elytra with angulate transverse band at apical third; postscutellar area infuscate; Fiji: Viti Levu muiri Van Zwaluwenburg Elytra with anterior margin of apical black area semicircularly emarginate; integument with apical triangular area piceous; Fiji: TEA, GU ss Meee ele, ete moor ue cys taveuni Van Zwaluwenburg Elytra with anterior margin of apical black area cordately emar- ginate; integument with apical triangular area rufous; Fiji: Wakai wakayensis Van Zwaluwenburg Elytra with base flavous between humerus and scutellum, and with angulate transverse, brunneous bands at midlength and apical SEDGE Seeleege us ts le get ds MUR Me a ance Seballe | re fee VN Sol wes ceed dde 8 164 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(3) 7b. Elytra concolorous, with an oblique series of spots of pallid pu- bescence at midlength and a transverse band of pallid setae at ADP LCA lel TeG! Sees Paces es Aa ee A og lies ba elle MME Sos 10 8a (7a). EE lytral intervals flat, midlength macula forming a band reaching suture; pronotum infuscate, hind angles piceous; Fiji: Viti Levu Sb. Elytral intervals shallowly convex, midlength macula reaching me- dially to interval 5 or 6; pronotum pale with infuscate maculae on disc, hind angles pale; Tonga: Eua ...... eua NEW SPECIES 9a (8a). Length 11.0 mm; pronotum dark infuscate to piceous on disc; el- ytra with piceous integumental color beneath pubescence of an- terior band and apical third ............. beaveri NEW SPECIES Ob. Length 12.9—14.2 mm; pronotum flavous with infuscate highlights; elytra with brunneoflavous integument beneath pubescence of anterior band and apical third ............... flexuosus Fairmaire 10a (7b). Larger, 13-14 mm in length; scutellum and elytral bases dark; el- ytra with anterolateral midlength spot circular and discrete from intervals 6 to 8; Fiji: Viti Levu ............ guttulatus Fairmaire 10b. Smaller, 10-12 mm in length; anterior one-half of scutellum and base of elytral interval 3 flavous; elytra with anterolateral mid- length spot transverse from interval 6 and reaching costal mar- gin; Fiji: Ovalau, Viti Levu ..... ovalauensis Van Zwaluwenburg DISCUSSION Candéze (1891) assigned Dioxypterus to his broadly inclusive “‘Ludiites.”’ Schwarz (1902) placed the genus in his ‘“Chalcolepidiites’’ (= Hemirhipini), based on the fusion of the meso- and metasterna and lack of a sutural trace between the mesocoxae. On this same character, Van Zwaluwenburg (1959) ar- ranged the genus in Campsosterninae, the latter a synonym of Oxynopterini. How- ever, the relative degree of fusion and disappearance of surface traces of the meso- metasternal suture is highly variable within many elaterid lineages (e.g., Laurent 1961, Casari-Chen 1985), and is undoubtedly convergently derived. In general, relationship extrapolation and suprageneric taxonomic assignment of taxa within the family is most reliably based on larval structure (e.g., Hyslop 1917, Ohira 1962, Dolin 1978, Calder et al. 1993). Unfortunately, larvae attrib- utable to any Dioxypterus species remain unknown. Based on salient adult traits given in the generic redescription above and notably the lack of setae on the tarsal claws, morphology of mesonotal sclerites (Gurjeva, 1974), and wing venation (Dolin, 1976), Dioxypterus properly belongs in Elaterinae. A further assignment to Dicrepidiini is based on the head capsule having a convex frons, the frontal carina conjoint with the supra-antennal carinae, closed pronotosternal sutures, and the tarsi with ventral setal pads and an ventroapical extension of segment 4. Assignment to Elaterinae, Dicrepidiini, is a new classificatory arrangement. Described species of Dioxypterus can be placed in two taxonomic groups based on coloration patterns and generally on indigenous distribution. The first group, Group I, including the new Tongan species, contains those species that are cryp- tically colored of dingy browns and yellows, often forming bands and maculae. 1997 JOHNSON: DIOXYPTERUS FROM TONGA & FIJI 165 Structurally, these species possess an incomplete frontal carina, the meso-metas- ternal suture is incompletely connate and there remains a traceable intercoxal suture line or groove. These species have a more southerly distribution and are found throughout Fiji, Tonga and Vanuatu, generally being endemic to either a single island or a local archipelago. Species of this group are known from the Solomon Islands, but only on Guadalcanal. The second species group, Group II, exhibits contrasting bright red to orange on black patterns that may be aposematic coloration. In contrast to the species in Group I these tend to possess derived characteristics such as a complete frontal carina, and connate meso-metasterna with the intercoxal suture absent and usually untraceable at the surface. Most of these species are each found on one or more islands throughout the Solomon and Bismarck archipelagos, and northeastern- most Papua New Guinea, but there is one species in each of Fiji and Vanuatu. In general, the diagnostic traits of the first species group are relatively ancestral in their expression, while those of the second species group are relatively derived. The pattern of generalized relationship and distribution is noted here because of its additive importance with that of other taxa noted below to Fijian regional biogeography. The species of Group I indicate that ancestral character states are found in those species endemic to the Fijian Region. This region contains an unusually high concentration of isolated taxa and ancestral characteristics within Elateridae. Further, the few Group II taxa in Fiji and their absence from Tonga suggest some degree of parapatric speciation between the two species groups. The biotic similarity of Tonga with Fiji and the shared geological history (Ewart, 1988) of both archipelagos made the discovery of Dioxypterus species from Tonga predictable. Previous insect faunal conclusions of a largely Fiji-de- rived biota in Tonga were made for cicadas (Duffels 1988), barklice (Thornton 1981a-b), and butterflies (Miller & Miller 1993). Similarly, Tonga shares with Fiji and Vanuatu a number of click beetle genera, such as Dioxypterus, that are either endemic (Photophorus Candéze, Hifo Candéze, Conobajulus Van Zwalu- wenburg) to the Fijian region, or have numerous species endemic to the region with species pairs shared between islands and archipelagoes (i.e. Simodactylus Candéze, Pacificola Van Zwaluwenburg, Tetrigus Candéze). The species of this latter set of taxa express character states that are ancestral in the Fijian region, relative to congenerics elsewhere in the South Pacific region. As noted above, the species of Dioxypterus follow this latter pattern with those species expressing the most intragenerically derived character states found in the Solomon Islands and Papua New Guinea. Overall, the distributions of Dioxypterus species and other endemic taxa correspond quite well with proposed phytogeographic segregation of the Fijian region (e.g., Takhtajan, 1986) and generalized Outer Melanesian Island Arc distributions (e.g., Holloway 1984, Polhemus 1995). Though there is not a phylogenetically established sister genus for Dioxypterus, potential candidates are Symphostethus Schwarz (7 species in the Solomon Is. and Papua New Guinea) and the monobasic and neotropical Ypsilosthetus Candéze. All three genera share traits involving unique prosternal and mesosternal mor- phology, with Dioxypterus and Ypsilosthetus being more similar in elytral struc- ture and pubescence style. A potential sister group for these three genera remains unresolved. This latter genus pairing is similar to four other South Pacific/neotropical ge- 166 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(3) neric pairs of click beetles: the Tongan Hifo Candéze with the Brasilian Crypto- lamprus Costa (Costa 1984), Photophorus Candéze from Fiji and Vanuatu with Meso-American [gnelater Costa (Costa 1975), the Fijian Propsephus Candéze with neotropical Dipropus Eschscholtz and the Fijian Conobajulus Van Zwalu- wenburg with the neotropical Chalcolepis Candéze. The Hifo/Cryptolamprus and Photophorus/Ignelater associations are particularly intriguing as the Fijian Region species of Hifo and Photophorus are the only bioluminescent elaterids known outside of the neotropics. With Indomalesian and Australian affinities lacking for any of the Fijian endemic genera, their unusual apparent neotropical phylogenetic associations demand further attention as they suggest that these biogeographically uniques are most likely Gondwanian faunal relics. ACKNOWLEDGEMENT Thanks are extended to G. A. Samuelson, Bishop Museum, Honolulu, and R. A. Beaver, University of the South Pacific, Suva, for the loan of specimens; and to the faculty and staff of the B. P. Bishop Museum, especially S. E. Miller, G. A. and S. Samuelson, and G. Nishida, for their support and assistance during museum visits. Travel and support was provided by the Ernst Mayr Grant Com- mittee, Museum of Comparative Zoology, Harvard University, and the Valentine Property Fund, B. P. Bishop Museum. Authorized as South Dakota Agricultural Experiment Station article No. 3004. LITERATURE CITED Calder, A. A., J. E Lawrence and J. W. H. Trueman. 1993. Austrelater, gen. nov. (Coleoptera: Elater- idae), with a description of the larva and comments on elaterid relationships. Invert. Taxon., 7: 1349-1394. Campbell, J. M. & J. D. Marshall. 1964. The ocular index and its application to the taxonomy of the Alleculidae (Coleoptera). Coleopts. Bull., 18: 42. Candéze, E. 1891. Catalogue methodique des Elaterides connus en 1890. H. Vaillant-Carmanne, Liege. Casari-Chen, S. A. 1985. Sistematica e evolugao dos Hemirhipini neotropicais (Pyrophorinae, Elater- idae, Coleoptera). Rvta. bras. Entomol., 29: 383-423. Costa, C. 1975. Systematics and evolution of the tribes Pyrophorini and Heligmini, with description of Campyloxeninae, new subfamily (Coleoptera, Elateridae). Arq. Zool., 26: 49-190. Costa, C. 1984. On the systematic position of Hifo Candéze, 1881 (Elateridae, Coleoptera). Rvta. bras. Entomol., 28: 399—402. Dolin, V. G. 1976. Wing venation in click beetles and its significance for the taxonomy of the family. Zool. Zhur., 54(11): 1618-1633. (1976 Engl. transl. by Dept. Secretary of State., Transl. Bur., Ottawa). Dolin, V. G. 1978. Table for identification of click beetles of the USSR fauna. Urozhai, Kiev, 127p. (1981 Engl. transl. for U.S.D.A. and N.S.E by Al-Ahram Center Sci. Transl.) Duffels, J. P 1988. The cicadas of the Fiji, Samoa and Tonga Islands, their taxonomy and biogeography (Homoptera, Cicadoidea). Entomonograph, Vol 10. E. J. Brill, Leiden. Ewart, A. 1988. Geological history of the Fiji-Tonga-Samoan region of the S. W. Pacific, and some palaeogeographic and biogeographic implications. Jn J. P. Duffels (see above). Fairmaire, L. 1881. Diagnoses de Coléoptéres de la Mélanesie. Naturaliste, 1: 406—407. Gurjeva, E. L. 1974. The thorax of click beetles (Coleoptera, Elateridae) and significance of its char- acters for the system of the family. Entomol. Obozr., 53: 96-113. Holloway, J. D. 1984. Lepidoptera and the Melanesian Arcs. Jn EK J. Radovsky, P. H. Raven, & S.H. Sohmer (eds.). Biogeography of the Tropical Pacific. Bish. Mus. Spec. Publ., No. 72. Hyslop, J. A. 1917. The phylogeny of the Elateridae based on larval characters. Ann. Entomol. Soc. Amer., 10: 241-256. 1997 JOHNSON: DIOXYPTERUS FROM TONGA & FIJI 167 Hyslop, J. A. 1921. Genotypes of the elaterid beetles of the world. Proc. U.S. Nat. Mus., 58: 621- 680. Kukalova-Peck, J. & J. EF Lawrence. 1993. Evolution of the hind wing in Coleoptera. Can. Entomol., 125: 181-258. Laurent, L. 1961. Sur la suture méso-métasternale chez les Elateridae (Coleoptera). Bull. Soc. Roy. Sci., Liége, 30: 68-71. Lawrence, J. E & E. B. Britton. 1991. Coleoptera (Beetles), pp. 543-683. Jn CSIRO, Insects of Australia (2nd ed.), Vol. 2. Melbourne University Press, Melbourne. Miller, J. Y. & L. D. Miller. 1993. The butterflies of the Tonga Islands and Niue, [and] Cook Islands, with the descriptions of two new subspecies. Bish. Mus. Occas. Pap. No. 34. Motteler, L. S. 1986. Pacific island names. Bish. Mus. Misc. Publ. 34. Ohira, H. 1962. Morphological and taxonomic study on the larvae of Elateridae in Japan. Entomo- logical Laboratory, Aichi Gakugei University, Ikaxaki City. Polhemus, D. A. 1995. Two new species of Rhagovelia from the Philippines, with a discussion of zoogeographic relationships between the Philippines and New Guinea (Heteroptera: Veliidae). J. N. Y. Entomol. Soc., 103: 55-68. Schwarz, O. 1902. Neue Elaterides aus dem tropischen Asien, den malayischen Inseln und den Inseln der Siidsee. Deuts. Entomol. Zeits., 1902(Heft IT): 305-350. Takhtajan, A. 1986. Floristic regions of the world. University of California Press, Berkeley. Thornton, I. W. B. 1981a. Psocoptera of the Fiji Islands. Pac. Ins. Mon. No. 37: 1-105. Thornton, I. W. B. 1981b. Psocoptera of the Tongan Archipelago. Pac. Ins. Mon. No. 37: 106-135. Van Zwaluwenburg, R. H. 1933. New Elateridae (Col.) from Melanesia. Stylops, 2: 176-185. Van Zwaluwenburg, R. H. 1940. New species and new records of elaterid beetles from the Pacific. Occas. Pap. Bern. P. Bish. Mus., 16: 91-130. Van Zwaluwenburg, R. H. 1959. Some type designations, with notes on Pacific Elateridae (Coleoptera). Pac. Ins., 1: 347-414. Received 29 Apr 1996; Accepted 12 Mar 1997. PAN-PACIFIC ENTOMOLOGIST 73(3): 168-171, (1997) A NEW SOUTHERN NEVADA SPECIES OF AEGIALIA (AEGIALIA) (COLEOPTERA: SCA RABAEIDAE: APHODIINAE) ROBERT D. GORDON! AND RICHARD W. RUST? PO. Box 65, Willow City, ND 58384 2 Biology Department, University of Nevada, Reno, NV 89557 Abstract—A new species, Aegialia (Aegialia) knighti Gordon and Rust is described and illus- trated. Aegialia knighti is from an isolated sand dune in southern Nevada and constitutes the fifth member in a flightless, inland clade of North American Aegialia. Key Words.—Insecta, Scarabaeidae, Aegialia, psammophilous, Southern Nevada. The genus Aegialia is represented by 30 species in North America (Gordon & Cartwright 1988, 1977; Gordon 1990). Aegialia are cold-adapted fossorial detri- tivores with both adult and larval stages active in the winter. They are psammo- philes, found on coastal dune systems, inland dunes, or wherever the substrate is generally sandy such as river banks, lake shores and deltas (Brown 1931; Jerath & Ritcher 1959; Jerath 1960; Stebnicka 1977; Gordon & Cartwright 1977, 1988; Rust & Hanks 1982; Gordon 1990). Here we describe a new species of Aegialia (Aegialia) from an isolated sand dune in southern Nevada. AEGIALIA (AEGIALIA) KNIGHTI, NEW SPECIES Types.—Holotype male (Figs. la, 2a) deposited in United States National Mu- seum (USNM), Washington, DC, data: NEVADA. CLARK Co.: Logandale-Ov- erton Exchange, R67E-T14S, 28 Dec 95, J. B. Knight, S. O. Cichowlaz. Allotype, female (Fig. 2b) deposited USNM, same data as holotype. Paratypes, 22 speci- mens—same data as holotype; 27 specimens data: NEVADA. CLARK Co.: Lo- gandale-Overton Exchange, R67E-T14S, 17 Dec 1995, J. B. Knight, S. O. Ci- chowlaz. Ten paratypes are in the USNM, 4 in the Nevada State Department of Agriculture collection in Reno, Nevada, 10 in the California Academy of Sci- ences, San Francisco, 10 in the Natural History Museum, London, England, 10 in the University of Nebraska collection in Lincoln, Nebraska and 5 in the Cal- ifornia State Department of Food and Agriculture collection in Sacramento, Cal- ifornia. Description.—Holotype. Male (Figs. la, 2a) length 4.3 mm, greatest width 2.4 mm. Form oval, convex, broad posteriorly. Color pale yellowish red except mouthparts and tibial spurs reddish brown to black and ventral surfaces lighter in color, exoskeleton translucent. Pubescence straw-yellow to golden-yellow, setae on legs bright golden-yellow. Head shiny, weakly alutaceous, some small, widely separated punctures in central area; clypeus weakly emarginate; gena slightly produced. Pronotum shiny, smooth, some shallow widely separated punctures throughout, posterior marginal line well developed. Elytron smooth, shiny, intervals nearly flat, finely punctate, punctures separated by twice a punctures width. Metasternum shiny, with fine punctures centrally becoming alutaceous laterally, lacking medial impunctate area. Hindwing reduced to very small lobe. Front tibia with middle and basal teeth narrowly acute, middle longer than apical tooth (Fig. 1a). Middle tibia with two well developed transverse ridges, without surface denticles, tibial spurs narrow, acute, outer spur longer and subequal to first three tarsal segments. Hind tibia with two transverse ridges, basal ridge only 1997 GORDON AND RUST: NEW AEGIALIA SPECIES 169 oe E Figure 1. Fore tibia of inland, flightless North America Aegialia species. A. A. knighti. B. A. magnifica. C. A. concinna. D. A. crescenta. E. A. hardyi. Measurement line = 0.5 mm. formed half way across tibia, transverse ridges and apical ridge with small broad setae; tibial spurs broadly spatulate, outer spur longer and subequal to first and second tarsal segments. Male genitalia as in Fig. 2a. Allotype——Similar to male except length 4.4 mm, greatest width 2.5 mm. Female genital plate as in Fig. 2b. Diagnosis:—Aegialia knighti is extremely similar to A. magnifica and will key to A. magnifica in Gordon & Cartwright (1988). Aegialia knighti differs from A. magnifica in being smaller (3.6 to 4.5 versus 4.4 to 5.9 mm in length and 2.1 to 2.4 versus 2.4 to 3.2 mm in greatest width), color pale yellow-red versus red, in having the basal and middle front tibial teeth acute (figs, la, 1b), in having broad setae on the hind tibial ridges versus lacking setae, and in lacking a median impunctate area on the central metasternum. The male and female genitalia are very different from A. magnifica (figs. 2a, 2b) and are more similar to A. crescenta (figs. 26, 29 in Gordon and Cartwright 1988). The key to Aegialia (Aegialia) (Gordon and Cartwright 1988) must be modified as follows: 11. Color pale red to pale yellow-red; head smooth, lacking tubercles, gran- ules, or coarse punctations (fig. 16) ..................0-200008- 11A Color dark brown to nearly black (except concinna); head rough, with coarse granules, punctations, or rugae ..............0. eee eee eee 12 11A. Color pale red; large, 4.4—5.9 mm long and 2.4—3.2 mm greatest width; front tibial teeth rounded, not acutely pointed ........... A. magnifica Color pale yellow-red; small, 3.6—4.5 mm long and 2.1—2.4 mm greatest width; front tibial basal and middle teeth acutely pointed ... A. knighti Figure 2. Genitalia of Aegialia knighti. A. Male. B. Female. Measurement lines = 0.25 mm. 170 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(3) Aegialia knighti is closely associated with and a member of the inland dune, reduced wing and flightless clade identified by Porter & Rust (1996). Its close resemblance to A. magnifica suggests they may be sister species, but a cladistic analysis is required to know the relationship of clade members, particularly be- cause the genitalic structures of A. knighti are not closely similar to those of A. magnifica but are strikingly like those of A. crescenta and A. hardyi. The male internal sac of A. knighti has the same number of sclerites as does A. crescenta and the form of these sclerites, especially the large primary sclerite, is little dif- ferent from that of A. crescenta and A. hardyi. Aegialia knighti has the female genital plate shaped like that of A. hardyi except with the apex obliquely angled rather than truncated as in A. hardyi. Variation.—Length ranges from 3.6 to 4.5 mm and greatest width from 2.1 to 2.5 mm. Habitat.—Aegialia knighti was collected from low, red sand hills and sand blow-outs in an area of approximately 12 km? that extends south of Mormon Mesa ridge and north and east of the Meadow Valley Wash-Wieser Wash-Muddy River drainage system from the Logandale-Overton exchange on Interstate 90 southward approximately 6 km to Logandale, Nevada. The Mojave Desert veg- etation in the area is characterized by Creosote Bush (Larrea tridentata Nuttall), Mojave Yucca (Yucca schidigera Roezl] ex Ort.), White Bur Sage (Ambrosia du- mosa Payne), Brittlebush (Encelia farinosa A. Gray), species of Opuntia cactus, and species of Atriplex. The area ranges in elevation from 550m at the Logandale- Overton exchange to 450m near Logandale. The area receives approximately 9.5 cm of precipitation per year, which may fall in any month. The area has a mean annual temperature of 18° C with a mean minimum winter temperature of —4° C and mean summer maximum temperature of 40° C (Houghton et al. 1975). Biology.—Aegialia knighti was collected by sifting sand from beneath Ephedra plants. No adults were observed on the surface sand during any of the three visits. Larvae were not obtained. Aegialia knighti appears to be very sensitive to soil moisture during the winter activity period. When the sand hills were first visited, the sand was moist a few centimeters below the surface and the individuals were very easy to collect. The sand hills were visited again on 28 Dec 1995 and a second series of specimens was easily obtained. The area was visited a third time on 16 Jan 1996 and only two individuals were collected by sifting sand. The sand hills had become dry and moist sand was not found until a depth of one meter was reached, much sand was sifted to procure the two individuals. Etymology.—We name this species for Jeff Knight who collected the new spe- cies and is continuing to study and collect the insects of Nevada. Material examined.—2 specimens data: NEVADA. CLARK Co.: 1 mi E of Logandale, 36°36'31.2" N, 114°27'44.3” W, Jan 12 1996, J. B. Knight, in the collection of the Nevada State Department of Agriculture. ACKNOWLEDGMENT We thank Jeff Knight and Scott Cichowlaz for their efforts in collecting Ae- gialia knighti, Dave Carlson, Sacramento, California for reviewing the manuscript, and Peter Brussard and the Nevada Biodiversity Initiative for providing partial funding. 1997 GORDON AND RUST: NEW AEGIALIA SPECIES 171 LITERATURE CITED Brown, W. J. 1931. A revision of the North American Aegialiinae (Coleoptera). Can. Entomol., 63: 9-19. Cornell, J. E 1967. Description of the larva of Aegialia browni Saylor. (Coleoptera: Scarabaeidae, Aphodiinae). Pan-Pac. Entomol., 43: 189-192. Gordon, R. D. & O. L. Cartwright. 1977. Four new species of Aegialia (s. str.) (Coleoptera: Scara- baeidae) from California and Nevada sand dunes. J. Wash. Acad. Sci., 67: 42—48. Gordon, R. D. & O. L. Cartwright. 1988. North American representatives of the tribe Aegialiini (Coleoptera: Scarabaeidae: Aphodiinae). Smithsonian Contr. Zool. 461: 1-37. Gordon, R. D. 1990. A new species of Aegialia (Aegialia) (Coleoptera: Scarabaeidae: Aphodiinae) from southern Arizona and new data on North American Aegialia. Coleopt. Bull., 44: 271-273. Houghton, J. G., C. M. Sakamoto & R. O. Gifford. 1975. Nevada’s weather and climate. Nevada Bur. Mines Geol. Spec. Publ. 2: 1-78. Jerath, M. L. 1960. Notes on the larvae of nine genera of Aphodiinae in the U.S. (Coleoptera: Scar- abaeidae). Proc. U.S. Nat. Mus. 3245. Jerath, M. L. & P. O. Ritcher. 1959. Biology of Aphodiinae with special reference to Oregon. Pan- Pac. Entomol., 35: 169-175. Porter, J. L. & R. W. Rust. 1996. Genetic variation within populations of five species of Aegialia (Coleoptera: Scarabaeidae). Ann. Entomol. Soc. Amer. 89: 710-721. Rust, R. W. & L. M. Hanks. 1982. Notes on the Biology of Aegialia hardyi Gordon and Cartwright (Coleoptera: Scarabaeidae). Pan-Pac. Entomol., 58: 319-325. Stebnicka, Z. 1977. A revision of the Tribe Aegialiini (Coleoptera: Scarabaeidae, Aphodiinae). Acta Zool. Crac., 22: 397-506. Received 10 Oct 1996; Accepted 21 Jan 1997. PAN-PACIFIC ENTOMOLOGIST 73(3): 172-183, (1997) FORAGING ACTIVITY, TRAILS, FOOD SOURCES AND PREDATORS OF FORMICA OBSCURIPES FOREL (HYMENOPTERA: FORMICIDAE) AT HIGH ALTITUDE IN COLORADO JOHN R. CONWAY Department of Biology, University of Scranton, Scranton, Pennsylvania 18510 Abstract.—The thatching ant, Formica obscuripes Forel, was studied at high altitude in Colorado by marking workers and flagging trails. Mounds had 1-5 trails up to 53.6 m long. Seventeen mounds had trails going to a Douglas fir tree (Pseudotsuga sp.). Activity at trail checkpoints varied from 0-171 ants per minute during the day. Ants marked on one mound were found on as many as 24 other mounds up to 77.9 m away. Ants were observed on 12 plant species and tended aphids on nine of them. Leaf clusters on mountain snowberry (Symphoricarpos rotun- difolius A. Gray) and Saskatoon serviceberry (Amelanchier alnifolia Nuttall) contained up to 1163 aphids/cluster and predaceous insect larvae. Ants also tended treehoppers, scale insects, mealybugs and galls on plants. Ants were seen feeding on an owl carcass, but usually scavenged dead insects. A bear cub was observed excavating mounds. The results are compared to other studies of this species. Key Words.—Insecta, Formica obscuripes, thatching ant, Colorado, foraging, trails. Formica obscuripes Forel, is a species of the Formica rufa group (Weber 1935) that ranges from northern Indiana and Michigan westward across the United States and southern Canada. It is an abundant ant in western North America, especially in semi-arid sagebrush areas and has been found at altitudes up to 3,170 m in Nevada (Wheeler & Wheeler 1986) and 2,896 m in Colorado (Gregg 1963). My study compared F. obscuripes predation, trails, foraging activity, and food sources at high altitude in Colorado with studies in other areas (McCook 1884; Jones 1929; Cole 1932; Weber 1935; King & Sallee 1953, 1956; Talbot 1959; 1972; Gregg 1963; Kannowski 1963; Windsor 1964; Bradley 1972, 1973a, 1973b; Clark & Comanor 1972; Knowlton 1975; Herbers 1977, 1978, 1979a, 1979b; Inouye & Taylor 1979; Wheeler & Wheeler 1983, 1986; Henderson & Akre 1986; O’Neill 1988; Seibert 1992; McIver & Loomis 1993; McIver & Steen 1994). MATERIALS AND METHODS The study site was in Gunnison County, Colorado, N of Blue Mesa Reservoir and W of Soap Creek road (N 38 30.350’, W 107 19.602’) at an altitude of 2560 m. Field observations were conducted from 5—6 Aug 1990; 20 Jun—11 Oct 1992; 28 Jun—16 Aug 1993; 29 Jun—31 Jul and 14-16 Aug 1994; 3, 29-31 Jul and 15- 16 Aug 1995, and 1—4, 18-19 Aug 1996. Eighty-five mounds were mapped in a study area (64.6 m X 114 m) using a surveyor’s transect and compass. The area, dominated by big sagebrush (Artemesia tridentata Nuttall), is adjacent to a quak- ing aspen grove (Populus tremuloides Michaux). Other plants in the study area were Chrysothamnus nauseosus (Pallas) Britton (rubber rabbitbrush), Purshia tri- dentata (Pursh) De Candolle (antelope bitterbrush), Lupinus argenteus Pursh (sil- very lupine), Symphoricarpos rotundifolius A. Gray (mountain snowberry), Rosa woodsii Lindley (Woods rose), Urtica gracilis Aiton (stinging nettle), Penstemon 1997 CONWAY: FORMICA OBSCURIPES FORAGING 173 strictus Bentham (Mancos penstemon), Jpomopsis aggregata (Pursh) Grant ssp. aggregata (trumpet gilia), one Saskatoon serviceberry tree, Amelanchier alnifolia var. pumila, and one Douglas fir tree, Pseudotsuga sp. Hundreds of workers were individually marked on eight mounds and five plants in 1992-1993 with a fine-tipped brush and model airplane paint. Thousands more were marked by spraying five mounds in 1994 and one mound in 1996 with acrylic enamel. These marking techniques did not seem to adversely affect many workers (O’Neill 1988). Trails were delineated with sprinkler flags. The terms “nest”? and “‘mound”’ are used synonymously. RESULTS Carnivorous, Insectivorous, and Herbivorous Habits.—A\though workers were observed feeding on a small owl carcass on 6—8 Jul 1993, they usually scavenged dead arthropods from late June to October. Beetles were common prey. Workers occasionally attacked and carried live insects, but did not pursue some observed on their mounds, such as aphids, a mealybug, a beetle, a spider, and another ant species. Workers were seen with plant material at three mounds: a sagebrush leaf, a sagebrush gall, and a red flower. Predation.—Seventeen mounds were found disturbed or excavated in the sum- mers of 1993-1995. On 6 Jul 1993, a bear cub was observed digging about 25 cm into one mound and about 15 cm into another mound, presumably feeding on workers and brood. Nests recovered and one mound was largely rebuilt two weeks later. Trails.—Well-defined trails lead from mounds to plants and/or to other mounds. Workers carried twigs, insects and spiders, nestmates, callows, wingless queens, larvae, and pupae on the trails. Each of the mounds studied (n = 10) had 1—5 main trails. Three mounds had branching trails: two mounds had a single branch off a trail; the third mound had two branching trails. Trails (n = 35) from these mounds ranged from 0.6—44.8 m long (mean = 7.1 m). The greatest decline in trail number for a mound over the years was from four in 1992 to one in 1994. The mound was abandoned in 1995. Marking Experiments.—Marking experiments and trails suggest that some mounds are related. Mounds #8 and #9 were connected by a trail and workers marked on mound #9 were found on mound #8. A trail ran between mounds #80 and #81 and workers marked on each mound were found on both. In addition, workers marked on a nearby Saskatoon serviceberry appeared on both mounds and both mounds had trails to the same sagebrush. Workers marked on a particular mound were found on vegetation and other mounds. For example, mound #9 workers were found on nine mounds 4.3—14.6 m away and on vegetation up to 8.2 m away. Circadian Activity.—Workers were seen leaving mounds as early as 06:40 and as late as 20:45 h in June 1992, but mound activity fluctuated greatly during the day. Mounds were active from 07:48 to approximately 11:00 h from 1 Jul—16 Aug 1993, although activity sometimes subsided as early as 10:45 and sometimes not until noon. Mound activity increased again from 13:45 to 18:40 h. It was not 174 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(3) determined if mound activity was related to changes in temperature, sunlight, or other environmental factors. Trail activity also changed during the day. Ant activity monitored for seven days in July 1994 at four checkpoints along two trails to the Douglas fir, fluctuated from O—171 ants/min. Trail activity at the checkpoints was generally high in the morning from 07:31—11:17 h, but declined from 10:12—17:05 h. Activity increased and was high again later in the day from 17:07—18:59 h. At three of four check- points, the highest trail counts (165—171 ants/min) occurred from 17:17—18:07 h. The highest count at the fourth checkpoint (117 ants/min) occurred in the morning at 09:03 h. Trails to Douglas fir.—Trails to the Douglas fir tree varied over the years. Workers marked on the trunk in 1993 were observed on or near five mounds 4.0— 15.1 m away. In 1994, 17 mounds, 3.8-53.6 m away (mean = 15.3 m), were connected to the Douglas fir by nine trails. Four of these trails led to single mounds, four led to two mounds, and one trail went to four mounds and branched to a fifth mound. Ants marked on the most distant mound from the tree, #70, were found on 24 other mounds up to 77.9 m away (mound O) (Fig 1). In 1996, ten trails led to the Douglas fir from 17 mounds 4.1-52.6 m away (mean = 15.7 m). Six trails went to single mounds, three led to two mounds, and one trail led to five mounds. Six 1994 mounds (#68, 70, 94, J, M, O) no longer had trails to the tree, but six different mounds did. The longest trail came from a mound (#65) 52.6 m away that passed through two unidentified mounds and two old mounds (#84, #48) on its way to the tree (Fig 1). Ants marked on mound #65 were subsequently found on four mounds; three with trails leading to the tree. Every year ants went into thatch at the base of the Douglas fir tree and onto the branches to tend aphids. Some workers coming down the trunk had swollen gasters and one was carrying an aphid. Ants collected insects on the tree: one was observed pursuing a small beetle and others carried small flies. Ants died while foraging; over a 20 minute period, six ants were seen carrying dead co- workers or their remains on the trunk. A number of dead ants were also found mired in tree resin. Foraging on Plants—Workers were observed on 12 plants in the area from late June to October: quaking aspen, Douglas fir, Rocky Mountain penstemon, silvery lupine, rubber rabbitbrush, big sagebrush, Saskatoon serviceberry, moun- tain snowberry, stinging nettles, Woods rose, russian thistle, and redroot buck- wheat. Ants tended aphids on nine of these plants (Table 1), but were seen most commonly on big sagebrushes which were heavily infested with aphids and visited by large numbers of ants. Some aphid locations were unusual. One aphid site was inside a curled leaf gall produced by Eriophyid mites on a quaking aspen. Large numbers of aphids were also in leaf clusters on Saskatoon serviceberry (Table 2) and mountain snowberry (Table 3). Mountain snowberry leaf clusters were 2—3 cm in diameter and one bush had at least eight leaf clusters. Workers also tended other insects, such as psylloidea on Saskatoon service- berry, treehoppers on rubber rabbitbrush and stinging nettles, scale insects on rubber rabbitbrush and big sagebrush, dipteran galls on big sagebrush, and mealy- bugs on big sagebrush and stinging nettles (Table 1). 1997 CONWAY: FORMICA OBSCURIPES FORAGING 175 QO W-SW ee © 70 Trails from Formica obscuripes mounds to a Douglas fir tree in the summers of 1994 Figure 1. and 1996 at Soap Creek, Colorado study site. xxxx 1994 trails, 1996 trails, Scale: 1 cm = 2.2 m. DISCUSSION Carrion feeding appears rare in this species. Colorado ants fed on a small owl carcass and Weber (1935) reported ants feeding on Richardson ground squirrel carcasses, Citellus richardsonii (Sabine), placed on a mound. 176 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(3) Ants carried a variety of arthropods back to nests in Colorado as noted by others (Cole 1932; Weber 1935; Bradley 1973a, 1973b; O’ Neill 1988; McIver & Steen 1994). Beetles were common prey in Colorado, but others reported the most common prey to be orthopterans (Weber 1935) or terrestrial isopods, leafhoppers, lepidopteran larvae and ants (O’Neill 1988). O’ Neill (1988) noted ants bringing back living arthropods as I observed. Colorado ants brought dead and live aphids to their mounds. Seibert (1992) also noted ants commonly carried dead aphids to their colonies, but Weber (1935) reported no aphids were taken to the nests. Workers were observed with plant material at three Colorado mounds, but were not seen carrying seeds into mounds as reported by Cole (1932). Nor were yucca seeds found in the thatch and refuse piles as noted by Windsor (1964). Weber (1935) found no evidence of ants using plants as food. Bradley (1973b) reported bear excavation of mounds as observed in this study. Other predators that have been recorded are kingbirds (Tyrannus tyrannus (L.) and 7. verticalis Say), flickers (Colaptes auratus borealis Ridgw.), the common crow (Corvus brachyrhynchos Brehm), toads (Bufo hemiophrys Cope and B. woodhousei Girard) (Weber 1935), and six species of spiders (McIver & Loomis 1993). Colorado mounds had one to five trails. Some trails branched and some trails changed during the season and from year to year as noted by O’Neill (1988). The longest trails, 44.8 m (between mounds) and 53.6 m (between a mound and a plant), went farther than those (1—21.5 m) reported by Weber (1935), Her- bers (1977), Henderson & Akre (1986), and McIver & Loomis (1993), but were shorter than the 135 m trail connecting nests reported by O’ Neill (1988). The ten trails to the Douglas fir tree from 17 different mounds were greater than previously reported to a plant. Henderson & Akre (1986) and O’Neill (1988) found that trails from several mounds frequently overlapped. O’Neill (1988) reported a system of two long parallel trails and branches serving 23 nests. In general, this did not occur in Colorado, except for some trails to the Douglas fir tree. One such trail connected five mounds. Colonies are known to be polydomous and to reproduce by budding. I found small secondary mounds around plants along the trails from large primary mounds as noted by Herbers (1979b). Secondary mounds may become a new primary nest, serve as a refuge for aphids and ant tenders from summer heat and/or ra- diation, or provide a place where tenders transfer honeydew to larger ants for transport back to the primary nest (Weber 1935, Seibert 1992, McIver & Steen 1994). Henderson & Akre (1986) saw one colony relocate to a new nest site (presumably a secondary mound) 1 meter away over a period of about two weeks. They also found a secondary mound 20 m from the primary nest, which is about 5 m farther than the ones I located in Colorado. Although Colorado workers generally followed the same trail each day as re- ported by Herbers (1977), marked ants were sometimes found off trails and on different trails. The maximum distance traveled by a marked worker between mounds was 77.9 m, farther than the 47.6 m reported by Weber (1935), but less than the 135 m of O’Neill (1988). I recovered ants marked on a single mound 1997 CONWAY: FORMICA OBSCURIPES FORAGING 177 Table 1. Insects and growths tended by Formica obscuripes on plants at Soap Creek, Colorado study site (*) compared to reports in literature. Aphids (Homoptera—EF Aphididae) Big sagebrush—Artemisia tridentata Nutt Pleotrichophorus pseudoglandulosus (Palmer) Aphis sp. = Aphis artemisicola Will. Aphis hermistonii Wil. Aphis oregonensis Wil. Macrosiphum frigidae Oest. Jones (1929) Aphis spp. Aphis minuta Macrosiphum spp. Knowlton (1975) Unident. spp. Mclver & Loomis (1993); Mclver & Steen (1994) Quaking aspen—Populus tremuloides Michaux Unidentified spp. Pterocomma populifoliae (Fitch) i Chaitophorus sp. Chaito phorus populicola Thos. Jones (1929) Neothasmia po pulicola (Thos.) Weber (1935) Lupine—Lupinus ar genteus Pursh Aphis lupini Gillette & Palmer e Mountain snowberry—Symphoricarpos rotundifolius Gray Unident. spp. Cedoaphis incognita Hottes & Frison Brevicoryne symphoricarpi (Thomas) a Saskatoon serviceberry—Amelanchier alnifolia var. pumila Unident. spp. Aphis sp. Nearctaphis sensoriata (Gillette & Bragg) * Stinging nettles—Urtica gracilis Aiton Unident. spp. Aphis sp. Bs Woods rose—Rosa woodsii Lindley Maculolachnus submacula (Walker) Unident. spp.—E Aphididae (probable) a Douglas-fir—Pseudotsuga sp. Cinara pseudotaxifoliae Palmer x Lachnus splendens Gill. & Pal. on P. taxifolia (Poir.) Jones (1929) Rubber rabbitbrush—Chrysothamnus nauseosus (Pallas) Britton Unident. spp. 7 Psylloidea (Homoptera—E Triozidae) Saskatoon serviceberry—Amelanchier alnifolia var. pumila Probable Trioza sp. ‘9 178 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(3) Table 1. Continued. Treehoppers (Homoptera—E Membracidae) Rubber rabbitbrush—Chrysothamnus nauseosus (Pallas) Publilia modesta Uhler—nymphs and adults * Unident. spp. Mclver & Loomis (1993); Mclver & Steen (1994) Stinging nettles—Urtica gracilis Aiton Unident. spp. = Scale insects (Homoptera—F. Coccidae) Rubber rabbitbrush—Chrysothamnus nauseous (Pallas) Unident. spp. i Big sagebrush—Artemisia tridentata Nutt Parthenolecanium sp. * Mealybugs (Homoptera—E Pseudococcidae) Big sagebrush—Artemisia tridentata Nutt Amonosterium lichtensioides (Cockerell) - Stinging nettles—Urtica gracilis Aiton Unident. spp. - Dipteran galls—(Diptera—F Cecidomyiidae) Big sagebrush—Artemisia tridentata Nutt Rhopalomyia pomum Gagne - Two unidentified growths (one containing aphids and other insects) Rubber rabbitbrush—Chrysothamnus nauseosus (Pallas) = on as many as 24 surrounding mounds. O’Neill (1988) found ants from eight nests on 29 other nests. Colorado trail and mound activity varied during the day. Activity at trail check- points varied from 0-171 ants/minute; the latter being the highest rate reported for this species. Activity was generally high in the morning and later in the afternoon and decreased from 10:12—17:05 h, a somewhat longer duration than the 11:00—15:00 h reported by Weber (1935). As he noted, high temperatures and direct sunlight probably curtail summer midday activity. The greatest Colorado trail activity was from 17:17—18:07 h at three checkpoints; a fourth checkpoint had the greatest activity in the morning as reported by Weber (1935). Henderson & Akre (1986) noted a different circadian pattern; fairly constant foraging from 05:00—23:00 h, but little activity from 23:00—05:00 h. I observed ants on 12 plant species and tending aphids on nine of them. Gregg (1963) also reported aphid-tending on a variety of Colorado plants. Jones (1929) listed 9 genera and 31 species of aphids on 22 plant genera in Colorado, but many of the plants and aphids differed from the ones in our study. Colorado ants tended aphids, Aphis sp. and Pleotrichophorus pseudoglandu- losus (Palmer), on big sagebrush. Although others observed ants on sagebrush (Cole 1932, Weber 1935, McIver & Steen 1994) and tending Aphis spp. (Jones 1929, Knowlton 1975), different aphids were also reported, such as Macrosiphum 1997 CONWAY: FORMICA OBSCURIPES FORAGING 179 Table 2. Four Saskatoon Serviceberry (Amelanchier alnifolia var. pumila) leaf clusters tended by Formica obscuripes at Soap Creek, Colorado study site (2560 m). Cluster 1: Cluster 2: Cluster 3: Cluster 4: 10 leaves 24 workers 403 wingless aphids (Homoptera) FE Aphididae Unidentified immature specimens Nearctaphis sensoriata (Gillette & Bragg) Ladybird larvae (Coleoptera) E Coccinellidae Small wasp (Hymenoptera) E Charipidae Lytoxysta brevipalpis Kieffer Parasitic hymenopteran larva Dipteran larvae (Diptera) E Chamaemyiidae 7 leaves and 12 berries 7 workers 24 aphids (Homoptera) FE Aphididae Unidentified immature specimens E Triozidae Probable Trioza sp. 8 leaves 5 workers 63 wingless aphids (Homoptera) FE Aphididae Unidentified immature specimens Nearctaphis sensoriata (Gillette & Bragg) 1 ladybird larva (Coleoptera) E Coccinellidae Scymnus sp. 8 leaves + 4 buds 5 workers 190 aphids (1 winged) Dipteran larvae (Diptera) E Cecidomyiidae Unidentified larvae Bremia sp. spp. (Jones 1929, Knowlton 1975) (Table 1) and Bipersona sp. (on an unidentified sagebrush species) (Weber 1935). I saw ants tending the aphid, Pterocomma populifoliae (Fitch) on quaking aspen in Colorado, but others reported species in the genera Chaitophorus (Jones 1929) and Neothasmia (Weber 1935) (Table 1). Colorado ants tended the aphid, Cinara pseudotaxifoliae Palmer, on Douglas fir, but Jones (1929) reported Lachnus splendens Gillette & Palmer on Pseudo- tsuga taxifolia (Poiret) (Table 1). Colorado ants tended aphids on plants not reported in the literature, such as silvery lupine, stinging nettle, Saskatoon serviceberry, mountain snowberry and 180 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(3) Table 3. Three Mountain snowberry (Symphoricarpos rotundifolius Gray) leaf clusters tended by Formica obscuripes at Soap Creek, Colorado study site (2560 m). Cluster 1: 93 leaves 11 workers 1163 aphids (50 winged) (Homoptera) FE Aphididae Unidentified immatures Cedoaphis incognita Hottes & Frison Brevicoryne symphoricarpi (Thomas) Thrips (2 adults and larvae ) (Thysanoptera) FE Thripidae 1st instar larva Frankliniella occidentalis (Perande) adults Frankliniella sp. larva Ladybird larvae (Coleoptera) FE Coccinellidae Dipteran larvae (Diptera) E Chamaemyliidae and possible F Chamaemyiidae E Cecidomyiidae Lestodiplosis sp. Cluster 2: 84 leaves 40 workers 707 aphids (28 winged) (Homoptera) E Aphididae Unidentified immatures Cedoaphis incognita Hottes & Frison adults Ladybird larvae (Coleoptera) FE Coccinellidae Scymnus sp. Dipteran larvae and pupae (Diptera) E Chamaemyiidae—pupae; poss. larvae EF Cecidomyiidae Lestodiplosis sp. Cluster 3: 101 leaves 9 workers 906 aphids (36 winged) (Homoptera) E Aphididae Unidentified immatures Cedoaphis incognita Hottes & Frison Ladybird larvae (Coleoptera) FE Coccinellidae Scymnus sp. Small wasp (Hymenoptera) FE Figitidae or FE Charipidae Dipteran larvae (Diptera) E Cecidomyiidae Lestodiplosis sp. FE Syrphidae Syrphinae 1997 CONWAY: FORMICA OBSCURIPES FORAGING 181 Woods rose. McIver & Loomis (1993) and McIver & Steen (1994) reported work- ers foraging for honeydew on different lupine species (Lupinus caudatus Kellogg), but the insect was not identified. Others noted ants tending aphids on the same genera as the Woods rose (Rosa) and mountain snowberry (Symphoricarpos) in Colorado (Weber 1935). Wheeler & Wheeler (1986) even noted the same two species of aphids on another species of Symphoricarpos that were on mountain snowberry in Colorado. Knowlton (1975) reported the same aphid genus, Pleotrichophorus sp., on Chrysothamnus, that I found on big sagebrush in Colorado. This is not surprising, as it is known that one ant species may tend the same aphid species on different host plants (Jones 1929). A new finding in Colorado was the large numbers of aphids (up to 1163) and other insects in Saskatoon serviceberry and mountain snowberry leaf clusters (Tables 2 and 3). Aphids have been reported to cause leaves to curl and have been found under curled leaves (Jones 1929, Weber 1935), but it is unclear if leaf cluster formation is due to aphid or ant activity. The role of other insects in Colorado leaf clusters, such as a psylloidea, small wasps (FE Charipidae and possible EF Figitidae), and a parasitic hymenopteran larva are unclear. Dipteran larvae (E Chamaemylidae, EK Cecidomyiidae, FE Syrphidae), ladybird larvae (EK Coccinellidae), and thrip larvae and adults (EK Thripidae) in the clusters probably preyed on aphids (Tables 2 and 3). Jones (1929) suggested that ants may protect aphids from many natural enemies, such as chalcids, syr- phids, coccinellids and chrysopids. Workers tended treehopper nymphs and adults of Publilia modesta Uhler on rubber rabbitbrush in Colorado. Others have also noted ants tending Membracids (Wheeler 1910, Cole 1932) and collecting honeydew on rubber rabbitbrush (McIver & Loomis 1993, McIver & Steen 1994) (Table 1). O’Neill (1988) re- ported the membracids, Campylenchia latipes Say and Publilia modesta (Uhler), being tended on Canada thistle (Cirsium arvense) and chokecherry (Prunus vir- giniana). Colorado ants also tended scale insects on rubber rabbitbrush and big sage- brush, dipteran leaf galls on big sagebrush, mealybugs on big sagebrush and stinging nettles, and two unidentified growths on rubber rabbitbrush (one housing aphids and other insects) (Table 1). They probably collected honeydew from all the above, except the dipteran galls. Further research is needed to determine if they harvest emerging flies from the latter. Thus, as Seibert (1992) noted, F. obscuripes is not dependent on any one mutualistic partner. It has many nectar sources including coccids (Bradley 1973a, Seibert 1989, McIver & Steen 1994) and extrafloral nectar (Tilman 1978, Inouye & Taylor 1979, Seibert 1989). ACKNOWLEDGMENT Support for this research was provided by grants in 1993-1994 from the How- ard Hughes Medical Institute through the Undergraduate Biological Sciences Ed- ucation Program to the following University of Scranton students: John Bridge, Tom Sabalaske, Anthony Musingo, and Jeanne Rohan. I thank the Systematic Entomology Laboratory in Beltsville, Maryland and John T. Sorensen, Insect Tax- onomy Laboratory, California Department of Food & Agriculture, Sacramento, 182 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(3) California for identification of insect specimens. The Forest Service in Gunnison, Colorado identified plants. LITERATURE CITED Bradley, G. A. 1972. Transplanting Formica obscuripes and Dolichoderus taschenbergi (Hymenoptera: Formicidae) colonies in Jack Pine stands of southeastern Manitoba. Can. Ent., 104: 245-249. Bradley, G. A. 1973a. Effect of Formica obscuripes (Hymenoptera: Formicidae) on the predator-prey relationship between Hyperaspis congressis (Coleoptera: Coccinellidae) and Toumeyella num- ismaticum (Homoptera: Coccidae). Can. Ent., 105: 1113-1118. Bradley, G. A. 1973b. Interference between nest populations of Formica obscuripes and Dolichoderus taschenbergi (Hymenoptera: Formicidae). Can. Ent., 105: 1525-1528. Clark, W. H. & P. L. Comanor. 1972. Flights of the western thatching ant, Formica obscuripes Forel, in Nevada. Great Basin Nat., 32: 202—207. Cole, A. C., Jr. 1932. The thatching ant, Formica obscuripes Forel. Psyche, 39:30-33. Conway, J. R. 1996. Nuptial, pre-, and postnuptial activity of the thatching ant, Formica obscuripes Forel, in Colorado. Great Basin Nat., 56: 54—58. Gregg, R. E. 1963. The ants of Colorado. University of Colorado Press. Boulder, Colorado. Henderson, G. & R. D. Akre. 1986. Biology of the myrmecophilous cricket, Myrmecophilia manni (Orthoptera: Gryllidae). J. Kans. Ent. Soc., 59: 454-467. Herbers, J. M. 1977. Behavioral constancy in Formica obscuripes. Ann. Ent. Soc. Am., 70: 485-486. Herbers, J. M. 1978. Trends in sex ratios of the reproductive broods of Formica obscuripes. Ann. Ent. Soc. Am., 71: 791-793. Herbers, J. M. 1979a. Caste-biased polyethism in a mound-building ant species. Amer. Midl. Nat., 101: 69-75. Herbers, J. M. 1979b. The evolution of sex-ratio strategies in hymenopteran societies. Amer. Nat., 114: 818-834. Inouye, D. W. & O. R. Taylor. 1979. A temperate region plant-ant-seed predator system: consequences of extra floral nectar secretion by Helianthella quinquenervis. Ecology, 60: 1-7. Jones, C. R. 1929. Ants and their relation to aphids. Bull. Colo. Agr. Exp. Sta., 341: 1-96. Kannowski, P. B. 1963. The flight activities of formicine ants. Symp. Genet. Biol. Ital., 12: 74-102. King, R. L. & R. M. Sallee. 1953. On the duration of nests of Formica obscuripes Forel. Proc. lowa Acad. Sci., 60: 656-659. King, R. L. & R. M. Sallee. 1956. On the half-life of nests of Formica obscuripes Forel. Proc. lowa Acad. Sci., 63: 721-723. Knowlton, G. F 1975. Ants of Curlew Valley, Utah and Idaho. Utah State University Ecology Center. Terrestrial Arthropod Series No. 13: 1-10. McCook, H. C. 1884. The rufous or thatching ant of Dakota and Colorado. Proc. Acad. Nat. Sci. Phila. Part 1: 57-65. Mclver, J. D. & C. Loomis. 1993. A size-distance relation in Homoptera-tending thatch ants (Formica obscuripes, Formica planipilis). Insect. Soc., 40: 207-218. Mclver, J. D. & T. Steen. 1994. Use of a secondary nest in great basin desert thatch ants (Formica obscuripes Forel). Great Basin Nat., 54: 359-365. O’Neill, K. M. 1988. Trail patterns and movement of workers among nests in the ant Formica obscu- ripes (Hymenoptera: Formicidae). Psyche, 95: 1-13. Seibert, T. EF 1989. Foraging on insect mutualists by the thatching ant Formica obscuripes: interspecific and intraspecific mechanisms and implications. Ph.D. dissertation, University of Illinois, Ur- bana. Seibert, T. E 1992. Mutualistic interactions of the aphid Lachnus allegheniensis (Homoptera: Aphid- idae) and its tending ant Formica obscuripes (Hymenoptera: Formicidae). Ann. Ent. Soc. Am., 85: 173-178. Talbot, M. 1959. Flight activities of two species of ants of the genus Formica. Amer. Midl. Nat., 61: 124-132. Talbot, M. 1972. Flights and swarms of the ant Formica obscuripes Forel. J. Kans. Ent. Soc., 45: 254-258. Tilman, D. 1978. Cherries, ants and tent caterpillars: timing of nectar production in relation to sus- ceptibility of caterpillars to ant predation. Ecology, 59: 686-692. 1997 CONWAY: FORMICA OBSCURIPES FORAGING 183 Weber, N. A. 1935. The biology of the thatching ant Formica obscuripes Forel in North Dakota. Ecol. Monogr., 5: 165-206. Wheeler, W. M. 1910. Ants; their structure, development and behavior. Columbia University Press. Wheeler, G. C. & J. N. Wheeler. 1983. The ants of North Dakota. Nat. Hist. Mus. Los Ang. Cty., Los Angeles, CA. Wheeler, G. C. & J. N. Wheeler. 1986. The ants of Nevada. Nat. Hist. Mus. Los Ang. Cty., Los Angeles, CA. Windsor, J. K. 1964. Three Scarabaeid genera found in nests of Formica obscuripes Forel in Colorado. Bull. South. Calif. Acad. Sci., 63:205—209. Received 28 Sep 1996; Accepted 17 Apr 1997. PAN-PACIFIC ENTOMOLOGIST 73(3): 184-185, (1997) Scientific Note AN OUTBREAK OF THE MOTH ACHAEA SERVA (FABR.) ON THE MANGROVE EXCOECARIA AGALLOCHA (L.) There have been several reports of lepidopteran larvae causing damage to the foliage of mangroves. One noteworthy case occurred in 1983 when between 5— 10 km? of Excoecaria agallocha (L.) near Belawan in Northern Sumatra, Indo- nesia, was almost completely defoliated by caterpillars of the noctuid moth Ophiu- sa melicerta (Fabr.) (Whitten A. J. & S. J. Damanik, 1986. Biotropica, 18: 176). Whitten & Damaniks’ report is intriguing because E. agallocha has also been recorded as having one of the lowest levels of leaf damage among mangroves and it has been suggested that E. agallocha is relatively resistant to attack by herbivorous insects because it produces a toxic sap (Robertson, A. I. 1991. Aust. J. Ecol., 16: 433-443). Excoecaria agallocha is common within the landward margins of mixed man- grove stands along the banks of the Fitzroy River, Central Queensland, Australia, as far inland as Rockhampton (23°23'S, 150°31’E). During April 1995 we noticed that the E. agallocha within a strip at least 10 km long downstream from Rock- hampton were almost completely defoliated. We surveyed the forest in April 1995 and found the mud beneath the E. agallocha trees littered with chewed fragments of leaf and numerous caterpillars of the noctuid moth Achaea serva (Fabr.) eating the few leaves remaining on the trees. Within the area of defoliation we also found 3 separate patches, each of which was less than 60 m?’, where E. agallocha showed little or no leaf damage or defoliation. Fresh groundwater (salinity less than 2%o) was running into the man- grove forest from the landward edge of each patch, the boundary of which fol- lowed the extent of obvious freshwater seepage and was sharply defined; trees with almost undamaged foliage occurred within 1—2 m of completely defoliated ones. No other patches of freshwater input were found within the defoliated area. Caterpillars of A. serva were present on the undefoliated trees, but stopped feeding and often fell to the ground within a few seconds of starting to feed; the leaves from these trees were rigid and oozed a copious watery latex when snapped across the midrib. In contrast, caterpillars feeding on leaves remaining on almost completely defoliated trees usually consumed the entire leaf apart from the midrib; the leaves were limp and produced little or no latex when snapped. This may be another example of the outbreak of an herbivorous insect on plants which have become more attractive or susceptible to attack due to drought stress (e.g. White, T. C. R. 1993. The inadequate environment. Springer-Verlag, Berlin). From 1993-1995 there was a severe drought in Central Queensland coinciding with the 1993—95 El Nino Southern Oscillation event (Anonymous. 1995. Month- ly weather review for Queensland. Australian Bureau of Meterology, Canberra), and we suggest this drought resulted in reduced toxin production by E. agallocha and hence the outbreak of A. serva. Furthermore, the outbreak of Ophiusa meli- certa on E. agallocha in Indonesia reported by Whitten & Damanik (1986) oc- 1997 SCIENTIFIC NOTE 185 curred in January 1983; the year following one of the most extreme El Nino- Southern Oscillation events ever recorded, during which there was severe and widespread drought in Indonesia (Gill A. E. & E. M. Rasmusson. 1983. Nature, 306: 229-234). The current El Nino event appears to have ended and rainfall during the south- ern hemisphere winter—spring of 1995 (June—November) in Rockhampton was significantly greater than in the same seasons of the previous year (Anonymous 1995). The defoliated E. agallocha have grown new leaves and no adults, pupae or caterpillars of A. serva were found during subsequent surveys in September 1995 or April 1996. We think our observations are noteworthy for two reasons. First, they have resulted in an hypothesis which can explain two outbreaks of noctuids on E. agallocha. Second, considering that A. serva has been reported on castor, Ricinus communis L. (Common, I. E B. 1990. Moths of Australia. Melbourne University Press, Melbourne), the outbreak on E. agallocha emphasises that species which have become more susceptible to herbivores due to drought stress may temporarily increase the effective host range (and thus perhaps the population density) of polyphagous pests of commercial crops. We intend to continue monitoring E. agallocha in Rockhampton because a future drought may provide an opportunity to test the above-mentioned hypothesis by experimentally irrigating parts of the Mangrove swamp. Acknowledgment.—The Rockhampton City Council allowed access to the man- grove forest and Dr. Bob Newby identified Achaea serva. Stephen C. McKillup and Ruth V. McKillup, Department of Biology, Central Queensland University, Rockhampton, Queensland 4702 Australia. Received I Jul 1996; Accepted 30 Sep 1996. PAN-PACIFIC ENTOMOLOGIST 73(3): 186-189, (1997) Scientific Note HESPEROPSIS GRACILIAE (MacNEILL) (LEPIDOPTERA: HESPERIITIDAE) FLIGHT BETWEEN HOSTPLANTS AND PROSOPIS GLANDULOSA TORREY MacNeill’s sootywing skipper, Hesperopsis graciliae (MacNeill), is a small (wingspread ~ 23 mm) dark-gray butterfly found along the lower Colorado River and near the river along its tributaries in southeastern California, western Arizona, southern Nevada, and southern Utah (Scott, J. A. 1986. The butterflies of North America: a natural history and field guide. Stanford Univ. Press, Stanford, Calif.). Flights of H. graciliae occur from April to October in two (Emmel, T. C. & J. EF Emmel. 1973. The butterflies of southern California. Nat. His. Mus. Los Angeles Co., Sci. Series, 26: 1-148.) or three (Austin, G. T. & A. T. Austin. 1980. J. Res. Lepidoptera, 19: 1-63.) generations. Larvae of H. graciliae feed only on Atriplex lentiformis (Torrey) (Chenopodiaceae), a shrub found in dense clumps along lower Colorado River drainages (Emmel & Emmel 1973) that is halophytic and typically dioecious (Turner, R. M., J. E. Bowers & T. L. Burgess. 1995. Sonoran desert plants: an ecological atlas. Univ. Ariz. Press, Tucson.). Hesperopsis graciliae, however, is only sporadically encountered compared with the distribution of its larval host (Austin & Austin 1980). The apparent rarity of this butterfly, and concern over habitat destruction due to urban and agricultural development, has afforded H. graciliae the global rank of ‘G3?’ (L. Jaress, Ariz. Game & Fish Dept., Phoenix, personal communication), indicating that the conservation status of the species is uncertain but currently considered as rare or uncommon but not imperiled (Master, L. L. 1991. Conservation Biol., 5: 559-—563.). The rarity of H. graciliae compared with that of A. lentiformis suggests that factors other than larval-host availability influence its occurrence. One such factor may be the availability of a food source for H. graciliae adults. Hesperopsis graciliae adults must forage on other plant species, because A. lentiformis is wind- pollinated (Turner et al. 1995) and does not produce nectar, and foraging flights between plant species would be expected. However, H. graciliae adults are re- ported as seldom straying from the cover of their hostplants (Howe, W. H. [ed.]. 1975. The butterflies of North America. Doubleday & Co., Garden City, New York.). The present study describes and analyzes flights by H. graciliae between A. lentiformis and honey mesquite, Prosopis glandulosa Torrey (Fabaceae), a riparian plant of the lower Colorado River not used by H. graciliae larvae. The study site was located on the upper floodplain along the southern edge of the Bill Williams River 3 km east of Lake Havasu, a Colorado River reservoir, at the northern boundary of La Paz County, Arizona. The site lies within the Sonoran Desert biome (Turner, R. M. & D. E. Brown. 1982. Sonoran desertscrub. pp. 181-221. In Brown, D. E. [ed.]. Biotic communities of the American South- west—United States and Mexico. Desert Plants 4 [1—4].) at an elevation of 150 m. Precipitation (measured 19 km south near Parker, Ariz.) during the previous 12 mos (20 mm) had been 21% of the annual average (97 mm, Turner & Brown 1982). Principal vegetation at the site is A. lentiformis, Acacia greggii Gray, 1997 SCIENTIFIC NOTE 187 Pluchea sericea (Nutt.), Cercidium sp., P. glandulosa, and Salix gooddingii Ball. Hesperopsis graciliae flights were observed between a 14m X 5.5m X 3 m high closed-canopy patch of A. lentiformis male and female plants and a 16 m X 13 m X 6 m high closed-canopy patch of clumped P. glandulosa. The edges of the A. lentiformis and P. glandulosa canopies were 4.0 m apart and separated by a dirt and gravel road that traversed the site from east to west. The north edge of the road transected 4.6 m of the A. lentiformis canopy, and the south edge of the road transected 9.8 m of the P. glandulosa canopy. The P. glandulosa was not in flower. Hesperopsis graciliae flights between the plant canopies were observed, begin- ning at approximately 0930 MST, for 4 hrs on 17 and 20 Sep and for 2.5 hrs on 24 Sep 1996. A line of 5-cm diam. rocks 2 m apart was placed along the center of the road, and the time was recorded when a H. graciliae was observed flying across the line of rocks en route to either the A. lentiformis or P. glandulosa. Air temperature at the site was recorded hourly and averaged 30.8, 30.6, and 34.4° C on the three dates, respectively. Two H. graciliae adults (1 male and 1 female) were collected from the A. lentiformis on 26 September, verified as to species (G. Austin, Nev. St. Mus., Las Vegas, personal communication), and deposited as voucher specimens at the University of Arizona Insect Museum, Tucson. The number of H. graciliae flights to A. lentiformis and to P. glandulosa on each date was compared against an expected proportion of 1:1 using a x? test. The sequence of flight directions observed on each date was analyzed with a runs test (Sokal, R. R. & E J. Rohlf. 1981. Biometry [2nd ed]. W. H. Freeman & Co., New York.), a nonparametric method that uses the ¢,-distribution to test if an observed sequence of dichotomized events, in this case flight to A. lentiformis or to P. glandulosa, departs from random. Significant (P < 0.05), positive values of t, (©1.960) indicate flight directions that alternate in sequence, whereas negative values of t, (<—1.960) indicate sequences with each flight direction tending to repeat. In addition to performing runs tests on observed sequences of flight di- rection, runs tests were performed on sequences of flight direction after shifting the observed times of flight to A. lentiformis earlier by 1, 2, 3, 4, and 5 min. Shifting the time of flight allowed determining if flights to A. lentiformis were delayed behind those to P. glandulosa by a constant interval on each date. A total of 267 flights by H. graciliae was observed between the P. glandulosa and A. lentiformis canopies during the 10.5 hrs that the site was monitored (Table 1). Hesperopsis graciliae flew close to the ground, rarely exceeding a height of 20 cm, when traversing the open area between the plants and exhibited the flut- tering, bouncy pattern as has been previously described (MacNeill, C. D. 1970. Entomol. News, 81: 177-184, Ferris, C. D. & E M. Brown [eds.]. 1981. Butter- flies of the Rocky Mountain states. Univ. of Okla. Press, Norman.). Flights to P. glandulosa were slower and with more wandering than those to A. lentiformis, and therefore more easily observed. Although H. graciliae flying to A. lentiformis frequently landed near the periphery of the plant canopy, few adults flying to P. glandulosa similarly landed; most adults instead continued their flight into the interior of P. glandulosa’s comparatively less-dense canopy. The number of flights to A. lentiformis compared with those to P. glandulosa did not differ significantly (P > 0.05) within each date (Table 1). Across all three dates, the median time interval between successive flights to P. glandulosa was 188 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(3) Table 1. Frequency and interval of Hesperopsis graciliae flights between Prosopis glandulosa and Atriplex lentiformis. Interval between successive flights (min) Plant species No. of Date flown to flights x? Median Range 17 Sept. P. glandulosa 74 1.692 2.42 0.08—11.42 A. lentiformis 59 , 3.21 0.08-15.92 20 Sept. P. glandulosa 51 3.408 Dd: 0.08—18.17 A. lentiformis 34 i 4.50 0.08—21.50 24 Sept. P. glandulosa 29 1.65 3715 0.08—21.92 A. lentiformis 20 ; 4.17 0.17—28.00 4No. of flights compared; n.s. (1 df, P > 0.05). 2.83 min, and the median time interval between successive flights to A. lentiformis was 3.58 min. The median time interval between successive flights to either plant Species was 1.33 min across all three dates. The observed sequence of H. graciliae flight direction, to A. lentiformis or to P. glandulosa, did not depart from random on any of the dates sampled (Table 2). Alternating sequences in flight direction were detected after the observed times of flight to A. lentiformis were shifted earlier by 2 min on 17 September, by 4 min on 20 September, and by 1 min and 2 min on 24 September. Observed flights to A. lentiformis therefore were delayed behind observed flights to P. glandulosa by a constant interval on each date. The interval corresponding to the delay, however, varied between dates. Flights to A. lentiformis occurred approximately 3 min later than flights to P. glandulosa after averaging the delays (2, 4, and 1.5 min) across dates. A maximum of 6, 5, and 4 flights consecutively in the same direction was observed on the three dates, respectively. In contrast to previous descriptions of H. graciliae as a weak flyer preferring to remain within foliage (MacNeill 1970, Howe 1975), the present study found the species frequently (approx. once every 1.3 min) flying across a 4 m span of open terrain that separated larval-host and non-host plants. The frequent flights by H. graciliae between plant canopies, lack of significant difference between the number of flights to A. lentiformis compared with those to P. glandulosa, and sequences of alternating flight direction detected between the plant canopies, in- dicate that individual H. graciliae adults were repetitively flying back and forth between the two plant species. The maximum number of flights consecutively in Table 2. Values of ft, testing if sequences of Hesperopsis graciliae flight direction, to Prosopis glandulosa or to Atriplex lentiformis, deviate from random. Flight direction sequence after shifting times ' of flight to A. lentiformis earlier Observed flight Date direction sequence Shift of 1 min Shift of 2 min Shift of 3 min Shift of 4 min Shift of 5 min 17 Sept. 1.648 1.295 2.177" 0.414 1.472 0.061 20 Sept. 1.638 1.865 0.273 1.410 2.320° 0.955 24 Sept. 0.996 2.790° 2.192" 0.996 —0.798 0.398 4 Runs test for dichotomized data (Sokal & Rohlf 1981). > Alternating sequence in flight direction (df = », P < 0.05). ¢ Alternating sequence in flight direction (df = ~™, P < 0.02). 1997 SCIENTIFIC NOTE 189 the same direction (4-6 across dates) therefore provides an estimate of the insect’s minimum population density on the A. lentiformis, and the average 3-min delay between flights to and from the P. glandulosa represents the time spent by H. graciliae at that plant species. The benefit gained by H. graciliae repetitively flying from hostplants to P. glandulosa is not clear. For example, the insects may have been flying into P. glandulosa’s shaded interior for thermoregulation, the flights may have been as- sociated with mate finding and reproduction, or the insects may have been search- ing for additional hostplants. I suggest that the observed flights represent foraging behavior by H. graciliae, an hypothesis supported by the presence of extrafloral nectaries on P. glandulosa (Pemberton, R. W. 1988. Madrojfio, 35: 238—246.). I found one nectary located between each of the paired, primary leaflets and several nectaries along the rachis between the secondary leaflets (see Hickman, J. C. [ed.]. 1993. The Jepson manual: higher plants of California. Univ. of Calif. Press, Berkeley.). Leaf-rachis nectaries from the P. glandulosa examined under magni- fication were exuding nectar, and H. graciliae were observed landing on the sec- ondary leaflets. However, these observations were infrequent due to the difficulty of tracking the insects once they entered the P. glandulosa canopy, and the insect’s small size and tendency to land on interior foliage prevented observing if their proboscises were extended and in contact with leaf-rachis nectar. The hypothesis that H. graciliae were foraging at P. glandulosa extrafloral nectar is supported by the insect’s opportunistic feeding as an adult. They have been reported to feed at flowers of an exotic weed, Tamarix pentandra Pallas (= T. ramosissima) (Tamaricaceae), a crop (alfalfa), Medicago sativa L. (Fabaceae), and a native plant, Heliotropium curassavicum L. (Boraginaceae) (Austin & Aus- tin 1980). During the present study, I observed H. graciliae feeding at flowers of Bebbia juncea (Benth.) (Asteraceae) ~ 0.25 km from the study site and the only insect-pollinated plants in flower in the vicinity. Adult Lepidoptera will feed on extrafloral nectar, and extrafloral nectar has been suggested to provide nectar- feeding insects an alternative food source when other sources are rare (Rogers, C. E. 1985. Bull. Entomol. Soc. Amer., 31: 15—20.). Alternative food sources may be important to H. graciliae, because floral nectar would be less abundant during the insect’s fall flight. Additional effort is needed to determine if H. graciliae were foraging when repetitively flying to P. glandulosa or if the observed flights were due to other physiological demands, the significance of P. glandulosa ex- trafloral nectar as a food source to H. graciliae adults, and the influence of P. glandulosa on the insect’s distribution and abundance. Acknowledgment.—The author is grateful to the U.S. Fish and Wildlife Service for granting permission allowing the study to take place on their lands. W. D. Wiesenborn, U.S. Bureau of Reclamation, P.O. Box 61470, Boulder City, Nevada 89006-1470. Received 20 Nov 1996; Accepted 26 Feb 1997. PAN-PACIFIC ENTOMOLOGIST 73(3): 190-191, (1997) Scientific Note HOST RANGE EXPANSION OF HELCOSTIZUS RUFISCUTUM CUSHMAN (HYMENOPTERA: ICHNEUMONIDAE) TO PHORACANTHA SEMIPUNCTATA FABR. (COLEOPTERA: CERAMBYCIDAE) IN CALIFORNIA The eucalyptus longhorned borer, Phoracantha semipunctata Fabr., was first discovered in southern California in 1984, and during the last decade has deci- mated eucalyptus stands as it has spread throughout the southern counties, north- wards through the San Joaquin and Sacramento Valleys and up the California coast (Scriven, G. T., E. L. Reeves & R. E Luck. 1986. Calif. Agric., 40: 4-6; Paine, T. D., J. G. Millar & L. M. Hanks. 1995. Calif. Agric., 49: 34-37). Prior to this introduction, there were no North American Phoracantha species. As part of a biological control program aiming to limit damage caused by P. semipunc- tata, we have imported and introduced from Australia several species of braconid parasitoids of Phoracantha larvae (Paine et al. 1995). In particular, the parasitoid Syngaster lepidus Brullé showed promise of establishing at a site in Santa Barbara County, with several hundred wasps emerging from a single borer-infested tree a few months after release of 4,500 lab-reared wasps in summer 1994. On 11 Nov 1995, we monitored establishment of this parasitoid by looking under the bark of Eucalyptus globulus LaBillardiére logs for the symmetrically oblong cocoons at the ends of P. semipunctata feeding galleries, but instead discovered a consider- able number of very different, irregularly-shaped cocoons (one per borer gallery). Nine ichneumonid parasitoids of the species Helcostizus rufiscutum Cushman emerged from these cocoons in the laboratory (5 females and 4 males). Helcos- tizus rufiscutum is native to California and has apparently expanded its host range to become either a primary parasitoid of P. semipunctata or a secondary parasitoid of S. lepidus. Voucher specimens of H. rufiscutum have been placed in the En- tomology Teaching and Research Museum at the University of California, Riv- erside. Helcostizus rufiscutum belongs to the subfamily Cryptinae and tribe Hemitelini, other species of which are parasitic on cocoons of small insects in several insect groups, including Coleoptera and braconids (Townes, H. 1983. Mem. Am. En- tomol. Inst., 35: 1-281). That the natural hosts of H. rufiscutum are apparently wood-boring larvae or their parasitoids is indicated by emergence of adult para- sitoids from logs of the coniferous trees Monterey Cypress (Cupressus macro- carpa Gordon) and Douglas Fir (Pseudotsuga menziesii [Mirbel] Franco) as well the woody angiosperm poison oak (Toxicodendron diversilobum [Torrey & A. Gray] E. Green) (Townes 1983). It is clear that P. semipunctata was the host in Santa Barbara because S. lepidus was too rare at the site to account for the abundance of AH. rufiscutum. For example, an intensive search of the study site on 11 Nov 1995 yielded only a single empty S. Jepidus cocoon among hundreds of borer larvae. On 6 Aug 1996, we saw no evidence of S. lepidus, but counted 47 H. rufiscutum cocoons and 11 borer larvae (a parasitism rate of 81%). More- 1997 SCIENTIFIC NOTE 191 over, we have tested the ability of H. rufiscutum to develop on P. semipunctata larvae in the laboratory; caged adult females readily oviposited on borer larvae through the bark of eucalyptus logs, and adult parasitoids emerged about 1 month later. Although all the parasitoids that emerged were males, this sex bias was likely due to either inhibition of mating behaviors in lab cages or to the size of host larvae that were provided. Field-collected cocoons yielded both sexes. Host range expansion by H. rufiscutum is probably the result of a combination of factors. First, adult H. rufiscutum may search for a potentially broad range of borer hosts in a variety of habitats, as indicated by its being collected from a diversity of woody plant types. Second, location of the cryptic host insects is apparently mediated primarily by acoustic or vibrational cues that are detected by organs in the swollen front tibiae of females (J. Luhman, pers. comm.), and these cues may be similar for different borer species. Third, natural hosts and habitats were in close proximity to eucalyptus trees and slash infested with P. semipunc- tata larvae; all of the woody plant species from which this parasitoid has been reared (Monterey Cypress, Douglas fir, Poison Oak) occur in Santa Barbara Coun- ty (Hickman, J. C. 1993. The Jepson Manual. Univ. of California Press). However, it is not known which species of wood borers are the native hosts. One possibility is Xylotrechus nauticus (Mannerheim) which commonly infests a wide variety of woody hosts (Linsley, E. G. 1964. Univ. Calif. Pub. Entomol., 22: 1-197). Xy- lotrechus nauticus larvae also feed in eucalyptus logs (Solomon, J. D. 1995. U.S.D.A. For. Serv. AH-706), and may occur in the same host logs as P. semi- punctata (LMH, pers. obs.), suggesting a simple behavioral mechanism leading to parasitism of this introduced borer. Feeding galleries of X. nauticus may be distinguished from those of later instar P. semipunctata by their narrower breadth. In Santa Barbara, H. rufiscutum apparently parasitizes a significant proportion of P. semipunctata larvae and so may play an important role in reducing borer populations. This parasitoid may also attack a congeneric borer species, P. re- curva, which has been recently discovered in the state, but only in Orange, Riv- erside and San Bernardino Counties to date (C. Campbell & L. Hanks, unpub- lished data). Because H. rufiscutum is a native parasite and is broadly distributed in California (Townes 1983), fortuitous host range expansion may provide addi- tional mortality of Phoracantha throughout the state. Acknowled gment.—We thank Dr. John Luhman, Minnesota Department of Ag- riculture, for identifying the parasitoid specimens and reviewing the manuscript. Lawrence M. Hanks, Department of Entomology, University of Illinois, 320 Morrill Hall, 505 South Goodwin Ave., Urbana, IL 61801. Christopher Campbell, Timothy D. Paine, and Jocelyn G. Millar, Department of Entomology, University of California, Riverside, CA 92521. Received 17 Dec 1996; Accepted 26 Feb 1997. PAN-PACIFIC ENTOMOLOGIST 73(3): 192-196, (1997) PROCEEDINGS OF THE PACIFIC COAST ENTOMOLOGICAL SOCIETY, 1996 FIVE HUNDRED TWENTY-THIRD MEETING The 523rd meeting of the Pacific Coast Entomological Society was held on 19 Jan 1996 at 8:00 PM in the Morrison Auditorium of the California Academy of Sciences in Golden Gate Park, San Francisco with President-Elect Warren E. Savary presiding. Ms. Barbara Deutsch presented a note on the migration of Monarch butterflies in North America and Mr. Curtis Y. Takahashi of the California Department of Food and Agriculture announced seasonal employment opportunities with the County of San Mateo and State of California Department of Food and Agriculture. The featured speaker, Dr. Norman D. Penny of the California Academy of Sciences presented a slide lecture entitled ‘““The Neuropterists’ Contribution to the Costa Rican All Biotic Inventory’’. Dr. Penny discussed the tremendous diversity and species counts of Neuroptera that have been inventoried in over five years of extensive collecting in Costa Rica. The inventory reveals that Costa Rica has disproportionately high numbers of species diversity per area when compared with other areas outside of the country. The meeting was concluded at 9:05 PM and was followed by a social hour in the Department of Entomology Conference Room. The following 39 persons were present: (30 members) PH. Amaud Jr., C.B. Barr, T.S. Briggs, R.M. Brown, R.L. Cable, H.K. Court, C.K. Griswald, A. Hom, A.S. Hunter, D.H. Kavanaugh, R.L. Langston, V.E Lee, J.E Parinas, D.R. Parks, A.M.L. Penny, D.A. Piechnik, K.A. Reynolds, J.M. Ribardo, K.J. Ribardo, D.C. Russel, L.S. Saul-Gershenz, W.E. Savary, K.A. Schick, N.M. Schiff, J. Schweikert, M. Sharp, C.Y. Takahashi, D. Ubick, and S.E. Vaughn; (9 guests) G.R. Almany, M.M. Amaud, J.E. Court, M. Hurley, T. Pape, W.E. Rauscher, J. Schick, S. Smelter, and 1 illegible signature. FIVE HUNDRED TWENTY-FOURTH MEETING The 524th meeting of the Pacific Coast Entomological Society was held on 16 Feb 1996 at 8:00 PM in the Morrison Auditorium of the California Academy of Sciences in Golden Gate Park, San Francisco with President-Elect Warren E. Savary presiding. Mr. Vincent FE Lee of the California Academy of Sciences announced that the first issue of the Pan- Pacific Entomologist was published on 12 Feb 1996 and is back on schedule. Mr. Lee also announced that the Pacific Branch of the American Association for the Advancement of Science will be held at San Jose State University on June 23 through 27. Dr. Charles Griswald of the California Academy of Sciences announced that the International Con- gress of Arachnology will meet in July of 1998 in Chicago, Ilinois. Mr. Warren E. Savary of the Department of Agriculture announced that workshops on the future of San Francisco’s Glen Canyon Park is requesting participation from the Pacific Coast Entomological Society, and Ms. Theresa Meickle presented a slide note illustrating the aesthetics of macro photo- graphed sawfly frass. The featured speaker, Dr. Nathan Schiff of the US Department of Agriculture-ARS, Albany pre- sented a slide lecture entitled ‘““Sawfly Biology; Dabblings of a Naturalist Turned Entomologist’. Dr. Schiff provided entertaining insights into the phylogeny of sawflies as well as detailing the biology of the only true phytophagous Hymenoptera. The meeting was adjourned at 9:35 PM and was followed by a social hour in the Department of Entomology Conference Room. The following 37 persons were present: (29 members) PH. Amaud Jr., C.B. Barr, T.S. Briggs, R.M. Brown, R.L. Cable, H.K. Court, C.K. Griswald, A. Hom, A.S. Hunter, D.H. Kavanaugh, R.L. Langston, V.E Lee, J.-E Parinas, D.R. Parks, A.M.L. Penny, D.A. Piechnik, K.A. Reynolds, J.M. Ribardo, K.J. Ribardo, L.S. Saul-Gershenz, W.E. Savary, K.A. Schick, N.M. Schiff, J. Schweikert, M. Sharp, C.Y. Takahashi, D. Ubick, and S.E. Vaughn; (8 guests) G.R. Almany, M.M. Amaud, J.E. Court, M. Hurley, T. Pape, W.E. Rauscher, and J. Schick. 1997 PROCEEDINGS 193 THE FIVE HUNDRED TWENTY-FIFTH MEETING The 525th meeting of the Pacific Coast Entomological Society was held on 15 Mar 1996 at 8:00 PM in the Morrison Auditorium of the California Academy of Sciences in Golden Gate Park, San Francisco with President-Elect Warren E. Savary presiding. Dr. Norman D. Penny of the California Academy of Sciences Announced that the auditor of the Society, H. Vannoy Davis found all business matters in order for 1995. Dr. Penny also announced that the Insect Fair at the Los Angeles Arboretum will be held on 18 May 1996. This is the largest insect fair in the United States. Mr. Curtis Y. Takahashi of the California Department of Agriculture announced that the Insect Fair at Sanborn Park in Santa Clara County will be held on 18 May 1996 and Ms. Leslie Saul-Gershenz of the San Francisco Insect Zoo announced that a training workshop entitled “‘“Managing and Working in Insectariums’”’ will be held on the same day in Denver, Colorado. Dr. Nathan Schiff of the Department of Agriculture-ARS, exhibited a living specimen of Timema californica. The featured speaker, Mr. Norman E. Gershenz of the Center for Ecosystems Survival, presented an engaging slide lecture entitled ““Conservation Strategies for the 21st Century’’. Mr. Gershenz de- scribed how conservation efforts have historically concentrated on a single species and how current strategies have evolved to include systemic habitat conservation. With this in situ approach, Mr. Gershenz outlined the socio-economic factors that necessitate a conservation partnership in country with residents forming a more effective conservation effort. The meeting was adjourned at 9:05 PM and was followed by a social hour in the Department of Entomology Conference Room. The following 30 persons were present: (25 members) PH. Amaud Jr., L.G. Bezark, T.S. Briggs, S.V. Fend, C.E. Griswold, W. Hamersky, A. Hom, J. Honda, R.L. Langston, D.L. Mead, T. Meikle, J.E Parinas, D.R. Parks, A.M.L. Penny, N.D. Penny, S. Renkes, R.G. Robertson, E.S. Ross, W.E. Savary, N.M. Schiff, J. Schweikert, EA.H. Sperling, C.Y. Takahashi, D. Ubick, and S.E. Vaughn; (5 guests) M.M. Amaud, B. Landry, K. Norton, W.E. Rauscher, and D.T. Wyatt. FIVE HUNDRED TWENTY-SIXTH MEETING The 526th meeting of the Pacific Coast Entomological Society was held on 19 Apr 1996 at 8:00 PM in the Morrison Auditorium of the California Academy of Sciences in Golden Gate Park, San Francisco with President Wojciek J. Pulawski presiding. Mr. Vincent F Lee of the California Academy of Sciences announced the availability of The Insect Study Source Book published by the Young Entomologists’ Society. The Source book catalogs in- structional videos and other entomological listings. Ms. Jeanette McNicol of San Jose State University announced that the annual Arroy Seco overnight collection trip sponsored by the SJSU Entomology Club will be held on 27 Apr 1996. The featured speaker, Dr. David H. Kavanaugh of the California Academy of Sciences presented a slide lecture entitled ‘“‘In Search of Old World Relatives of Nearctic Carabid Beetles in Central Siberia, Lake Baikal Region’’. Dr. Kavanaugh detailed the logistical difficulties and the cultural experiences encountered while traveling to the Lake Baikal region and surrounding mountain ranges as well as demonstrating the phylogenetic similarities of carabids found in Siberia with those found in Western Europe and North America. The meeting was adjourned at 9:45 PM and was followed by a social hour in the Department of Entomology Conference Room. The following 30 persons were present: (25 members) PH. Amaud Jr., L.G. Bezark, TS. Briggs, S.V. Fend, C.E. Griswold, W. Hamersky, A. Hom, J. Honda, R.L. Langston, D.L. Mead, T. Meikle, J.E Parinas, D.R. Parks, A.M.L. Penny, N.D. Penny, S. Renkes, R.G. Robertson, E.S. Ross, W.E. Savary, N.M. Schiff, J. Schweikert, EA.H. Sperling, C.Y. Takahashi, D. Ubick, and S.E. Vaughn; (5 guests) M.M. Amaud, B. Landry, K. Norton, W.E. Rauscher, and D.T. Wyatt. FIVE HUNDRED TWENTY-SEVENTH MEETING The 527th meeting of the Pacific Coast Entomological Society was held on 17 May 1996 at 8:00 PM in the Morrison Auditorium of the California Academy of Sciences in Golden Gate Park, San Francisco with President Wojciek J. Pulawski presiding. Mr. Patrick Craig presented a slide of an unidentified arthropod, encased in Lebanese amber for identification. After much speculation, inconclusive identification resulted. 194 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(3) The featured speaker, Dr. Charles Griswald of the Califomia Academy of Sciences presented a slide lecture entitled ““Spider Webs Through Space and Time’’. Dr. Griswald presented an overview of spider biology involving silk production and web formation as a means of determining phylogenetic relationships of arachnids. The convergence hypothesis explained by Dr. Griswald differs from pre- vious systematic theory in that the webs of spiders are viewed as behavioral extensions rather than architectural or structural compositions. The meeting was adjourned at 9:25 PM and was followed by a social hour in the Department of Entomology Conference Room. The following 46 persons were present: (34 members) R.A. Aalbu, PH. Amaud Jr., C.B. Barr, L.G. Bezark, J. Brandriff, TS. Briggs, K.W. Brown, R.M. Brown, H.K. Court, PR. Craig, B. Deutsch, C.E. Griswald, R.E. Hill, A.S. Hunter, B. Keh, C.Y. Kitayama, R.L. Langston, V.E Lee, D.L. Mead, T. Meikle, W.J. Pulawsk, A.E. Rackett, K.M. Reynolds, R.G. Robertson, W.E. Savary, K.N. Schick, N.M. Schiff, J. Sweikert, H.I. Scudder, E.L. Smith, FA.H. Sperling, D. Ubick, S.E. Vaughn, and D.T. Wyatt; (12 guests) D. Aalbu, V. Ahrens-Pulawski, M.M. Amaud, J. Baker, I. Brown, J.E. Court, D. Kitayama, K. Kitayama, R. Kitayama, K. Norton, W.E. Rauscher, and S. Stone. FIVE HUNDRED TWENTY-EIGHTH MEETING The 528th meeting of the Pacific Coast Entomological Society was held on 20 Sep 1996 at 8:00 PM in the Goethe Room of the California Academy of Sciences in Golden Gate Park, San Francisco with President Wojciek J. Pulawski presiding. Dr. Nathan Schiff of the Department of Agriculture-ARS presented an exhibit of a sawfly (Euro- cerus), a buprestid (Melanophila), and a cerambicid that were collected while posing as a Department of Forestry firefighter in Northern California and Mr. Ron Robinson presented a slide note of noctuid moths collected “‘in photo” during the summer. The featured speaker, Dr. Thomas Briggs, Research Associate at the California Academy of Sciences presented a slide lecture entitled ‘““The New Melones Cave Invertebrate Transplant’. Dr. Briggs chron- icled the details of the search, selection, and transplantation of invertebrates prior to the construction of the New Melones dam and subsequent flooding of the area. The New Melones Reservoir now covers the habitat from which the invertebrates transfer took place. The meeting was adjourned at 9:15 and concluded with a social hour held in the Department of Entomology Conference Room. The following 46 persons were present: (34 members) R.A. Aalbu, PH. Amaud Jr., C.B. Barr, L.G. Bezark, J. Brandriff, TS. Briggs, K.W. Brown, R.M. Brown, H.K. Court, PR. Craig, B. Deutsch, C.E. Griswald, R.E. Hill, A.S. Hunter, B. Keh, C.Y. Kitayama, R.L. Langston, V.E Lee, D.L. Mead, T: Meikle, W.J. Pulawsk, A.E. Rackett, K.M. Reynolds, R.G. Robertson, W.E. Savary, K.N. Schick, N.M. Schiff, J. Sweikert, H.I. Scudder, E.L. Smith, EA.H. Sperling, D. Ubick, S.E. Vaughn, and D.T. Wyatt; (12 guests) D. Aalbu, V. Ahrens-Pulawski, M.M. Amaud, J. Baker, I. Brown, J.E. Court, D. Kitayama, K. Kitayama, R. Kitayama, K. Norton, W.E. Rauscher, and S. Stone. FIVE HUNDRED TWENTY-NINTH MEETING The 529th meeting of the Pacific Coast Entomological Society was held on 18 Oct 1996 at 8:00 PM in the Morrison Auditorium of the California Academy of Sciences in Golden Gate Park, San Francisco with President Wojciek J. Pulawski presiding. Ms. Cheryl Barr of the University of Califomia, Berkeley announced that she has a companion ticket available for half price to the Entomological Society of America National Meeting to be held in Louisville, Kentucky. Mr. Vincent F Lee of the California Academy of Sciences announced that the books ‘Australian Orthpteroids” and “Insect Musicians and Cricket Champions’’ will be on display during the social hour following the meeting. Dr. Nathan Schiff of the US Department of Agriculture-ARS presented a larval specimen of Nu- diprion collected on white fir in the Yuba Pass of California, and Dr. Kirby Brown displayed a basket decorated with images of butterflies obtained from the Pitt River region of northeastern California. The featured speaker, Dr. Bernard Landry of the University of California, Berkeley presented a slide lecture entitled ‘““The Lepidoptera of the Galapagos Islands’’. In highlighting the zoogeographical history of the islands, Dr. Landry described probable patterns of insect colonization and described the low diversity of fauna encountered while collecting. The meeting was adjourned at 9:15 PM and was followed by a social hour held in the Department of Entomology Conference Room. 1997 PROCEEDINGS 195 The following 56 persons were present: (41 members) R.A. Aalbu, PH. Amaud Jr., C.B. Barr, TS. Briggs, K.W. Brown, R.M. Brown, M.S. Caterino, L.W. Currie Jr., B. Deutsch, W.A. Doolin, E.M. Fisher, C.E. Griswald, W. Hammersky, L.A. Irons, M.A. Isaak, B. Landry, R.L. Langston, FEF Lee, D.L. Mead, T:.C. Meikle, M.H. Niehoff, L.A. Norton, S.T. O’ Keefe, J.E Parinas, A.M.L. Penny, N.D. Penny, J.-A. Powell, W.J. Pulawski, A.E. Rackett, J.L. Rasgon, W.E. Savary, N.M. Schiff, J.S. Schwei- kert, H.I. Scudder, FA.H. Sperling, R.E. Stecker, CY. Takahashi, D. Ubick, S.E. Vaughn, J.D. Wells, and R.L. Zuparkko; (15 guests) V. Ahrens-Pulawski, M.M. Arnaud, S.A. Brown, S.M. Covarrubias, B. Deutsch, D.D. Giuliani, D.E. Kain, J.J. Kruse, J. Myatt, S.G. Nguyen, R.D. Reed, A. Schedlock, R.M. Shelly, L.A. Swenson, and L. Volpe. FIVE HUNDRED THIRTIETH MEETING The 530th meeting of the Pacific Coast Entomological Society was held on 15 Nov 1996 at 8:00 PM in the Morrison Auditorium of the California Academy of Sciences in Golden Gate Park, San Francisco with President Wojciek J. Pulawski presiding. Dr. Pulawski announced the appointments of Dr. Paul H. Arnaud Jr., Dr. Norman D. Penny, and Ms. Cheryl B. Barr to the Nominations Committee and Mr. H. Vannoy Davis, Ms. Helen Court, and Mr. Jere Schweikert to the Audit Committee. Dr. Edward L. Smith presented a detailed note on the evolutionary modifications of cerci in some Apteragota, Ephemeroptera, and Odonata and Mr. Ron Robinson presented a slide note capturing wood-boring craneflies in copula. The sequence of slides show the female dragging the engaged male into a bored hole of a dead Bay tree. The featured speaker, Dr. Wojciek J. Pulawski of the California Academy of Sciences presented a slide lecture entitled ‘““The Biological Evolution of Hymenoptera’’. Dr. Pulawski discussed the struc- tural and behavioral development of Hymenoptera from the most basal vegetarian Symphyta up through to the origin of sociality of the Apocrita. The development of the acuneates was illustrated especially well by Dr. Pulawski’s slides. The meeting was adjourned at 10:15 PM and was followed by a social hour held in the Department of Entomology Conference Room. The following 37 persons were present: (27 members) R.A. Aalbu, C.B. Barr, L.G. Bezark, K.W. Brown, A.I. Cognato, H.K. Court, C.E. Griswald, W. Hamersky, A. Hom, R.L. Langston, V.F Lee, T.C. Meikle, D.R. Parks, A.M.L. Penny, N.D. Penny, J.A. Powell, W.J. Pulawski, A.E. Rackett, J.L. Rasgon, K.M. Reynolds, K.J. Ribardo, J.S. Schweikert, E.L. Smith, D. Ubick, S.E. Vaughn, J.D. Wells, and R.L. Zuparko; (10 guests) V. Ahrens-Pulawski, TJ. Bernot, D. Bunn, J.E. Court, C.D. Dailey, B. Kepner, D. Povolny, D. Ross, V. Van Wly, and S. Wilson. FIVE HUNDRED THIRTY-FIRST MEETING The 531st meeting of the Pacific Coast Entomological Society was held on 13 Dec 1996 at 8:00 PM in the Blakeslee Conference Room of San Francisco State University, San Francisco with President Warren E. Savary presiding. Dr. Paul H. Arnaud Jr. of the California Academy of Sciences announced the acquisition, by the Academy, of specimens from the estate of Dr. Donald Denning. Dr. Arnaud also reported on behalf of the Nominations Committee. The following members were voted on and approved as officers for 1997: Mr. Warren Savary, President Dr. William Shepherd, President-Elect Ms. Julieta F Parinas, Treasurer Mr. Vincent E Lee, Managing Secretary Mr. Stanley E. Vaughn, Recording Secretary Mr. Vincent EF Lee of the California Academy of Sciences reported that membership totaled 372 and Dr. Susan B. Opp of California State University, Hayward reported that all issues of the Pan- Pacific Entomologist were published on schedule in 1996. Ms. Leslie Saul-Gershenz of the San Francisco Insect Zoo announced that the Entomological Society of America has a page on the World Wide Web as well as an entomology site by the Iowa State Teachers’ Association. Ms. Saul-Gershenz also announced the completion of Dynastinae in the All Taxa Survey of Costa Rica. 196 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(3) Mr. Curtis Y. Takahashi of the California Department of Food and Agriculture presented, as an exhibit, some towels designed with images of insects and flowets. The featured speaker, Dr. John E. Hafernik of San Francisco State University presented a slide lecture entitled ‘“The Re-introduction of the Damselfly Jshnera gemina in Glen Canyon’’. Dr. Hafernik described the natural history of Glen Canyon and detailed past and present biological conservation techniques in reintroducing the damsilfly to the area. Initially, during re-introduction, populations of Ishnera exhibited increased mortality. Currently, however the emergence of new cohorts is approaching previous recapture levels, indicating the possibility of a reestablished population. The meeting was adjourned at 9:45 PM. The following 30 persons were present: (17 members) M.M. Amaud, PH. Amaud Jr., B. Deutsch, J.G. Edwards, N.E. Gershenz, J.E. Hafernik, B. Landry, V.E Lee, S.B. Opp, J.L. Rasgon, K.M. Rey- nolda, L. Saul-Gershenz, W.E. Savary, N.A. Schiff, C.Y. Takahashi, S.E. Vaughn, and J.D. Wells; (13 guests) M. Brewer, J. Conner, R.A. Cruib, T. Dockery, J. Gulbranser, G. Hannon, A. LeMon, S. LeMon, D.A. Piechnik, W.A. Rauscher. C. Risden, R. Rozman, and E. Valdivia. PAN-PACIFIC ENTOMOLOGIST 73(3): 197-198, (1997) PACIFIC COAST ENTOMOLOGICAL SOCIETY NOTES TO THE FINANCIAL STATEMENTS YEAR ENDED SEPTEMBER 30, 1996 SUMMARY OF SIGNIFICANT ACCOUNTING POLICIES Accounting Method Income and expenses are recorded by using the cash basis of accounting. Note from the Treasurer The Pan-Pacific Entomologist, the journal of the Pacific Coast Entomological Society, is published quarterly. However, due to editorial delays, the issues are often not published and charged to the Society on schedule. This explains the abnormal fluctuation in publishing costs. Capital Expenditures Annual capital expenditures of $5,000 or less are charged to expense. Marketable Securities American Telephone & Telegraph Co., Pacific Telesis Group and Air Touch Com- munications common stocks are carried at market value. Increases and decreases in value are reflected in income. As of Sept. 30, 1996 AT & T Corp. issued one share of Lucent Technologies Inc. capital stock for every 3 shares of AT & T Corp. shares owned plus cash for fractional shares. This distribution of 25 shares of Lucent Technologies Inc. is reflected in these statements as an increase in value of captial stock owned of $1147. Income Tax The Society is exempt from Federal Income and California franchise tax. As Chairman of the Accounting and Tax Committee, and in accordance with the Society’s bylaws, I have reviewed the financial records of the Society but have not made an audit of them. During the course of this review, nothing was noted which indicated any material inaccuracy in the financial statements. H. Vannoy Davis Chairman of the Accounting and Tax Committee 198 THE PAN-PACIFIC ENTOMOLOGIST PACIFIC COAST ENTOMOLOGICAL SOCIETY STATEMENT OF INCOME, EXPENDITURES AND CHANGES IN FUND BALANCES YEARS ENDED SEPTEMBER 30, 1996 AND 1995 Income Dues and subscriptions Interest Total Income ........................ Expenditures Publication costs—Pan-Pacific Entomologist RRepriatt COStS. inh. ccna s Aner needy 24 Postage, newsletter and miscellaneous expenses Cash in bank Commercial account ................ Special Funds: General Fund—Wells Fargo Bank C. P. Alexander Fund—Capital Preservation Fund Fall Memoir Fund—Wells Fargo Bank Total cash in bank and special funds Capital Stock (at market value) Reprints. and miscellaneous. oc. soe ne ehh tne ele ele eek oe ewte wt gee ngs wees Increase (Decrease) in value of capital stock: American Telephone & Telegraph Company Pacific Telesis Group and Air Touch Communications Lucent Technologies (see note) STATEMENT OF ASSETS AS OF SEPTEMBER 30, 1996 AND 1995 American Telephone & Telegraph Co., 80 shs. .......................0 200s Lucent. Technologies, 25- SHS. 0220.52 ferent Merit Dn ccacedgare tan ate a datas Pacing. Telesis: Gramps 264 Sissi Pe ced ea take sree abe are eae Pet ae toate Air Touch Communications, 264 ShS. ......... 0. c cece ccc cece cece ee eeeeees Total Assets 1996 $ 17,808 15,681 3,815 700 (1,080) (33) 1,147 $ 38,038 $ 30,478 3,098 1,215 $ 34,791 $ 3,247 133,258 $136,505 1996 $ 17,475 5,060 57,413 35,060 $115,008 4,180 1,147 8,877 W299 21,497 $136,505 Vol. 73(3) 1995 $ 17,066 10,421 3,730 681 940 528 $ 33,366 $ 25,590 2,614 1,274 $ 29,478 $ 3,888 129,370 $133,258 1995 $ 17,914 4,942 54,711 34,228 $111,795 5,260 8,118 8,085 21,463 $133,258 PAN-PACIFIC ENTOMOLOGIST 73(3): 199, (1997) 1996 SPONSORING MEMBERS OF THE PACIFIC COAST ENTOMOLOGICAL SOCIETY Ernest Anderson Richard L. Penrose Paula & Robert Buickerood Albert E. Rackett Bryan K. Eya Norman E. Gershenz & Leslie S. Saul E. Eric Grissell Warren E. Savary Teresa C. Meikle & Charles E. Harvey I. Scudder Griswold Frank E. Skinner John E. Hafernik Jr. Edward L. Smith Frank T. Hovore Thomas J. Zavortink Calvert E. Norland Harry W. Oswald PAN-PACIFIC ENTOMOLOGIST 73(3): 200, (1997) Editorial Policy Change Beginning with manuscripts submitted after 1 January 1998, The Pan-Pacific Entomologist will require that voucher specimens for all articles be deposited in a properly maintained collection accessible to other scientists. This policy repre- sents a departure in tradition for non-taxonomic studies that is necessitated by the rapid discovery of numerous cryptic species and species complexes. These situ- ations make it imperative that future scientists be able to confirm exactly what Species was studied in past papers. The uncertainty surrounding whether older studies used Bemisia tabaci (Gennadius) or Bemisia argentifolii (Bellows & Per- ring) is a good example of a situation that could have been avoided if voucher specimens were available. Similar confusion surrounds older studies of Bactro- cera dorsalis (Hendel) from numerous locations, several Rhagoletis species and numerous aphids. The location at which the voucher specimen(s) have been de- posited and the coding necessary to access them shall be noted in the Materials and Methods section of the article or the text of the scientific note. PAN-PACIFIC ENTOMOLOGIST Information for Contributors See volume 66(1): 1-8, January 1990, 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 11 in, nonerasable, high quality paper. THREE (3) COPIES of each manuscript must be submitted, EACH INCLUDING REDUCTIONS OF ANY FIGURES TO THE 85 xX 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 corner. 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 KEY WORDS. 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. E 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. FE Eastop. 1987. Problems in pest aphid taxonomy: can chromosomes plus morphometrics 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 (5S) 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. THE PAN-PACIFIC ENTOMOLOGIST Volume 73 JULY 1997 Number 3 Contents BARTHELL, J. F, T. L. GRISWOLD, G. W. FRANKIE, & R. W. THORP—Osmia (Hyme- noptera: Megachilidae) diversity at a site in central coastal California _______- 141 FITZGERALD, S.—A new species of Enicoscolus (Diptera: Bibionidae) from Brazil, with additional distributionercconds fat the Scmuk _ pee vee ee 8k Re 152 JOHNSON, P. J.—New species of Dioxypterus Fairmaire from Tonga and Fiji, with new dis- tribution records, a tribal reassignment, and key to the species of the region (Coleoptera: Elateridac) =a ook See Bs De ee eA Be 156 GORDON, R. D. & R. W. RUST—A new southern Nevada species of Aegialia (Aegialia) (Coleoptera: Séanaiaeidae: Aphid nn acy 8 eet a ee 168 CONWAY, J. R.—Foraging activity, trails, food sources and predators of Formica obscuripes Forel (Hymenoptera: Formicidae) at high altitude in Colorado Le SCIENTIFIC NOTES McKILLUB, S. C. & R. V. McKILLUP—An outbreak of the moth Achaea serva (Fabr.) on the MAN SLOVErEAROCOOIALa pe LeCME AL ye Bene ee, 184 WIESENBORN, W. D.—Hesperopsis graciliae (MacNeill) (Lepidoptera: Hesperiidae) flight between hostplants and Prosopis elandulpsa Teimey 28) 186 HANKS, L. M., C. CAMPBELL, T. D. PAINE, & J. G. MILLAR—Host range expansion of Helcostizus rufiscutum Cushman (Hymenoptera: Ichneumonidae) to Phoracantha semi- punctate Fabr. (Coleoptera: Cerambycidae) im California, 190 Pacific Coast Entomological Saciety, Procecdimesmor 996. 0 192 Pacific Coast Entomological Society, financial statements for 1995, 1996 __ 197 Pacific Coast Entomological Society, Sponsoring Members 1996 _ Boe The Pan-Pacific Entomologist Editorial policy change -_ 2 200 The PAN-PACIFIC ENTOMOLOGIST Volume 73 October 1997 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. V. Dowell, Editor R. M. Bohart R. L. Penrose, Associate Editor J. T. Doyen R. E. Somerby, Book Review Editor J. E. Hafernik, Jr. Julieta E Parinas, Treasurer Warren E. Savary Published quarterly in January, April, July, and October with Society Proceed- ings usually appearing in the October issue. All communications regarding non- receipt of numbers should be addressed to: Vincent EK Lee, Managing Secretary; and financial communications should be addressed to: Julieta F Parinas, Treasurer; at: Pacific Coast Entomological Society, Dept. of Entomology, California Acad- emy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599. Application for membership in the Society and changes of address should be addressed to: William Hamersky, Membership Committee chair, Pacific Coast Entomological Society, Dept. of Entomology, California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599. Manuscripts, proofs, and all correspondence concerning editorial matters (but not aspects of publication charges or costs) should be sent to: Dr. Robert V. Dowell, Editor, Pan-Pacific Entomologist, California Dept. of Food & Agriculture, 1220 N St., Sacramento, CA 95814. See the back cover for Information-to-Con- tributors, and volume 73(4): 248—255, October 1997, for more detailed informa- tion. Information on format for taxonomic manuscripts 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. Pacific Coast Entomological Society OFFICERS FOR 1997 Warren E. Savary, President Vincent E Lee, Managing Secretary Julieta E Parinas, Treasurer Stanley E. Vaughn, Recording Secretary 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 2 October 1997 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 73(4): 201-203, (1997) Obituary: Richard F. Wilkey (1925-1995) DOUGLASS R. MILLER! AND JOHN A. DAVIDSON? 'Systematic Entomology Lab, ARS, USDA, Beltsville, MD 20705 ?Department of Entomology, University of Maryland, College Park, MD 20742 Richard F. Wilkey in 1993. Richard (Dick) E Wilkey was a talented and innovative preparator of insect and mite specimens on microscope slides. The more than 1,000,000 specimens that he prepared during his career are highly prized by researchers and identifiers alike because they are the best for studying the minute structures that are so important in diagnosing species. Richard also was dedicated to 4-H (a training program for children and young adults) and had a major positive impact on the more than 400 students that he taught about insects over the 40 years that he served as part of the 4-H program. Richard Wilkey died 29 Oct 1995 in Bluffton, Indiana. He was born 14 Aug 1925 in Providence, Rhode Island, to Frank K. and Laura A. Plummer Wilkey. He made his first insect collection when he was six, creating an interest in things entomological that lasted a lifetime. He moved to Indianapolis, Indiana, when he was 12 and following high school graduation served three years during World War II with U.S. Armed Forces in the Pacific Campaign. Following the war he married Dorothy Weber and moved to Lafayette, Indiana, where he earned a B.S. degree in entomology at Purdue University in 1950. He received his M.S. degree in entomology at Colorado State University in 1951. During this period he developed a life-long interest in micro- 202 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(4) scopic organisms and their preparation. While at Purdue he did an undergraduate project on Collembolla. His Master’s thesis continued in this vein as an analysis of the springtails of Larmier County, Colorado. When in later years his focus changed to groups other than springtails, he turned over his collection, including specimens of nearly 50 undescribed species and a dozen new genera, to Kenneth Christiansen, Grinnell College, Grinnell, Iowa. Wilkey began his professional career in 1951 with the Mexican Cotton Com- pany in Baja California, Mexico, working on pests of cotton. Later that year he was hired by the Santa Clara County, County Agricultural Commissioner’s Office where he worked for a year. He then was promoted and moved to San Diego where he did quarantine identifications for three years for the State of California, Department of Agriculture. In Sacramento he finally obtained the job he wanted most: taxonomist responsible for Homoptera identification (except aphids) also working for the State of California. In this position he became recognized world wide for his knowledge of scale insect systematics. Wilkey often helped colleagues in other states such as Florida, Washington, and Arizona with difficult identifications. Following the death of Howard Mc- Kenzie he served as the point person for visitors working with the ‘“‘Ferris- McKenzie”’ scale insect collection at the University of California at Davis. In 1971, after a distinguished career with the California Department of Agriculture, he retired early and returned to Bluffton, Indiana. Here Wilkey began a new career. He started Arthropod Slidemounts, a mail order business that supplied arthropods to clientele needing a diverse array of well prepared specimens. The primary clientele were high schools and colleges, but he frequently provided specimens at a nominal fee for Extension short courses, pest control operator training, the food industry, and 4-H programs. During this period he developed a set of tools for micro-manipulation of specimens under the dissecting microscope, which most users believe are the best of their kind. After another 20 year career, in 1991 he retired and sold his business to BioQuip Products. Richard E Wilkey was an energetic, enthusiastic, and caring human being who was most comfortable working behind the scenes. Within entomology it is difficult to pick one area among his many accomplishments as the most important, and when consideration is given to his commitment to entomology in 4-H, and his leadership in other community activities the choice becomes even more difficult. We suspect that after all is said and done, the nearly 1,000,000 perfect, or nearly perfect, arthropod slide mounts that Richard prepared will be the accom- plishment that will have the most permanent and important impact. Certainly in our own research it is the beautifully cleared and stained Wilkey preparations that we seek out whenever available. Studying them, it is easy to correctly ascertain the position and structure of morphological features and make accurate determi- nations, illustrations, and descriptions. The largest concentrations of Wilkey slides are in the collections of the Cali- fornia Department of Food and Agriculture in Sacramento and the Bohart Mu- seum at the University of California at Davis. He also took on the task of slide mounting the dry scale insect type material of Gordon Floyd Ferris and distrib- uting it to the major museums of the world. This material is especially useful because it is of better quality than preparations made by Ferris himself and is present in collections such as the Smithsonian’s National Museum of Natural 1997 MILLER & DAVIDSON: WILKEY OBITUARY 203 History at Beltsville, Maryland and The Natural History Museum in London, where only limited Ferris material was available previously. Wilkey also contributed significantly as a teacher. He taught others his slide mounting techniques and was a behind-the-scenes coorganizer and teacher in all seven Coccidology short courses at the University of Maryland. In this role he taught over 100 students from around the world how to properly slide-mount scale insects. He also used his broad entomological knowledge and extraordinary enthusiasm to encourage hundreds of 4-H’ers to learn about insects. He was in- volved in 4-H for 40 years and during that time taught more than 400 students about insects. Through his teaching and enthusiasm, approximately 10 of these students have pursued careers in entomology. Recently, in recognition of contri- butions to 4-H and the community, he and wife Dorothy were chosen Grand Marshals of the Wells County 4-H parade, an important annual affair in the Wells County area, and he was the Wells County, Indiana, Citizen of the Year in 1994. He was a past board member of the Wells County Society for Crippled Children and Adults, the Bluffon Park Board, the Rivergreenway Project, Public Library Board, and member of Friends of the Library and Bluffon Lions Club. He is survived by wife, Dorothy Weber Wilkey of Bluffon, Indiana; three sons, John R. and Frank J. of Hayden Lake, Idaho, and David A. of Washington, D.C.; a sister Nancy Thut of Fairfield, Connecticut, and two grandchildren. Received 10 Feb 1997; Accepted I Apr 1997. PAN-PACIFIC ENTOMOLOGIST 73(4): 204-212, (1997) SEASONAL FLIGHT PATTERNS OF BARK AND AMBROSIA BEETLES (COLEOPTERA: SCOLYTIDAE) IN NORTHEASTERN OREGON ROBERT W. PECK,!':? ARMANDO EQUIHUA-MARTINEZ** AND DARRELL W. Ross? 'Department of Forest Science, Oregon State University, Corvallis, Oregon 97331 3Department of Entomology, Oregon State University, Corvallis, Oregon 97331 ‘Department of Forest Science, Oregon State University, Corvallis, Oregon 97331 Abstract—The abundance and phenology of scolytid beetles collected in multiple-funnel traps baited with the Douglas-fir beetle (Dendroctonus pseudotsugae Hopkins) pheromones frontalin, seudenol, MCOL, and ethanol in NE Oregon are reported. Other than D. pseudotsugae, Den- droctonus ponderosae Hopkins, and Dendroctonus rufipennis (Kirby), a total of 17,612 beetles from 44 species were collected between 5 May and 21 Sep 1993. Dendroctonus brevicomis LeConte and Hylastes nigrinus (Mannerheim) were most abundant (comprising 44.5% and 31.7% of the total, respectively), followed by Pityophthorus confertus Swaine (8.5%), Dendroctonus valens LeConte (4.2%), Hylastes longicollis Swaine (3.4%), and Hylastes ruber Swaine (2.7%). Most species were rare; the combined number of individuals of the 26 least common species comprised <1% of the total. Pityophthorus deletus LeConte and Pityophthorus grandis Black- man are reported from Oregon for the first time. Flight activity for most species began after a seasonal increase in temperature in mid-May and subsided by late July. Seasonal flight patterns are shown for the 14 most abundant species. It is unknown how each species was affected by the lure, but ethanol may have been an important attractant for many species. Key Words.—Insecta; Scolytidae; Oregon; Phenology; Pheromones; Trapping The Scolytidae are a diverse group of beetles that live primarily within phloem and cambium (bark beetles) or xylem (ambrosia beetles) tissues of freshly dead, dying, physiologically stressed, or sometimes healthy trees (Rudinsky 1962, Fur- niss & Carolin 1977). Scolytids contribute to forest processes such as providing food for insectivorous birds (Knight 1958, Otvos 1965), facilitating the decom- position of dying trees (Schowalter et al. 1992), vectoring tree pathogens (Graham 1967, Hessburg et al. 1995), and providing fuel for wildfires (Geizler et al. 1980), thus influencing the structure and species composition of forest communities (Schmid & Hinds 1974, Veblan et al. 1991). Although the biology of some eco- nomically important species is well known (Stark & Dahlsten 1970, Thatcher et al. 1980, Wood, S. L. 1982, Christiansen & Bakke 1988, Raffa 1988), consider- ably less is known about the distribution and life-history of other species. Cham- berlin (1917) first summarized the scolytids found in Oregon. S. L. Wood (1982) and Furniss et al. (1992) subsequently added a number of species to those iden- tified in the earlier report. However, distribution records for most species are from a small number of localities. Information on the flight behavior and seasonal distribution of some scolytids in western Oregon has been provided by Rudinsky * Current Address: USDA Forest Service, Pacific Northwest Research Station, 3200 Jefferson Way, Corvallis, Oregon 97331 4 Current Address: Programa de Entomologia y Acarologia, Instituto de Fitosanidad, Colegio de Postgraduados, Montecillo, Edo. de Mexico, C.P. 56230 Mexico 1997 PECK ET AL.: BARK AND AMBROSIA BEETLE FLIGHT 205 & Daterman (1964), Daterman et al. (1965), and Zethner-M@ller & Rudinsky (1967). Pheromones and host-tree derived chemicals are useful for studying the biology and behavior of some scolytid species. For example, pheromones have been used to identify communication patterns regulating host-tree colonization, mate-finding, and reproduction of various species (Furniss et al. 1972, Wood & Bedard 1977, Birch 1978, Wood, D. L. 1982). This understanding has led to the development of pheromone applications to reduce damage caused by pest species (Bakke 1982, McGregor et al. 1984, Ross & Daterman 1994, 1995). However, because individ- ual pheromones, or their combinations, are generally narrow in specificity, most studies have been restricted to only one or a few species. In contrast, chemicals released from stressed or decaying trees, such as ethanol (Kimmerer & Kozlowski 1982, Byers 1992, Lindelow et al. 1992, Kelsey 1994) and terpenes (Vité & Gara 1962), are attractive to a variety of scolytids (Rudinsky 1966, Moeck 1970, 1971, Moeck et al. 1981, Montgomery & Wargo 1983, Klimetzek et al. 1986, Chenier & Philogene 1989, Schroeder & Lindelow 1989, Byers 1992). As a result, studies using traps baited with these chemicals have added significantly to our under- standing of scolytid distribution and activity patterns (Roling & Kearby 1975, Turnbow & Franklin 1980, Atkinson et al. 1988). Life-history information for many scolytid species found in Oregon is lacking, particularly east of the Cascade Mountain Range. This study describes the sea- sonal flight patterns of scolytids caught in funnel traps baited with Douglas-fir beetle (Dendroctonus pseudotsugae Hopkins) pheromones in the Blue Mountains of NE Oregon. Of the pheromones used, frontalin (Pitman & Vité 1970), seudenol (Vité et al. 1972) and MCOL (Libbey et al. 1983) are released by the female beetle during the initial stages of tree colonization, and ethanol can be released by either sex, the host tree, or by associated microorganisms (Pitman et al. 1975). Data presented here were obtained by sorting scolytids from a subset of trap samples from a study designed to test the effectiveness of attractant-baited traps for area-wide management of Douglas-fir beetle populations (Ross & Daterman £997,)3 METHODS AND MATERIALS Traps were placed within the NE portion of the Wallowa Valley Ranger District of the Wallowa-Whitman National Forest in NE Oregon. All traps were located at elevations between 1470 and 1690 m. Forests in the area are mixed conifer, comprised largely of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), pon- derosa pine (Pinus ponderosa Dougl. ex Laws.), grand fir (Abies grandis (Doug}.) Lindl.), western larch (Larix occidentalis Nutt.) and lodgepole pine (Pinus con- torta Dougl.), with scattered Engelmann spruce (Picea engelmannii Parry ex En- gelm.) and alpine fir (Abies lasiocarpa (Hook.) Nutt.). Forest structure in the area is variable reflecting a history of natural and human-caused disturbances. Tem- perature data for the trapping period were obtained from a weather station located at 1281 m elevation approximately 26 km from the trap sites. Within the study area, three 259-ha plots were established for trapping (Ross & Daterman 1997). Approximate latitude and longitude for each plot were 45°51 N/117°07' W, 45°50’ N/117°03' W, and 45°44’ N/116°52’ W. Within each plot, 12-14 trap sites were chosen that were a minimum of 0.25 km apart. Three traps, 206 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(4) spaced approximately 10 m apart, were located at each site. Traps were placed in clearings or young stands away from potential scolytid breeding material. Due to disturbance from cattle and/or wildlife, some traps had missing samples on at least one occasion during the study. Twenty-one traps without missing samples (seven from each plot) were identified for analysis. No site was represented by more than one trap. The traps were a 16-unit multiple-funnel type (Lindgren 1983) suspended from an eight-foot metal pole, with a DDVP-impregnated piece of plastic in the col- lecting cup to kill insects. Each trap was baited with 250 ul of frontalin in an eppendorf vial, 200 mg of MCOL in a bubble capsule, and 15 ml of ethanol in a plastic pouch (Phero Tech Inc., Delta, BC, Canada). Each trap also contained either 400 mg of frontalin and 200 mg of seudenol, or 200 mg of frontalin and 100 mg of seudenol, in 5% PVC formulations (Daterman 1974). All formulations released pheromone or kairomone throughout the trapping period (DWR, unpub- lished data). Traps were deployed by 5 May 1993. Samples were collected on or within one day of the following dates: 18 and 25 May; 7, 22 and 29 Jun; 6, 14 and 27 Jul; 10, 18 and 24 Aug; 3 and 21 Sep 1993. Samples were initially sorted only for Dendroctonus pseudotsugae and selected predators. However, because Dendroc- tonus ponderosae Hopkins and Dendroctonus rufipennis (Kirby) look similar to D. pseudotsugae, and were relatively rare in the samples, they were probably combined with D. pseudotsugae during the sorting process. Remaining scolytids were removed from the samples at a later date for the present study. Species were identified and compared to reference material within the USDA Forest Service Hopkins Forest Insect Collection. Voucher specimens from this study were de- posited in the Oregon State University Systematic Entomology Laboratory Col- lection. RESULTS AND DISCUSSION Other than D. pseudotsugae, D. ponderosae and D. rufipennis, a total of 17,612 scolytid beetles within 44 species were identified from the 21 funnel traps (Table 1). Dendroctonus brevicomis LeConte and Hylastes nigrinus (Mannerheim) dom- inated the samples numerically, comprising 44.5% and 31.7% of the total, re- spectively. All other species were much less common, with Pityophthorus con- fertus Swaine (8.5%), Dendroctonus valens LeConte (4.2%), Hylastes longicollis Swaine (3.1%), and Hylastes ruber Swaine (2.7%) being the next most abundant species. Overall, most species were relatively rare; the 26 least common species collectively comprised <1% of the total. Specimens of Pityophthorus deletus LeConte and Pityophthorus grandis Blackman represent first records of occur- rence for Oregon. For P. deletus, this record fills a distributional gap, with pre- vious collections in British Columbia, Idaho and California, and the known dis- tribution of P. grandis is extended northward from California. All species col- lected are known to utilize coniferous trees present within the study area (Bright & Stark 1973, Wood, S.L. 1982). Flight patterns of many species showed a strong seasonal trend, with flight initiation corresponding to the first warm days of spring. Temperatures showed a marked seasonal increase on 10 May, rising from a previous two-week average of 6.7° C, to 24° C by 12 May (Fig. 1). Hylurgops porosus (LeConte), Hylurgops L66I Table 1. Number of scolytid beetles collected in funnel traps baited with frontalin, seudenol, MCOL and ethanol between 5 May and 21 Sep 1993 in NE Oregon. Taxon Number trapped Taxon Number trapped Hylastini Scolytus piceae (Swaine) 3 Hylurgops porosus (LeConte) 54 Scolytus tsugae (Swaine) 3 Hylurgops reticulatus Wood 29 Scolytus unis pinosus LeConte 83 Hylurgops subcostulatus (Mannerheim) 90 Scolytus ventralis LeConte 6 Hylastes gracilis LeConte 4 Dryocoetini Hylastes longicollis Swaine 547 Dryocoetes sechelti Swaine ] Hylastes macer LeConte 110 Ipini Hylastes nigrinus (Mannerheim) 5583 Pityogenes carinulatus (LeConte) 49 Hylastes ruber Swaine 483 Pityogenes fossifrons (LeConte) 2 Hylastes tenuis Eichoff 2 Pityogenes knechteli Swaine 1 Tomicini Pityokteines elegans Swaine a Pseudohylesinus dis par Blackman 8 Pityokteines ornatus (Swaine) 2 Pseudohylesinus nebulosus (LeConte) 153 Orthotomicus caelatus (Eichhoff) 8 Pseudohylesinus granulatus (LeConte) 3 Ips emarginatus (LeConte) 1 Dendroctonus brevicomis LeConte 7830 Ips latidens (LeConte) 2 Dendroctonus ponderosae Hopkins — Ips pini (Say) 24 Dendroctonus pseudotsugae Hopkins — Xyloterini Dendroctonus rufipennis (Kirby) — Trypodendron lineatum (Olivier) 2 Dendroctonus valens LeConte 745 Corthylini Phloeotribini Pityophthorus confertus Swaine 1490 Carphoborus intermedius Wood 1 Pityophthorus confinis LeConte 3] Phloeotribus lecontei Sched] 3 Pityophthorus deletus LeConte 1 Polygraphini Pityophthorus grandis Blackman 2. Polygraphus rufipennis (Kirby) 17 Pityophthorus nitidulus (Mannerheim) 4 Scolytini Gnathotrichus retuses (LeConte) 192 Scolytus laricis Blackman 5 Gnathotrichus sulcatus (LeConte) 5 Scolytus opacus Blackman 28 Total number of beetles 17,612 4D. ponderosae and D. rufipennis were probably grouped with D. pseudotsugae during the sorting process. Data for these three species are presented elsewhere (Ross and Daterman 1997), LHO'Id AILadad VISOUPNV GNV MV “IV LA MOdd LOC 208 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(4) O i Ss Y 5 20 5 2 | 2 fi) z= 10 mo) E uo) ) : 0 Apr May Jun Jul Aug Sep Date Figure 1. Median daily temperature near the study site. Solid arrows indicate sample collection dates and dashed arrow shows date traps were placed in the field. subcostulatus (Mannerheim), Hylastes macer LeConte, H. nigrinus, Pseudohyle- sinus nebulosus (LeConte), Pityogenes carinulatus (LeConte), and Gnathotrichus retusus (LeConte) were collected at or near their greatest abundance during one of the two sampling periods in May (Fig. 2). Similar early season activity patterns were reported by Daterman et al. (1965), who found a maximum daily temperature of 14° C necessary to initiate flight for most of the scolytid species they collected in western Oregon. Although temperatures were consistently low prior to the time traps were set out, some scolytids may have been flying earlier. For example, P. nebulosus was most abundant on the first sample date, with a precipitous drop in activity thereafter (Fig. 2), and was previously found to be one of the earliest species to fly in other regions (Walters & McMullen 1956, Daterman et al. 1965). Pityophthorus confinus LeConte was most abundant in the 7 June samples, while five species (H. longicollis, H. ruber, D. brevicomis, D. valens, Scolytus unispi- nosus LeConte) were most abundant in the 22 or 29 June samples (Fig. 2). Da- terman et al. (1965) also found H. ruber and S. unispinosus to be most active slightly later than other species, suggesting their need for higher temperatures to initiate flight. Although most species showed a single peak in activity, P. confertus was abundant in May, June and July, and D. brevicomis and D. valens were trapped in relatively large numbers into September. The individuals collected dur- ing mid- to late season were likely re-emergent parent adults searching for new breeding substrates or newly emerging early-season brood adults (Miller & Keen 1960, Stark & Dahlsten 1970). For most other species, flight activity had de- creased by 14 July. Daterman et al. (1965) also found diminished activity levels by the end of July. More individuals of many species were collected than expected by chance, suggesting that these species were attracted to the lure rather than being passively intercepted by the traps. However, because lure composition was not experimen- tally manipulated, it is not possible to determine which components were attrac- tive to each species. Of the attractants used, frontalin is known to be a component of the aggregation pheromone of D. brevicomis and may have contributed to its attraction (Bedard et al. 1980). Ethanol has been shown to be a strong attractant 1997 PECK ET AL.: BARK AND AMBROSIA BEETLE FLIGHT 209 Hylurgops porosus Pseudohylesinus nebulosus ON fF OD CO Hylurgops subcostulatus 150 Dendrostonns brevicomis Hylastes longicollis Dendroctonus valens Hylastes macer Scolytus unispinosus Total number of beetles / day oO - NY W Hylastes ruber 1.0 Pityogenes carinulatus 0.5 0.0 Gnathotrichus retusus Hylastes nigrinus O-pANWHA Pityophthorus confinis 40 Pityophthorus confertus May = Jun Jul Aug Sep Date Date Figure 2. Seasonal flight patterns for the 14 most frequently trapped scolytid species within the study area. for ambrosia beetles, such as G. retusus, (Moeck 1970, 1971, Roling & Kearby 1975, Turnbow & Franklin 1980, Kelsey 1994) as well as for H. nigrinus (Wit- cosky et al. 1987). Ethanol was possibly responsible for the attraction of some of the abundant scolytids, because only D. brevicomis is known to be attracted to any of the Douglas-fir beetle pheromones used in the lure. Trapping in the same area in 1994 with frontalin and seudenol lures captured very few scolytids other 210 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(4) than D. pseudotsugae (DWR, unpublished data), further suggesting the importance of ethanol as an attractant. The number of scolytids collected in each trap varied greatly even though the surrounding forests were similar in elevation and tree composition. For most spe- cies, a relatively small number of traps caught a large proportion of the total number of individuals. For example, 95.0% of P. confertus were from one trap, 69.8% of H. nigrinus were collected from one trap and 87.3% from two traps, 86.5% of D. brevicomis were collected from four traps, 75.4% of D. valens were collected from three traps and 81.4% of H. longicollis were collected from four traps. Although the numbers collected were concentrated in a few traps for these species, they occurred frequently, being collected in 11, 20, 19, 20, and 12 of the 21 traps, respectively. In contrast, many species were only collected in a few traps. For example, 16 species were caught in only one or two traps and 17 other species were collected in five traps or less. This study has identified a large number of bark and ambrosia beetles captured in multiple-funnel traps baited with Douglas-fir beetle pheromones from a poorly studied part of Oregon, and has shown seasonal flight patterns for many of these species. It is not known which components of the lure were most attractive to the beetles, but ethanol may have been important because it is a by-product of tree decomposition as well as a pheromone. Because many species are patchily dis- tributed across the landscape, a large number of traps in various forest stand types will be necessary to adequately sample scolytid communities. ACKNOWLEDGMENT We thank G.E. Daterman and R.L. Livingston for reviewing an earlier draft of this manuscript. 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PAN-PACIFIC ENTOMOLOGIST 73(4): 213-224, (1997) NATURE OF GALLERIES, DURABILITY OF BORING SCARS, AND DENSITY OF XYLOTRECHUS VILLIONI (VILLARD) LARVAE (COLEOPTERA: CERAMBYCIDAE), ON CONIFEROUS TREE TRUNKS' RYOTARO IWATA’, FUSAO YAMADA?, HIROFUMI KATO?, HIROSHI MAKIHARA}?, KuUNIO ARAYA‘, HISASHI ASHIDA> AND MASASHI TAKEDA® *Department of Forest Science and Resources, College of Bioresource Sciences, Nihon University, Kameino, Fujisawa 252, Japan 3Forestry and Forest Products Research Institute (MAFF), Matsunosato, Kukizaki 305, Japan 4Graduate School of Human and Environmental Studies, Kyoto University, Sakyo-ku, Kyoto 606-01, Japan °KDK Corporation, Higashi-Kuj6-Nishi-Aketaché, Minami-ku, Kyoto 601, Japan *Ryokusei Kenkytisho Co., Inc., Muromachi 1-chéme, Ikeda 563, Japan Abstract——Spatial distributions and shapes of ‘“‘whirl-like” scars on the trunks, made by gallery formation of mature larvae of Xylotrechus villioni (Villard) (Coleoptera: Cerambycidae), a pri- mary borer of Abies and Picea coniferous trees in Japan, were investigated at an Abies firma Sieb. et Zucc. plantation in Hachidji, Tokyo Pref., an A. firma natural stand in Miyama, Kyoto Pref. and an A. sachalinensis (Fr. Schm.) Mast. plantation in Imakane, Hokkaid6. Although all the forests investigated showed cumulative “whirl-like”’ scars on the tree trunks, a low density of existing larvae was inferred from the analyses of the locations and shapes of these scars. Mortality throughout the larval stages, as well as between the final phase of larva and the adult emergence, was suggested. Trunk analysis of a damaged A. firma tree showed that a “‘whirl- like’’ scar can remain on the trunk surface for as long as 27 years after the formation of the larval gallery. The most susceptible class of Abies trees had a diameter at the breast height of 35—45cm. “Whirl-like”’ scars were distributed more densely in the lower part of the trunks. Key Words.—Insecta, Cerambycidae, Xylotrechus villioni, larval gallery, spatial distribution, co- nifers, Abies Xylotrechus villioni (Villard), a cerambycid beetle endemic to Japan (Fig. 1), is a primary borer of coniferous tree species, mostly of the genera Abies and Picea (Iwata et al. 1990). This species is probably the largest member of the tribe Clytini (subfamily Cerambycinae), with its adult body length being 20—26 mm in males and 25—30 mm in females. Although this species, insofar as historically recorded, once had an outbreak and its larvae caused serious damage on Abies sachalinensis (Fr. Schm.) Mast. plantations in Hokkaidé (Kamijo et al. 1970, Kamijo & Suzuki 1973), adult beetles, as well as larvae, are found only rarely. This low density makes direct observations difficult, and little is known about the bionomics and ecology of this species (Iwata et al. 1990). Larvae bore under the bark of the trunk and branches, and larval galleries cause cicatricial scars, which become more evident on the trunk as the tree cures the ' Presented at the 102nd Annual Meeting of Japanese Forestry Society at Nagoya (April, 1991), the 105th Ann. Mtg. of Jpn. For. Soc. at Fuchii (April, 1994), the 106th Ann. Mtg. of Jpn. For. Soc. at Sapporo (April, 1995) and 20th International Congress of Entomology at Firenze (August, 1996). 214 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(4) aa Figure 1. Xylotrechus villioni adults from Mt. Kasuga, Nara. Reproduced from Iwata (1991). left, male; right, female. damage through cicatrization with phloem recovery. These scars remain recog- nizable for a long period of time because the complete recovery of outer phloem takes many years after the appearance of a scar. The larval boring consists of 5 phases (Kamijo & Suzuki 1973, Iwata et al. 1990), of which Phase 4, attributed to mature larva, is characterized by the “‘whirl- like”’ gallery (Fig. 2), which is peculiar to this species and presumably made as a guard against resin exudation (Iwata et al. 1990, Iwata 1991). Every scar with a straight, sinuate or “‘whirl-like’’ appearance found on Abies and Picea trunks in Japan is almost always accompanied by a gallery with tightly packed fine frass, which characterizes boring by a clytine cerambycid. As no other clytine species that attacks living conifers is known from Japan, the presence of such scars on coniferous trunks in Japan always indicates attacks by X. villioni. Formation of a “‘whirl-like” scar (Fig. 3) on the trunk surface is caused through “‘whirl-like’’ gallery formation, and their distribution reflects the historical spatial distribution of the insect. Studying the “‘whirl-like’’ scars caused by X. villioni larvae may contribute to its enigmatic biology. This paper reports the nature and the spatial distribution of X. villioni larval galleries on coniferous tree trunks. Figure 2. ‘‘Whirl-like’ gallery (arrowed) made by a X. villioni mature larva on a dead Abies mariesii tree-trunk, Hinoemata, Fukushima Pref. 1997 IWATA ET AL.: XYLOTRECAUS IN CONIFEROUS TREES 7A is) Figure 3. ‘‘Whirl-like’’ scar (arrowed) made through gallery formation of a X. villioni mature larva on a living Abies firma tree-trunk, Mt. Kasuga, Nara. Reproduced from Iwata (1991). MATERIALS AND METHODS Study sites.—Investigations were carried out at three sites, (A): an Abies firma Sieb. et Zucc. plantation in Tama Forest Science Garden, FFPRI (MAFFP), Hachi- 6ji, Tokyo Pref. (about 0.5 ha in extent, northeast-inclined, alt. 220 m) in Sep— Nov 1990; (B): an A. firma natural stand in Ashiu Kyoto University Forest (Sec- tion 33), Miyama, Kyoto Pref. (about 0.6 ha in extent, southwest-inclined, alt. 450m) in Aug 1990; and (C): an A. sachalinensis plantation in Kanahara, Ima- kane, Hokkaid6 (about 0.3 ha in extent, flatland, alt. 80 m) in Aug 1991. The large-scale damage in the A. sachalinensis plantations in Imakane (C) and the damage in the A. firma plantation in Hachidji (A), caused by X. villioni, have previously been documented (Ganda et al. 1986, Makihara et al. 1995, respec- tively). Spatial distributions and morphology of ‘“‘whirl-like’’ scars.—The following were recorded for each tree at all sites: (1) diameter at the breast height (DBH), (2) presence of resin exudations (a possible index of larval existence), (3) presence of straight and sinuate scars, and (4) presence of “‘whirl-like’’ scars on the trunk. These give the spatial distribution of “‘whirl-like’’ scars in relationship to tree’s DBH, as well as the density of the existing insects. In sites (A) and (B), (5) number of “‘whirl-like’’ scars in each tree was also recorded, and additional parameters were measured for each of the “‘whirl-like”’ scars: (6) rotation (either clock- or counterclock-wise), (7) up-and-down direction of the straight (or sinuate) scar connected with “‘whirl-like’’ one, (8) height above the ground, (9) size (as expressed by that divided by (7/4)), (10) compass direction and (11) exposure of xylem and the gallery through peeling-off of the phloem above it. Then, the trees were categorized with regard to parameters 2—5, and the ‘“‘whirl-like’’ scars were categorized with regard to parameters 6—11. Spatial distribution of ‘“‘whirl-like’’ scars is quantitatively expressed with Lloyd’s (1967) ‘“‘mean crowding”’ and “‘patchiness’’. Whole larval gallery observation.—In Hinoemata, Fukushima Pref., a blighted A. mariesii Mast. tree was felled on 24 Jun 1990 to expose the whole gallery made by a single larva. Also, at the A. firma plantation in Tama Forest Science Garden, FFPRI (MAFFP), Hachidji, one living tree (DBH 19 cm, 12 m high) was felled and cross-cut on 23 Aug 1993 to record the appearances, dimensions and 216 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(4) Table 1. Categorizations of Abies trees with regard to the parameters concerning the damage by X. villioni larvae in three forest sites, Tama Forest Science Garden, Hachidji (A), Ashiu Kyoto Uni- versity Forest (Section 33), Miyama (B), and a plantation in Kanahara, Imakane (C). Locality (a) Hachi6ji (B) Miyama (C) Imakane Tree species A. firma A. firma A. sachalinensis Resin exuded 19 (50%) 12 (43%)? 16 (17%)? Resin little or not exuded 19 (50%) 16 (57%) 77 (83%) Straight and/or sinuate scar(s) present 31 (82%) 17 (61%)? Straight and/or sinuate 31 (82%) 25 (89%) scar(s) indistinctly present 0 (0%) 8 (29%) Straight or sinuate scar scars present: absent 7 (18%) 3 (11%) 25 (27%) With no ‘‘whirl-like” scars 14 (37%) 14 (50%)? scars absent: With | ‘“‘whirl-like’’ scar 15 (39%) 5 (18%) 68 (73%) With 2 ‘‘whirl-like’’ scars 6 (16%) 8 (29%) With 3 ‘‘whirl-like’’ scars 2 (5%) 1 (4%) With 4 ‘“‘whirl-like’” scars 0 (0%) 0 (0%) With 5 “‘whirl-like’’ scars 1 (3%) 0 (0%) Total 38 28 93 4Percent of trees examined. shapes of all the “‘whirl-like’’ scars, as well as all the larval galleries within the trunk. The ages of the ‘“‘whirl-like galleries’’ were estimated by counting the numbers of annual rings between each gallery and the cambium. RESULTS Spatial distributions of ‘‘whirl-like’’ galleries.—For the three forest sites in- vestigated, the trees were categorized with regard to the presence of resin exu- dation, the presence of straight and sinuate scars, and the presence or the number of “‘whirl-like’’ scars on their trunks (Table 1, Fig. 4). The spatial distributions of ‘‘whirl-like’’ scars in these forest sites in relation- ship to trees’ DBH are shown in Tables 2 to 4. We found that 17-50% of the trees investigated showed resin exudation, 82— 89% of the trees possessed straight and/or sinuate scar(s) and 50-63% possessed ‘“‘whirl-like’’ scar(s) (Tables 1-3). Although the ages of the investigated forest stands vary considerably, in all the highest ratio of the trees with ‘‘whirl-like’’ scars was found in trees with 35-55 cm DBH (Tables 2—4), suggesting a definite tree diameter preference by X. villioni Ovipositing females independent of forest age. The mean crowding value of the “‘whirls-like’’ scars in each DBH-class is low (Tables 2—3), except for the 2.31 value in the trees with 25-35 cm DBH in Hachidji. Trees with 35-45 cm DBH had the highest value of mean crowding in Miyama. Lloyd’s patchiness values ranged from 0.40 to 2.48 (Tables 2—3). The mean crowding value, and the ratio of trees with “‘whirl-like’’ scars suggest that the most susceptible trees have 35— 45 cm DBH although there is a time lag of a few years between beetle oviposition and the appearance of scars. 1997 IWATA ET AL.: XYLOTRECAUS IN CONIFEROUS TREES 217 © O @ O © Figure 4. Overview of the distribution of Abies sachalinensis trees damaged by X. villioni larvae in a plantation, Kanahara, Imakane, Hokkaidé (C). The sign O represents a tree without scars or resin exudation, @ with resin exudation and without scars, © without resin exudation and with scars, © with both resin exudation and scars, and X represents a dead tree. The ‘“‘whirl-like’’ scars were almost evenly observed to have either clock- or counterclock-wise rotation and to be either upward- or downward-connected. The size and the manner of presence of the “‘whirl-like’’ scars were shown to be highly variable (Table 5). The mean ‘“‘whirl’’ size was larger in Hachidji ((251 X m/4) cm’) than in Miyama ((179 X 7/4) cm’), although the mean DBH of the trees (31 cm and 49 cm, respectively) shows the opposite trend. The correlation between the presence of straight and/or sinuate scars and “‘whirl-like’”’ scars on the trunk surface is shown in Table 6. The data show a positive significant correlation (Fisher’s exact probability test, P < 0.0005): ‘“‘whirl-like’” scars are always accompanied by straight and/or sinuate scar(s), suggesting that a straight and/or sinuate scar without a “‘whirl-like’’ scar is in- dicative of larval mortality. Statistical analyses found that most of the parameters measured are independent of each other at Hachidji and Miyama, although the presence of straight and/or sinuate scar(s) are correlated to the presence and the abundance of “‘whirl-like”’ scar(s) (Fisher’s exact probability test, P < 0.01 for all), and the presence of resin exudation to the presence of ‘“‘whirl-like’”’ scar(s) (x-test, P < 0.01 for two sites). Also, there were no correlations among the presence of scars, presence of resin exudation, and trees’ DBH at Imakane. Whole larval gallery observation.—The gallery made by a single larva, found in the trunk of a blighted A. mariesii at Hinoemata (Fig. 5), had a length of 218 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(4) Table 2. Spatial distribution of “whirl-like’’ scars made by X. villioni larvae on Abies firma tree trunks in relationship to trees’ diameter at breast height (DBH) in Tama Forest Science Garden, Hachidji (A). DBH range (cm) 5-15 15-25 75-35 35-45 45-55 55-65 Total Number of trees (N) 0 11 14 10 5 0 38 Number of trees with ‘‘whirl- like’’ scars (N’) 0 8 5 8 2) 0 24 Ratio of trees with ‘“‘whirl-like” scars (N’/N) — 0.73 0.36 0.80 1.00 — 0.63 Total number of ‘‘whirl-like”’ scars ({w; = W)* 0 10 13 10 5 0 38 Median value of the DBH range as expressed in meter (8) 0.10 0.20 0.30 0.40 0.50 0.60 -— Total surface area of all trees as expressed in arbitrary area unit (N87)? 0 0.44 1.26 1.60 0.75 0 4.05 Number of ‘‘whirl-like’’ scars per arbitrary area unit (W/Nd’) —_ DO] 10.3 6.3 6.7 — 9.38 Number of “‘whirl-like”’ scars per a tree (W/N = w) — 0.91 ' 0.93 1.00 1.67 — 1.00 Mean crowding of ‘“‘whirl-like” scars (Sw2/Sw, — 1 = Wy == 0.40 2.31 0.40 1.20 — 1.16 Lloyd’s “‘patchiness”’ (Wiw) — 0.44 2.48 0.40 0.72 — 1.16 4 Let w; be the number of “‘whirl-like’ scars on each tree. > Here, the real total area must be aNé&* in square meter, with the non-dimensional constant a representing a factor related to the taperness of the trees, supposing all the trees have the same a value. 112 cm, including 25 cm after the entrance into the xylem for pupation. The boring initiation point of the Ist instar larva and the top of the ‘‘whirl-like”’ gallery were situated 205 cm and 295 cm high above the ground, respectively. A living A. firma, felled at Hachi6ji, had seven independent larval galleries, of which four had ‘“‘whirl-like’’ scars followed by pupal chambers. The appearances, dimensions, shapes and estimated ages of these larval galleries are summarized in Table 7. The formation of the “‘whirl-like” gallery, preceding pupation, can considerably damage the meristem, and a “‘whirl-like’’ scar can remain on the trunk surface for as long as 27 years after the formation of the larval gallery. The bark discs over some of the ‘“‘whirl-like’ galleries were peeled off, exposing the xylem surface. DISCUSSION Lloyd’s “‘patchiness”’ values (0.40—2.48; Tables 2—3) indicate that the ‘‘whirl- like’’ scars are distributed randomly and sparsely with the tree regarded as the sample-unit. Because the “‘whirl-like’’ scars represent a cumulative spatial distri- bution of mature larvae for as long as 27 years, the present distribution of existing larvae within the trees must be even sparser: the existing individuals of this spe- cies are distributed in an extraordinarily low density. This is not incompatible with the ratio of trees with ‘‘whirl-like’ scars of 63% at Hachidji or 50% at Miyama (Tables 2 and 3). The low density of existing beetles is supported by the 1997 IWATA ET AL.: XYLOTRECHUS IN CONIFEROUS TREES 219 Table 3. Spatial distribution of ‘‘whirl-like” scars made by X. villioni larvae on Abies firma trees in relationship to trees’ diameter at breast height (DBH) in Ashiu Kyoto University Forest (Section 33), Miyama (B). (For footnotes a and b, see Table 2.) DBH range (cm) 15225 25-35 35-45 45-55 55-65 65-75 Total Number of trees (N) 1 4 6 Ps 10 5 28 Number of trees with ‘“‘whirl-like’’ scars (N’) 0 1 4 1 5 3 14 Ratio of trees with ‘“‘whirl-like”’ scars (N’/N) 0.00 0.25 0.67 0.50 0.50 0.60 0.50 Total number of “‘whirl-like”’ scars (Sw, = W)2 0 iD 8 2; of 5 24 Median value of the DBH range as expressed in meter (8) 0.20 0.30 0.40 0.50 0.60 0.70 —- Total surface area of all trees as expressed in arbitrary area unit (N87)? 0.04 0.36 0.96 0.50 3.60 2.45 7.91 Number of “‘whirl-like’”’ scars per arbitrary area unit (W/N6’) 0 5.56 8.33 4.00 1.94 2.04 21.87 Number of “‘whirl-like”’ scars per a tree (W/N = w) 0 0.50 1.33 1.00 0.70 1.00 0.86 Mean crowding of “whirl-like”’ scars (Sw2/Sw, — 1 = W) = 1.00 1.25 1.00 0.57 1.00 0.96 Lloyd’s “‘patchiness’’ (W/w) — 2.00 0.94 1.00 0.81 1.00 1.12 failure to capture any adult beetles during the field surveys conducted during the adult emergence season using either traps baited with kairomones and genus- specific sex pheromones (Iwata et al. 1991, 1992, 1993) or by visual searching. Because resin exudation indicates that the coniferous tree is inhabited by boring larvae, as has been demonstrated by Kobayashi (1982) for Semanotus japonicus (Lacordaire) infesting Cryptomeria japonica D. Don, just half of the trees in the forest site and 17-43% of the trees in other sites seemed to contain beetle larvae (Table 1). The plantation site included open gaps, as well as many stumps made by cut- ting, suggesting that many young trees had been removed by felling due to lethal damage inflicted by X. villioni (Ganda et al. 1986). The damage by X. villioni took place on the edge of the plantation, reconfirming the observation of Kamijo & Suzuki (1973). This is not incompatible with the random and sparse distribution of ‘“‘whirl-like’”’ scars on trees because tree damage is viewed not within a single tree but over a forest section. Table 4. Scars made on Abies sachalinensis trunks through boring activity of X. villioni larvae in relationship to trees’ diameter at breast height (DBH) in Kanahara, Imakane (C). DBH range (cm) 5-15 15-25 25-35 35-45 Total Number of trees (N) 16 51 24 2 93 Number of trees with ‘‘whirl-like, straight and/or sinuate scars (N’) 4 12 8 1 25 Ratio of trees with scars (N’/N) 0.25 0.24 0.33 0.50 0.27 220 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(4) Table 5. Categorizations of “‘whirl-like” scars made by X. villioni larvae with regard to geometrical parameters in two Abies firma forest sites, Tama Forest Science Garden, Hachidji (A), and Ashiu Kyoto University Forest (Section 33), Miyama (B). The numbers in some categories, when summed up, are inconsistent to the total number due to lack of data. (A) Hachidéji (B) Miyama Connected straight or sinuate downward 19 5 scar®: upward 19 17 (Height above the ground) = h: 800 cm Sh < 900 cm 1 0 700 cm = h < 800 cm 0 0 600 cm Sh < 700 cm 5 0 500 cm Sh < 600 cm 5 0 400 cm = h < 500 cm 1 2 300 cm Sh < 400 cm 6 0 200 cm Sh < 300 cm 7 5 100 cm =h < 200 cm 11 9 0cm Sh < 100 cm 1 8 (Size)/(m/4) = S: 50 cm? S S < 100 cm? 4 4 100 cm? = S < 150 cm? 6 3 150 cm? = S < 200 cm? 5 8 200 cm* S$ S < 250 cm? 4 1 250 cm* = S < 300 cm? 4 2 300 cm? S S < 350 cm? 6 2 350 cm? = S < 400 cm? 1 0 400 cm? S S < 450 cm? 3 0 450 cm? = S < 500 cm? 2 0 500 cm? = S < 550 cm? 1 0 550 cm? = S < 600 cm? 1 0 Compass direction: North 3 1 Northeast 6> 11° East z. 1 Southeast 5 1 South 6 2 Southwest Oe ae West 4 1 Northwest 2 4 Phloem disc above the “‘whirl-like’’ gallery: Wholly peeled off (“‘whirl-like’’ gallery exposed) 8 —4 Partly peeled off (“‘whirl-like”’ gallery half exposed) 3 — Recovered (“‘whirl-like”’ gallery covered) 26 — Total 38 24 ‘Not significantly biased (Binomial test, P > 0.05) in (A), but biased to “‘upward”’ (Binomial test, P < 0.05) in (B). > The study area is inclined to this direction. ‘ Not significantly biased to this (Kolmogorov-Smirnov’s test, P > 0.05). 4 Not checked. 1997 IWATA ET AL.: XYLOTRECHUS IN CONIFEROUS TREES 221 Table 6. Correlation between the presence of straight and/or sinuate scar(s) and the presence of “‘whirl-like”’ scar(s) made by X. villioni larvae on each of the tree trunks at two Abies firma forest sites, Tama Forest Science Garden, Hachidji (A) and Ashiu Kyoto University Forest (Section 33), Miyama (B). h y (A) Hachiéji ‘“‘whirl-like”’ scar(s) (B) Miyama “‘whirl-like’’ scar(s) Straight and/or sinuate scar(s) Present Absent Present Absent Present 24 (63%) 7 (18%) 13 (46%) 4 (14%) Absent 0 (0%) T (18%) 0 (0%) 11 (39%) ‘Including 8 trees only with very indistinct straight and/or sinuate scars. Radial section Tree surface INAS NE Are ae OTAN | Reeds Pp = < 3 bo rah ~y Pupal chamber ro s Poke 7 ‘ E = 5 = | a > i] aa! 3 a0 * = » 3 Be a7 a » Fay B 3 - a0 © —s oI 3 a= a (3) Ss =e = ry 1 e a a 2 U << ® t) 0 —_—_——_—S——— >" lcm 10cm Figure 5. Whole gallery made by a single larva of X. villioni on a dead Abies mariesii tree-trunk. The tree, of 18cm DBH, was found in 1985, harvested on 24 Jun 1990 at the end of Toyasu Tributary Trail, Funamata Valley, Hinoemata, Fukushima Pref. and presumed to have been blighted in 1982. 222 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(4) Table 7. Four ‘“‘whirl-like” scars made by X. villioni larvae on a living Abies firma tree sampled for the whole gallery observation, at Tama Forest Science Garden, Hachidji (A). Scar A Cc D G Height above the ground (m) mel 355 3H 11.6 Horizontal diameter X vertical diameter of the ‘‘whirl-like”’ scar (cm) 7 & 12 6X 6 9x9 6 X 6 Appearance of the “whirl-like”’ scar obscure evident evident evident Phloem disc over the ‘“‘whirl- wholly peeled recovered partly peeled recovered like” scar off off Total length of the connected straight and sinuate gallery (cm) 85 DT 45 68 Presumed site of the initial larval branch branch branch branch gallery (lost) (lost) (lost) (lost) Depth of the pupal chamber (cm) 9 6 8 0) Presence of the adult emergence exit hole* oa “ +t os Time elapsed since the formation of the gallery (years) 27-28 14-15 ] 11 4+, present; —, absent. The “‘whirl-like’’ scars tend to be distributed more densely in the lower part of the trunks although some in very high positions might have been overlooked to some degree (Table 5). This tendency in X. villioni infesting A. sachalinensis (Kamijo & Suzuki 1973) is known also in S. japonicus infesting C. japonica. In the latter species, this tendency has been ascribed to the change of bark roughness (Kobayashi & Yamada 1982). However, in Abies firma, by our visual inspection, the roughness of bark does not seem to be related to tree height. The predominance of upward gallery connections to the ‘‘whirl-like”’ scars in Miyama (Table 5) is ascribed to the concentration of the “‘whirl-like’”’ scars toward the bottom portion of the trunk. Although not significant, the compass direction of ‘“‘whirl-like’”’ scars on the tree-trunk is biased to the direction opposite to that toward which the study area is inclined, suggesting a possible strategy to avoid direct attack by natural enemies, such as picid woodpeckers (Iwata et al., in prep.). The condition of phloem disc above the “‘whirl-like’’ gallery (Table 5) suggests that Abies trees are partly successful in protecting the severe wounds by cicatri- zation with phloem recovery. However, in some cases they fail in curing the “‘whirl-like’’ wound exposing the xylem, which presumably allows insects and fungi to invade the trunk. Kamijo & Suzuki (1973) inferred a high mortality of X. villioni larvae boring within sound coniferous trees. The data on the presences of straight and/or sinuate scars and ‘‘whirl-like’’ scars on the tree trunk surface (Table 6) suggest that im- mature larvae, which are responsible for straight and sinuate scars, are not always successful in developing into mature larvae, which are responsible for ‘‘whirl- like’ scars. At most only 77% of immature larvae formed ‘‘whirl-like’”’ galleries. Further, of the four ‘“‘whirl-like’”’ scars within the tree sampled at Hachidji (Table 7), only one possessed an adult emergence exit hole, suggesting a rather low L997 IWATA ET AL.: XYLOTRECAUS IN CONIFEROUS TREES 225 probability of successful pupation to adult emergence. Further investigations are needed to clarify mortality factors acting during the immature stages of this spe- cles. The low density of larvae and adults, and the low Lloyd’s patchiness value suggest that little interference occurs among beetle adults or larvae. Although X. villioni larvae cause damage to branches of host trees by boring (Iwata et al. 1990, Adachi 1995), the lack of branches and the presence of a complete gallery of one individual in the sampled dead tree in Hinoemata indi- cates that this species can complete its development only in the tree trunk. Herein we have described the gross nature and spatial distribution of X. villioni larvae within tree trunks. Further investigations on the use of branches versus trunks by ovipositing females, movement of immature larvae from the branches into the trunk and a sensus of adult beetles are needed. ACKNOWLEDGMENT We thank Dr. Kazuaki Kamijo (formally, Hokkaido Forest Experiment Station) for his kind advice during our field work in Hokkaid6, Dr. Ko-ichi Soné, Tama Forest Science Garden, FFPRI (MAFF), Hachidyji (presently, Faculty of Agricul- ture, Kagoshima University), for his help in felling the sample tree at Hachidji, Messrs. Takahisa Maro, Itaru Suda, Yasuyuki Koma, Seiji Fujii and the other students of Nihon University, and Dr. AleS Smetana, Biosystematics Research Centre, Ottawa, for their help during the surveys in Hachidji, Hinoemata and Imakane. This work is supported by a Grant-in-Aid for Scientific Research (No.02660163) from the Ministry of Education, Science and Culture of Japan. LITERATURE CITED Adachi, K. 1995. Discovery of Xylotrechus villioni (Villard) (Coleoptera, Cerambycidae) from the mainland of Kyushu, Southwest Japan. Gekkan-Mushi, Tokyo, (293): 3-8 (in Japanese). Ganda, K., S. Shida & H. Konno. 1986. Forest damage caused by Xylotrechus villioni (Villard) in Imakane, Hokkaid6. Shinrin-Hogo (Forest Protection), Sapporo, (193): 19-21 (in Japanese). Iwata, R. 1991. Bionomics of a primary borer of coniferous trees, Xylotrechus villioni, a review. The Insectarium, Tokyo, 28: 108-113 (in Japanese). Iwata, R., I. Suda, EF Yamada & K. Nagata. 1993. Capture surveys of beetles by attracting chemicals at coniferous forests. (IV) Further survey at Abies firma grove in Tama Forest Science Garden (FFPRI, MAFF), Hachidji, Tokyo, by using kairomones. Trans. 44th Mtg. Kanto Branch, Jpn. For. Soc.: 119-122 (in Japanese). Iwata, R., EF Yamada, H. Ashida, K. Araya & A. Kawabata. 1992. Capture surveys of beetles by attracting chemicals at coniferous forests. (II) Survey at Abies firma grove in Kyoto University Forest, Ashiu, Kyoto, by using kairomones and pheromones. Trans. 103rd Mtg. Jpn. For. Soc.: 537-538 (in Japanese). Iwata, R., E Yamada, I. Suda, H. Makihara, K. Iwabuchi, & K. Nagata. 1991. Capture surveys of beetles by attracting chemicals at coniferous forests. (I) Survey at Abies firma grove in Tama Forest Science Garden (FFPRI, MAFF), Hachidji, Tokyo, by using kairomones and pheromones. Trans. 102nd Mtg. Jpn. For. Soc.: 261-264 (in Japanese). Iwata, R., E Yamada, M. Yagi, A. Kitayama, T. Kinoshita, K. Hosokawa, K. Kitayama, K. Iwabuchi & H. Makihara. 1990. Studies on Xylotrechus villioni (Villard) (Coleoptera: Cerambycidae), a primary borer of coniferous trees in Japan. (I) General bionomics. Trans. 101st Mtg. Jpn. For. Soc.: 525-528 (in Japanese). Kamijo, K., O. Ishizaka & S. H6j6. 1970. Damage on plantations of Abies sachalinensis by Xylotrechus villioni (Villard) (Cerambycidae). Hokkaidé Ringyé Kenkya Happyé6-kai Ronbunshi (Proc. For- estry Res. Symp. Hokkaid6), Sapporo, (fiscal 1969): 302-314 (in Japanese). Kamijo, K. & S. Suzuki. 1973. Damage to Abies sachalinensis Masters by Xylotrechus villioni Villard 224 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(4) (Cerambycidae). Hokkaid6 Ringy6é Shiken-j6 Hékoku (Bull. Hokkaido Forest Exp. Sta.), Bibai, (11): 113-119, plts. 1-4 (in Japanese). Kobayashi, K. 1982. Resin exudation of Cryptomeria japonica in relation to the attacks of Semanotus japonicus (Coleoptera: Cerambycidae). Trans. 33rd Mtg. Kansai Branch, Jpn. For. Soc.: 272- 275 (in Japanese). Kobayashi, K. & E. Yamada. 1982. Semanotus japonicus. In F. Kobayashi (ed.). Boring insects of Japanese cedar and cypress. Introduction to their ecology and control. S6bun Co. Ltd., Tokyo: 11—57 (in Japanese). Lloyd, M. 1967. ‘Mean crowding’. J. Anim. Ecol., 36: 1-30. Makihara, H., Y. Buyo & Y. Oka. 1995. Host trees and distribution of large tiger longicorn beetle, Xylotrechus villioni (Villard) in Tama Forest Science Garden. Trans. 46th Mtg. Kanto Branch, Jpn. For. Soc.: 107-110 (in Japanese). Received 17 Sep 1996; Accepted 6 Jan 1997. PAN-PACIFIC ENTOMOLOGIST 73(4): 225-230, (1997) A NEW JAPONICA (LEPIDOPTERA: LYCAENIDAE: THECLINAE) FROM SOUTHWESTERN CHINA YU-FENG Hsu Department of Biology, Taiwan Normal University, Taipei 117, Taiwan, Republic of China Abstract——Japonica bella Hsu, NEW SPECIES, is described and illustrated based on material from Guizhou Province, southwestern China. The new species has an unusual brachium and papilla analis, both are unique to the genus Japonica and the rest of Theclini species and con- sidered autopomorphies of J. bella. The discovery of the new species brings the number of species in genus Japonica to five. Key Words.—Insecta, Lepidoptera, Lycaenidae, Theclini, China, Japonica Theclini lycaenid butterflies of the genus Japonica generally inhabit oak forests in Old World temperate regions. The butterflies are active at twilight or during cloudy conditions (Fukuda et al. 1984). The members of Japonica have extensive orange scaling on the wings and possess many pnmitive characters (Shir6zu & Yamamoto 1956, Hsu & Lin 1994). Fujioka (1993) recently reviewed the genus Japonica and recognized four species. He also reported the presence of a characteristic “vaginal membrane’’, which is torn during copulation, on the ductus bursae in the female genitalia. Life histories and host associations of all four Japonica species have been docu- Figures 1-4. Japonica bella Hsu, NEW SPECIES. Fig. 1. Holotype male upperside. Fig. 2. Ho- lotype male underside. Fig. 3. Paratype female upperside. Fig. 4. Paratype female underside. 226 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(4) Figures 5—6. Venation of Japonica bella. Fig. 5. Forewing. Fig. 6. Hindwing (scale = 1 mm). mented: larvae of J. lutea (Hewitson) utilize both deciduous and evergreen Quercus (Fagaceae) species in Japan (Shiré6zu 1961) and deciduous Q. mongolica Fischeri in Far East Russia (Fujioka 1993); larvae of the closely related J. adusta (Riley) feed only on deciduous Q. dentata Thunberg in Japan (Inomata 1990) and Far East Russia (Fujioka 1993); larvae of J. saepestriata (Hewitson) are associated with deciduous Quercus species in most parts of Japan (Fukuda et al. 1984), except for a population in southern Ki-i Peninsula where evergreen Q. phillyraeoides Gray is utilized (Sai- gusa 1993); J. patungkoanui Murayama of Taiwan is known to use evergreen Quer- cus stenophylloides Hayata as the larval host (Uchida 1991). According to Fujioka (1993), three of the four known Japonica species have been found in southwestern China. An undescribed species was recognized from the above region and is described here. This new species is distinct from the previously de- scribed species in wing pattern and genitalia of both sexes. MATERIAL AND METHODS Genitalic dissections were made by removing the entire abdomen, which was placed in 10% KOH at room temperature for 24 hours, then transfered to cellu- L997 HSU: NEW JAPONICA FROM CHINA 227 Figures 7-12. Male genitalia of Japonica bella excluding phallus. Fig. 7. Lateral view of sclerites of 9 + 10 genitalic segments with left brachium and valva attached. Fig. 8. Dorsum of sclerites of 9 + 10 genitalic segments. Fig. 9. Posterior view of left brachium. Fig. 10. Ventral view of left brachium. Fig. 11. Posterior view of juxta. Fig. 12. Dorsal view of left valva. solve for another 24 hours for descaling, and finally placed in 70% ethyl alcohol for dissections. During examination, a few drops of xylene were placed on the wings to improve the contrast between veins and the covering scales. Japonica bella Hsu, NEW SPECIES (Figs. 1-17) Types.—Holotype, male; data: CHINA. GUIZHOU PROVINCE: Tongren Pre- fecture, Mt. Fanjing. 1000-1350 m, 18/19 Jun 1995; deposited: Zoological Insti- tute, Academia Sinica, Beijing. Paratypes: same data as holotype, 4 males; de- posited: Insect Museum, National Taiwan University, Taipei; California Academy of Sciences, San Francisco; 1 female; same locality as holotype, 23/24 Jun 1996; deposited: Zoological Institute, Academia Sinica, Beijing. Description.—Male (Figs. 1-2). Length of forewing 17.0-19.2 mm (mean = 17.96 + 0.84 mm, n = 5); Length of antenna 6.0—6.7 mm (mean = 6.38 + 0.28mm, n = 5). Head. Hairy, clothed with erect, dark brown hairs on vertex and frons, a white, narrow rim surrounding eye; eye semi-oval, sparsely hairy; labial palpus hairy, porrect, pointed, projecting ahead of plane of front; maxillary 228 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(4) Figures 13-15. Phallus of Japonica bella. Fig. 13. Lateral view of phallus. Fig. 14. Dorsal view of phallus. Fig. 15. Anterior portion of venter of phallus (scale = 1 mm). reduced, invisible; proboscis unscaled; antenna smooth-scaled, with projecting setae at nudum. Thorax. Pale brown clothed with chrome orange scaling dorsally; white ventrally, legs white, banded with dark brown on tarsi. Forewing. Eleven veins, R4+5, M1 forked with R3 (Fig. 5); termen, costa curved; dorsum straight; ground color of upperside chrome-orange with underside markings visible by trans- parency, margin, apex black; underside ground color pale chrome-orange with dark brown markings. Submarginal band parallel to termen, outlined by white scaling proximally and distally, intersected by orange scaling along veins. Discal band straight, slightly tilted inwards, marking in Cu2 indented. Two bars present in discoidal cell: distal one rectangular, basal one triangular. Fringe dark brown. Hindwing. Nine separate veins (Fig. 6). Termen produced at distal ends of veins, forming zig-zag outline. Ground color of upperside chrome-orange with dark brown outline. A patch of black scaling present at anterior corner along termen. A small, black dot present near distal end of Cul. Anal area slightly lobed, covered with metallic blue and black. Underside ground color chrome-orange with distal half darkened. Four transverse bands outlined with narrow, white lines; submarginal band sharply narrowed poste- riorly, forming a slender ‘“W’’-shaped marking around tornal area. Tornal area bright orange, a distinct, black dot present in cell Cul, a patch of black scaling mixed with metallic blue present at tornal lobe. Slender tail-like projection extending from Cu2, black with a white distal tip, approximately 6.5 mm in length. Abdomen. Chrome orange dorsally, white ventrally. Male genitalia (Figs. 7-15). Sclerites of 9th and 10th segments fused, forming a complete ring, width 0.60 X height. Tegumen 9 + 10 with dorsum fairly flat, slightly concave posteriorly with medial bump; uncus absent; socii folded deeply inwards; brachium double-articulated with tegumen, smooth, flattened, enlarged at base, abruptly nar- rowed, tapering to a posteriorly directed, hooklike process; saccus produced, approximately 0.48 X height of tegumen; phallus elongate, upcurved posteriorly, slightly asymmetrical with caudal end produced along right side; aedeagus 1.95 X phallobase; cornutus present, forming an elongate, trian- gular plate with sharp end near caudal end of aedeagus; valva semicircular with inward-curved, digitate posterior process ending with a terminal club; juxta narrow, U-shaped. Female (Figs. 3—4).—Forewing length 16.0 mm (n = 1); antennal length 6.0 mm (n = 1). Head. and Thorax. Structure, color pattern as described for male. Wings. Shape similar to male, but with straight termen; color pattern as described for male. Abdomen. Color as described for male. Female genitalia (Figs. 16-17). Apophyses posteriores elongate, slender, down-curved, ending with club- shaped anterior ends. Papillae anales with terminal, heavily sclerotized, bifid processes. Apophyses anteriores short, with blunt terminal ends, approximately one-third x length of apophyses posteriores. Sternite 8 divided, forming oval sclerites with postrior edges straight, serrate. Ductus bursae elongate, 1997 HSU: NEW JAPONICA FROM CHINA 229 Figures 16-17. Female genitalia of Japonica bella. Fig. 16. Ventral view. Fig. 17. Lateral view; arrow indicates the caudal processes at papilla analis (scale = 1 mm). forming heavily sclerotized tube at the posterior end, with point of origin of ductus seminalis just anterior to the sclerotized tube. Corpus bursae oval, bearing a pair of small, flattened, amoeboid- shaped signa. Diagnosis.—The pattern of four prominent bands or bars on the underside of the wings is unique among the species of Japonica. The enlarged brachium has a form not found in any members of the Theclini species. The caudal end of the valva is curved and clubed (Fig. 12) in J. bella, whereas it is straight and bifid in J. saepestriata and not clubed in J. lutea, J. adusta, and J. patungkoanui. The signa of J. bella are shallow and amoeboid-shaped, whereas those of the other Japonica species are invaginated and oval-shaped. The posterior edge of the sternite 8 is serrate, bearing numerous prom- inent teeth in J. bella, but smooth, without teeth in the other Japonica species. The terminal processes (Fig. 17) on papillae anales of J. bella are absent in the other Japonica members. Geographical Distribution.—Currently only known from Guizhou Province, southwestern China. Etymology.—An adjective of latin, from bella = beautiful. Discussion.—The most unusual character found in J. bella is the enormously basally enlarged brachium, which is not present in the other members of Japonica. Hsu & Lin (1994) considered “‘smooth and simple’”’ as the most plesiomorphic state for the shape of the brachium in the Theclini and assigned this state to the 230 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(4) genus Japonica in their phylogenetic analysis of a subgroup of Theclini. The discovery of J. bella could put such an assignment in jeopardy, but probably will not affect their overall decision on the polarity of character states in the shape of brachium in Theclini. The enlarged part of the brachium in J. bella is flattened along a plane perpendicular to the axis of the terminal narrow portion (Figs. 9— 10). Such a form of enlargement is unique among members of Theclini and is clearly an autopomorphy of J. bella. The sole possession of bifid caudal processes in female genitalia of J. bella is evidently also an autopomorphy of J. bella. ACKNOWLEDGMENT I thank James E. Baxter and Douglas E. Kain, Division of Insect Biology, University of California, Berkeley and C. Don MacNeill, Department of Ento- mology, California Academy of Sciences, San Francisco for critically reading the manuscript. LITERATURE CITED Fujioka, T. 1993. Zephyrus (Theclini butterflies) in the world (4). -Genus Japonica-. Butterflies, 5: 13-31. Fukuda, H., E. Hama, T. Kuzuya, A. Takahashi, M. Takahashi, B. Tanaka, H. Tanaka, M. Wakabayashi, & Y. Watanabe. 1984. The life histories of butterflies in Japan. Vol. III. Hoikusha, Osaka. Hsu, Y. E & M. Y. Lin. 1994. Systematic position of Sibataniozephyrus and description of a new species from Taiwan (Lycaenidae: Theclini). J. Lepid. Soc., 48: 128-147. Inomata, T. 1990. Keys to the Japanese butterflies in natural color. Hokuryukan, Tokyo. Saigusa, T. 1993. A study on new subspecies of Tribe Theclini from eastern Asia. Zephyrus Research, 1: 12-21. Shirézu, T. 1961. Evolution of the food-habits of larvae of the Thecline butterflies. Ty6 to Ga, 12: 144-162. Shirézu, T. & H. Yamamoto. 1956. A generic revision and the phylogeny of tribe Theclini (Lepidop- tera; Lycaenidae). Sieboldia, 1: 329-421. Uchida, H. 1991. Charms of Formosa, Island of Everlasting Summer. Self-published, Numazu City. PAN-PACIFIC ENTOMOLOGIST 73(4): 231-235, (1997) XYELA (PINICOLITES) LATA SMITH (VESPIDA: XYELIDAE), A LIVING FOSSIL SAWFLY FROM WESTERN NORTH AMERICA ALEXANDR P. RASNITSYN Paleontological Institute, Russian Academy of Sciences, Moscow, 117647 Russia Abstract.—Xyela lata Smith is an extant member of Pinicolites Meunier, 1920, a subgenus previously known from a single fossil species from Tertiary deposits of Germany. The species, known from the highlands of Colorado, Nevada, and Oregon, is first recorded from California (Sierra Nevada Mountains, altitude 2,560—2,800 m). Key Words.—Insecta, Xyelidae, Tertiary relict, living fossil Xyelidae is one of the oldest insect families, whose fossil record starts as early as at the Middle or Late Triassic. The genus Xyela Dalman is also an ancient as it first appeared in the Lower Cretaceous (Rasnitsyn 1969). The fossil history of Xyela was recently reviewed by Rasnitsyn (1995) who recognized three subgenera: Mesoxyela Rasnitsyn (1965) (with one Early Creta- ceous species from Zaza lake sediments in Transbaikalia), Pinicolites Meunier (1920) (one species from the mid-Tertiary (Aquitanian) of Germany near Bonn), and Xyela s.s. (five mid-Tertiary and 31 extant species). Pinicolites is found now to persist since mid-Tertiary until the present. Recently I have examined two specimens of X. Jata in the collection of the California Academy of Sciences, San Francisco, and found the species to be a member of Pinicolites. The species was not recognized as a member in the orig- inal description of this subgenus, so it is redescribed here to include important characters not visible on the fossil specimens. Discovery of the living fossil was presented to the general public in the Sep 1995 edition of the San Francisco Chronicle. The reporter contacted Donald J. Burdick, the collector, who provided details about the habitat. GENUS XYELA DALMAN SUBGENUS PINICOLITES MEUNIER Finding of an extant species made my previous diagnosis of Pinicolites obso- lete. A new diagnosis follows. Diagnosis.—Pinicolites differs from Xyela s.str. and Mesoxyela in having an Ovipositor that is both flat (saw-like) and upcurved. In addition, RS either touches M in a point, or is connected to it by short lr-m crossvein (fused with M fora distance in Mesoxyela and a majority of Xyela s.str.). Within Xyela s.1., Pinicolites has a unique combination of a lightly colored mesoscutum (with black spots that indicate muscle attachments) a short antennal flagellum (shorter than or subequal to the article III and about 0.7 as long as the head width). In Mesoxyela, the mesoscutum is black and the antennal flagellum short, and in Xyela s.str. the antennal flagellum is longer and the mesoscutum is usually lightly colored. Pin- icolites is similar to Mesoxyela and differs from Xyela s.str. in having free stalk of SC. Judging from the extant species only, Pinicolites differs from Xyela s.str. 232 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(4) Figure 1. Xyela lata Smith, specimen from Kaiser Pass. and Mesoxyela also in having antennal flagellum 11-segmented, mesonotum sur- face smooth, unsculptured (except for minute, sparse punctures), and, for the Ovipositor blades, in the dorsal valve lacking visible structures, and the ventral valve having a narrow, acute apical projection that is abruptly separated from the main valve portion. Xyela (Pinicolites) lata Smith (Figs. 1-4) Xyela lata Smith, 1990: 9. Description—Female. Length of forewing 4.4 mm, of sawsheath 2.3-2.5 mm. Integument shiny, asetose, unsculptured (except for scattered, minute punctures and pubescent abdominal apex). Body yellow with usual dark pattern (evanescent in part in one specimen); brown (sometimes with red tint) on head: antennomere III and flagellomeres, mandibular apex, lines along and spot between ocellar- antennal furrows, interocellar and postocellar area, spots laterad of postocellar sutures, posterior ocular orbits, and posterior head surface above; on thorax: pronotum, propleuron (except yellow caudally along thoracic midline), mesopleuron anterad of pseudosternal suture, pseudosternum except rostrally, laterally, mesonotum narrowly along notauli, medial, and scutellar furrows and along posterior scutellar margin, lateral and sublateral spots of mesonotum; additionally brown are dorsal body surface posterior of mesonotum disc (except for whitish cenchri, and yellowish metascutellum and abdominal tergal margins), and sawsheath apex. Pterostigma and veins pale yellow, membrane hyaline. Maxillary palp with article III as wide as and almost as long as antennal article III, with article V bearing 4 curved setae submedially, and membranous flap adaxially, subapically. Wing venation (Fig. 2), with several irregularities, viz., one specimen has an extra A, stub on left hindwing, another a supernumerary rs,- rs, crossvein in right forewing, its rudiment in left forewing, and r-rs crossvein in left hindwing. Sawsheath slightly upcurved in basal half, almost straight distally, widest subbasally, tapering slightly both basally and distally, tapering abruptly toward apex that is narrowly rounded, almost symmetrical. Basal plate of ovipositor (2nd valvifer) short: externally scarcely longer than maximum height of sheath. Ventral valve (V,) high, membranous except for simple dorsal longitudinal thickening, abruptly 1997 RASNITSYN: XYELA LATA, FOSSIL SAWFLY 233 Figure 2. Xyela lata Smith, line drawing of Kaiser Pass specimen (legs omitted). Scale line = 1 mm. Figure 3. Xyela lata Smith, wings and external view of the ovipositor of Bodie specimen. Lon- gitudinal veins are capitalized, crossveins are hyphenated; cr—cercus, h—hamuli, pt—pterostigma, to, t;> abdominal terga, v,—sawsheath, vr,—second valvifer (basal plate of ovipositor). Scale line = 1 mm. 234 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(4) Figure 4. Xyela lata Smith, ovipositor blades of Kaiser Pass specimen (v, bottom, v, top). narrowed toward needle-like apex armed with 4 oblique, toothed ribs, which are facing cephalad. Dorsal valve (V,) also high, membranous and lacking visible structure (except for narrow ventral longitudinal rib not reaching apex of valve); narrowing gradually toward simple, hardly visible apex. Male.—Unknown. Diagnosis.—Xyela lata differs from X. graciosa Meunier 1920, in being slight- ly larger (forewing length 4.0—4.4 mm) and in having the following: antennal flagellum somewhat longer than article III, forewing with SC not reaching the level of RS base, sawsheath more upcurved, widest subbasally, and shorter (equal to distance from forewing R base to lr-rs). In X. graciosa the forewing length is 3.5 mm, antennal flagellum shorter than article III, forewing with SC reaching the level of RS base, sawsheath less curved, widest basally, and as long as the forewing from R base to 2r-rs. Biology.—One of females was collected ‘“‘flying around the [male] cone of a white bark pine tree [Pinus albicaulis Engelm.] at an elevation of 9,500 feet [= 2,900 m]’’ (observation by Donald J. Burdick, cited in the San Francisco Chron- icle, Sept. 4, 1995, p. 16). Material Examined—CALIFORNIA. FRESNO Co.: Kaiser Pass, 2900 m, 13 Jun 1966, D. J. Bur- dick, 1 female. MONO Co.: Bodie [2560 m], 12 Jun 1937, W. C. Bush. ACKNOWLEDGMENT My visit to California was made possible through a grant from the International Society of Hymenopterists, and also through friendly hospitality by Wojciech J. Pulawski and Veronica Pulawski, San Francisco, California. Wojciech J. Pulawski, Curator, California Academy of Sciences, San Francisco, California, also offered research facilities in the Department of Entomology, California Academy of Sci- ences. The photograph of the ovipositor blades is through the courtesy of Charles E. Griswold and Darrel Ubick of the same Department. The photograph of the whole specimen was provided by Susan Middleton, formerly of California Acad- emy of Sciences. I sincerely thank David R. Smith of the Systematic Entomology 1997 RASNITSYN: XYELA LATA, FOSSIL SAWFLY 235) Laboratory, USDA, who helped identify the species. An anonymous reviewer pointed out several important deficiences in the early version of the article. LITERATURE CITED Meunier, F 1920. Quelque insectes de |’ Aquitanien de Rott, Sept.-Monts (Prusse rhénane). Koninklijke Akademie van Wettenschappen te Amsterdam, Proceedings, Section of Science, 22: 891-898. Rasnitsyn, A. P. 1969. Origin and evolution of Lower Hymenoptera. Trans. Paleontological Institute, Acad. Sci. U.S.S.R., 123 (in Russian; translated into English in 1979 by Amerind Co., New Delhi). Rasnitsyn, A. P. 1971. Evolution of Xyelidae (Hymenoptera). Jn Current problems in paleontology. Trans. Paleontological Institute, Acad. Sci. U.S.S.R., 130: 187-196 (in Russian). Rasnitsyn, A. P. 1995. Tertiary sawflies of the Tribe Xyelini (Insecta: Vespida = Hymenoptera: Xye- lidae) and their relationship to the Mesozoic and modern faunas. Contributions in Science, Natural History Museum of Los Angeles County, 450: 1-14. Smith, D. R. 1990. A new Xyela (Hymenoptera, Xyelidae) from the Western United States. Entomol. News, 101: 9-12. Sphon, G. G. 1973. Additional type specimens of fossil Invertebrata in the collections of the Natural History Museum of Los Angeles County. Contributions in Science, Natural History Museum of Los Angeles County, 250: 1-75. Statz, G. 1936. Uber alte und neue fossile Hymenopterenfunde aus den tertiiren Ablagerungen von Rott am Siebengebirge. Decheniana, 93: 256-312. Received 26 Apr 1996; Accepted 30 Dec 1996. PAN-PACIFIC ENTOMOLOGIST 73(4): 236-242, (1997) Scientific Note INTRODUCTION OF WESTERN ASIAN EGG PARASITOIDS INTO CALIFORNIA FOR BIOLOGICAL CONTROL OF BEET LEAFHOPPER, CIRCULIFER TENELLUS In North America, beet leafhopper, Circulifer tenellus (Baker), is the only known vector of curly top virus which affects a wide variety of crop species, including tomatoes, sugar beets, peppers, melons, spinach, and beans. Curly top virus has been a major economic problem in sugar beet and vegetable production in the western United States for almost 100 years (Bennett, C. W. 1971. The Curly Top Disease of Sugarbeet and Other Plants. Amer. Phytopath. Soc. Monogr. 7). To reduce losses caused by curly top virus, the California Department of Food and Agriculture (CDFA) has been conducting a control program since 1943 to reduce regional populations of the beet leafhopper vector in California. In addition to beet leafhopper’s importance as the sole vector of curly top virus in North America, it is also a vector of citrus stubborn disease, a serious plant pathogen affecting citrus (Oldfield, G. N., Kaloostian, G. H., Pierce, H. D., Calavan, E. C., Granett, A. L., & Blue, R. L. 1976. Calif. Agric. 30[6]: 15). Beet leafhopper has a strong migratory habit and is polyphagous with over 30 plant species from several plant families serving as reproductive hosts, and per- haps 60 or more additional species on which the leafhopper can feed (Severin, H. H. P. 1933. Hilgardia 7: 281-360, Cook, W. C. 1967. U.S.D.A. Agr. Res. Service Tech. Bull. 1365, Bennett 1971, and personnel in CDFA’s Curly Top Virus Control Program). Beet leafhopper is believed to be native to the arid and semi- arid areas of central Asia. Both its migratory and polyphagous habits are adap- tations to arid and semi-arid regions, where its success as an herbivore is largely a result of its ability to abandon areas undergoing seasonal drying and to move to locations where the vegetation is still green. Location of green vegetation at different times of the year is a daunting challenge in these arid and semi-arid habitats even without having to locate specific plant species; thus, the leafhopper’s polyphagous habits complement its migratory habits to make this species a suc- cessful nomad in areas characterized by ephemeral host plant patches that occur in different locations throughout the year. In North America, beet leafhopper is a serious pest only in the western part of the United States where the leafhopper’s migratory and polyphagous habits make it well suited for the generally arid and semi-arid climate of this region (Cook 1967, Johnson, C. G. 1969. Migration and Dispersal of Insects by Flight. Methuen & Co. pub.). In California’s San Joaquin Valley, beet leafhopper’s annual migratory cycle is summarized as follows (Cook 1967, Johnson 1969, and personnel in CDFA’s Curly Top Virus Control Program). Adults in reproductive diapause congregate for overwintering in the foothills of the coastal range on the west side of the valley. Here they subsist on a wide variety of perennial plants that are still green after the long hot dry summer, but they do not reproduce on these perennial “holdover hosts.’’ When winter rains stimulate germination of annual plants in 1997 SCIENTIFIC NOTE 237 the foothills, the leafhoppers move to these annuals and begin to oviposit in them, generally in February. The most important of the winter/spring annual host plants for beet leafhopper reproduction are filaree (Erodium sp.), plantain (Plantago sp.), and peppergrass (Lepidium sp.). Winter/spring rains are generally erratic and usu- ally cease by the end of March, causing the winter/spring annual host plants in the foothills to die, and thus forcing the leafhoppers to migrate down into the valley where a new set of host plants are now growing. The uncultivated host plants now include London rocket (Sisymbrium irio L.), mustards (Brassica spp.), goosefoot, pigweed, and lamb’s quarters (Chenopodium spp.), Russian thistle (Sal- sola tragus L.), annual saltbushes (Atriplex spp.), Bassia sp., and Kochia sp. These plants are generally concentrated in disturbed habitats along roadsides, in culti- vated fields, and along irrigation ditches. Many of these late-spring hosts such as Sisymbrium, Brassica and Chenopodium die during the dry summer unless they occur in irrigated fields, but some such as Russian thistle, a few annual saltbushes, Bassia sp., and Kochia sp. survive even in non-irrigated locations throughout the summer and into the early fall. In early summer, beet leafhoppers abandon the dying short-lived uncultivated spring hosts and move to the remaining longer- lived uncultivated host plants or migrate into cultivated host plants, bringing curly top virus with them. In the fall, the uncultivated summer hosts (Russian thistle, annual saltbushes, Bassia, and Kochia) finally mature and die, and the cultivated hosts (as well as weed hosts in cultivated fields) are removed by harvesting or disking, forcing the leafhoppers to migrate back to the perennial holdover hosts in the foothills where they overwinter and complete the annual cycle. Beet leaf- hopper has similar types of migratory cycles in other regions of western North America. Because of the migratory nature of beet leafhopper, control strategies on a field- by-field basis are not effective at reducing the spread of curly top virus. Crops become inoculated with virus from leafhoppers migrating into the fields from surrounding wild vegetation. Thus, a regional control strategy rather than a local field-by-field strategy is the most efficient way to reduce the incidence of curly top virus in the affected agricultural crops. The primary strategy of CDFA’s Curly Top Virus Control Program is to reduce regional populations of beet leafhopper by spraying malathion in the leafhopper’s overwintering and spring breeding grounds in wild vegetation (mostly rangeland in the San Joaquin Valley and desert in the Imperial Valley). CDFA is currently funding attempts to utilize biological control agents as an alternative regional control strategy in the beet leafhopper’s wild vegetation habitats. As a component of the biological control strategy, we have made three foreign exploration trips to Asia in search of natural enemies of beet leafhopper, two in the former Soviet republic of Turkmenistan in May-June 1992 and September 1994, and one to Iran in June-July 1995 (Fig. 1). The focus of the exploration trips was on egg parasitoids. This report documents the im- portation of at least 11 different species of egg parasitoids of leafhoppers from Asia, the release of seven of these in California, and the establishment of at least one of these species. The migratory nature and broad host plant range of beet leafhopper make it a particularly difficult target for biological control. Because the leafhopper does not stay in one place, successful natural enemies will have to either 1) remain in the same habitat and aestivate during the times of year when beet leafhopper is not 238 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(4) Figure 1. Map of the Iran—Turkmenistan region where parasitoids of beet leafhopper were collected and shipped to California during foreign exploration in May—June 1992 (Turkmenistan), September 1994 (Turkmenistan), and June—July 1995 (Iran). Collection sites of the species listed in Tables 1 and 2 are marked with asterisks on the map. present; 2) remain in the same habitat and locate an alternative leafhopper host during the times of year when beet leafhopper is not present; or 3) follow the beet leafhopper on its migratory route. Another difficulty for biological control of beet leafhopper relates to the diversity of host plants that the leafhopper utilizes throughout its migratory cycle. Natural enemies that are effective on one of the host plants are not necessarily effective on a different host plant. For example, the native mymarid parasitoid Anagrus nigriventris Girault is an effective para- sitoid of beet leafhopper eggs in unsprayed sugar beet fields in California (Mey- erdirk, D. E. & Hessein, N. A. 1985. J. Econ. Entomol. 78: 346-353, Meyerdirk, D. E. & Moratorio, M. S. 1987. J. Econ. Entomol. 80: 362—365); however in our studies, this parasitoid was ineffective when it was mass-reared and released in beet leafhopper habitats in the foothills of the west side of the San Joaquin Valley, where the beet leafhopper host plant complex consisted of filaree, plantain, and peppergrass, and in Oildale, California, where the beet leafhopper host plant com- plex consisted of Russian thistle and an annual saltbush species (Triapitsyn, un- published data). 1997 SCIENTIFIC NOTE 239 Table 1. List of egg parasitoids emerging from the plant samples collected in Turkmenistan (1992), the host plants from which they emerged, and locations from where they were collected. Host plants from which ey Parasitoid species they were collected Nearest point Province Family Mymaridae Anagrus atomus red beet Bayram-Ali Mary red beet Giami Ashgabat red beet Enev Ashgabat Erythmelus mar gianus Atriplex sp. Old Nisa Ashgabat Atriplex sp. and Salsola sp. Bayram-Ali Mary Atriplex sp. Ashgabat Ashgabat Gonatocerus sp(p). Atriplex sp. and Salsola sp. Bayram-Ali Mary Polynema sp(p). Atriplex sp. and Salsola sp. Bayram-Ali Mary Atriplex sp. Old Nisa Ashgabat Family Trichogrammatidae Aphelinoidea turanica* Atriplex sp. Ashgabat Ashgabat Atriplex sp. Old Nisa Ashgabat Atriplex sp.* Bayram-Ali Mary Salsola sp. Bayram-Ali Mary Ufens sp. Atriplex sp. and Salsola sp. Old Nisa Ashgabat Atriplex sp. Bayram-Ali Mary Multiple lines for a species indicate that the species was collected from a number of different host plants and/or locations. * Only the Aphelinoidea turanica collected from Atriplex sp. in Bayram-Ali was successfully prop- agated in quarantine, released and established in the San Joaquin Valley; all of the other species listed in Table 1 died in quarantine before they could be released. Our objective is to establish biological control of beet leafhopper in desert and semi-arid rangeland habitats of beet leafhopper, where the host plant complex consists of a broad range of non-cultivated plants. Therefore, we focused our foreign explorations in arid and semi-arid climatic zones, and more on non-cul- tivated vegetation than on agricultural crops. This strategy was intended to obtain natural enemies adapted to the less succulent wild vegetation rather than to irri- gated crop species. Also, we collected egg parasitoids from a diversity of plant species (Tables 1 and 2) in order to obtain a complex of parasitoids that would attack beet leafhopper eggs over a range of host plants. Leafhoppers lay their eggs embedded in host plant tissue. To obtain parasitized beet leafhopper eggs, host plants or parts of host plants of beet leafhopper were collected from areas where beet leafhoppers were present (determined by sam- pling nymphs and adults) in various locations in Turkmenistan and Iran (Fig. 1). Collected plant materials were sealed in plastic bags or in boxes within plastic bags, refrigerated, and then shipped to the entomology quarantine facility at the University of California, Riverside (UCR). In the quarantine facility, many insects emerged from the plant samples. Newly emerged adult wasps that were from taxonomic groups likely to be leafhopper egg parasitoids (mostly Mymaridae and Trichogrammatidae) were then collected alive and placed in small cages contain- ing sugar beet plants with large numbers of beet leafhopper eggs embedded in the plant tissue as described by A. K. Al-Wahaibi and G. P. Walker (in press. Bull. Entomol. Res.). There, the parasitoids had an opportunity to oviposit in the Table 2. List of egg parasitoids emerging from the plant samples collected in Iran (1995), the host plants from which they emerged, and locations from where they were collected. Identification Host plants from which rs Parasitoid species number* they were collected Nearest city Province Family Mymaridae Anagrus atomus** 45-An sugar beet Karaj Tehran Polynema sp. 1 10-P sugar beet Karaj Tehran Polynema sp. 2 41,42,43-P Chenopodium sp. & Salsola sp. Karaj Tehran 45-P sugar beet Karaj Tehran Gonatocerus sp. 1A 36-GL Kochia sp. Atar, Neyshabur Khorasan Gonatocerus sp. 1B** 15-GLX Atriplex sp. Shams-abad, Neyshabur Khorasan Gonatocerus sp. 2 11-GD Salsola sp. Karaj Tehran 23-—GD Artemisia sp. between Mashhad and Khorasan Ghuchan; 35 km from Mashhad Family Trichogrammatidae Aphelinoidea turanica 15,25-Ap Atriplex sp. & Chenopodium sp. Shams-abad, Neyshabur Khorasan Aphelinoidea anatolica 3-Ap Chenopodium sp. Akbar-abad, Kavar Fars Oligosita sp. 36-O Kochia sp. Atar, Neyshabur Khorasan 45-O sugar beet Karaj Tehran * Identification numbers refer to separate cultures of parasitoids and to the voucher specimens (deposited in the Entomology Research Museum at the University of California, Riverside) for those cultures. Three of the parasitoid species (Polynema sp. 2, Gonatocerus sp. 2, and Oligosita sp.) have two cultures each, with each culture originating from a different host plant and/or location. ** Died before they could be released; no longer available. OVC LSIDO'TONOLNA OWIOVd-NVd AHL (PEL TOA 1997 SCIENTIFIC NOTE 241 beet leafhopper eggs and initiate a quarantine culture. Parasitoid cultures then were maintained on beet leafhopper-infested sugar beets kept at approximately 25 + 2° C and ~ 60% relative humidity. The first trip to Turkmenistan in May-June 1992 yielded the egg parasitoids Anagrus atomus (L.) (Mymaridae), one or more species of Gonatocerus (My- maridae), one or more species of Polynema (Mymaridae), a species of Ufens (Trichogrammatidae), and two previously unknown species, Aphelinoidea turan- ica Trjapitzin (Trichogrammatidae) and Erythmelus margianus Trjapitzin (My- maridae). Most of the collecting sites were in oases, the Kara Kum desert, and the foothills of low mountains along the Iranian border (Fig. 1). The collections were focused on Chenopodiaceae, especially Atriplex and Salsola spp., and ad- ditional collections were made from Plantago sp. and cultivated red beets (Beta vulgaris L.) (Table 1). Voucher specimens for the parasitoids have been deposited in the Entomology Research Museum at the University of California, Riverside. Of the parasitoids listed in Table 1, only A. turanica survived through the quar- antine process. Aphelinoidea turanica was successfully reared on beet leafhopper eggs, and was released between 30 Apr and 11 Jun 1993 in an abandoned gravel quarry alongside Highway 65 near Oildale, California where the beet leafhopper host plant complex consisted of Russian thistle, annual saltbush, filaree, and mus- tards. The release area was sampled for A. turanica from 24 Jul (at least two generation times after the last release date) to 19 Aug 1993 and again from March through May 1994. Aphelinoidea turanica were recovered from the 1993 and 1994 samples, indicating that it successfully established and overwintered at this location. The second trip to Turkmenistan in September 1994 was not as successful. Only a few A. turanica emerged from samples collected from drying Salsola and Atriplex spp. Unfortunately, these were all males, and consequently, this collection of A. turanica died in quarantine. The third trip to Iran in June—July 1995 was very successful, and nine species of leafhopper egg parasitoids emerged from the plant samples and reproduced on beet leafhopper eggs in sugar beet plants in the UCR entomology quarantine facility (Table 2). The host plants that were collected in Iran and shipped to the quarantine facility at UCR were: Amaranthus sp., Artemesia sp., Atriplex sp., Beta vulgaris (sugar beet), Chenopodium sp., Kochia sp., Plantago sp., Polygonum sp., Salsola sp., and Zantium sp. Of the listed parasitoid species, A. atomus died in quarantine and was never released. It reproduced poorly on beet leafhopper eggs in the sugar beet plants, suggesting that beet leafhopper may not be a good host for A. atomus, and that it may have emerged from other species of leafhopper eggs present in the plant samples from Iran. Gonatocerus sp. 1B also died in quarantine and, like A. atomus, it did not reproduce vigorously on beet leafhopper eggs and also may have emerged from other species of leafhopper eggs present in the plant samples from Iran. The remaining parasitoid species, Gonatocerus sp. lA & 2, Polynema sp. 1 & 2, A. turanica, Aphelinoidea anatolica Nowicki, and Oligosita sp. (Trichogrammatidae), all reproduced well on beet leafhopper eggs in quarantine. The two surviving Gonatocerus species are thelytokous and all other species are arrhenotokous. The two Polynema species originated from different collection locations and were originally thought to represent a single species. However, scanning electron microscopy revealed some subtle morpho- 242 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(4) logical differences between the two, and subsequent cross-mating tests revealed an inability to cross-breed, hence establishing the two Polynema as separate spe- cies. Unfortunately, there are no taxonomists working on Gonatocerus, Polynema, or Oligosita, and thus species identifications are not possible at this time. Voucher specimens have been deposited in the Entomology Research Museum at UCR. Reference numbers for these voucher specimens are given in Table 2. We have been releasing the seven surviving parasitoid species in beet leafhop- per-infested sugar beets grown at UCR’s Agricultural Operations in Riverside since December 1995, in wild vegetation breeding areas of beet leafhopper in the San Joaquin Valley since February 1996, in wild vegetation breeding areas of beet leafhopper in Fresno County, California since September 1996, and in wild vegetation breeding areas of beet leafhopper near Hemet, California since October 1996. To date, we have recovered Oligosita sp. from sugar beets and Polynema sp. from Chenopodium in Riverside, but it is premature to determine whether or not they have established a viable population. We have not yet made any recov- eries of the introduced species from any of the other release areas. Releases of the introduced species and sampling to determine establishment is planned to continue for at least another year. Acknowled gment.—The authors thank Bob Peterson and Rod Clark of the Cal- ifornia Department of Food and Agriculture for critically reviewing the manu- script and for assistance in choosing release sites for the parasitoids. This project is funded by a grant from the California Department of Food and Agriculture Curly Top Virus Control Program. G. P. Walker, N. Zareh, I. M. Bayoun & S. V. Triapitsyn, Department of En- tomology, University of California, Riverside, California 92521 Received 9 Dec 1996; Accepted 26 Feb 1997. PAN-PACIFIC ENTOMOLOGIST 73(4): 243-244, (1997) Scientific Note NOTES ON BRACHYCAUDONIA ASHMEAD SPP. (HYMENOPTERA: PTEROMALIDAE) Two species have been described in the genus Brachycaudonia (Hymenoptera: Pteromalidae), B. californica Ashmead and B. cyaniceps Boucek. In the descrip- tion of the latter, Boucek (1993. J. Nat. Hist. 27: 1239-1313), noted there were no host records available for either species. I have examined Brachycaudonia material from the California Academy of Sciences, San Francisco, California, Bohart Museum of Entomology, University of California, Davis, Essig Museum of Entomology, University of California, Berkeley, California, Oregon State Uni- versity, Corvallis, Oregon, United States National Museum, Washington, D.C., and my personal collection. Specimens from California exhibit a range of mor- phological characters and may represent a complex of two or more species (S. Heydon, pers. comm.). However, for the purposes of this paper, I regard them as belonging to a single species, B. californica. Specimens of B. californica have been reared from oak galls formed by several species of Cynipidae (Hymenoptera). These include galls of Dryocosmus dubiosus (Fullaway) on Quercus agrifolia Neé (both leaf galls formed by the unisexual generation, and flower galls formed by the bisexual generation), and galls of Andricus occultatus (Weld) on Q. lobata Neé. These specimens were collected in every month (except December) from September through May, from the following California counties: Alameda, Contra Costa, Fresno, Los Angeles, Marin, Mon- terey, Napa, Riverside, San Joaquin, San Mateo, Santa Barbara, Santa Clara, So- lano and Tulare. A male B. californica was also reared in April from a gall of Dros pedicellatum (Kinsey), and a female from an unknown gall, from Benton County, Oregon. In March 1995 I collected three adult B. californica females from the foliage of a Q. agrifolia, and exposed them to nine D. dubiosus leaf galls (both light and dark galls) on leaves collected from the same tree, 2 days later. The first female antennated both dark and light galls. On five of these, she stood on top of the gall and inserted her ovipositor about midway along the center. The ovipositor did not appear to be deeply inserted into the gall, but the time of insertion varied from a few seconds to 1—2 min. The second female explored the galls with the tip of her ovipositor sheath and antennae, but never drilled. The third female evinced no interest in the galls. One day later I dissected all nine galls. Two of the five galls drilled by the first female each contained a small larva with an egg laying on them. None of the other three drilled galls contained eggs—two each contained a large larva and the other was empty. Of the undrilled galls, one contained a torymid pupa. I collected additional D. dubiosus leaf galls from the same tree during 1996. A male B. californica emerged between 14-18 Feb 1997, from a gall collected on 13 Jul 1996. I also found specimens of B. cyaniceps from Kansas, Missouri, Maryland and Pennsylvania (representing a range extension for this species that was formerly 244 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(4) reported only from New York and Ontario) but rearing data are not available. The specimen from Maryland is a male, 1.4 mm long, and largely resembles the female, except for the following characteristics: antennae only slightly clavate; propodeal plicae absent; relative length of marginal vein 13, stigmal vein 10, postmarginal vein 13; gaster slightly longer than mesosoma; entire dorsum of gaster non-metallic light brown; head and mesosoma concolorous, maculation of wing lighter. This specimen was taken from a malaise trap, so the true colors may be different from those reported here. Two Brachycaudonia specimens from Minnesota and Michigan resemble B. cyanice ps, but are without rearing data. Thus, in the western United States Brachycaudonia is associated with cynipid galls from Quercus species, probably as an ectoparasitoid. Interestingly, one spec- imen each of B. californica and B. cyaniceps was collected from a peach tree or orchard. However, it is unknown if these species are primary or secondary par- asitoids, or if they attack gall-makers or inquilines. I have also reared Torymus fullawayi (Huber) (Hymenoptera: Torymidae), a ?Brasema sp. (Hymenoptera: Eu- pelmidae), two Aprostocetus spp. (Hymenoptera: Eulophidae), and a Synergus sp. (Hymenoptera: Cynipidae) from D. dubiosus leaf galls on Q. agrifolia. Of special interest is the rearing of B. californica from galls formed by both the unisexual and bisexual generations of D. dubiosus. Askew (1961. Trans. Soc. Brit. Entomol. 14: 237-268) noted that the same parasitoid species seldom at- tacked alternating generations of the same cynipid host, as the gall’s structure, placement on tree and the season of growth were the most important factors in determining its parasitoid complement, and these varied between alternating gen- erations. However, Doutt (1959. Ann. Entomol. Soc. Am. 52: 69-74), noted that D. dubiosus flower galls are found from February-May in California, and I have noted leaf galls in March and April. Presumably, this seasonal overlap in gall occurrence facilitates the transfer of B. californica between the alternating gen- erations of the cynipid. Acknowledgment.—I thank Eric Grissell of the Systematic Entomology Labo- ratory, % United States National Museum, Jim Lebonte of Oregon State Univer- sity and Steve Heydon of the University of California, Davis for their assistance in providing specimens, and Ken Hagen of the University of California, Berkeley for reviewing the manuscript. Robert L. Zuparko, Center for Biological Control, 201 Wellman Hall, Univer- sity of California, Berkeley, California 94720. Received 31 Oct 1996; Accepted 5 Feb 1997. PAN-PACIFIC ENTOMOLOGIST 73(4): 245-247, (1997) Scientific Note NATURALLY OCCURRING INFESTATIONS OF DRYWOOD TERMITES IN BOOKS Drywood termites of the family Kalotermitidae (Isoptera) occur throughout the tropics and subtropics (Snyder, T. E. 1949. Smith. Misc. Coll. 112: 1-490) with a few species occurring 1n warmer temperate regions of the world. All members of the Kalotermitidae are essentially wood dwellers with the exception of Para- neotermes simplicicornis (Banks), which exhibits a semi-subterranean habit (Gul- mahamad, H. 1995. Pan-Pacif. Entomol. 71: 105-109). Incisitermes fruticavus Rust, which occurs in California, has been reported to infest living plants (Rust, M. K. 1979. Pan-Pacific Entomol. 55: 273-278). Recent evidence suggests that I. fructicavus also attacks structures in San Diego County (Michael K. Rust, per- sonal communication). Incisitermes minor (Hagen) is the most common drywood termite in western North America. Its distribution is mainly confined to California, portions of Ar- izona and northern Mexico. Isolated infestations of J. minor have also been re- ported from Washington, Utah and many parts of Canada. In many areas where itis endemic, J. minor is the most destructive drywood termite species. In southern California, where infestations of J. minor are very common in wooden structures, more money is spent controlling this species than subterranean termites. Natural infestations of J. minor largely occur in sound, dry, wood. In California, infestations are commonly found in dead trees and shrubs and in dead portions of living trees and shrubs. Infestations are also found in wooden buildings and other wooden structures. Over the past 19 years of field work in southern California, I have found infestations of 7. minor in mobile homes, recreational vehicles, two classic auto- mobiles (a 1932 Ford Huckster and a Woody), trucks, boats, ships, poo! tables, planos, spas, gazebos, wooden water tanks, power poles, fence posts, furniture, wooden pallets, wooden crates, wooden tool handles, decorative wooden statues, wood carvings, totem poles, wooden crosses, picture frames, firewood, scrap lum- ber, and wooden desks. Many of these infestation records have not been previ- ously reported for this species. It is common to find infestations and damage of subterranean termites in books and other paper products. These are well documented in the literature. In fact, subterranean termite damage to books, magazines, journals, written and printed records, and other paper documents, has been so great in the tropics that it was once thought that termites were responsible for delaying or at least slowing the progress of intellectual development in tropical areas (Light, S. KF, M. Randall & E G. White. 1930. Univ. Cal. Agric. Exp. Stn. Circ. 318: 1-64). However, I was unable to locate any reference in the literature pertaining to drywood termite infestations and damage to books. Here I document naturally occurring infestations of J. minor in books. Case 1.—This incident involved a paperback book which was taken from a bookshelf in a home in Riverside, California. The top of this book had a single 246 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(4) Figure 1. Paperback book showing chamber excavated by a pair of alates of I. minor. entry hole. Figure 1 shows this book when it is opened at about midway through the entry hole. The only termites discovered in the book were two live dealate J. minor. It seems that this pair of swarmers excavated a chamber in this book to begin colony initiation. Case 2.—Figure 2 shows a coloring book which was found on a built-in book- Figure 2. Coloring book showing a chamber and tunnel excavated by pseudergates of J. minor. 19977 SCIENTIFIC NOTE 247 shelf in a home in West Covina, California. Note the main chamber at the right side of this book and the gallery extending to the shoulder of the boy carrying a pumpkin. Live pseudergates of /. minor were present in the main chamber of this book. Here, a colony of J. minor had infested the wall and built-in bookcase and it extended its infestation into this book. Case 3.—Case 3 involved a hard cover book which was found in a home in Ontario, California. Three small holes and accompanying tunnels were excavated from one cover of the book to the other. Some small chambers were found about midway through the book. Drywood termite pellets, with six concave sides, were present in the holes and chambers within this book but no live termites were found. Apparently alates tunnelled through this book and, not finding it suitable went elsewhere. Acknowled gement.—I thank Stoy Hedges, John Chapman, Rusty Bracho, Mike Rust, and two anonymous reviewers for manuscript reviews. Hanif Gulmahamad, Terminix International, 1501 Harris Court, Anaheim, Cal- ifornia. 92806. Received 19 Dec 1996; Accepted 26 Feb 1997. PAN-PACIFIC ENTOMOLOGIST 73(4): 248-255, (1997) THE PAN-PACIFIC ENTOMOLOGIST: FORMAT INFORMATION FOR CONTRIBUTORS The Pan-Pacific Entomologist is published quarterly by the Pacific Coast Entomological Society, in cooperation with the California Academy of Sciences. The journal serves as a refereed publication outlet and accepts original manuscripts on all aspects of the biosystematics of insects and closely related arthropods, especially articles dealing with their taxonomy, biology, behavior, ecology, life history, biogeography and distribution. Articles with either a natural, descriptive orientation or a technical and analytical emphasis are welcome. Articles that deal with the strictly economic aspects of insects, however, are inappropriate for submission to the Pan-Pacific Entomologist. Manuscripts must be in English, but foreign language summaries are permitted. As of 1 January 1990, beginning with volume 66, number 1, the Pan-Pacific Entomologist will employ changes in format which will be consistently applied. The format changes will require con- tributing authors to closely observe these guidelines. Because of comments by peer reviewers, but more particularly copy-editing, very few manuscripts are ultimately accepted without being redrafted to incorporate changes. FORMAT All manuscripts must be typed or printed on one side of 8.5 X 11 in nonerasable, high quality paper. Three (3) copies of each manuscript must be submitted: an editor’s copy (original or high quality photocopy), and two high quality review copies each including reductions of any figures to an 8.5 X 11 in page. Manuscripts must either be double or triple spaced in a legible typeface (pref- erably a serif font) that makes identification of letters and numbers distinct, especially characters such as the number 1, lower case L, and upper case I. All margins should be 1.5 in, all pages must be numbered and the senior author’s name should appear in the upper right corner of each page following the title page. Do not break words with hyphenation at the right margin of the text. The pages must be ordered and numbered separately in sequence 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, original figures (unnumbered). TITLE PAGE E The upper right corner must include the statement “‘Send galleys to:” and list the corresponding author’s name, address including ZIP code, and phone number. Type the title of the manuscript in all upper case letters in the center of the page. Use a title that is informative and specific, but as brief as possible. Compound titles are permitted and are separated by a colon; do not include a serial number in the title. The title should not include both the common name and scientific binomial of an insect. Titles must list the order and family involved within parentheses and separated by a colon. The name of each author should be typed in upper case letters several lines below the title. In no case should the professional or academic title of authors be noted. Several lines below the author’s name, the institutional affiliation should be typed in upper and lower case letters, as the sequence: department, institution, city, state or province, postal ZIP code or equivalent, and country if different from the United States or Canada. Abbreviations must not occur within affiliations or addresses, spell out all words in the affiliation, but omit building and room numbers (i.e., Department of Entomological Sciences, University of California, Berkeley, California 94720); do not use two letter postal abbrevi- ations for states. In institutional addresses use a street number (i.e., 2258 Howard Avenue) or building (i.e., Wellman Hall) only if mail would not be likely to ultimately reach its destination without that addition. If the address on an article is different from the author’s current correspondence address, use a footnote number after the address associated with the title and supply the current correspondence address on a separate footnote page. If multiple authors are associated with different institutions or addresses, separate these addresses with a semicolon; place a footnote marker after each author’s name, and before the respective address. ABSTRACT PAGE The second page of the manuscript should contain only the abstract and key words. It is essential that the abstract be concise, not exceeding 250 words. The abstract should be an informative digest 1997 FORMAT INFORMATION FOR CONTRIBUTORS 249 of the significant findings or main conclusions of the article and is often the only information made available by the abstracting publications. The abstract must be in a left indented paragraph format and begin with the word Abstract in upper and lower case letters, underlined, and followed by a period and two hyphens (i.e., Abstract.--). Begin the abstract to the right of the hyphens. When a taxonomic binomial is first mentioned in the abstract, the author (unabbreviated) of the name must be stated. Reference citation is not permitted in the abstract. The key words paragraph (again, left indented) follows several lines below the abstract. Its lead is treated similarly to the abstract (i.e., Key Words.--). This is followed by five to seven key words or concise phrases, the first of which should be “‘Insecta”’ (or the class of related arthropod if applicable). The key words are aids to abstracting services that will index the article and as such should be chosen wisely. You may repeat words or phrases from the title. TEXT PAGES The text should generally follow the guidelines established in a recent edition of the Council of Biological Editors Style Manual (CBE Style Manual Committee. 1983. CBE style manual: a guide for authors, editors, and publishers in the biological sciences. Sth ed. Council of Biological Editors, Bethesda, Maryland.). In the introduction avoid statements such as ‘““The purpose of this paper is...” and use instead simply “‘This paper . . .”” followed by what is accomplished. Throughout the article use simple and concise phrasing. Major sections are delimited by centered headings using upper and lower case letters, such as: Methods and Materials, Results, Discussion, Results and Discussion, Acknowledgment, Literature Cited. Do not use a heading for the introduction. In taxonomic manuscripts the major headings might include: Taxonomy, Biology, Behavior, etc. Minor headings are delimited as are the abstract and key words paragraphs. 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Sequences of references in the text must be cited chronologically (i.e., Nelson 1978, Nieukoop & Sutasurya 1983, Raff 1987). In cases where sequential citations occur within the text and note multiple papers by the same author, the dates of articles should be separated by commas and the authors separated by semicolons (i.e., Weber 1932, 1936, 1941; Whitcomb 1950, 1952; Henderson 1978, 1979). Citations within the text that include reference to pages, figures or tables should be cited as: (Smith 1983: 149-153, Price 1985: fig. 7a, Nothwith 1987: table 3). Note that citations within the text of illustrations or tables from other works should begin with a lower case letter (i.e., fig. 2, table 2), to distinguish them from the figures and tables of the manuscript which begin with upper case letters. All (and only) articles cited in the text must occur within the Literature Cited section. Cite as “in press’’ (i.e., Hawksworth in press), any manuscript which has been formally accepted for publication by a journal (not which is in preparation, or that merely has been submitted, acknowl- edged, or is in review by a journal!); do not estimate a date of publication in either the text reference or Literature Cited listing for articles that are in press. Data which does not exist in press (in this strict sense) or in publication and read and are not being presented in the manuscript under consid- eration can be cited as unpublished. If the unpublished data are from the sole author or all authors of the manuscript under consideration, then cite simply as “‘(unpublished data)”’ but if the data are from only one of multiple authors of the manuscript under consideration cite by initials (i.e., JAC, unpub- lished data). Cite as ‘‘unpublished data’ any manuscript in preparation or which has been submitted for publication but has not yet been formally accepted after review. If the unpublished data are from a source other than the author(s) of the manuscript under consideration, cite the data as a personal communication. Personal communications should be kept to an absolute minimum, but where neces- sary should be cited without abbreviation as: (D. Hille Ris Lambers, personal communication). Per- sonal communications and unpublished data citations are not listed in the Literature Cited section. 250 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(4) Measurements——All measurements must be in metric units. Measurements in U.S. equivalents are permissible only within quoted data (as from the label of a holotype), or within parentheses after their metric counterpart if such display has a practical value. All fractions must be in decimal format (i.e., 0.2, 0.33, 0.67, 0.125); if such numbers are less than one, place a zero before the decimal (i.e., 0.05, not .05). When using measurement or count data and citing its range, mean and standard deviation, use the format: low-high (mean + SD) (i.e., 337-388 [361 + 16]). Numbers.—Numerals are expressed as their word equivalents if between one (1) and nine (9), unless part of a sequence including values of 10 or greater; values greater than 10 are expressible as their numbers. Always begin a sentence with the word equivalent of a numeral (i.e., ““Four specimens were ...’). When a large number begins a sentence it is usually preferable to modify the beginning of the sentence to avoid awkwardness (i.e., ““We examined 175 taxa. ..”’ rather than “One hundred and seventy-five taxa were .. .”’). Usage can be confusing, and acceptable examples are: ‘‘Six genera were examined which contained a total of 26 species... ,” “.. . 10 males among the seven groups could be classified into three categories ... ,” ““The number of synapomorphies associated with each respective internodal segment in the phyletic sequence is: 3, 5, 13, 11, and 8.’ Values representing counts or distances should use their numerical rather than word equivalents, even if under 10; for example, in the Material Examined section of a taxonomic paper use the data format: “...3 km E of Wilsonville, hwy 17, 6 males, 10 females,....” Comparative Ratios ——Use a slash (/) rather than “‘per’’ to indicate standard units in relation to other standard units (i.e., mg/liter, not mg per liter). Use “‘per’’ to indicate counts in relation to nonstandard units (i.e., 10 insects per leaf). Indicate ratios using a colon (i.e., 10 males : 1 female). Use a lower case ‘“‘x’’ to indicate ‘‘times”’ in comparative ratios (i.e., mesothoracic width 0.8 x width). Dates and Time.—Dates must be expressed in the format: day month year, without punctuation; not as: month day, year. Months are abbreviated as their first three letters in upper and lower case: Jan, Feb, Mar, Apr, May, Jun, Jul, Aug, Sep, Oct, Nov, Dec. Do not shorten the notation for the year, cite in full (i.e., 1989, not 89 or ’89). Examples of acceptable dates are: 4 Jul 1990, 17 May 1753. Time is expressed in 24 hour format followed by h. If appropriate, time zones can be noted following the time, as a standard, three letter, upper case abbreviation. An example is 06:30 h (PDT), representing 6:30 AM, Pacific Daylight Time. Italics.—Italics are indicated by underlining words. It is preferable not to use actual italicized words in a manuscript, even if the capability exists in word processing, but rather to use a single underline to denote words to be italicized; this promotes clarity and reduces the chance of error during the editing and type-setting stages of publication. Italics are used only for taxonomic names, for denoting minor subheading sections in the text, for highlighting categorical descriptors or ‘‘flags” within a block of text, and for denoting mathematical variables as in formulae. Italics may be used to denote a word which should stand out in a block of text. Examples are: the word ‘““‘Thorax”’ when used within text comprising a taxonomic description; the name of a county (or equivalent) when used in a Material Examined section; the word ‘In’? denoting a contribution to a larger work, as used in a bibliographic citation in the Literature Cited section. Do not use italics to emphasize a word for effect in a sentence (i.e., ““Only some of the insects were . . .”’). Do not indicate italics for Latin abbreviations such as: 1.€., €.g., Ca., Sic., et al., etc. Abbreviations.—Reference to tables should not be abbreviated in the text, whereas reference to figures should be (i1.e., Table 4, Figs. 1-3, Fig. 6A); capitalize such references when they are part of the manuscript but not in citations from other articles (i.e., Hamilton 1983: fig. 2, Rolling 1975: table 2). Abbreviations for conventional units are in lower case letters and not followed by a period. Ex- amples are: h, min, sec, km, m, mm, wm, cc, ml, g, mg, wg, kg. Spell liter completely, do not abbreviate as “‘l.”” Symbols for male (d) and female (2) may be used. Avoid the use of ca. by using approximately instead. States should be spelled out in full, do not use their two letter postal abbreviations (except possibly in tables where space is at a premium). Do not abbreviate morphs, castes or life forms as their symbols. Do not include periods after, or spaces between, the letters of abbreviations of institutions (i.e., use USDA, NMNH and BM[NH] for U.S. Department of Agriculture, National Museum of Natural History and British Museum [Natural His- tory], respectively). Abbreviate percent as %. Abbreviate feet as ft, not ('), when necessary in ele- vations, after altitude in meters. Trademarks and registered brand names are noted as ® and ®, re- spectively. Compass directions are presented entirely in upper case without periods (i.e., N, S, E, W, NW, SE, NEE, SSW). Direction from a source, as in data presented in the Material Examined section of 1997 FORMAT INFORMATION FOR CONTRIBUTORS 251 taxonomic papers, should have the word “‘of’’ between the direction and source to avoid potential confusion in locality names (i.e., 3 km E of Palo Alto, instead of 3 km E Palo Alto, which should be taken as East Palo Alto rather than East of Palo Alto). TAXONOMIC STUDIES AND CITATIONS Taxonomic works present special editorial problems in maintaining relative consistency within and between articles on systematics within the journal. Because of these inherent problems, there is nec- essarily less flexibility for style and format in taxonomic works in the journal than for articles on solely biology. For detailed instructions see Pan-Pac. Ent. 69: 194-198. Taxonomic Citations——Upon the first mention of a species or lower level taxon in both the abstract and text, the author of an animal taxon must be cited using the International Commission of Zoological Nomenclature convention; botanical names must be so cited using the ICBN. Do not abbreviate the generic name of a taxon upon first mention. The names of authors of taxa must not be abbreviated, except for Linnaeus (as L.) and Fabricius (as Fabr.). If more than a single taxonomic author with the same last name worked in the taxonomic group considered, the author’s initials should be used to avoid confusion (i.e., Papilio bairdii W. H. Edwards, to avoid confusion with H. Edwards who was also a lepidopterist). Multiple authorship for taxa should substitute an ampersand (&) for ‘‘and’”’ between the names (1.e., Gillette & Palmer). When citing authors of taxa, citation of the year of description is optional; if used, however, the year must be enclosed within parentheses and the citation must be considered a reference citation within the article and be listed in the Literature Cited section (see below for an exception). Examples are: Chaitophorus populellus Gillette & Palmer (1928) (where populellus was originally described in Chaitophorus), and Chaitophorus populifolii (Essig 1912) (where populifolii was not originally de- scribed in Chaitophorus). In these cases Gillette & Palmer (1928) and Essig (1912) must be listed in the Literature Cited section. If no year citation is noted for a species when citing its author, a literature citation is not invoked. The only exception to treating the author-year citation as a literature citation is in checklists and synonymies, where year citations may be more appropriate than in the text, and where the numbers of such citations would make literature citation unwieldy. In such cases, cite years as C. populellus Gillette & Palmer, 1928, and C. populifolii (Essig), 1912. All nomenclature erected in taxonomic studies must follow the rules established in the most recent nomenclatural code by the ICZN. When mentioning the ICZN code, refer to it as such, rather than just “‘the code.” At any mention of a particular article in the ICZN code, cite the publication date of the code version used, to avoid confusion (i.e., “‘. . . due to ICZN article 49 [ICZN 1985].’’) and list the code citation in the Literature Cited section (see the Literature Cited examples for citation of the ICZN code). New taxa or synonymies that are erected should be clearly and appropriately marked, or “‘flagged,”’ in upper case letters after mention in the text (.e.,. NEW GENUS, NEW SYNONYMY, NEW STA- TUS, etc.), and notation should also occur in the abstract. New taxa must be listed with the name of the describing author(s), even if it is the same as the manuscript author, after the binomial but before any taxonomic “‘flag’”’ (i.e., Diodontus retiolus Eighme, NEW SPECIES). Descriptions and Diagnoses.—Taxa being described must have the following: a description (pref- erably with appropriate illustrations), and a separate diagnosis. The diagnosis must occur as a separate paragraph delimited as “Diagnosis.--’’ and must be concise but not in telegraphic style. Comment briefly and only on those attributes required to separate the taxon from related taxa in the diagnosis. The description must be delimited by a minor subheading, as is the diagnosis; it need, however, not be labeled as ‘“‘description’’ but rather can be labeled as a sex, life stage or form (i.e., male, egg, larvae, viviparous apterae, etc.). Descriptions must be concise and be in telegraphic style. It is pref- erable to separate major components with periods (i.e., legs), minor components with semicolons (1.e., tibiae, tarsi) and the details of minor components with commas (i.e., color of tibiae and the forms of setae on them). Avoid conjunctives as much as possible in telegraphic descriptions; also avoid “‘-ish”’ adjective suffix forms of colors, such as ‘“‘head yellow with brownish” using instead “‘head yellow with slight brown”’; avoid latin color descriptors (i.e., testaceous). An example is: “*. . . Genitalia (Fig. 5A): valve breadth greatest ventrally, filling entire vinicular area, bilobed and constricted near inden- tation, caudal extension tapered, blunt terminally; saccus diminutive, lobate with rounded margins; aedeagus robust, ceacum two-fifths aedeagal length.” Types.—Descriptions and revisions also require comments on the types involved. Comments on types are to be in a separate paragraph delimited as ““Types.--’’ that lists the type of type(s) (holotype, 22 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(4) paratype, etc.) erected, and their data and deposition. An example is: ‘‘Holotype, male (Figs. 2E, 5A) deposited BM(NH), data: PERU. Cajamarca, 2800 m, Simons collection. Paratype, 1 male deposited NMNH, (poor condition with tails broken off), data: same as holotype but 3800 m, O. T. Baron collection, ex Hamilton collection 1919.’ When citing data from the labels of types it is permissible to quote within quotation marks. If not directly quoting from type labels, use the data format listed below for material examined. The deposition of types must be noted appropriately and be in accord * with ICZN requirements. Deposition of types in private collections should be avoided for reasons of ‘professionalism,’ but when deposition is so designated, the ultimate intended deposition after the death of the author(s) should be stated; neotypes require institutional deposition (ICZN 1985: article 75-d-6). Keys.—Keys are not required in taxonomic works, but are highly recommended. When presented, keys must be concise, clear, easy to follow and have reversibility provisions. Keys must also be in adjacent couplet style, and each couplet should contain preferably more than a single, nonoverlapping attribute. It must be clear to which life stage, sex, caste, morph, etc., the key pertains; keys requiring more than a single such life form are discouraged, and it is recommended that separate keys be provided in such cases. Material Examined.—Data for material studied in taxonomic manuscripts must be listed under a separate paragraph delimited as ‘‘Material Examined.--.’”’ This taxonomic section should have the countries, as well as major (1.e., state, province) and minor (i.e., county or equivalent) political units spelled out in upper case, with the minor political units also underlined for italics. Use the following format, with modification as appropriate: USA, ARIZONA, APACHE Co.: 10 km N of Lupton, hwy 12, 2070 m, 11 Sep 1978, J. T. Sorensen (JTS 78118), P. ponderosa, 6 females. COCHISE Co.: nr Rustler Park, Chiricahua Mts, 2500 m, 16 Sep 1978, J. T. Sorensen (JTS 78147), P. ponderosa, 12 females. COLORADO. ARCHULETA Co.: 25 km W of Pagosa Springs, hwy 160, 2140 m, 8 Aug 1978, J. T; Sorensen (JTS 78H5O), P. ponderosa, 1 female. CANADA. BRITISH COLUMBIA. Fair- mont Hotsprings, hwy 93, 17 Jul 1978, J. T. Sorensen (JTS 78G91), P. ponderosa, 25 females. ACKNOWLEDGMENT PAGE Begin this section on a separate page and spell the heading as Acknowledgment, not as AcknowI- edgement, Acknowledgements or Acknowledgments. The acknowledgment should be concise, thank- ing people first, institutions second where necessary, and grant or contract support third where appro- priate. Do not use the professional or academic titles of those being acknowledged. If the affiliations of those acknowledged are included, do not abbreviate institutional names but do include their loca- tions. Do not abbreviate number as “‘No.”’ or as ““#” in citing grant or contracts; rather, for example, cite as ““NSF grant BSR-8908456.” LITERATURE CITED PAGES Begin this section on a separate page, titled Literature Cited, not References, References Cited or Bibliography. All paragraphs should be hanging format (left block first line and indented thereafter). Do not list references which are not cited in the text. Do not list unpublished data, personal com- munications, or works in preparation in the Literature Cited section. Citations listed should be in alphabetical order first and then in chronological order; if multiple citations bearing the same author(s) and year are cited, they should be listed using lower case letters after the year and be in the sequence in which they appear in the text. Do not use Ibid. Authors cited are listed as last name first followed by initials for sole or senior authors and initials followed by last name for subsequent authors. Omit reference to number (issue) in citations after the volume. Omit the number of total pages in books, separates, pamphlets, etc. Abbreviate journal titles as listed in the International Serials Catalogue: Part I: Catalogue (International Council of Scientific Unions Abstracting Board, 1978). Do not abbreviate single word journal names (i.e., Evolution, Ecol- ogy). All citations must fully cite the authors, even when more than one article is present for any author(s). When listing articles or books, all letters in the title should be lower case, except the first letter of the title’s first word and any proper nouns. Examples of acceptable citation formats follow: One author articles: Arnold, R. A. 1983. Speyeria callippe (Lepidoptera: Nymphalidae): application of information— theoretical and graph-clustering techniques to analyses of geographic variation and evaluation of classifications. Ann. Entomol. Soc. Am., 76: 929-941. 1997 FORMAT INFORMATION FOR CONTRIBUTORS 253 Two author articles: Ferrari, J. A. & K. S. Rai. 1989. Phenotypic correlates of genome size variation in Aedes albopictus. Evolution, 42: 895-899. Articles with more than two authors: Atchley, W. R., E. V. Nordheim, E C. Gunsett & PL. Crump. 1982. Geometric and probabilistic aspects of statistical distance functions. Syst. Zool., 31: 445-460. Manuscripts in press: Sorensen, J. T. (in press). Three new species of Essigella (Homoptera: Aphididae). Pan-Pacif. En- tomol. Books: Anderson, T. W. 1984. An introduction to multivariate statistical analysis (2nd ed.). John Wiley & Sons, New York. Parts of larger works: Klecka, W. R. 1975. Discriminant analysis. Chapter 23. pp. 434-465. In Nie, N. H., C. H. Hull, J. G. Jenkins, K. Steinbrenner & D. H. Brent. 1975. SPSS: statistical package for the social sciences (2nd ed.). McGraw-Hill, New York. Blackman, R. L., PB. A. Brown & V. FE Eastop. 1987. Problems in pest aphid taxonomy: can chro- mosomes plus morphometrics provide some answers? pp. 233-238. In Holman, J., J. Pelikan, A. G. E 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. Proceedings of meetings: Philbrick, R. N. (ed.). 1967. Proceedings of the symposium on the biology of the California islands. Santa Barbara Botanic Garden, Santa Barbara, California. Wilson, M. R. & L. R. Nault (eds.). 1987. Proceedings of the second international workshop on leafhoppers and planthoppers of economic importance, held In Provo, Utah USA, 28th July— Ist August 1986. CAB International Institute of Entomology, London. Governmental or institutional publications: Little, E. L. Jr, & W. B. Critchfield. 1969. Subdivisions of the genus Pinus (Pines). U.S. Dept. Agric., Forest Serv. Misc. Publ., 1144. Hafernik, J. E. Jr. 1982. Phenetics and ecology of hybridization in buckeye butterflies (Lepidoptera: Nymphalidae). Univ. Calif. Publ. Entomol., 96. Anonymous institutional or organizational publications: International Code of Zoological Nomenclature. 1985. (3rd ed.) International Trust for Zoological Nomenclature (BM[NH]). University of California Press, Berkeley, California. California Department of Food & Agriculture. 1987. Environmental assessment of gypsy moth and its eradication in California, 1987 program. California Department of Food & Agriculture, Division of Plant Industry, Sacramento, California. Theses and dissertations: Sorensen, J. T. 1983. Cladistic and phenetic analysis of Essigella aphids: systematics and phylogeny in relation to their Pinaceae host plants (Homoptera: Aphididae, Lachninae). Ph.D. Thesis, University of California, Berkeley. Computer programs: Felsenstein, J. 1984. PHYLIP—Phylogeny inference package (Version 2.5). (A phylogenetics com- puter program package distributed by the author). J. Felsenstein, Dept. of Genetics, University of Washington, Seattle, Washington. Pimentel, R. A. & J. D. Smith. 1985. Biostat II. (A multivariate computer program package dis- tributed by the authors). Sigma Soft, Placentia, California. 254 THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(4) FOOTNOTE PAGES All title and text footnotes must appear on a separate and numbered footnote page and be indicated by consecutive superscript numbers. Author-line footnotes that denote a change of affiliation or current address must also appear on the separate footnote page. Only multiple author and affiliation address footnotes (as described under the title page section) should not be on the footnote page. Text footnotes provide additional information and are usually discouraged; such information, if short, may preferably be enclosed within parentheses in the text. Footnotes cannot be used for acknowledgments, list these in the Acknowledgment section. FIGURE CAPTION PAGE Figure captions should briefly interpret the figures but need not be complete sentences. Captions should not unnecessarily repeat or explain information provided in the text. Be sure, however, to adequately explain any graphics that employ symbols (i.e., graphs, charts, scattergrams) in their figure captions. The caption for each plate or block of illustrations should be represented by a separate left block paragraph. If several figures are illustrated on a single plate, each figure may be cited separately as a sequential Arabic number. Alternatively, each individual figure making up a plate may be cited as a subcomponent and labeled sequentially and alphabetically (preferably as an upper case letter) when the plate itself bears a single Arabic number. In the former case, appropriate citation for a plate containing several figures would be the figure caption paragraph: “Figures 1-3. Thoracic morphology of Rhyacophila chordata (lateral views). Figure 1. Prothorax. Figure 2. Mesothorax. Figure 3. Meta- thorax.’’ In the latter case, the paragraph would state: “Figure 1. Thoracic morphology of Rhyacophila chordata (lateral views). A. Prothorax. B. Mesothorax. C. Metathorax.”’ Do not use “multiple nested’”’ explanations when multiple figures of different subjects appear on the same figure plate. For example, the following format is unacceptable: “‘Figures 1-4. Aphid tax- onomic characters. Figures 1-2. Myzus persicae. Figure 1. Siphunculi. Figure 2. Cauda. Figures 3-4. Myzus cerasi. Figure 3. Siphunculi. Figure 4. Cauda.’’ Instead use a format such as: ‘‘Figures 1—4. Aphid taxonomic characters. Figure 1. Myzus persicae, siphunculi. Figure 2. Myzus persicae, cauda. Figure 3. Myzus cerasi, siphunculi. Figure 4. Myzus cerasi, cauda.”’ ILLUSTRATIONS Illustrations must be of high quality and large enough to stand reduction. Authors must plan their illustrations to reduce to a 117 X 181 mm (approximately 4.5 X 7.0 in) galley bed, allowing space for the figure caption below the figure. They are strongly encouraged, however, to provide illustrations which are not larger than 8.5 X 11 in format for easy handling; damage to larger illustration formats is the responsibility of the author(s). Do not attach legends to the illustrations. Figures must be numbered in the order presented. Multiple figures on a plate may be numbered or lettered (preferably upper case). All illustrations must be mounted; the editor will not handle loose illustrations. Line drawings must be done in black, waterproof ink (i.e., India ink). The original drawings need not necessarily be submitted, but copies submitted must be of equal quality. Photostats (from nega- tives), the highest quality photocopies, or laser-printer output may be acceptable, provided the black lines are uniformly and absolutely black and the background is absolutely and uniformly white. Mul- tiple figures on a plate must be grouped closely to eliminate unnecessary large spaces between the illustrations. Photographs must be trimmed and mounted, abutting each other; they must not be less than the width of the printed page. Lettering must be consistent throughout, and appropriately sized for the potential reduction so that it does not reduce to less than 1 mm. Lettering must be of quality appearance; hand lettering or typewriter lettering is not acceptable for illustrations. If return of original illustrations is desired, request it when the manuscript is originally submitted. To avoid confusion and loss at the press, all original illustrations submitted should be labeled with nonreproducible, light blue pencil on the back noting the following: (1) figure number, (2) direction of top as indicated by an arrow, (3) author’s name, (4) title of the manuscript, and (5) journal. For guidance in preparing illustrations for optimal presentation, consult the following books: Hodges, E. R. S. (ed.) 1989. The guide handbook of scientific illustration. Van Nostrand Reinhold, New York. Papp, C. S. 1976. Manual of scientific illustration, with special chapters on photography, cover design and book manufacturing. American Visual Aid Books, Sacramento, California. 1997 FORMAT INFORMATION FOR CONTRIBUTORS 255 TABLES Because of the expense involved in reproducing tables, their use should be kept to a minimum. Tables should have a short title and be numbered in Arabic numerals (i.e., Table 1). Each table must be double-spaced and can be continued on additional sheets of paper as necessary. Do not reduce tables. Double-space all segments of the table, including title, heading, body and any footnotes. Use standard abbreviations for column heads. Nonstandard abbreviations may be used sparingly and must be defined in footnotes. Table footnotes are indicated by consecutive use of lower case letters. Tables must be numbered in the order presented. SCIENTIFIC NOTES As an alternative format, the Pan-Pacific Entomologist also publishes scientific notes to allow an outlet for contributions which are not judged suitable for full length articles. Scientific notes use an abbreviated format. The title is in full upper case and as a centered paragraph. No abstract, key words, footnotes or major and minor section headings are permitted. Minimal use of figures and tables is permitted. A Literature Cited section is not allowed. Reference citations are kept to a minimum, but when necessary are cited initially in the format (Bohart, R. M. 1989. Pan-Pacif. Entomol., 65: 156—161.), without title. After being cited initially, if it becomes necessary to cite a reference again, simply cite it as author and year, in this instance: (Bohart 1989) or Bohart (1989). Avoid citing contributions to larger and edited works or those which entail lengthy citations, to maximize reading ease. An Acknowledgment paragraph is permitted, but is handled as a paragraph with a left indented, minor subheading (i.e., Acknowledgment.--). Authors and affiliations are cited as the last left indented paragraph of the note, with the affiliation underlined for italics as: John T. Sorensen, Insect Taxonomy Laboratory, California Department of Food & Agriculture. Sacramento, California 95814. VOUCHER SPECIMENS Where appropriate, manuscripts must name a public repository where specimens documenting the identity of the studied organisms can be found (see Pan-Pac. Ent. 73: 200). Please put in Materials and Methods section of the manuscript or in the text of scientific notes. PAN-PACIFIC ENTOMOLOGIST 73(4): 256, (1997) Pan-Pacific Entomologist Reviewers Volume 73 Anderson, R. Andrews, E Austin, JG. Baranowski, R. Becker, E. Berenbaum, M. Berlocher, S. Bezark, L. Brower, L. Brownbridge, M. Burdick, D. Byers, G. Campbell, J. Chalfant, R. Clark, D. Cromartie, W. Deyrup, M. Dobson, H. Ehler, L. Fisher, E. Garrison, R. Gelhaus, J. Gilbert, A. Gill, B. Gill, R. Gordon, R. Greenberg, B. Hamilton, S. Hardy, A. Henry, T. Hoffman, K. Hovore, E Hunter, A. Johnson, K. Kimsey, L. Krafsur, E. Landolt, P Latheef, M. Lee, V. Lewis, V. Luck, R. Marsh, P. McMillen, C. Merickel, FE Oswald, J. Pickett, C. Poole, R. Price, P. Ratcliffe, B. Roubik, D. Rust, R. Rust, M. Savary, W. Shapiro, A. Shepard, W. Siebert, C. Sikes, D. Sorensen, J. Thomas, D. Waller, D. Ward, P. Wasbauer, M. Wehling, W. Weller, S. Westcott, R. Whaley, W. Whitfield, J. Wood, D. Wood, S. Yokoyama, V. Young, C. Zera, A. PAN-PACIFIC ENTOMOLOGIST 73(4): 257-259, (1997) The Pan-Pacific Entomologist Contents for Volume 73 ADAMS, J., see MURPHY, B. C............ 4 AHN A, K-J.—A review of Liparoce phalus Maklin (Coleoptera: Staphylinidae: Aleocharinae) with descriptions of larvae ............. 79 ARAY, K., see IWATA, R. ........0..02. 213 AREFINA, T. I—A new species of the genus Ceraclea Stephens (Trichoptera: Leptoceridae) from Zelyoni Island (South Kuril Islands) . . Oe lee Ok Cee a eee ge 100 ASHIDA, H., see IWATA, R. ........... 213 BAKER, T. C., see VETTER,R.S. ........ 28 BARTHELL, J. F, T. L. GRISWOLD, G. W. FRANKIE & R. W. THoRP—Osmia (Hymenoptera: Megachilidae) diversity at a site in central coastal California .............. 141 Bayoun, I. M., see WALKER, G.P...... 236 BRAILOVSKY, H.—Sibuyanhygia, a new genus of Colpurini from the Philippine Republic, with descriptions of three new _ species (Heteroptera: Coridae) ............ 70 Brown, J. W. & D. K. FAULKNER—A new species of Litoprospus (Lepidoptera: Noctuiidae) from Baja California, Mexico...... 122 BURQUEZ, A.—Distributional limits of Euglossine and Meliponine bees (Hymenoptera: Apidae) in northwestern Mexico .... 137 CAMPBELL, C., see HANKS, L. M. ....... 190 CHEMSAK, J. A.—BLEUZEN, P. 1994. Les Coleopteres du Monde (The Beetles of the World). Prioninae 1: Macrodontini: Macrodontia, Chalcoprionus, Ancistrotus, Acanthinodera, Acalodegma: Prioni: Titanus, Braderochus. Vol. 21. Sciences Nat. 92 pp. 16: plates) 2. 12 Ae eee ee ees 60 Conway, J. R—Foraging activity, trails, food sources and predators of Formica obscuripes Forel (Hymenoptera: Formicidae) at high altitude in Colorado ............. 172 CRoFT, B. A., see DRAPEK, R. J. ......... 9 DAVIDSON, J. A., see MILLER, D. R. ..... 201 DRAPEK, R. P, B. A. Crorr & G. FISHER—An examination of spatial input parameters in order to improve corn earworm (Lepidoptera: Noctuidae) damage predictions for a pheromone trap catch regression model... 4... eee ees 9 EQUIHUA-MARTINEZ, A., see PECK, R. W. ... 204 FAULKNER, D. K., see BROWN, J. W. ..... 122 FINSTON, T. L., S. B. PEcK & R. B. PERRY— Population density and dispersal ability in Darwin’s darklings: flightless beetles of the Galapagos Islands .............. 110 FISHER, G., see DRAPEK, R. J. FITZGERALD, S.—A new species of Eniocoscolus (Diptera: Bibionidae) from Brazil, with additional distribution records for the genus Ae ere ee Re nee ee eee ewe ee 152 FRANKIE, G. W., see BARTHELL, J. EF ..... 141 Furniss, M. M., see KEGLEY, S.J. ....... 40 GorpDon, R. D. & R. R. RuST—A new southern Nevada species of Aegialia (Aegialia) (Coleoptera: Scarabaeidae: Aphodiinae) . . SW OT, Oe Na oe Seer ay eh Re Ae 168 GREGOIRE, J., see KEGLEY, S.J. ......... 40 GRISWOLD, T. L., see BARTHELL, J. EF .... 141 GULMAHAMAD, H.—Ecological studies on Cardiocondyla ectopia Snelling (Hymenoptera: Formicidae) in southern Galifornia feist oo2-seuk alee ss, meses a | GULMAHAMAD, H.—Naturally occurring infestations of drywood termites in books 245 Hanks, L. M., C. CAMPBELL, T. D. PAINE & J. G. MILLAR—Host range expansion of Helcostizus rufiscutum Cushman (Hymenoptera: Ichneumonidae) to Phoracantha semipunctata Fabr. (Coleoptera: Cerambycidae) in California ae ey ee ee i 190 Hsu, Y-F—A new Japonica (Lepidoptera: Lycaenidae: Theclinae) from southwestern GR aa ei era ieee ee ke 225 Hynes, C. D.—The immature stages and biology of the craneflies Toxorhina caledonica and Elephantomyia garrigouana_ (Diptera: Limoniidae) IwaTA, R., E YAMADA, H. Kato, H. MAKIHARA, K. AraydA, H. AsHipaA & M. TAKEDA— Nature of galleries, durability of boring scars and density of Xylotrechus villoni (Villard) larvae (Coleoptera: Cerambycidae), on coniferous tree trunks 213 JOHNSON, P. J.—New species of Dioxypterus Fairmaire from Tonga and Fiji, with new distribution records, a tribal reassignment, and key to the species of the region (Coleoptera: Elateridae) .......... 156 KATO SH ty SC@eIWATIAC IR. sao cutest faces 258 Kaya, H. K., see LEoNG, K. L.H........ 49 KEGLEY, S. J, M. M. Furniss & J. GREGOIRE— Electrophoretic comparison of Dendoctonus punctatus LeConte and D. micans (Kugelann) (Coleoptera: Scolytidae)............ 40 KURAHASHI, H., see WELLS, J. D. ... 2... 16 LeonG, K. L. H., M. A. YOSHIMURA & H. K. KayA—Occurrence of a _ neogregarine protozoan, Ophryocystis elektroscirrha McLaughlin and Meyers, in populations of monarch and queen butterflies ...... 49 MAKIHARA, H., see IWATA, R. 213 MARTINEZ, M. J.—The first record of the ant Pheidole moerens Wheeler from the western United States (Hymenoptera: Formicidae) . . ee ee een oe 46 McKiLuup, R. V., see McKiLtup, S.C. ... 184 McKiLLup, S. C. & R. V. McKiILLup—An outbreak of the moth Achaea serva (Fabr.) on the mangrove Excoecaria agallocha (L.) Be Sinemet: eile hase es eee ge Pl UR 184 MILLAR, J. G., see HANKS, L.M. ....... 190 Miter, D. R. & J. A. DAvmson—Obituary: Richard FE Wilkey (1925-1995) 201 MucHMorE, W. B.—The identity of Chelanops serratus Moles (Pseudoscorpionida: Chiemetidae) = a8. Bao Se 55 Murpuy, B. C., J. ADAMS & M. P. PARRELLA— Experimental arena for confirming thrips and other small arthropods in the laboratory OPA eed Wears 7 eee gee BoM eer bal: evens 4 NELSON, G. H.—A new Poecilonota from southern California (Coleoptera: Buprestidae) j. 2. Ale te Perea et 1 NICHOLS, B. J., see SITES, R. W. ....... 127 O’BRIEN, C. W. & R. S. ZAcK—Weevils new to the State of Washington (Coleoptera: Curculionidae) PACIFIC COAST ENTOMOLOGICAL SOCIETY—Financial statements for 1995, 1996 ......... 197 PaciFIC COAST ENTOMOLOGICAL SOCIETY— Proceedings for 1996 ............ 192 PaciFIC COAST ENTOMOLOGICAL SOCIETY— Sponsoring Members 1996 ........ 199 PAINE, T. D., see HANKS, L. M. ........ 190 PARRELLA, M. P., see MurpHy, B.C. ...... 4 Peck, R. W, A. EQUIHAU-MARTINEZ & D. W. Ross—Seasonal flight patterns of bark and ambrosia beetles (Coleoptera: Scolytidae) in northwestern Oregon ............ 204 PECK, .S..B;-see FINSTON,, Th. 2 pa a 110 PENNY, N. D.—Four new species of Costa Rican Ceraeochrysa (Neuroptera: Chrysopidae) . . PERRY, R. B., see FINSTON, T. L. PERMKAM, S., see SITES, R. W.......... 127 RASNITSYN, A. P—Xylea (Pinicolites) lata Smith THE PAN-PACIFIC ENTOMOLOGIST Vol. 73(4) (Vespida: Xyelidae), a living fossil sawfly from western North America 231 Rocers, D. C.—Aphodius alternatus Horn (Aphodiinae: Scarabaeidae), first record of a semi-aquatic scarab beetle ........ 135 Ross, D. W., see PEcK, R. W. ......... 204 Rust, R. R. see GORDON, R. D. ........ 168 SHAW, S. R.—The Costa Rican species of Wesmaelia Foerster with a description of a new species (Hymenoptera: Braconidae: Buphoritae)* 2... .ou. eee 2 ae 103 SHEPPARD, W. D.—Lilioceris sp. (Coleoptera: Chrysomelidae) herbivory on Cycas siamensis Miguel (Tracheophyta: Cycadales) POR RAs eae: Bes wells Be RR gee. Ales 36 SITES, R. W., B. J. NICHOLS & S. PERMKAM—The Naucoridae (Heteroptera) of southern Nivea anc. 222 er Pace reer eS Aas oe 127 SORENSEN, J. T. & K. H. SORENSEN—Aggregations of Thaumatomyia glabra (Meigen) (Diptera: Chloropidae) on Wisteria flowers (Fabacae) NR eae RS el Re eck veh Sicge er 47 SORENSEN, K. H., see Sorensen, J.T. ..... 47 STEFFAN, S. A.—Flower-visitors of Baccharis pilularis De Candolle subsp. consanguinea (De Candolee) C. B. Wolf (Asteraceae) in Berkely, Califomia’ .....°. > DOS... 52 TAKEDA, M., see IWATA, R. ........... 213 TATEVOSSIAN, S., see VETTER, R. S. ...... 28 The Pan-Pacific Entomologist—Editorial Change 200 The Pan-Pacific Entomologist—Format information for contributors ................ 248 The Pan-Pacific Entomologist—Index for Volumes 73) eee. A eee ee 259 The Pan-Pacific Entomologist—Reviewers for AZe)I! 1) sae ee a ee Re 256 The Pan-Pacific Entomologist—Table of Contents for Volume 73 .......... 257 THORP, R. W. see BARTHELL, J. FE ...... 141 TRIAPITSYN, S. V., see WALKER, G.P. .... 236 VETTER, R. S., S. TATEVOSSIAN & T. C. BAKER—Reproductive behavior of the female carob moth (Lepidoptera: Pyralidae) dou, © Pape ice FS el pa OU FRc te 28 WALKER, G. P, N. ZAREH, I. M. BAYOUN & S. V. TRIAPITSYN—Introduction of western asian egg parasitoids into California for biological control of beet leafhopper, Circulifer tenellus ee eee Ba ah ue SRR lig ed 236 WELLS, J. & H. KURAHASHI—Chrysomya megacephala (Fabr.) is more resistant to attack by Ch. rufifacies (MacQuart) in a laboratory arena than is Cochliomyia macellaria (Fabr.) (Diptera: Calliphoridae) Re SEN Bog AA ee ee 16 WIESENBORN, W. D.—Hesperopsis graciliae 1997 CONTENTS FOR VOLUME 73 259 (MacNeil) (Lepidoptera: Hesperiidae) flight Zack, R. S., see O'BRIEN, C.W. ........ 58 between hostplants and Prosopis glandulosa ZAREH, N. see WALKER, G.P ......... 236 POLLO Ya ee DS ne ee i eae 186 ZupaRKO, R. L.—Notes on Brachycaudonia YAMADA; Fo scesWATA, IR] weet. 2 hs 2 e 213 Ashmead spp. (Hymenoptera: Pteromalidae) YOSHIMURA, Mi Ac, see“-LEONG, Keb He e049 Rie pee ee ee Eide Bos dha 243 PAN-PACIFIC ENTOMOLOGIST 73(4): 260-261, (1997) The Pan-Pacific Entomologist Index to Volume 73 (title and key words) Abies, cerambycid in 213 Achaea serva 184 Aegialia knighti NEW SPECIES 168 Ammophorus 110 ant foraging trails 172 Anthonomus cycliferus 58 Anthonomus sphaeralciae 58 Aphodius alternatus 135 Apidae 137 aposomatic coloration 36 Australia, moth outbreak in 184 Baccharis pilularis consanguinea 52 Baja California, noctuid moth from 122 beet leafhopper, egg parasitoids 236 Bibionidae 152 biological invasion 16 Blapstinus 110 Brachycaudonia 243 Braconidae 103 Brazil, bibionid from 152 Buprestidae 1 California bee diversity in 141 cerambycid parasite from 190 new buprestid in 1 Calliphoridae 16 Cardiocondyla ectopia 21 carob moth 28 Ceraclea valentinae NEW SPECIES 100 Ceraeochrysa inbio NEW SPECIES 61 Ceraeochrysa tauberae NEW SPECIES 61 Ceraeochrysa nigripedis NEW SPECIES 61 Ceraeochrysa costaricensis NEW SPECIES 61 cerambycid galleries 213 density 213 boring scars 213 Cerambycidae 190, 213 Ceutorhynchus erysimi 58 Chelanops serratus 55 Chernetidae 55 China, new lycaenid 225 Chloropidae 47 Chrysoma rufifacies 16 Chrysoma megacephala 16 Chrysomelidae 36 Chrysopidae 61 Circulifer tenellus, egg parasitoids 236 Cleonidius erysimi 58 Cochliomyia macellaria 16 Coleoptera 190 Colorado, ant foraging in 172 competitive displacement 16 Coreidae 70 corn ear worm 9 corn damage predictions 9 Costa Rica Braconidae 103 lacewings 61 Curculionidae 58 cycad 36 Cycadales 36 Cycas siamensis 36 Darwin’s darklings 110 Dendroctonus micans 40 Dendroctonus punctatus 40 Dinocheirus sicarius 55 Dioxypterus tonga NEW SPECIES 156 Dioxypterus eua NEW SPECIES 156 Dioxypterus beaveri NEW SPECIES 156 Diptera 152 distributional limits of bees 137 drywood termites in books 245 Ectomyelois ceratoniae 28 Elateridae 156 electrophoretic comparison 40 Elephantomyia garrigouana 93 Enicoscolus hardyi NEW SPECIES 152 Euglossa viridissima 137 Euglossine 137 Eulaema polychroma 137 Euphorinae 103 Excoecaria agallocha 184 experimental arena 4 Fabaceae 47 Fiji, elaterid from 156 flower visiting insects 52 fly aggregations 47 foraging behavior 21 Formica obscuripes 172 Formicidae 21, 46 Frankliniella occidentalis 4 Galapagos Islands, darkling beetles 110 1997 Gymentron pascuorum 58 Helcostizus rufiscutum 190 Helicoverpa zea 9 Hesperiidae 186 Hesperopsis graciliae 186 Hymenoptera 141, 190 Ichneumonidae 190 Incisitermes minor 245 isoenzyme electrophoresis 40 Japan, cerambycid in 213 Japonica bella NEW SPECIES 225 Lepesoma remota 58 Lepidoptera 184, 186 Leptoceriidae 100 Lilioceris sp. 36 Limoniidae 93 Line pithema humilie 21 Liparocephalus tokunagai 79 Liparocephalus cordicollis 79 Liparoce phalus brevipennis 79 Litoprosopus bajaensis NEW SPECIES 122 Lobularia maritima 21 Lycaenidae 225 MacNeill’’s sootywing skipper 186 mangrove 184 mating periodicity 28 Mecinus pyraster 58 Megachilidae 141 Meliponine 137 Mesocheira bicolor 137 Mesoplia 137 Mexico, bibionid from 152 monarch butterfly 49 moth outbreak 184 Nannotrigona perilampoides 137 Naucoridae 127 neogregarine protozoan 49 Nevada, skipper flight in 186 Noctuidae 9, 122, 184 North America, living fossil sawfly in 231 Ophryocystis elektroscirrha 49 Oregon, scolytid flight in 204 Osmia 141 Pacific Coast Entomological Society—Financial statement for 1995, 1996 197 Pacific Coast Entomological Society—Proceedings for 1996 192 Pacific Coast Entomological Society—Sponsoring members 1996 199 parasite host shift 190 Pheidole moerens 46 phenology of scolytids 204 INDEX FOR VOLUME 73 261 pheromone trap for corn earworm 9 pheromones of scolytids 204 Philippine Republic, corids from 70 Phorcantha semipunctata 190 Poecilonota bridwelli | Poecilonota viridicyanea NEW SPECIES 1 population structure 110 Prosopis glandulosa 186 psammophilous 168 Pseudoscorpionida 55 Ptermalidae 243 Pyralidae 28 queen butterfly 49 reproductive behavior 28 Scarabaeidae 135, 168 Scolytidae 40, 204 semiaquatic scarab 135 Sibuyanhygi sibulana NEW SPECIES 70 Sibuyanhygi atra NEW SPECIES 70 Sibuyanhygi calle jai NEW SPECIES 70 Sibuyanhygia NEW GENUS 70 skipper flight 186 southern Nevada, scarabid from 168 Staphylinidae 79 Stomion 110 Tenebrionidae 110 Thailand, Naucoridae in 127 thatching ant 172 Thaumatomyia glabra 47 The Pan-Pacific Entomologist—Editorial statement 200 The Pan-Pacific Entomologist—Format information for contributors 24 The Pan-Pacific Entomologist—Index to Volume 73 259 The Pan-Pacific Entomologist—Reviewers for Volume 73 256 The Pan-Pacific Entomologist—Table of Contents to Volume 73 257 thrips 4 Tonga, elaterid from 156 Toxorhina caledonica 93 Washington, new weevils in 58 Wesmaelia lizanoi NEW SPECIES 103 Wesmaelia pendula 103 western U. S., new ant 46 Wilkey, Richard F, obituary 201 Wisteria 47 Xyela lata 231 Xyelidae 231 Xylocopa muscaria 137 Xylocopa guatemalensis 137 Xylotrechus villoni 213 Zelyoni Island, Tricoptera from 100 PAN-PACIFIC ENTOMOLOGIST 73(4): 262, (1997) Announcement Publications of the Pacific Coast Entomological Society PROCEEDINGS OF THE PACIFIC COAST ENTOMOLOGICAL SOCIETY Vol. 1 (16 numbers, 179 pages) and vol. 2 (9 numbers, 131 pages). 1901-1930. Price $5.00 per volume. THE PAN-PACIFIC ENTOMOLOGIST Vol. 1 (1924) to vol. 51 (1975), price $10.00 per volume of 4 numbers, or $2.50 per single issue. Vol. 52 (1976) to vol. 57 (1981), price $15.00 per volume or $3.75 per single issue, except for vol. 57, no. 1, $10.00. Vol. 58 (1982) to vol. 66 (1990), $20.00 per volume or $5.00 per single issue. Vol. 67 (1991) to vol. 69 (1993), $30.00 per volume or $7.50 per single issue. Vol 70 (1994) and sub- sequent issues, $40.00 per volume or $10.00 per single issue. MEMOIRS OF THE PACIFIC COAST ENTOMOLOGICAL SOCIETY Volume 1. The Sucking Lice by G. E Ferris. 320 pages. Published October 1951. OUT-OF-PRINT. Volume 2. A Revision of the Spider Mite Family Tetranychidae by A. Earl Prich- ard and Edward W. Baker. 472 pages. Published July 1955. OUT-OF-PRINT. Volume 3. Revisionary Studies in the Nearctic Decticinae by David C. Rentz and James D. Birchim. 173 pages. Published July 1968. Price $4.00* (plus $0.75 postage and handling). Volume 4. Autobiography of an Entomologist by Robert L. Usinger. 343 pages. Published August 1972. SPECIAL PRICE $5.00 (plus $1.00 tax, postage, and handling for California orders, 0.70 postage and handling for non-California U.S. orders, or $1.70 for foreign orders). No members discount at this special price. Volume 5. Revision of the Millipede Family Andrognathidae in the Nearctic Region by Michael R. Gardner. 61 pages. Published January 21, 1975. Price $3.00* (plus $0.75 postage and handling). *For California orders only, add appropriate sales tax currently in effect in your country. If requested, members of the Society can receive a 20% discount on the price of the memoirs, except vol. 4. Send orders to: Pacific Coast Entomological Society % California Academy of Sciences Golden Gate Park San Francisco, California 94118-4599 U.S.A. PAN-PACIFIC ENTOMOLOGIST 73(4): 263, (1997) Editorial Policy Change Beginning with manuscripts submitted after 1 January 1998, The Pan-Pacific Entomologist will require that voucher specimens for all articles be deposited in a properly maintained collection accessible to other scientists. This policy repre- sents a departure in tradition for non-taxonomic studies that 1s necessitated by the rapid discovery of numerous cryptic species and species complexes. These situ- ations make it imperative that future scientists be able to confirm exactly what Species was studied in past papers. The uncertainty surrounding whether older studies used Bemisia tabaci (Gennadius) or Bemisia argentifolii (Bellows & Per- ring) is a good example of a situation that could have been avoided if voucher specimens were available. Similar confusion surrounds older studies of Bactro- cera dorsalis (Hendel) from numerous locations, several Rhagoletis species and numerous aphids. The location at which the voucher specimen(s) have been de- posited and the coding necessary to access them shall be noted in the Materials and Methods section of the article or the text of the scientific note. 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 11 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 |), abstract and key words page (page 2), text pages (pages 3+), acknowledgment page, literature cited pages, footnote page, tables, figure caption page; place orginal figures last. List the corresponding author’s name, address including ZIP code, and phone number on the title page in the upper right corner. 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 KEY WORDS. 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. FE 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. — A]] 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 morphometrics provide some answers? pp. 233—238. In Holman, J., J. Pelikan, A. G. EF 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. — I]lustrations 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. THE PAN-PACIFIC ENTOMOLOGIST Volume 73 October 1997 Number 4 Contents MILLER, R. EF & J. A. DAVIDSON—Obituary: Richard F Wilkey (1925-1995) _____W.--- PECK, R. W.,, A. EQUIHUA-MARTINEZ & D. W. ROSS—Seasonal flight patterns of bark and ambrosia beetles (Coleoptera: Scolytidae) in northeastern Oregon IWATA, R., EF YAMADA, H. KATO, H. MAKIHARA, K. ARAYA, H. ASHIDA & M. TA- KEDA—Nature of galleries, durability of boring scars and density of Xylotrechus villioni (Villard) larvae (Coleoptera: Cerambycidae) on coniferous tree trunks HSU, Y-F—A new Japonica (Lepidoptera: Lycaenidae: Theclinae) from southwestern China _. RASNITSYN, A. P—Xylea (Pinicolites) lata Smith (Vespida: Xyelidae), a living fossil sawfly from western North America SCIENTIFIC NOTES WALKER, G. P, N. ZAREH, I. M. BAYOUN & V. TRIAPITSYN—Introduction of western Asian egg parasitoids into California for biological control of beet leafhopper, Circulifer tenellus ZUPARKO, R. L.—Notes on Brachycaudonia Ashmead spp. (Hymenoptera: Pteromalidae) -... GULMAHAMAD, H.—Naturally occurring infestations of drywood termites in books The Pan-Pacific Entomologist: Format Information for Contributors The Pan-Pacific Entomologist: Reviewers for Volume 73 The Pan-Pacific Entomologist: Table of Contents for Volume 73 The Pan-Pacific Entomologist: Index for Volume 73 204 213 Zen 231 236 243 245 248 256 237 260