Vol. 28, No.3 &4 Fall/Winter 1995 THE GREAT LAKES ENTOMOLOGIST

PUBLISHED BY

THE MICHIGAN ENTOMOLOGICAL SOCIETY THE GREAT LAKES ENTOMOLOGIST

Published by the Michigan Entomological Society

Volume 28 No.3 & 4

ISSN 0090-0222

TABLE OF CONTENTS

Temperature effects on development of three cereal aphid porasitoids {Hymenoptera: Aphidiidael N. C. Elliott,J. D. Burd, S. D. Kindler, and J. H. Lee...... 199

Parasitism of P/athypena scabra (Lepidoptera: Noctuidael by Sinophorus !eratis (Hymenoptera: Ichneumonidae) David M. Pavuk, Charles E. Williams, and Douglas H. Taylor ...... 205

An allometric study of the boxelder bug, Boiseo Irivillata (Heteroptera: Rhopolidoe) Scott M. Bouldrey and Karin A. Grimnes ...... 207

S/aferobius insignis (Heleroptera: Lygaeidael: association with granite ledges and outcrops in Minnesota A. G. Wheeler, Jr...... 213

A note on the sympotric collection of Chymomyza (Dipiero: Drosophilidael in Virginio's Allegheny Mountains Henretta Trent Bond ...... 217

Economics of cell partitions and closures produced by Passa/oecus cuspidafus (Hymenoptera: Sphecidael John M. Fricke...... 221

Distribution of the milliped Narceus american us annularis (Spirabolida: Spirobolidae) in Wisconsin Dreux J. Watermolen...... 225

Adult female Mydas clavafus (Diptera: Mydidae) feeding on flowers in Wisconsin Andrew H. Williams...... 227

A Michigan record for Clyrus marginicollis (Coleoptera: Cerambycidoe: Clytini) James E. Zablotny ...... 231

Uraphoraquadrifasciala {Diptera: TephritidaeL an introduced seedhead fly new to midwestern North America A. G. Wheeler, Jr...... 235

First county records for Acarapis woodi (Acari: TarsonemidaeJ in Michigan Murray Hanna and Sharon Pratt Anzolduo ...... 237

Three new food plonts and first Wisconsin record of Pub/ilia reticulala {Hemiptera: Membracidael Andrew H. Williams ...... c ...... 243

COVER PHOTOGRAPH elytus marginicol/is (Coleoptera: Cerambycidae). Drawing by 1. E. Zablotny. THE MICHIGAN ENTOMOLOGICAL SOCIETY

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Copyright © 1995. The Michigan Entomological Society 1995 THE GREAT LAKES ENTOMOlOGIST 199

TEMPERATURE EFFECTS ON DEVELOPMENT OF THREE CEREAL APHID PARASITOIDS (HYMENOPTERA: APHIDIIDAE)

N. C. Elliott1, J. D. Burd, S. D. Kindler, and J. H. lee

ABSTRACT Temperature is an important climatological variable that influences the biology and ecology of insects. Poor climatic adaptation can limit the effec­ tiveness of parasitic insects in biological control. Two exotic parasites (Syrian Diaeretiella rapae (M'Intosh) and Argentinean Aphidius colemani Viereck) imported for biological control of the Russian wheat aphid, Diuraphis noxia (Mordvilko), and one native parasite (Diaeretiella rapae) were reared in growth chambers in three fluctuating temperature regimes with average daily temperatures of 12, 18, and 24°C. Estimates of temperature thresholds for immature development were 3.3, 3.5, and 2.8°C, for Oklahoman D. rapae, Syrian D. rapae, and A. colemani, respectively. Estimates of thermal require­ ments for development from egg to adult were 297, 278, and 301 degree-days for the three parasitoids. Dry weights of adults reared in different fluctuating temperature regimes did not differ significantly among sexes, but adults from regimes with low average temperatures of 12 and 18°C had significantly greater weights than those reared in a regime with an average temperature of 24°C. Results suggest that developmental response to temperature will not limit the effectiveness of the exotic parasites in biological control.

Diaeretiella rapae M' Intosh is a cosmopolitan aphidiid that commonly parasitizes a wide range of hosts in agroecosystems (Mackauer and Stary 1967). Aphidius colemani Viereck is widely distributed in the southern hemi­ sphere and broadly oligophagous on Aphididae (Stary 1975). Both species par­ asitize several economically important aphid pests of small grain crops in­ cluding the greenbug, Schizaphis graminum (Rondani), and Russian wheat aphid, Diuraphis noxia (Mordvilko) (Mackauer and Stary 1967, Stary 1975). Both A. colemani and D. rapae were imported into the United States for clas­ sical biological control of the Russian wheat aphid in the Southern Great Plains and elsewhere in the western US (Gould and Prokrym 1994). Temperature is an important abiotic variable that influences insect popu­ lations in the field. Temperature can influence insect population growth through its effects on development rate, survival, fecundity, and dispersal (Ratte 1984, Rankin and Singer 1984). Temperature may partially determine the effectiveness of a parasitoid as a biological control agent in a particular re­ gion. For example, a parasitoid with a higher developmental threshold tem­ perature than its host may have limited effectiveness in regions where tem­ peratures below the parasitoids developmental threshold but above the hosts

1 USDA, ARS, SPA, Plant Science Research Laboratory, 1301 N. Western St., Still­ water, OK 74075. 200 THE GREAT LAKES ENTOMOLOGIST Yol. 28, No.3 & 4 threshold occur often, because development from birth to maturity, and hence population growth rate would lag behind that of its host (Campbell et al. 1974). Thus, in addition to factors such as searching ability, host preference, and host suitability, knowledge of developmental responses to temperature should be considered when deciding whether to release an exotic parasitoid for biological control (Bernal and Gonzalez 1993). Estimates of developmental thresholds for D. rapae populations from dif­ ferent regions range from 2.1 to 7.0'C (Campbell et al. 1974, Bernal and Gon­ zalez 1993). The estimates may result from adaptation to geographic differ­ ences in climate (Campbell et al. 1974). To the best of our knowledge, the developmental threshold of A. colemani has not been reported. The first ob­ jective of this study was to determine developmental thresholds and temper­ ature requirements for development of three cereal aphid parasitoids. Two of the parasitoids were imported into the US; Diaeretiella rapae was imported from Syria, while A. colemani was imported from Argentina. A native Okla­ homa population of D. rapae was also examined. Because this parasitoid is en­ demic to the Southern Great Plains, and presumably well adapted to the cli­ mate there, estimation of temperature effects on its development may serve as a benchmark against which to assess the exotic parasitoids. The second ob­ jective was to determine the influence of temperature during immature stages on adult size. Body size is related to fecundity in aphidiids (Hofsvang 1991) and is therefore an important factor influencing parasitoid population dy­ namics.

MATERIALS AND METHODS The parasitoids used in this study were obtained from three sources. Di­ aeretiella rapae was reared from Aphis gossypii Glover on cantaloupe from fields near Lane, Oklahoma in 1991 and collected from D. noxia in wheat fields in Syria in 1990. The Syrian collection was designated T90026 at the Texas A&M University quarantine facility to distinguish it from other collec­ tions. A colony of A. colemani was established from cereal aphids collected in wheat fields in Argentina in 1990. Parasitoid colonies were maintained in plant growth chambers at 20-24°C and 16L:8D using D. noxia as host. Development time from oviposition to adult eclosion was determined for cohorts exposed to three fluctuating temperature regimes in programmable plant growth chambers. Temperature in each of the three regimes oscillated through a sine wave with a 24-h period and an amplitude of lOoC. Mean daily temperatures in the three regimes were 12, 18, and 24°C. Relative humidity was maintained at 50±5% and photoperiod was 16L:8D. Parasitoid cohorts were established by placing 50 1st-3rd instar D. noxia on barley seedlings growing in 10-cm diameter pots (approximately 4 seedlings per pot). Seedlings and nymphs were then covered with a vented plastic cage. Nymphs were allowed to settle on plants for approximately 4-h after which time 10 mated female parasitoids were introduced into the cage. The caged plants, aphids, and parasitoids were held in the laboratory at room temperature (20-22°C) for 4-h to allow ample time for oviposition. Afterwards, adult parasitoids were removed and the caged plant with aphids was placed in a growth chamber programmed for one of the three temperature regimes. Plants were checked daily for mummies which were removed, placed individ­ ually in labeled petri dishes (5-cm diameter by 1A-cm height) and returned to the growth chamber. Petri dishes were shaded to eliminate the effect of irra­ 1995 THE GREAT LAKES ENTOMOLOGIST 201 diation from chamber lights on temperature within the dishes. Dishes were checked daily for emergence of adult parasitoids. Adult parasitoids from a co­ hort were identified to sex, pooled according to sex, and freeze dried using a speed-vac. Dried parasitoids from a cohort ofa particular sex were weighed as a group. Approximately 7 cohorts were established per temperature regime for each parasitoid. Lower developmental thresholds and thermal (degree-day) requirements for immature development were estimated using methods of Campbell et al. 1974. Dry weights and developmental times were subjected to analysis of variance to test for differences among sexes and temperature regimes. All sta­ tistical hypotheses were tested at the 0.=0.05 level of significance.

RESULTS AND DISCUSSION The number of days required for development decreased with increasing average temperature for each of the three parasitoids (Table 1). Development times for the three parasitoids were similar at each temperature. For exam­ ple, development times ranged from 34.6 days for Syrian D. rapae andA cole­ mani to 35.9 days for Oklahoman D. rapae reared at 12°C; while at 24°C de-

Table 1. Mean number of days (:t SE) required for immature development and mean adult dry weight (:tSE) of three parasites reared in three fluctuating temperatures regimes. Temp. No. Dry Weight Species/LocationfSex (OC) Individuals Days (mg) D. rapae - Oklahoma Female 12 30 35.9 (0.3S) 0.046 (0.0012) IS 31 1S.9 (0.41) 0.041 (0.0017) 24 29 14.5 (0.26) 0.037 (0.0011) Male 12 27 35.9 (0.59) 0.040 (0.0019) IS 32 1S.7 (0.33) 0.042 (0.0049) 24 34 14.S (0.35) 0.036 (0.0021) D. Rapae - Syria Female 12 23 34.6 (0.65) 0.049 (0.0073) IS 37 18.9 (0.2S) 0.039 (0.0067) 24 41 14.0 (0.28) 0.036 (0.0062) Male 12 32 33.5 (0.38) 0.045 (0.0074) 18 27 17,4 (0.32) 0.041 (0.0048) 24 26 13.5 (0,43) 0.034 (0.0065) A. colemani ­ Argentina Female 12 13 34.6 (0.68) 0.042 (0.0076) IS 19 19.3 (0.37) 0.038 (0.0032) 24 26 14.7 (0.15) 0.028 (0.0066) Male 12 15 34.3 (0.95) 0.043 (0.0082) 18 14 17.9 (0.30) 0.034 (0.0033) 24 15 14.5 (0.25) 0.029 (0.0066) 202 THE GREAT LAKES ENTOMOLOGIST Vol. 28, No.3 & 4

velopment times ranged from 13.5 days for Syrian D. rapae to 14.8 days for Oklahoman D. rapae. Females generally required slightly longer to develop than males, but the differences were small and not significant for any of the parasitoid species. Therefore, development times of the sexes were pooled for each species for the purpose of determining developmental thresholds and thermal requirements. Developmental thresholds were 3.3, 3.5, and 2.8°C for Oklahoman D. rapae, Syrian D. rapae, and A. colemani, respectively. Thermal requirements for immature development were 293, 273, and 300 degree-days for Okla­ homan D. rapae, Syrian D. rapae, and A. colemani, respectively. Previously reported estimates of the developmental threshold tempera­ ture for D. rapae from various locations range from 2.1 to 7.0°C (Campbell et al. 1974, Bernal and Gonzalez 1993). Our estimates for D. rapae from Okla­ homa and Syria are within the range of those previously reported. A parasitoid with a thermal threshold that differs markedly from that of its host may be ineffective in biological control due to poor temporal synchrony in population growth rates of host and parasitoid caused by differences in de­ velopment rates from birth to maturity (Campbell et al. 1974, Bernal and Gonzalez 1993). Estimates of developmental thresholds for known cereal aphid host species of D. rapae and A. colemani range from 0.9-6.1°C (Elliott et al. 1988, Elliott and Kieckhefer 1989, Kieckhefer and Elliott 1989, Kieckhefer et al. 1989, Honek and Kocourek 1990). Thus, it appears that developmental thresholds for Syrian D. rapae and A. colemani are within the range of devel­ opmental thresholds previously reported for their hosts in cereal agroecosys­ terns, suggesting that biological control of cereal aphids in the Great Plains by these exotic parasitoids may not be limited by differential developmental re­ sponses to temperature. It should be mentioned that thermal thresholds were estimated from pop­ ulations of aphid species from widely divergent regions, and while several es­ timates were from cereal aphid populations from the Northern Great Plains, none were from aphids collected from the Southern Great Plains. Some stud­ ies of geographical variation in thermal thresholds of aphids indicate that populations from cooler climates have lower thresholds (Campbell et al. 1974, Hutchison and Hogg 1984), while others fail to detect differences in thresh­ olds among geographic populations (Lamb et al. 1987). Thus, we cannot pro­ vide conclusive evidence that these parasitoids are well adapted for exploiting cereal aphids in the Southern Great Plains in terms oftheir developmental re­ sponses to temperature. However, the similarity of developmental thresholds and thermal development requirements for D. rapae native to Oklahoma, which is presumably well adapted to the climate there, and Syrian D. rapae and A. colemani suggests that these exotic parasitoids are adapted to the cli­ mate of the Southern Great Plains, at least with respect to temperature ef­ fects on immature development. Dry weights of adult females were generally slightly greater than males at a given temperature (Table 1), but differences in weights did not differ sig­ nificantly among sexes for any of the parasitoids. Dry weights generally de­ creased with increasing average rearing temperature from approximately 0.045 mg at 12°C to approximately 0.035 mg at 24°C for D. rapae from both locations, and from approximately 0.042 mg at 12°C to 0.029 mg at 24°C for A. colemani (Table 1). For Oklahoman D. rapae and A. colemani, weights of individuals reared at 12°C were significantly greater than those reared at 24°C; for Syrian D. rapae significant differences in weight were evident among all temperature regimes. Larger body size is generally associated with 1995 THE GREAT LAKES ENTOMOLOGIST 203 greater fecundity in aphidiid species CHofsvang 1991). Thus, temperature may influence fecundity, and therefore rates ofpopulation growth, of D. rapae and A colemani indirectly through its affect on adult size. However, the size of all three parasitoids appears to be affected similarly by variation in temperature, so that reduced size at higher temperatures is a characteristic of both the ex­ otic and native aphidiids studied. Our study suggests that the two exotic aphidiids studied are adapted to the climate of the Southern Great Plains, at least with respect to the influence of temperature on immature development. Furthermore, normal fluctuations in climate, such as seasonal variation in temperature, may influence popula­ tion growth rates of the aphidiids in an obvious way, through effects on im­ mature development, and in a less obvious way through effects on fecundity related to variation in adult size.

ACKNOWLEDGMEJI.;'TS We thank Wade French, Tim Johnson, and Perry Shelby for technical as­ sistance. Mention of a proprietary product does not constitute endorsement for its use by the USDA We also thank Dave Reed and Keith Pike for foreign explorations resulting in collection of the exotic parasitoids used in this study, and Bob Cartwright for supplying native parasitoids.

LITERATURE CITED Bernal, J., and D. Gonzalez. 1993. Temperature requirements of four parasites of the Russian wheat aphid Diuraphis noxia. Entomol. Exp. AppL 69:173-182. Campbell, A., B. D. Frazier, N. Gilbert, A. P. Gutierrez, and M. Mackauer. 1974. Tem­ perature requirements of some aphids and their parasites. J. Appl. Ecol. 11:431-438. Elliott, N. C., R. W. Kieckhefer, and D. D. Walgenbach. 1988. Effects of constant and fluctuating temperatures on developmental rates and demographic statistics for the corn leaf aphid (Homoptera; Aphididae). J. Econ. Entomol. 81:1383-1389. Elliott, N. C., and R. W. Kieckhefer. 1989. Effects of constant and fluctuating tempera­ tures on immature development and age-specific life tables of Rhopalosiphum padi (L.) (Homoptera; Aphididae). Can. Entomol. 121:131-140. Gould, J. R. and D. Prokrym.. 1994. APHIS project summary; Russian wheat aphid bi­ ological control project. Pages 9-15, Proc. 6th Russian Wheat Aphid Workshop, Jan­ uary 23-25, Ft. Collins, Colorado. 268 pp. Hofsvang, T. 1991. Fecundity of aphid parasitoids in the family Aphidiidae (Hy­ menoptera). (A review). in; L. Polgar, R. J. Chambers, A. F. G. Dixon, and I. Hodek (Eds.), Behaviour and Impact of Aphidophaga. SPB Academic Publishingbv, The Hague. 350 pp. Honek, A. and F. Kocourek. 1990. Temperature and development time in insects: A gen­ eral relationship between thermal constants. Zool. Jb. Syst. 117:401-439. Hutchison, W. D., and D. B. Hogg. 1984. Demographic statistics for the pea aphid (Ho­ moptera: Aphididae) in Wisconsin and a comparison with other populations. Env. En­ tornol. 13;1173-1181. Kieckhefer, R. W., and N. C. Elliott. 1989. Effect of fluctuating temperatures on devel­ opment of immature Russian wheat aphid (Homoptera: Aphididae) and demographic statistics. J. Econ. Entomol. 82;119-122. Kieckhefer, R. W., N. C. Elliott, and D. D. Walgenbach. 1989. Effects of constant and 204 THE GREAT LAKES ENTOMOlOGIST Vol. 28, No.3 & 4

fluctuating temperatures on developmental rates and demographic statistics of the English grain aphid (Homoptera: Aphididae). Ann. EntomoL Soc. Amer. 82:701-706. Lamb, R. J., P. A. MacKay, and G. H. Gerber. 1987. Are development and growth of pea aphids, Acyrthosiphon pisum, in North America adapted to local temperatures. Oe­ cologia 72:170-177. Mackauer, M. and P. Stary. 1967. Hym. Ichneumonoidea, World Aphidiidae. in: V. Deluc­ chi and G. Remaudiere (Eds.), Index of Entomophagous Insects. Le Francois, Paris, 167 pp. Rankin, M. A. and M. C. Singer. 1984. Insect movement: mechanisms and effects. in: C. B. Huffaker and R. L. Rabb (Eds.), Ecological Entomology. John Wiley & Sons, New York. 844 pp. Ratte, H. T. 1984. Thmperature and insect development. In. K. H. Hoffmann (Ed.), En­ vironmental Physiology and Biochemistry of Insects. Springer Verlag, Berlin. 296 pp. Stary, P. 1975. Aphidius colemani Viereck: its taxonomy, distribution and host range (Hymenoptera, Aphidiidae). Acta Entomol. Bohemoslov.72:156--163. 1995 THE GREAT LAKES ENTOMOLOGIST 205

PARASITISM OF PLATHYPENA SCABRA (LEPIDOPTERA: NOCTUIDAEI BY SINOPHORUS TERATIS (HYMENOPTERA: ICHNEUMONIDAEI

Daniel M. Pavukl, Charles E. Williams2, and Douglas H. Taylor3

A study was conducted at the Ecology Research Center, Miami University, Butler County, Ohio, during the summer of 1990 to examine the effects of strip intercropping sorghum and soybean on the occurrence ofparasitoids and incidence of disease in larvae ofthe green cloverworm, Plathypena scabm (F.) (Lepidoptera: Noctuidae), a sporadic pest of soybeans. The details of the ex­ perimental design and results are reported elsewhere (Williams et al. 1995). Ten species of larval parasitoids were reared from a total of 1,522 P. Bcabm larvae (Williams et al. 1995). One species, Sinophorus teratis (Weed), has apparently not been reared from P. scabra larvae previously (e.g., White­ side et al. 1967, Barry 1970, Lentz and Pedigo 1975, Roberts et al. 1977, Mueller and Kunnalaca 1979, McCutcheon and Turnipseed 1981, Hammond 1983, Pedigo et a11983, Pavuk and Barrett 1993). However, Hammond (1983) reported a single specimen of Sinophorus sp. from green cloverworm, Lentz and Pedigo (1975) reared one individual ofSinophorus validus (Cresson) from P. scabra, and Pedigo et al. (1983) observed a small proportion ofgreen clover­ worm larvae parasitized by S. validus. Sinophorus teratis was formerly a syn­ onym for S. validus; Sanborne (1984) removed S. temtis from synonymy with S. validus when he revised the world species of Sinophorus, and the two names now refer to two separate species. It is possible that records of S. validus from P. Bcabm may actually have been occurrences of S. temtis. In addition, rates of parasitism of P. scabra by Sinophorus spp. in other investigations were extremely low compared to our findings. Percentage of green cloverworm larvae parasitized by S. teratis pooled across the sampling period (i.e., 27 July to 14 September; 8 weekly samples) in the five different soybean agroecosystems ranged from 2.3 to 5.3% (Williams et al. 1995). Sur­ veys of green cloverworm parasitoids in the same study area but in different soybean agroecosystems in subsequent years failed to detect S. temUs (un­ published data). The occurrence of this parasitoid in this particular locality appears to be sporadic, and may be affected by varied plant community struc­ ture, among other factors (e.g., presence of alternate hosts). A survey on a larger scale and in widely separated, diverse soybean cropping systems would be valuable in determining the factors that may possibly influence parasitism ofP. scabra by S. temtis.

ACKNOWLEDGMENTS We thank the staff of the Ecology Research Center, especially M. Ben­ ninger-Truax, E. Bollinger, J. RaIley and R. Stander for field and laboratory

IDepartment of Biological Sciences, Bowling Green State University, Bowling Green, OH 43403. 2Department of Biology, Clarion University of Pennsylvania, Clarion, PA 16214­ 1232. 3Department ofZoology, Miami University, Oxford, OH 45056. 206 THE GREAT LAKES ENTOMOLOGIST Vol. 28, No.3 &4 assistance. We also thank R.W. Carlson, USDA-ARS, Taxonomic Services Unit, Beltsville, MD for identifying Sinophorus teratis. This study was sup­ ported in part by an Ohio Academic Challenge Grant in applied ecology to the Department of Zoology, Miami University.

LITERATURE CITED Barry, R.M. 1970. Insect parasites of the green c1overworm in Missouri. J. Econ. Ento­ mol. 63:1963-1965. Hammond, R.B. 1983. Parasites of the green cloverworm (Lepidoptera: Noctuidae) on soybeans in Ohio. Environ. Entomo1. 12:171-173. Lentz, G.L., and L.P. Pedigo. 1975. Population ecology of parasites of the green clover­ worm in Iowa. J. Econ. Entomol. 68:301-304. McCutcheon, G.S., and S.G. Turnipseed. 1981. Parasites oflepidopterous larvae in in­ sect resistant and susceptible soybeans in South Carolina. Environ. Entomol. 10: 69-74. Mueller, A.J., and S. Kunnalaca. 1979. Parasites of green cloverworm on soybeans in Arkansas. Environ. Entomol. 8:376-379. Pavuk, D.M. and G.W. Barrett. 1993. Influence of successional and grassy corridors on parasitism of Plathypena scabra (F.) (Lepidoptera: Noctuidae) larvae in soybean agroecosystems. Environ. Entomol. 22:541-546. Pedigo, L.P., E.J. Bechinski, and R.A. Higgins. 1983. Partial life tables of the green c1overworm (Lepidoptera: Noctuidae) in soybean and a hypothesis of population dy­ namics in Iowa. Environ. Entomol. 12:186-195. Roberts, S.J., W.K. Mellors, and E.J. Armbrust. 1977. Parasites oflepidopterous larvae in alfalfa and soybeans in central TIlinois. Great Lakes Entomol. 10:87-93. Sanborne, M. 1984. A revision of the world species of Sinophorus (Ichneumonidae). Mem. Am. Ent. Inst. 38:1-403. Whiteside, R.C., P.P. Burbutis, and L.P. Kelsey. 1967. Insect parasites of the green c1overworm in Delaware. J. Econ. Entomol. 60:326-328. Williams, C.E., D.M. Pavuk, D.H. Taylor, and T.H. Martin. 1995. Parasitism and dis­ ease incidence in the green c1overworm (Lepidoptera: Noctuidae) in strip-inter­ cropped soybean agroecosystems. Environ. Entomol. 24:253-260. 1995 THE GREAT lAKES ENTOMOLOGIST 207

AN ALLOMETRIC STUDY OF THE BOXELDER BUG, BO/SEA TRIV/TTATA (HETEROPTERA: RHOPALIDAE)

Scott M. Bouldrey and Karin A. Grimnes 1

ABSTRACT An allometric study was conducted on the boxelder bug, Boisea trivittata, to confirm the number ofinstars and to identify characteristics most useful for rapid instar identification of field samples. Analysis of field populations col­ lected throughout the 1990-92 seasons indicated that there were five instars, clearly defined on the basis of size and the presence of wing pads. This rmd­ ing is in contrast with the only other published study on stages ofthe boxelder bug, which claims there are six nymphal instars. Size data gathered from field populations were substantiated by labora­ tory growth studies. Head width and/or second antennal segment length were identified as the most useful parameters for instar identification.

Boisea trivittata (Say), the boxelder bug, is a pest closely associated with the pistillate boxelder tree (Acer negundo) throughout much of their shared range (Slater and Schaefer 1963). During hot, dry summers, the number of boxelder bugs can reach epidemic proportions, as evidenced by a long history of scattered reports in the literature (see the review in Wheeler 1982). Al­ though they rarely cause economic damage, boxelder bugs are considered a significant household nuisance. A recurring outbreak site in Gratiot County, Michigan, was discovered in the spring of 1990. As we be~an population studies, we realized that the only detailed report on this insect s life history identifies six nymphal instars after hatching (Smith and Shepherd 1937). According to these authors, minute wing pads did not appear until the fourth instar, and the authors did not iden­ tify any exuvia left in the egg after hatching (an egg-bound molt). The gen­ eral rule for hemipteran species appears to be five nymphal instars before the adult ecdysis, not counting any egg-bound molt left after hatching (Slater and Baranowski 1978). Subsequent investigators of this insect either avoided a direct challenge of the instar number (Tinker 1952) or divided the population into only adults and nymphs (Yoder and Robinson 1990). We became inter­ ested in the question of instar number for the boxelder bug because our pre­ liminary studies of field populations showed only five clearly discernable in­ stars that were not easily reconciled with results of Smith and Shepherd (1937). As a result, we began a detailed growth study and allometric analysis of our population of B. trivittata for use in subsequent seasonal studies.

1 Author to whom reprint requests should be sent: Department of Biology, Alma College, Alma, MI 48801. -

208 THE GREAT lAKES ENTOMOLOGIST Vol. 28, No.3 & 4

MATERIALS AND METHODS Insects were collected at Lang's Veterinary Clinic west ofAlma in Gratiot County, Michigan, biweekly from March emergence to October hibernation during 1990-1992. The site contains numerous staminate and pistillate box­ elder trees (Acer negundo) close to the residence. Insects were swept into 70% ethanol with a small brush from the same stretch of south-facing wall and west-facing fence on each collecting date. Samples were separated into five nymphal instars or adults on the basis of body size, presence and degree of wing pad growth, and head width. All measurements were made under a dissecting microscope with an ocular mi­ crometer. Body parts (see Tables 1 and 2) were measured at their maximum point with the exception of antennal segments (1-4) where only the melanized portions were measured. For each instar, and for adult males and females, thirty-five individuals were measured. Mated females were allowed to oviposit in the laboratory on filter paper in 100 mm petri dishes. Because mortality of individually raised animals was high, first instars were kept in groups of five in petri dishes containing moist­ ened filter paper. Maple syrup (5% v/v) was loaded into cotton-plugged tub­ ing and placed in the dish for food. The dishes were exposed to a 16:8 LD pho­ toperiod and maintained at 24°C ± 1°C in an incubator. Animals were observed daily. molts were recorded, and the newly molted animals were moved to new dishes (in groups of five if possible). Head widths and second antennal segment lengths were recorded for recently molted individuals. Means between morphological characteristics of male and female adults, and between field and laboratory-raised animals, were compared with stu­ dent's t-test. Growth curves were fit using linear regression techniques.

RESULTS Size of nymphs is highly correlated with age for all parameters measured in this study (Table 1). When growth data for the five instars were analyzed with simple linear regression, no parameter had an r2 value less than 0.90. Head width was most highly correlated to instar number, as was length ofthe second antennal segment. For pronotallength and width, visual inspection of the data indicated that a logarithmic curve might better describe the growth relationship. Subsequent analysis of the logarithmic data produced an r2 value of 0.96 (pronotum length) and 0.97 (pronotum width). All correlations reported in this study are somewhat time independent because the regression analysis is against instar number (which assumes equal duration of each stage) and not against actual growth in days. Adult females were significantly larger than adult males for all parame­ ters measured (Table 2). Difference in body length was easily discernable in the field. No attempt was made to sex fifth instars in this preliminary study, so some of the variation in fifth instar measurements could result from sex­ related growth differences. Although head width classically is used as an indicator ofgrowth for many insect species, we noticed that the length of the second antennal segment was equally predictive of instar number in B. trivittata (Table 1). This observation led us to compare growth rates (using instar number, and not actual time) of the four antennal segments (Fig 1). Clearly, the first segment changes little as the animal grows, but all other segments lengthen noticeably with instar. The second segment is the second shortest at hatching, yet by the fifth instar, '"~

Table 1. Measurements of nymphs of Boisea trivittata· (Say). Values are mean mm ± SE, n=35 (based on field collected individuals). Character 1st Instar 2nd Instar 8rd Instar 4th Instar 5th Instar r2 slope :c:--t m Body length 2.91 ± 0.04 3.80:!: 0.06 5.11:!: 0.06 7.01 :!: 0.09 9.60:!: 0.11 0.93 1.67 Q ;><:> m Head width 0.68 ± 0.01 0.92:!: 0.01 1.17 :!: 0.01 1.51 :!: 0.01 1.89 ± 0.02 0.97 0.30 ~ Interocular dist. 0.50:!: 0.01 0.63:!: 0.01 0.80:!: 0.01 1.04:!: 0.01 1.28:!: 0.01 0.96 0.20 ~ A m Pronotallength 0.29:!: 0.01 0.38:!: 0.01 0.57:!: 0.01 0.76:!: 0.01 1.51 ± 0.01 0.91 0.21 Vl m Z Pronotal width 0.80 ± 0.01 1.O1:!: 0.01 1.30:!: 0.01 1.69:!: 0.02 2.51:!: 0.03 0.90 0.41 --t 0 Antennal length ~ r­0 Segment 1 0.23:!: 0.01 0.28:!: 0.01 0.37:!: 0.01 0.49:!: 0.01 0.64 ± 0.01 0.93 0.10 0 Q Segment 2 0.45:!: 0.01 0.68:!: 0.01 0.93 ± 0.01 1.23 :!: 0.01 1.70:!: 0.01 0.97 0.31 Qj

Segment 3 0.46:!: 0.01 0.61 ± 0.01 0.81 ± 0.01 1.07 ± 0.01 1.44:!: 0.01 0.95 0.24

Segment 4 0.65:!: 0.01 0.85:l: 0.01 1.05 ± 0.01 1.28 ± 0.01 1.64:l: 0.01 0.95 0.24

N ~ 210 THE GREAT tAKES ENTOMOLOGIST Vol. 28, No.3 & 4

Table 2. Measurements of adults ofBoisea trivittata. Values are mean mm :!: SE, n=35 (field collected individuals). Adult Adult Character females males p valuesB

Body length 12.1 :!: 0.10 10.3 ± 0.08 <0.0001 Head width 2.14 ± 0.01 1.96 ± 0.02 <0.0001 Interocular dis. 1.34:!: 0.01 1.19 :!: 0.02 <0.0001 Pro notal length 1.93:!: 0.04 1.73 ± 0.02 <0.0001 Pronotal width 3.70 ± 0.03 3.09 ± 0.02 <0.0001 Antenna! length Segment 1 0.64:!: 0.01 0.62:!: 0.01 <0.01 Segment 2 2.09:!: 0.02 1.90 :!: 0.01 <0.0001 Segment 3 1.80 :!: 0.02 1.70:!: 0.02 <0.0001 Segment 4 1.90:!: 0.02 1.73 ± 0.02 <0.0001 a student's t-test.

2.5 E g "'-0--' antenna I seg 1 2.0 E --II- antennal seg II (]) -'--!r" antennal seg 111 E ..... ::~:::~ Ol ----0--- anlennal seg IV (]) en 1.5 .8···· til ...... 01' ..."'.. c: c: ...0...... K··· (]) 1.0 E .0,,- ", .-_••••­ til •••••• • ••• -l!: 15 0.5 .s:: c-oooooo_oooo::::::ooooC-oooo--ooooc-ooooo--ooo<>ooooooooooo t5l c: .!!!

insler 1 instar 2 inslar 3 inslar 4 inslar 5 adull Age

Figure 1. Mean length of antennal segments (mean ± SE) of field-collected Boisea tnvittata specimens. Thirty-five measurements were averaged for each nymphal data point; values for adults were obtained by averaging male and female means from Table 2.

it is the longest. In the regression analysis, the predictive growth line for the second antennal segment has the greatest slope (Table 1, Fig. 1). Growth studies in the laboratory were carried out on animals reared in groups of five, wherever possible. Animals reared as individuals perished quickly; only eight of 58 (13.8%) molted to the second instar, and none ofthose animals survived to the third instar. In contrast, 43.1% (148/343) of animals 1995 THE GREAT LAKES ENTOMOLOGIST 211 raised in groups offour or five survived to the second instar, and 17.0% (8/47) survived to the third instar. Mass rearing (even in groups of as few as five) increases the risk of death from cannibalism, a phenomenon reported first by Abbott (1948). In our study, cannibalism appeared to be rare. It was observed only in the absence of a water source and only when an animal emerged un­ usually helpless and unable to move away from the probosci of its fellow hatchlings. We found no evidence ofegg-bound exuvia after first instars hatched from the egg. Under our laboratory conditions, the first stadium averaged 5.5±0.2 days (n=148) and first instars had a mean head width ofO.64±0.01 mm (n=25) and a mean second antennal segment length of 0.42±0.01 mm (n=25). The second instars had a mean head width of 0.88±0.01 mm (n=20) and a mean second antennal segment length of 0.66±0.01 mm (n=20), while the mean in­ star duration was 6.5±0.8 days (n=8). For the third instars, the mean head width was 1.09±0.01 mm (n=6), while the mean length of the second anten­ nal segment was 0.92±0.01 mm (n=6). All third instar nymphs possessed wing pads, in contrast to the study of Smith and Shepherd (1937) where wing pads were not observed until the fourth instar. Although the mean values of all parameters measured were slightly smaller for insects raised in the labo­ ratory, neither head widths nor second antennal segment lengths were signif­ icantly different from field-caught instars (student's t-test). Unfortunately, all laboratory-reared animals were lost because of an incubator malfunction just before the majority ofinsects reached the third instar, resulting in a low num­ ber of data points (n=6) for measurements of that instar. Nevertheless, the presence of wing pads in lab-reared third instars clearly was established.

DISCUSSION There has not been a detailed growth analysis of the boxelder bug previ­ ous to the present study. Smith and Shepherd (1937) reported head width and body length data, but included only mean and range values. Also, as previ­ ously mentioned, they identified six, rather than five, instars for this insect. We believe we have clearly established the presence of five, and only five, in­ stars in B. trivittata, not only through our identification of discrete size classes for the parameters measured in field animals, but through the insects that were reared into the third instar. Although Smith and Shepherd report duration data for six instars, their sample numbers suggest that the same in­ dividuals were not followed throughout their entire life cycle. All of the third instars raised in our study possessed wing pads (often con­ sidered "the rule" for hemipteran species) gives the first clue toward explain­ ing the differences between our data and those ofSmith and Shepherd (1937). Comparing mean head widths for our third instar (1.17 mm) to their fourth (1.17 mm), our fourth instar (1.51 mm) to their fifth (1.51 mm), and our fifth instar (1.89 mm) to their sixth (1.83 mm) makes it clear that the discrepancy in instar identification must lie prior to our third instar. Also, mean head width for our second instar sample (0.92 mm) matches their third instar (0.90 mm), and if we average their values for first (0.64 mm) and second instars (0.78 mm), the head width of 0.71 mm compares nicely with our first instar value of 0.68 mm. No statistical differences were seen between our field and laboratory-reared populations for either head width or for length of the sec­ ond antennal segment for instars 1 - 3, further substantiating our proposed instar identification of field-caught animals. Based on these data, we believe that Smith and Shepherd may have inadvertently subdivided a highly vari­ able first instar population into smaller than average (instar 1) and larger 212 THE GREAT LAKES ENTOMOLOGIST Vol. 28, No.3 &4 than average (instar 2) samples and labelled them accordingly. We observed that slightly desiccated eggs may result in hatchlings that do not expand be­ fore sclerotization to the same extent as nondesiccated animals; thus, desic­ cation results in animals with smaller head widths and a wider variation in hatchling size. The values we report for adult head width, pronotum length and width, and length of the four antennal segments agree fairly well with the values presented by Schaefer (1975) for six populations of B. trivittata collected across the United States. His samples of adults were not identified b;V sex: slight differences in mean values, therefore, may represent variation In the sex ratios ofhis samples. Both male and female adults observed by Smith and erd (1937) possessed slightly larger dimensions for head width, body , and pronotal width than the adults in this study.

ACKNOWLEDGMENTS We thank Christina Krupp, University of Vermont, for editorial advice and Angie Williams, Alma College, for additional measurement data. This work was partially supported by a grant from the Kellogg Foundation enti­ tled: The Science Teacher Preparation Project.

LITERATURE CITED Abbott, C.E. 1948. Cannibalism in Leptocoris trivittatus Say. Bull. Brooklyn Entomol. Soc. 43:112-113. Schaefer, C.W 1975. A re-assessment of North American Leptocoris (Hemiptera-Het­ eroptera:Rhopalidae). Ann. Entomol. Soc. Am. 68:537-541. Slater, J.A. and RM. Baranowski. 1978. How to know the true bugs. Wm. C. Brown: Dubuque, Iowa. 256 pp. Slater, J.A and C.W Schaefer. 1963. Leptocoris trivittatus (Say) and Coriomeris humilis Uhl. in New England (Hemiptera: Coreidae). Bull. Brooklyn Entomol. Soc. 58:114-117. Smith, RC. and B.L. Shepherd. 1937. The life history and control of the boxelder bug in Kansas. Trans. Kansas Acad. Sci. 40:143-159. Tinker, M.E. 1952. The seasonal behavior and ecology of the boxelder bug Leptocoris trivittatus in Minnesota. Ecology 33:407-414. Wheeler, AG. 1982. Bed bugs and other bugs. pp. 313-345, in: A Mallis, ed. Handbook of Pest Control. Franzak and Foster Co., Cleveland, Ohio. Yoder, K.M. and WH. Robinson. 1990. Seasonal abundance and habits of the boxelder bug, Boisea trivittata (Say), in an urban environment. Proc. Entomol. Soc. Wash. 92:802-807. 1995 THE GREAT LAKES ENTOMOLOGIST 213

SLATEROBIUS INSIGNIS (HETEROPTERA: LYGAEIDAE): ASSOCIATION WITH GRANITE LEDGES AND OUTCROPS IN MINNESOTA

A. G. Wheeler, Jr. 1

ABSTRACT Adults and late-instar nymphs of the wide-ranging myrmecomorphic ly­ gaeid Slaterobius insignis were collected in northern Minnesota from cracks of granite outcrops and ledges, a habitat differing somewhat from that re­ ported elsewhere. At two of the four sites, S. ins ignis was observed in plant­ and litter-filled cracks with nymphs ofthe lygaeid Trapezonotus arenarius. All adults of the polymorphic S. insignis observed on granite were brachypters that belonged to the dark color morph of the species. Individuals occurred on rock surfaces with a black ant, Formica subsericea, which they resembled in appearance and behavior.

The myrmecomorphic lygaeid Slaterobius insignis (Uhler) belongs to the large subfamily Rhyparochrominae, tribe Myodochini. Formerly placed in the genus Sphaerobius (see Harrington 1980), this wide-ranging species is known from Newfoundland and Nova Scotia west to British Columbia, Alaska, the Northwest Territories, and the Yukon Territory south to New York, Iowa, Col­ orado, California, Texas, and Mexico (Slater 1964, Ashlock & Slater 1988, Scudder 1993, Slater et al. 1993). Its occurrence in the Northeast is thought to represent range expansion from western North America (Sweet 1964, Slater et aI. 1993). Habitat preferences are known only for a portion of this seed bug's exten­ sive range. In Connecticut and the Adirondacks of New York it occurs among clumps of little bluestem (Schizachrium scoparium) in open, xeric sites, in­ cluding railroad rights-of-way and roadsides. In these hot, barren areas this bivoltine lygaeid is associated with ants and feeds on fallen seeds of little bluestem and other grasses (Sweet 1964). The habitat reported for Nova Scotia is similar-a south-facing sandy slope (Scudder 1993)-but the habitat in Newfoundland differs by being wet­ ter: open, swampy ground harboring colonies of Carex and Sphagnum (Lind­ berg 1958). In the Midwest, S. insignis inhabits climax prairie dominated by Andropogon (or Schizachrium) and Stipa (Hendrickson 1930). In western Canada it occurs in "Mixed Prairie, Tallgrass prairie, and Fescue Grasslands" of the Prairie Provinces; in British Columbia it has been collected in the Pacific Northwest Bunchgrass ecosystem. Habitats in the Yukon include south-facing slopes characterized by Festuca grasslands and Artemisia (Scud­ der 1993). Sweet (1964) noted that fieldwork throughout the range of S. insignis was needed. Herein, I describe the sites where this lygaeid was observed in Min­ nesota, an area from which habitat preferences remain unrecorded. Notes are given on wing condition and color forms of this polymorphic species.

lBureau of Plant Industry, Pennsylvania Department of Agriculture, Harrisburg, PA 17110. 214 THE GREAT LAKES ENTOMOLOGIST Vol. 28, No.3 &4

COLLECTION SITES AND OBSERVATIONS All collections were made by the author in northern Minnesota during 25-27 June 1995. 1. Lake of the Woods Co., granite ledges (Fig.l) above Rapid River, east edge ofClementson; 4 adults and 2 late-instar nymphs in crevices with a thin layer of pine needles (Pinus strobus) and white pine "flowers" (microsporan­ giate strobili). 2. Koochiching Co., Wayside Rest, Rt. 11, 3 mi. W. of Indus; 1 adult from granite outcrop near Rainy River; in crack filled with moss, lichens, and grasses. 3. Koochiching Co., Tilson Bay, Rainy Lake; 5 adults, 1 fifth-instar nymph from grass-filled crevices of granite outcrop. 4. Lake Co., Bayside Park, Silver Bay; 9 adults, 1 fifth instar on granite cliffs along shore of Lake Superior; in cracks dominated by Potentilla triden­ tata (Fig.2) with occasional grass tufts.

DISCUSSION The habitats ofS. insignis observed in Minnesota-cracks and crevices of extensive granite outcrops and ledges-have not been reported in other areas of its range. Collection sites are somewhat similar, though, to many of those Sweet (1964) reported for New England-that is, open and xeric.

Figure 1. Granite ledges in Lake of the Woods Co., Minn., where S. insignis was found under white pine needles. 1995 THE GREAT LAKES ENTOMOLOGIST 215

Figure 2. Granite outcrop in Nova Scotia with Potentilla tridentata. Habitat is similar to that in Minnesota where S. insignis occurred in cracks colonized by the same plant species.

Individuals of S. insignis were occasionally seen running over the granite surface near plant-and litter-filled cracks, where they resembled coexisting black ants in their appearance and behavior. But the lygaeid was typically de­ tected only by scratching accumulated litter, which caused the bugs to leave the cracks and run onto the rock. The ant species observed at the Lake County site (no. 4) was Formica subsericea Say. At sites 1 and 3, late-instar nymphs of another rhyparochromine co-occurred with S. insignis; this gonianotine ap­ pears to be Trapezonotus arenarius (L.), a Holarctic species that in New Eng­ land is found mainly among sparse vegetation in dry, open areas of slopes (Sweet 1964). Slaterobius insignis was not encountered in cracks of smaller granite out­ crops along roadsides. Associated with these less expansive outcrops were adults and a few late instars of another myodochine, Ligyrocoris sylvestris (L.). This species was observed in litter at the edge of these outcrops, rather than in rock cracks and crevices. Further sampling in Minnesota is needed to determine ifhabitats occupied by these two lygaeids are consistently distinct. All 19 adults of S. insignis (7 males, 12 females) collected from granite outcrops in Minnesota were of the nearly black (dark) morpho In some areas, large populations of S. ins ignis consist of nearly equal numbers of the dark and of the bright tan or orange-red morph (Sweet 1964, Slater et aL 1993). Two brachypterous adults of the light-colored morph were collected from a dif­ ferent habitat in Minnesota at about the same time (21 June 1995); they were 216 THE GREAT LAKES ENTOMOLOGIST Vol. 28, No.3 & 4 swept by T. J. Henry from weeds in a gravel pit near Erhard, Otter Tail Co. All adults observed on granite outcrops were brachypterous. New England populations studied by Sweet (1964) were chiefly brachypterous with only about 20% macropterous. He remarked that it "would be interesting to see whether north midwestern populations in presumably widespread permanent habitats exhibit a lower frequency ofmacroptery." Although limited fieldwork in Minnesota did not yield any macropterous adults of S. insignis on granite outcrops, more extensive observations are needed to compare and contrast color and wing polymorphism of populations in this region with those in New England and other parts of its range. Further study is also needed to determine whether S. insignis feeds mainly on seeds of grasses, as it does in New England (Sweet 1964). It is pos­ sible that on rock outcrops the bugs are opportunists, feeding on seeds of grasses, Potentilla tridentata, and other plant seeds that fall or accumulate in cracks of the substrate.

ACKNOWLEDGMENTS I thank T. J. Henry (Systematic Entomology Laboratory, USDA, c/o Na­ tional Museum of Natural History, Smithsonian Institution, Washington, DC) for companionship in the field, for identifying adults ofL. syluestris and S. in­ signis, and the photograph used in Fig. 1; J. A. Slater (Department of Ecology & Evolutionary Biology, University of Connecticut, Storrs) for identifying nymphs of T. arenarius; W. L. Brown, Jr. (Department ofEntomology, Cornell University, Ithaca, NY) for identifying the ant F. subsericea; and E. R. Hoe­ beke (Department ofEntomology, Cornell University) for the photograph used in Fig. 2.

LITERATURE CITED Ashlock, P. D. &A. Slater. 1988. Family Lygaeidae Schilling, 1829 (=Infericornes Amyot and Serville, 1843; Myodochidae Kirkaldy, 1899; Geocoridae Kirkaldy, 1902. pp. 167-245 in: Henry, T. J. & R. C. Froeschner (eds.), Catalog ofthe Heteroptera, or true bugs, of Canada and the continental United States. E. J. Brill, Leiden. Harrington, B. J. 1980. A generic level revision and cladistic analysis ofthe Myodochini of the world (Hemiptera, Lygaeidae. Rhyparochrominae). Bull. Am. Mus. Nat. Hist. 167:45-116. Hendrickson, G. O. 1930. Studies on the insect fauna of Iowa prairies. Iowa State ColI. J. Sci. 4:49-179. Lindberg, H. 1958. Hemiptera Heteroptera from Newfoundland, collected by the Swedish-Finnish expedition of 1949 and 1951. Acta Zool. Fenn. 96:1-25. Scudder, G. G. E. 1993. Geographic distribution and biogeography of representative species ofxeric grassland-adapted Nearctic Lygaeidae in western North America (In­ secta: Heteroptera). pp. 75-113 in: Ball, G. E. & H. V. Danks (eds.), Systematics and entomology: Diversity, distribution, adaptation, and application. Mem. Entomol. Soc. Can. 165. Slater, J. A. 1964. A catalogue of the Lygaeidae of the world. 2 vols. University of Con­ necticut, Storrs. 1668 pp. Slater, J. A., M. H. Sweet & H. Brailovsky. 1993. Two new species of Slaterobius Har­ rington with comments on the ecology and distribution of the genus (Hemiptera: Ly­ gaeidae). Proc. Entomol. Soc. Wash. 95:590-602. Sweet, M. H. 1964. The biology and ecology of the Rhyparochrominae of New England (Heteroptera: Lygaeidae). Pts. I-II. Entomol. Am. 43:1-124, 44:1-201. 1995 THE GREAT LAKES ENTOMOLOGIST 217

A NOTE ON THE SYMPATRIC COLLECTION OF CHYMOMYZA (DIPTERA: DROSOPHIUDAEj IN VIRGINIA'S ALLEGHENY MOUNTAINS

Henretta Trent Band 1

ABSTRACT The attraction of two Chymomyza species, C. procnemoides and C. aldrichii, to the same damaged tree over 19 days in summer 1987 near Mt. Lake Hotel, Giles Co., Virginia is documented, confirming a previous report that Chymomyza species may be sympatric on the same fresh damaged tree/cut wood. A total of 17 males and 7 females were captured. An excess of males to females captured has been reported in Japan and Hungary.

Sturtevant (1916), Steyskal (1949), Wheeler (1952), Spieth (1957), Watabe (1985) and Bachli and Burla (1986) have all reported Chymomyza are at­ tracted to bleeding trees, tree trunks, cut wood, or freshly damaged trees. Wheeler (1952) also stated that C. aldrichii Sturtevant was collected at the same sites with C. coxata Wheeler on trees and peeled logs in Colorado and Wyoming. Watabe (1985), Band (1988, 1989, 1993) and Papp (1992) have sub­ sequently confirmed the sympatric capture of Chymomyza species on dam­ aged trees, logs or freshly cut wood. Papp (1992) included times of capture, numbers and sex, and species for Chymomyza collected on poplar (logs, trunks) in a Budapest forest over a 5-day interval in May and 2 days in June, 1990. For the collections in timberyards at Hokkaido (Watabe 1985) and the poplar forest near Budapest, Hungary (Papp 1992), it is not stated whether the different species occupied the same or different logs at the time of capture. Collections at the 1200 m elevation, Giles Co., Virginia in the vicinity of Mt. Lake Hotel have paralleled Wheeler's (1952) findings that two or more Chymomyza species are attracted to exactly the same damaged tree. Band (1988) collected on a wild cherry Prunus sp. in 1986 and a striped maple Acer pensylvanicum in 1987. Both trees are pictured in that paper. In 1986 the cap­ ture of a mating pair of C. aldrichii at 1640h on 22 July and a mating pair of C. procnemoides Wheeler at approximately the same time (1630h) on 25 July on the wild cherry where both species were sympatric over an extended period (Band 1988) demonstrated that both species used the same surface for mat­ ing. In 1987 species, sex and times of collection were noted for Chymomyza collected on the striped maple. Here I report the sympatric collection of C. procnemoides and C. aldrichii in 1987 between 18 July and 5 August, 1987 on the single damaged tree. In Virginia all flies were trapped in shell vials and transported to the lab­ oratory at the Biological Station within a half-hour of capture. In 1987 they were etherized, sexed, identified to species, and most specimens were affixed

lZoology Department, Michigan State University, East Lansing, MI 48824 and The University of Virginia, Mountain Lake Biological Station, Pembroke, VA 24136. 218 THE GREAT LAKES ENTOMOLOGIST Vol. 28, No.3 &4

to points on insect pins to establish a reference collection. Wheeler's (1952) key was used for species identification. Table 1 shows that males and females ofboth species were collected on the same damaged tree trunk for almost 3 weeks. This sympatric occurrence is reo inforced in Table 2 where the numbers of males and females of each species are grouped according to morning, afternoon or early evening captures. The low numbers also may reflect the small population sizes in nature. While not all attempted captures were successful, 1987 was the last year in which C. aldrichii outnumbered the numbers of C. procnemoides captured in that reo gion. Table 3 is constructed from the information supplied by Papp (1992) on the dates and location of new drosophilid species captured in Hungary in reo cent years. In 1986 males and females of C. amoena (Loew) and a pair of C. caudatula Oldenberg were captured sympatrically in an oak forest. Chy· momyza caudatula later was the most prevelant species among the sympatri. cally occurring Chymomyza in a poplar forest in 1990, as shown. Chymomyza caudatula is the only Holarctic species of the group (Wheeler 1981), C. amoena and the one specimen of C. procnemoides are introduced species of Nearctic origin; C. distincta (Egger) and C. fuscimana (Zetterstedt) have been

Table 1. Time of collections of Chymomyza procnemoides and C. aldrichii on damaged Acer pensylvanicum from mid·July into early August, 1987, Mt. Lake, VA. C. procnemoides C. aldrichii Month/date Time (hr) d'd' 22 d'd' 22 unknowns VII 18 1100 2 2 12 1600 3 1 3 1 VII 19 1100 1 1500 1 VII 22 1900 1 VII 27 1700 1 2 1 VII 28 1730 2 VIII 2 1730 1 VIII 3 ? 1 VI1I5 2000 1 Totals 7 2 10 5

Table 2. Comparisons ofcollections at different times of the day on Acer pensylvanicum. C. procnemoides C. aldrichii Period d'd' 2 d'd' 22 morning 2 1 2 afternoon 4 1 6 2 evening 1 1 2 1 Totals 7 2 9 5 1995 THE GREAT LAKES ENTOMOlOGIST 219

Table 3. Chymomyza collections in a Budapest, Hungary poplar forest in 1990, compiled from Papp (1992). Month May June day 20a 228 26b 28c 30c 3 11 time in hours 9-10 19-20 10 19-20 19·20 18-19 19-20

...... --~ C. amoena 19.10" 10" 10" C. caudatula 19,10" 40" 139,0" 20" 250" 90" 10" C. distincta 29,0" 20" 19 10" C. fuscimana 40' 60" 10" 49,0" 10" C. procnemoides 10" afresh cut poplar trunks hpoplar trunks "piles of poplar trunks

found in Europe and Japan (Watabe 1985). Steykal (1949) captured C. amoena in Michigan on poplar Populus deltiodes and black locust Robinia pseudoacacia in June. On the latter, they were among the Diptera feeding at frass. The excess numbers of males to females is typical. Hence, despite the small size ofthe collections, Virginia and Hungarian data agree with Watabe's (1985) findings in 4 Japanese timberyards where he collected a total of 693 males and 157 females representing 4 Chymomyza species.

ACKNOWLEDGMENTS I thank Gerhard Bachli for numerous reprints and L. Papp also for a reprint of his 1992 paper.

LITERATURE CITED Bachli, G. and H. Burla. 1986. Diptera Drosophilidae. Insecta Helvetica Fauna 7. Schweiz. Entomol. Ges., 116 pp. Band, H. T. 1988. Behavior and taxonomy of a chymomyzid fly (Chymomyza amoena). Intern. J. Compo Psycho!. 2:3--35. __. 1989. Behavior of the Chymomyza aldrichii species group (Diptera: Drosophili­ dae) in VlIginia's Allegheny Mountains. Va. J. Sci. 40:230-237. 1993. More on the mate recognition controversy. Michigan Academician XXV: 353

Papp, L. 1992. Nine drosophilid species new to Hungary (Diptera: Drosophilidae). Folia Entomol. Hungarica Rov. KDzl. 53:135-138. Spieth, H. T. 1957. Drosophila ofthe Itasca Park Minnesota region. N. Y. Entomol. Soc. 65:89-96. Steyskal, G. C. 1949. The Dipterous fauna of tree trunks. Papers Mich. Acad. Sci., Arts and Let. 35:121-134. Sturtevant, A H. 1916. Notes on North American Drosophilidae with descriptions of twenty-three new species. Ann. Entomol. Soc. Amer. 9:323-343. 220 THE GREAT LAKES ENTOMOLOGIST Vol. 28, No.3 &4

Watabe, H. 1985. A preliminary note on the drosophilid files collected at timberyards in northern Japan. Dros. Inf. Servo 61:183-184. Wheeler, M. R. 1952. The Drosophilidae of the Nearctic region exclusive of the genus Drosophila. Univ. of Texas Publ. No. 5204:162-218. __. 1981. The Drosophilidae: a taxonomic overview. pp. 1-99. in: M. Ashburner, H. L. Carson and J. N. Thompson, Jr. eds. The Genetics and Biology of Drosophila. Vol. 3a. Academic Press, London. 1995 THE GREAT LAKES ENTOMOLOGIST 221

ECONOMICS OF CEll PARTITIONS AI~D CLOSURES PRODUCED BY PA55ALOECU5 CU5PIDATU5 !HYMENOPTERA: SPHECIDAE)

John M. Fricke 1

ABSTRACT Sphecid wasps, Passaloecus cuspidatus were observed gathering fresh pine resin (Pinus strobus) which they used for creating partitions and closures in their nests. Based upon measurements taken of the cells, and an estimate of the load carried by the wasps, the number of collecting trips made by the wasps was correlated with the quantity of resin contained within the nest partitions and closures.

Passaloecus cupsidatus Smith is a small pemphredonine sphecid wasp that utilizes pre-existing tunnels and trap nests for its nests, which have cells within arranged linearly, each separated by a thin partition of pine resin. A final closure of the nest is also constructed of resin. Female wasps, therefore, must collect the resin from recently-oozing wounds on coniferous trees. This study examines the relationship between the amount of resin contained within the nest and the number oftrips required to collect the resin. Resin gathering activity of P. cuspidatus was observed at resin flows on Pinus strobus on 17 June 1987 in Ann Arbor, Michigan (see Fricke, 1992). Large fence staples (1.5 in) had been used to secure trap nest bundle carriers (described previously in Fricke, 1991) to trunks oftrees selected as trap-nest­ ing stations. Resin flows were produced in response to fence-staple wounds. Passaloecus cuspidatus selected resin flows with dimensions 9 by 4 mm. The wasp's mandibles were used in a scissors-like fashion to excise a drop of resin with a diameter the width ofthe wasp's head. When separation ofa resin drop was nearly complete, the wasp backed directly away from the resin flow. A thin strand ofresin, connecting the resin drop to the flow, was drawn into the excised drop by lateral and circular motions of the wasp's head. Any remain­ ing remnant of the resin strand was cut off by a continued backward move­ ment combined with an abrupt turning to the left or right. Twenty-two resin gathering trips were observed between 0947 and 1142 hr. Resin drops were carried on the ventral surface of the mandibles. Three separate resin flows were used during these resin gathering activi­ ties. Resin flow (I) was visited repeatedly and the wasp returned directly to the resin flow, landing within a few em ofthe flow and approaching it directly. After a number of resin gathering trips, the remaining portion of the resin flow was too small, or of improper consistency, and was abandoned as a resin source. The pine trunk was searched for another appropriate resin flow and resin gathering resumed. Resin flow (II) was then used repeatedly as a resin source.

lNaturaI Science and Mathematics Division, Concordia College, Ann Arbor, MI 48108. 222 THE GREAT LAKES ENTOMOLOGIST Vol. 28, No.3 &4

140 130

120 D Partition 110 • Closure 100 ('; 90 c Q) 80 :::s C" 70 ~ lJ.. 60 50 40 30 20 10 0 .1 .2 .3.4.5 2 3 4 5 10 Thickness (mm) Figure 1. Thickness ofresin partitions and closures in Passaloecus cuspidatus trap nests.

After three or four trips to resin flow (II) the wasp made a trip to resin flow (I), explored the resin mass, and returned to resin gathering at flow (II). Resin gathering was again observed 18 June (eight trips, 1631-1713 hr) and 24 July (11 trips between 0900 and 1000 hr, with no elapsed time recorded). Ringing of trap nest bore openings was also observed and the following ac­ tion pattern was noted. Passaloecus cuspidatus landed at or within a few cen­ timeters of the trap nest opening, entered head first, exited and re-entered gaster first. The wasp then appeared at the nest opening and with her mandibles spread a thin layer of resin on the face of trap nest at the margin of the opening. Krombein (1967) noted that resin partitions were usually 0.25 and occa­ sionally 4.0 mm thick, while closures, ranging from 0.25 to 4.0 mm, were usu­ ally 1.0 mm thick. Vincent (1978) noted that 4 drops of resin were used for nest closures. Data from my studies of 1987 were examined to determine the variability of partition and closure dimensions, their volumes, and the ener­ getics of resin gathering. It was noted during resin gathering described above that an excised drop of resin had a diameter approximately equal to the width ofthe wasp's head. Given a diameter of 1.5 mm, a resin drop carried by P. cus­ pidatus has a volume of L 77 mm3. The thickness of resin partitions of 295 pro­ visioned cells ranged from 0.1 to 5.0 mm in trap nests with bore diameters 2.4 1995 THE GREAT LAKES ENTOMOLOGIST 223

to 6.4 mm. Median and modal partition thicknesses were 0.5 mm; the mean partition thickness was 0.66 mm. Seventy-one closures had thicknesses rang­ ing from 0.25 to 4.0 mm with a mean of 1.69 mm. Based upon the bore diameters and thicknesses indicated above, the vol­ ume of resin required for partitions and closures ranged from 1.13 mm3 to 62.83 mms. The volumes ofresin required for partitions and closures in bores most frequently used by P. cuspidatus are respectively: 3.2 mm - 5.31 and 13.59; 4.0 mm - 8.29 and 21.24; and 4.8 mm - 11.94 and 30.58 mm3• Given a volume of 1.77 mm3 per resin drop, the numbers of resin gathering trips for partitions and closures for these respective bores are: 3.2 mm - 3 and 7.68 (8); 4.0 mm - 4.68 (5) and 12.0; 4.8 mm - 6.75 (7) and 17.28 (18). These data are rather conservative since they have not taken into consid­ eration the foundations for closures or partitions. Foundations, consisting of a resin ring on the wall of the trap nest bore, have base widths greater than that of their respective partition or closure. Resin partitions, closures, and their respective foundations, represent a significant energy investment. Resin volumes and bore diameters across which resin must be drawn possibly con­ tribute to the upper limits of the bore diameters used by P. cuspidatus. The distribution of partition and closure thicknesses from P. cuspidatus trap nests are given in Figure 1.

LITERATURE CITED Fricke, J. M. 1991. A trap-nest design for small trap-nesting Hymenoptera. Great Lakes Entomol. 24:121-122. 1992. Influence of tree species on frequency of trap-nest use by Passaloecus (Hymenoptera: Sphecidae). Great Lakes Eiltomol. 25:51-53. Krombein, K V. 1967. Trap-nesting wasps and bees: life histories, nests, and associates. Smithsonian Press, Washington DC. vi + 570 pp. Vincent, D. L. 1978. A Revision of the genus Passaloecus (Hymenoptera: Sphecidae) in America north of Mexico. Wasmann J. BioI. 36:127-198. 1995 THE GREAT LAKES ENTOMOLOGIST 225

DISTRIBUTION OF THE MILLIPED NARCEUS AMERICANUS ANNULARIS (SPIROBOLlDA: SPIROBOLIDAEI IN WISCONSIN

Dreux 1. Wotermolen1

The spirobolid milliped Narceus american us annularis Rafmesque 1820 is common and widespread throughout the eastern United States and Canada (Keeton 1960, Shelley 1988, Hoffman 1990). It has previously been recorded \ from Wisconsin. Cahn (1915) reported it from the Wingra Springs region in \ central Dane County, and Keeton (1960) reported it from Clark and Sauk Counties. General records include the "upper Mississippi Valley" by Hoffman (1951) and ''Wisconsin'' by Kevan (1985). Recent collections and examination of specimens at the Milwaukee Public Museum (MPM) and Illinois Natural History Survey (INHS) turned up additional locality records. On 6 August 1994, D.F. Guebken collected an adult male N. americanus annularis crawling on the floor of a pole building in Crawford County. I col­ lected two juvenile specimens on 28 September 1992 in an underground cor­ ridor at the University of Wisconsin-Green Bay in Brown County. Levi and Levi (1987) suggested N. americanus annularis is usually collected in forest logs. On 19 July 1992, I collected a male and a female specimen from within a rotting log (probably Thuja occidentalis) in Peninsula State Park, Door County. All five specimens have been deposited in the MPM invertebrate col­ lection. Additional records of N. americanus annularis obtained from museum col­ lections include specimens from Cedarburg Bog in Ozaukee County (May 1992; MPM), from Viroqua, Vernon County (August 1926; MPM), and from an unspecified locality in Milwaukee County (prior to 1897; MPM). A male spec­ imen collected at Wyalusing State Park, Grant County is maintained in the 'Iexas Memorial Museum collection (R.M. Shelley, pers. comm.). In addition, two juvenille Narceus specimens were collected at Devils Lake State Park, Sauk County (June 1950; INHS). . Although no comprehensive survey of Wisconsin millipeds has been com­ pleted, these records indicate that the distribution of N. americanus annularis is much more extensive than previously known (Figure 1). Indeed, I expect that this species is even more widely distributed in Wisconsin and probably occurs statewide.

ACKNOWLEDGMENTS J.P. Jass, Milwaukee Public Museum, and K.R. Methven, Illinois Natural History Survey, loaned specimens. R. Hammond assisted with the preparation of Figure 1. The specimen collected in Peninsula State Park was collected under Scientific Collectors Permit No. SCP-LM-036-C-9294 from the Wiscon­ sin Department of Natural Resources.

IBureau of Environmental Analysis and Review, Wisconsin Dept. of Natural Re­ sources, P.O. Box 7921, Madison, WI 53707-7921. 226 THE GREAT LAKES ENTOMOLOGIST Vol. 28, No.3 & 4

Figure 1. Distribution of Narceus american us annularis in Wisconsin. Circles are literature records plotted at the center of each county. Squares are new records reported herein.

LITERATURE CITED Cahn,A.R. 1915. An ecological survey ofthe Wingra Springs Region, near Madison,Wis­ consin, with special reference to the ornithology. Bull. Wisconsin Nat. Hist. Soc. 13(3):123-177. Hoffman, R.L. 1951. The name of the common eastern Spirobolid milliped. Florida En­ tornol. 34(1):15-16. Hoffman, R.L. 1990. Diplopoda. pp. 835-860 In Dindal, D.L. (ed.). Soil biology guide. John Wiley & Sons, New York. Keeton, W.T. 1960. A taxonomic study of the milliped family Spirobolidae (Diplopoda: Spirobolida). Mem. American Entomol. Soc. (17):1-146 Kevan, D.K. McE. 1983. A preliminary survey of known and potentially Canadian mil­ lipedes CDiplopoda). Canadian J. Zool. 61:2956-2975. Levi, H.W. and L.R. Levi. 1987. Spiders and their kin. Gulden Press, New York. 160 p. Shelley, R.M. 1988. The millipeds of eastern Canada (Arthropoda: Diplopoda). Cana­ dian J. Zool. 66:1638-1663. 1995 THE GREAT LAKES ENTOMOLOGIST 227

ADULT FEMALE MYDAS CLAVATUS (DIPTERA: MYDIDAE) FEEDING ON flOWERS IN WISCONSIN

Andrew H. Williams 1

ABSTRACT An adult female of Mydas clauatus was observed feeding on flowers. This confirms that females ofthis species feed as adults and that they feed on flow­ ers. This species is reported for the first time from Wisconsin and Minnesota.

-_...._-----­ On 29 July 1994, an adult female Mydas clauatus (Drury) was collected at Thomas Wet Prairie in Grant Co., Wisconsin. She was observed on flowers of Spiraea alba and Pycnanthemum uirginianum. She seemed to be nectaring, conceivably she was feeding on pollen, but her movement from flower to flower across the inflorescences showed that her interest was in the flowers themselves. She showed no interest in other insects. Pollinia ofAsclepias sp. were stuck on her mouthparts and on a protarsal spine, indicating that she had also visited milkweed flowers. Collected at noon, on a hot, sunny day, the author's specimen #1608 is now in the Insect Research Collection (IRC) ofthe University of Wisconsin - Madison. This confirms that adult females of M. clavatus feed on flowers.

DISCUSSION We know little about the life histories of most Mydidae. Whether adult mydids are predators of other insects or feed on flowers has long been dis­ cussed (Malloch 1915, Papavero and Wilcox 1968, Cole 1969, Papavero and Knutson 1975, Wilcox 1981, Richter and Zaitzev 1988), and it has been un­ clear whether or not adult females feed at all (Papavero and Knutson 1975, Richter and Zaitzev 1988). Zaitlin (1978) and Zaitlin and Larsen (1984) found the lack of piercing mouthparts in M. clavatus indicative of nectar feeding rather than predation. They found that males and females have identical head morphologies and can be seen feeding together on flowers, especially ofAsclepias syriaca, but also of A. verticillata, Monama punctata, Teucrium canadense, verbena hastata and Saponana officinalis. Hart and Malloch (Malloch 1915) collected adults on flowers ofAsclepias sp. Davis (1921) reported being bitten by M. clavatus, "... the bite being very painful for the moment." This would seem to indicate the presence of piercing mouthparts, but it's likely that Davis was pinched rather than bitten, as Malloch (1915) reported, "... that the insect can pinch rather severely with the hind femora and tibiae."

IDepartment ofEntomology, University ofWisconsin, Madison, WI, 53706. 228 THE GREAT LAKES ENTOMOLOGIST Vol. 28, No.3 & 4

Wilcox, Papavero and Pimentel (1989) mapped the distribution of M. clavatus using specimen data from several prominent collections. Their map shows the species to be widely distributed across the eastern United States and southernmost Ontario but absent from Wisconsin and Minnesota. Mydas clavatus specimens in the IRC, Milwaukee Public Museum, University of Michigan and Illinois Natural History Survey include single flies collected in Grant, Monroe, Pepin, Shawano and Waupaca counties in Wisconsin, and in Winona County, Minnesota, and two collected in Wood County, Wisconsin. The Field Museum, University of Minnesota and University of Kansas have none of these flies from this region in their collections. This extends the known range of this fly to the northwest, to include the southern half of Wisconsin and southeastern Minnesota. Mydas clavatus has rarely been collected in this region, a fact which, given the great size of this fly and its striking coloration, makes it likely that this species is rare at the northwest periphery of its range.

ACKNOWLEDGMENTS This paper results from the Prairie Insect and Spider Inventory of The Prairie Enthusiasts Southwest Chapter, basic biotic research being con­ ducted at Thomas Wet Prairie with support from The Prairie Enthusiasts ­ Southwest Chapter, the Citizens Natural Resources Association of Wisconsin, the Natural History Museums Council of UW - Madison and several private donors, support for which I am most grateful. I am also grateful to D. Young and S. Krauth of the Entomology Department of UW - Madison for their in­ terest in this research, and to G. Noonan, M. F. O'Brien, P. Parrillo, C. Reed, D. Webb and J. Welch-Jolly for their help with collection data at their respec­ tive institutions.

LITERATURE CITED Cole, F. R. 1969. The Flies of Western North America. University of California Press. Berkeley & Los Angeles. 693 pp. Davis, W. T. 1921. Proceedings of the Society. Meeting of February 10, 1921. BulL Brooklyn Entomol. Soc. 16:138-139. Malloch, J. R. 1915. Some Additional Records ofChironomidae for lllinois and Notes on Other Illinois Diptera. Bull. lllinois St. Lab. Nat. Hist. 11:305-363. Papavero, N. and L. V. Knutson. 1975. Family Mydidae. pp. 97-98. In: Delfinado, M. D. and D. E. Hardy (eds.). A Catalog of the Diptera of the Oriental Region. Vol. 2. Uni­ versity of Hawaii Press. Honolulu. 459 pp. Papavero, N. and J. Wilcox. 1968. Family Mydidae (Mydaidae, Mydasidae). 34:1-20. In: A Catalogue of the Diptera of the Americas South of the United States. Departmento de Zoologia, Secretaria da Agricultura, Sao Paulo. Richter, V. A. and V. F. Zaitzev. 1988. Family Mydidae. p. 181. In: Soos, A. and L. Papp (eds.). Catalogue of Palaearctic Diptera. Vol. 5. Elsevier Science Publishers. Amster­ dam. 446pp. Wilcox, J. 1981. Family Mydidae. Chapter 40. pp. 533-540. In: McAlpine, J. F., B. V. Pe­ terson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth and D. M. Wood. Manual of Nearc­ tic Diptera. Vol. 1. Research Branch. Agriculture Canada. Monograph No. 27. Ottawa. 674 pp. 1995 THE GREAT LAKES ENTOMOLOGIST 229

Wilcox, J., N. Papavero and T. Pimentel. 1989. Studies of Mydidae (Diptera). !Vb. Mydas and Allies in the Americas (Mydinae, Mydini). Museu Paraense Emilio Goeldi. Belem. 139 pp. Zaitlin, L. H. 1978. Morphology of the Head and Mouth Parts ofMydas clavatus Drury (Diptera: Mydaidae). PhD Thesis. University of l1linois at Urbana-Champaign. 216 pp. Zaitlin, L. M. and J. R. Larsen. 1984. Morphology of the Head of Mydas clavatus Drury (Diptera: Mydaidae). Int. J. of Insect Morphol. & Embryol. 13:105-136. 1995 THE GREAT LAKES ENTOMOLOGIST 231

A MICHIGAN RECORD FOR GYTUS MARGINICOLLIS (COLEOPTERA: CERAMBYCIDAE: CLYTINll

James E. Zablotnyl

Clytus marginicollis Castlenau and Gory is a small, rarely collected clytine cerambycid endemic to coniferous forests of Eastern North America (Knull 1943, Linsley 1964). Gosling (1973) did not mention any records for it in his survey of Michigan cerambycids. Knull (1943) reports that this species probably occurs in Ohio but he had seen no specimens to confirm its presence. With a standard sweep net, I collected a single specimen on 19 May 1985 and two specimens on 15 May 1986 at the Michigan State University tree dump in Lansing Township, Ingham County Michigan. They were sunning themselves on freshly cut pine and hardwood branches prior to capture. One of the 1986 specimens has been deposited in the MSU Insect Collection. Clytus marginicollis is similar to the ubiquitous C. ruricola (Olivier), but is more robust in appearance. The pale yellow antemedian pubescent band of C. marginicollis is oblique and arcuate while C. ruricola possesses aU-shaped pubescent band (Figure 1). Unlike the anthophilous C. ruricola, C. margini­ collis has never been recorded feeding on pollen or nectar. Also, while C. ruri­ cola oviposits on rotten hardwood logs, C. marginicollis relies on windblown or freshly cut pine branches for oviposition sites (Linsley, 1964) and additional records should be found near cultivated and wild Pinus species. This species' use of recently cut or windthrown small pine branches suggests that its lar­ vae have minimal or no economic impact on cut pine timber or cultivated nursery stock. Along with Phymatodes amoenus (Say), Megacyllene caryae (Gahan), and Cyrtophorus uerrucosus (Olivier), C. marginicollis is one of the first ceramby­ cids to appear in the spring. Its early appearance may account in part for its uncommonly collected status. Gosling (1973) and Gosling et al. (1976) compiled a comprehensive list of Michigan's cerambycid fauna and expected additional records from the south­ western Lower Peninsula and Western Upper Peninsula. I am pleased to re­ port that Michigan's 226th cerambycid species was not discovered in those boundary counties but in a highly urbanized area of Ingham County. Even though cerambycid beetles have been extensively studied, further work is still needed to discover additional species and to enhance distribution records for those species found within the state.

ACKNOWLEDGMENTS I thank Beth Bishop, Catherine Bristow, and Fred Stehr for their written comments and Jan Eschbach for assistance with computer graphics. I also thank Bob Androw for identifying the specimens.

leenter for Insect Diversity Study, Department ofEntomology, Michigan State Uni­ versity, East Lansing, MI 48824-1115. 232 THE GREAT LAKES ENTOMOLOGIST Vol. 28, No.3 & 4

Figure 1. Habitus of Clytus marginicollis Castelnau and Gory. 1995 THE GREAT LAKES ENTOMOLOGIST 233

LITERATURE CITED Gosling, D. C. L. 1973. An annotated list of the Cerambycidae of Michigan (Coleoptera) Part I, Introduction and the subfamilies Parandrinae, Prioninae, Spondylinae, Asem­ inae, and Cerambycinae. The Great Lakes Entomol. 6(3):65-84. Gosling, D. C. L. and N. M. Gosling. 1976. An annotated list of the Cerambycidae of Michigan (Coleoptera) Part II, the subfamilies Lepturinae and Lamiinae. The Great Lakes Entomol. lO(1):1-37. Knull, J. N. 1946. The long-horned beetles of Ohio. Ohio BioI. Surv., Bull. 39:133-354. Linsley, E. G. 1964. The Cerambycidae of North America, Part V. Taxonomy and classi­ fication of the subfamily Cerambycinae, tribes Callichromini through Ancylocerini. Univ. Calif. PubL in Entomol. 22:164-171. 1995 THE GREAT LAKES ENTOMOLOGIST 235

UROPHORA QUADRIFASCIATA (DIPTERA: TEPHRITIDAEL AN INTRO­ DUCED SEEDHEAD FLY NEW TO MIDWESTERN NORTH AMERICA

A. G. Wheeler, Jr. 1

ABSTRACT The Old World tephritid Urophora quadrifasciata, a gall-inducing seed­ head fly, was released in western and eastern North America for the biological control of knapweeds, Centaurea spp. (Asteraceae). Its establishment in the West (BC, CA, ID, MT, OR, WA) and in the East (CT, MA, MD, NH, NJ, NY, PA, RI, VA, VT, WV) has been previously reported. Collections from eastern Minnesota and western Michigan in 1995 are the first for the Northcentral re­ gion of North America.

The tephritid genus Umphora (about 100 spp.) includes eight species in­ digenous to the Nearctic Region and three Eurasian species that have been in­ troduced to North America for the biological control of adventive knapweeds, Centaurea spp. (Asteraceae) (White and Korneyev 1989, Turner et al. 1994). Umphora affinis (Frauenfeld) and U. quadrifasciata (Meigen), the best known of the introduced inducing, seedhead flies, were released in the early 1970s in British Co bia to help suppress infestations of diffuse knapweed (C. diffusa) and spotted knapweed (C. biebersteinii = C. maculosa ofAmerican authors). These weeds limit productivity in rangelands and displace native herbaceous vegetation. Larvae of both tephritids are able to coexist in heads of their host plants; despite negative interaction between these flies, their ef­ fects on knapweed seed production are complementary (Berube 1980, Harris and Myers 1984, see also Wheeler and Stoops, 1996). Establishment of U. quadrifasciata in the West is well documented. It is known to occur not only in British Columbia, where it was released, but it has also dispersed to, and become established in, California, Idaho, Montana, Ore­ gon, and Washington (Maddox 1982, Julien 1992, Foote et al. 1993). Although U. quadrifasciata was released in eastern North America -Quebec in 1979 and Maryland and New York in 1983-postrelease monitoring has been scant in comparison to that in the West. Establishment was not verified until the early 1990s when it was discovered in Connecticut, Massachusetts, New Hampshire, New Jersey, New York, Pennsylvania, Rhode Island, and Vermont (Hoebeke 1993). This tephritid has since been found in Maryland, Virginia, and West Virginia (Wheeler and Stoops, 1996). Recent fieldwork in Minnesota has resulted in the first records of U. quadrifasciata from the Midwest. Adults, including several mating pairs, were common on spotted knapweed growing along the railroad at Hinckley (Pine Co.) on 28 June 1995. Eighteen specimens (8 females, 10 males) were

IBureau of Plant Industry, Pennsylvania Department ofAgriculture, Harrisburg, PAl7110. 236 THE GREAT LAKES ENTOMOLOGIST Vol. 28, No.3 & 4 collected. In a much smaller population of the host (5-6 plants), two females were collected on the same date at Eden Prairie (Hennepin Co.) near Min­ neapolis. In addition, E.R. Hoebeke made available his record of U. quadri­ fasciata from spotted knapweed in Michigan: Kent Co., Grand Rapids, 11 Aug. 1995. Voucher specimens have been deposited in the insect collection of Cor­ nell University, Ithaca, NY. Urophora quadrifasciata disperses more readily than U. affinis, but the latter species tends to become dominant where they co-occur (Berube 1980, Harris 1980, Harris and Myers 1984, Story et a1. 1992). The record of U. quadrifasciata from Minnesota and Michigan are the first for an area between eastern Montana in the West and western New York and western Pennsylva­ nia in the East. Further collecting in this region is encouraged to document the continued spread of U. quadrifasciata and to detect the possible arrival of U. affinis.

ACK..~OWLEDGMENTS I thank E. R. Hoebeke (Department of Entomology, Cornell University, Ithaca, NY) for reviewing a draft of the manuscript, and allowing me to use his Michigan record of U. quadrifasciata.

LITERATURE CITED Berube, D. E. 1980. Interspecific competition between Urophora affinis and U. quadri­ fasciata (Diptera: Tephritidae) for ovipositional sites on diffuse knapweed (Centaurea diffusa: Compositae). Z. Angew. Entomol. 90:299-306. Foote, R. H., F. L. Blanc, and A. L. Norrbom. 1993. Handbook ofthe Fruit Flies (Diptera: Tephritidae) of America North of Mexico. Cornell University Press, Ithaca, N.Y. 571 pp. Harris, P. 1980. Establishment of Urophora affinis Frfld. and U. quadrifasciata (Meig.) (Diptera: Thphritidae) in Canada for the biological control of diffuse and spotted knapweed. Z. Angew. Entomol. 89:504-514. Harris, P. and J. H. Myers. 1984. Centaurea diffusa Lam. and C. maculosa Lam. s. lat., diffuse and spotted knapweed (Compositae), pp. 127-137. In: J.S. Kelleher and M. A. Hulme (eds.), Biological Control Programmes against Insects and Weeds in Canada 1969-1980. Commonwealth Agricultural Bureaux, Slough, England. Hoebeke, E. R. 1993. Establishment of Urophora quadrifasciata (Diptera: Tephritidae) and Chrysolina quadrigemina (Coleoptera: Chrysomelidae) in portions of eastern United States. Entomol. News 104:143-152. Julien, M. H. (ed). 1992. Biological Control of Weeds: A World Catalogue ofAgents and their Target Weeds, 3rd ed. CAB International, Wallingford, UK. 186 pp. Maddox, D. M. 1982. Biological control of diffuse knapweed (Centaurea diffusa) and spotted knapweed (C. maculosa). Weed Sci. 30:76-82. Story, J. M., K. W. Boggs, and W. R. Good. 1992. Voltinism and phenological synchrony of Urophora affinis and U. quadrifasciata (Diptera: Tephritidae), two seed head flies introduced against spotted knapweed in Montana. Environ. Entomol. 21:1052-1059. 'furner, C. E., R. Sobhian, D. B. Joley, E. M. Coombs, and G. L. Piper. 1994. Establish­ ment of Urophora sirunaseva (Hering) (Diptera: Tephritidae) for biological control of yellow starthistle in the western United States. Pan-Pac. Entomol. 70:206-211. Wheeler, A. G., Jr. and C. A. Stoops. 1996. Establishment of Urophora affinis on spot­ ted knapweed in Pennsylvania, with new eastern U. S. records of U. quadrifasciata (Diptera: Tephritidae). Proe. Entomol. Soc. Wash. 98:93-99. White, 1. M. and V A. Korneyev. 1989. A revision of the western Palearctic species of Urophora Robineau-Desvoidy (Diptera: Tephritidae). Syst. Entomol. 14:327-374. 1995 THE GREAT LAKES ENTOMOLOGIST 237

FIRST COUNTY RECORDS FOR ACARAPIS WOODI (ACARI: TARSONEMIDAEl IN MICHIGAN

Murray Hanna 1 and Sharon Pratt Anzaldua2

ABSTRACT Acarapis woodi is an internal parasite of Apis mellifera. Surveys con­ ducted by the Michigan Department of Agriculture in years 1986-1990 pro­ duced first county records for A. woodi in 63 of the 83 counties of Michigan.

Acarapis woodi (Rennie),the honey bee tracheal mite, is an exotic, inter­ nal, obligate parasite of the honey bee, Apis mellifera Linn. (Calderone and Shimanuki, 1993). The distribution ofA. woodi, therefore, is that ofits host. The host, A. mellifera, is an exotic, domesticated, social insect of particular value to agriculture in Michigan. Cook (1876) observed that cross-fertiliza­ tion of flowers, which can only be accomplished early in the season by the honey bee, is often necessary to a full yield offruit and vegetables. Cowan (1903) published a treatise on the culture of the honey bee for British beekeepers which stated that dysentery and foulbrood were the two most important diseases of the honey bee. Rennie et al.(1921) investigated the cause of Isle of Wight disease of the honey bee which derived its popular name from the island from which it was first recognized in 1904. The disease assumed epidemic proportions in honey bee colonies on Isle of Wight in 1905, and was reported from mainland England in 1909. Diagnosis of Isle of Wight disease from symptoms had been an unsatisfactory procedure. The most usual features of Isle ofWight disease recognizable by the beekeeper were in­ ability ofhoney bees to fly and continuous mortality ofadult honey bees. Dur­ ing the course ofinvestigation it was discovered that a mite in all stages ofde­ velopment occurred in prothoracic tracheae of honey bees exhibiting symptoms of Isle of Wight disease. Cumulative evidence indicated an invari­ able and clear association ofthe mite with diseased honey bees, and that there was a definite pathology in relation to infection in the individual honey bee. Rennie (1921) in a companion publication described tracheal mite, A. woodi. Bailey (1958) d:scussed the epidemiology of the infection of the honey bee by A. woodi and concluded that mortality of infected honey bees is only slightly greater than that of non-infected honey bees. He produced evidence to sug­ gest that Isle of Wight disease was due to factors other than, or supplemen­ tary to, infection with A. woodi. Jaycox (1958) cited the need for a plan to protect the apiary industry in the event A. woodi were to be discovered in the United States. He proposed a method of survey which employed dissection of adult honey bees to determine presence or absence ofA. woodi.

lMichigan Department of Agriculture, Lansing, MI 48909. 2Michigan Department of Public Health, Lansing, MI 48909. 238 THE GREAT lAKES ENTOMOLOGIST Vol. 28, No.3 & 4

The fIrst occurrence ofAcarapis woodi in the United States was reported from specimens ofA. woodi identifIed from adult honey bees collected in Texas (DelfInado-Baker [1984]). DelfInado-Baker stated that because of frequent movement of honey bee colonies throughout much of the United States, it could be expected that A. woodi would spread inexorably. Calderone and Shi­ manuki (1993) state thatA. woodi has caused a signifIcant reduction in both number and quality of honey bee colonies in the United States. The M" Apiary Law (Act No. 412, Public Acts of 1976, as amended 1985) enabl authorized representatives of the Michigan Department of Agriculture to conduct surveys on the premises of any property, private or public, to ascertain the existence of serious honey bee disease. In the years 1985-1989 A. meUifera specimens were collected at Michigan apiaries by Michigan Department of Agriculture apiary inspectors for dissection at the Michigan Department of Agriculture, Entomology Laboratory, to determine presence or absence of A. woodi (Figure 1). No survey was undertaken in ei­ ther Keweenaw or Schoolcraft counties. In 1990 arrangement was made for Michigan beekeepers voluntarily to collect A. mellifera specimens for dissec­ tion to determine presence or absence of A. woodi by Michigan State Univer­ sity, Department of Entomology, under contract to the Michigan Department of Agriculture. Results obtained by dissection of 16,700 honey bees collected in 54 coun­ ties in 1985 indicate, but provide no assurance, that A. woodi did not then occur in Michigan. The fIrst record of the occurrence of A. woodi in Michigan was obtained by dissection ofA. meUifera specimens collected on 19 May 1986 in Van Buren County by apiary inspector Jane Winkler. Results obtained by dissection of A. mellifera specimens collected in 81 of the 83 counties of Michi­ gan in years 1985-1990 indicate, but provide no assurance, that within a span of 5 years A. woodi had become distributed to 63 counties of Michigan (Table 1, Fig. 2). With the A. woodi population now apparently so widespread throughout the state, future surveys for the presence of A. woodi should focus on the role that the mites have played on the apparent decline in healthy feral honey bee populations, as well as the domesticated colonies. The lack of such numerous and effective pollinators would bring about signifIcant economic and ecologi­ cal repercussions in the agriculture industry.

Table 1. Chronology of first county records for Acarapis woodi in Michigan. YEAR COUNTIES 1986 Chippewa, Hillsdale, Jackson, Lenawee, Menominee, Oakland, VanBuren 1987 Alcona, Berrien, Branch, Genesee, Gratiot, Isabella, Kalkaska, Liv­ ingston, Mecosta, Monroe, Montcalm, Newaygo, Ogemaw, Ottawa, Tuscola 1988 Cass, Huron, Lapeer, Muskegon, Saginaw, Shiawassee, aw 1989 Alpena, Antrim, Barry, Bay, Benzie, Calhoun, Charlevoix, Clare, Clinton, Eaton, Ingham, losco, Iron, Kalamazoo, Kent, Leelanau, Macomb, Mason, Midland, Oceana, Osceola, St. Joseph, Sanilac, Wexford 1990 Cheboygan, Emmet, Gladwin, Ionia, Lake, Mackinac, Manistee, Ot­ sego, St. Clair 1995 THE GREAT LAKES ENTOMOLOGIST 239

Figure 1. Photomicrograph ofprothoracic trachea dissected from A. mellifera infected with A. woodi. Positive image produced by Michigan State Univer­ sity, Instructional Media Center. 240 THE GREAT LAKES ENTOMOLOGIST Vol. 28, No.3 & 4

Figure 2. Map indicating distribution of first county records for A. woodi in Michigan.

ACKNOWLEDGMENTS We thank reviewers and the editor for recommendations incorpoY'l.Lted into the manuscript; Howard Russell, Michigan State University, DepaI tment uf Entomology, for his involvement in 1990 laboratory operations; Rajagopal Sitaraman, Michigan Department of Agriculture. for access to survey records; and Robert W. Husband, Adrian College, Biology Departmer for encourage­ ment.

LITERATURE CITED Bailey, L. 1958. The epidemiology of the infestation of the honeybee, Apis mellifera L., by the mite Acarapis woodi Rennk and the mortality of infested bees.Parasitology 48:493-506. Calderone, N. W. and H. Shimanuki. 1993. Distribution of l;le tracheal mite, Arvapis woodi, among mesothoracic trunks ofthe honey bee, Apis meltifera. Exp. AppL ,:\earol. 17:663-672. Cook, A. J. 1876. The apiary. Fourteenth Ann. Rept. Mich.State Board Agric. 1375. pp. 315-357. 1995 THE GREAT LAKES ENTOMOLOGIST 241

Cowan, T. W. 1903. British Bee-keeper's Guide Book. Houlston and Sons, London. 180 pp. Delfinado-Baker, M. 1984. Acarapis woodi in the United States. Am. Bee J. 124. 805-806. Jaycox, E. R. 1958. Acarine disease of honey bees. Calif.Dept. Agric. Bull. 47:215-22l. Rennie, J. 1921. Isle of Wight disease in hive bees - Acarine disease: The organism as­ sociated with the disease - Tarsonemus woodi, n. sp. Trans. Roy. Soc. Edin. 52, pt. 4:768-779. Rennie, J., P. B. White, and E. J. Harvey. 1921. Isle of Wight disease in hive bees - The etiology of the disease. Trans.Roy. Soc. Edin. 52, pt. 1: 737-754. 1995 THE GREAT LAKES ENTOMOLOGIST 243

THREE NEW FOOD PLANTS AND FIRST WISCONSIN RECORD OF PUBLILIA RETICULATA (HEMIPTERA: MEMBRACIDAE)

Andrew H. Williams]

ABSTRACT Publilia reticulata was found feeding on the composites Silphium perfo­ liatum, S. integrifolium and Ambrosia trifida at eight sites in five Wisconsin counties in 1993-1995. This is the first report of P. reticulata using these plants and of its occurrence in Wisconsin.

In 1993, Publilia reticulata Van Duzee was found feeding on Silphium perfoliatum at Thomas Wet Prairie in Grant Co., Wisconsin. Because this treehopper is unreported from Wisconsin, it was sought widely around south­ western Wisconsin in the course of general prairie research during 1994­ 1995. Seven additional populations were found in Dane, Green, Iowa and Lafayette counties; at each site P. reticulata fed on either S. perfoliatum, S. integrifolium or Ambrosia trifida. This is the first report of this treehopper using these three composites.

METHODS AND MATERIALS All observations of P. reticulata were vouchered by specimens deposited in the Insect Research Collection (IRC) at the University of Wisconsin-Madison. Literature was reviewed for information on food plants and distribution ofP. reticulata, and regional museums were checked for possible Wisconsin speci­ mens. Plant nomenclature follows Kartesz (1994).

RESULTS Publilia reticulata was collected at eight sites in five counties in south­ western Wisconsin in 1993-1995. It was found in consecutive years at five sites and nymphs were found in the company of adults at four sites. It was more often observed on S. integrifolium than on S. perfoliatum, and was ob­ served only once on A. trifida. At two sites, it was observed on both S. inte­ grifolium and S. perfoliatum. It was most often found on or near the midvein on the underside of leaves, and these midveins were often damaged, even to the extent that the distal portion ofsome leaves folded down where damaged. This treehopper was also !bund on young stem tissue, and was occasionally observed elsewhere on the leaf surfaces. Almost always, ants attended P. reticulata, vigorously defending it from my attack. This treehopper and its at­ tending ants were observed during the day and night. At two sites, P. reticu­

1 Department of Entomology, University of Wisconsin, Madison, WI 53706. 244 THE GREAT LAKES ENTOMOLOGIST Vol. 28, No.3 & 4 lata was found on the same stems of S. integrifolium as was P. concava (Say), its more locally abundant congener.

DISCUSSION This is the first report of P. reticulata using S. perfoliatum, S. integri­ folium and A. trifida. Kopp and Yonke (1973) listed the composites ironweed, aster, and Vernonia baldwinii as food plants. Deay and Gould (1935) wrote, "Taken from burdock and iron weed and in general sweeping." Bristow (1984) listed "ironweed (Vernonia spp.) and several other closely related species of the Compositae," and reported use 0f V. noveboracensis in New Jersey. This treehopper appears to specialize on plants ofthe Asteraceae. Its occurrence on species in other families, such as alfalfa, black locust and post oak (Dennis 1965), should be considered incidental. On the sites included in this study, Vernonia fasciculata, the sole local member of this genus, occurs only at Thomas Wet Prairie, and P. reticulata has not been found using it here or elsewhere in the region. On three sites in this study, S. laciniatum grows with S. integrifolium and/or S. perfoliatum, but P. reticulata has not been found using S.laciniatum here or elsewhere in the region. This treehopper must be able to reproduce on both S. integrifolium and S. perfoliatum. It was observed on S. integrifolium in consecutive years at five sites, adults were observed with nymphs on these two plants at several and adults were reared in the lab from nymphs collected from among adults on each of these two plants. The sole observation of this treehopper using A. trifida two adults attended by four ants - occurred late in the course of this study on a site hosting large populations of S. integrifolium and P. retic­ uiata, but only a single A. trifida plant. Whether or not this treehopper can reproduce on A. trifida is unknown. This is the first report of P. reticulata in Wisconsin. The distribution and habits of Membracidae in Wisconsin were studied by Dennis (1951, 1952, 1969) and Dennis and Dicke (1953) who did not mention P. reticulata. Dennis' collection of Membracidae was given to the IRC and contains no Wisconsin specimens of this treehopper. Kopp and Yonke (1973) mapped its distribution; their map did not include Wisconsin but did include neighboring Illinois and Iowa. The only Wisconsin specimens of this treehopper in the IRC are the au­ thor's. The Milwaukee Public Museum and The Field Museum ofNatural His­ tory have no Wisconsin specimens of this treehopper.

ACKNOWLEDGMENTS This paper results from the Prairie Insect and Spider Inventory of The Prairie Enthusiasts-Southwest Chapter, basic biotic research being con­ ducted at Thomas Wet Prairie with support from The Prairie Enthusiasts­ Southwest Chapter, the Citizens Natural Resources Association of Wisconsin, the Natural History Museums Council of UW-Madison, and several private donors, support for which I am most grateful. I am also grateful to D. Young and S. Krauth ofthe Entomology Dept. ofUW - Madison for their interest in this research, to P. Johnson for bibliographic help, to S. Borkin and P. Parrillo for their help with collection data at their respective institutions, and to two anonymous reviewers. 1995 THE GREAT LAKES ENTOMOlOGIST 245

LITERATURE CITED Bristow, C. M. 1984. Spatial segregation between Aphis vernoniae (Aphididae) and Publilia reticulata (Membracidae), two species of colonial Homoptera on New York ironweed. Can. EntomoL 116:855-859. Deay, H. O. & G. E. Gould. 1935. An annotated list ofthe Membracidae of Indiana (Ho­ moptera). Proc. Ind. Acad. Sci. 44:236-243. Dennis, C. J. 1951. A list of the Wisconsin species of Membracidae. Can. EntomoL 88:183-184. __. 1952. The Membracidae of Wisconsin. Trans. Wisc. Acad. Sci., Arts and Lett. 41:129-152. 1965. Oklahoma treehoppers (Homoptera, Membracidae). Proc. Okla. Acad. Sci.

__. 1969. The treehoppers of Wisconsin in relation to the tension zone (Homoptera, Membracidae). Amer. MidL Nat. 81:236-242. __. and R. J. Dicke. 1953. The Membracidae of the University of Wisconsin Arbore­ tum. Trans. Wisc. Acad. Sci., Arts and Lett. 42:131-141. Kartesz, J. T. 1994. A synonymized checklist ofthe vascular flora of the United States, Canada, and Greenland. 2nd ed. VoL 1. Biota of North America Program of North Carolina Botanical Garden. Timber Press, Portland. 622 pp. Kopp, D. D. & T. R. Yonke. 1973. The treehoppers of Missouri: Part 2. subfamily Smili­ inae; tribes Acutalini, Ceresini, and Polyglyptini (Homoptera: Membracidael. J. Kans. Entomol. Soc. 46:233-276. INSTRUCTIONS FOR AUTHORS

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