GREAT LAKES ENTOMOLOGIST

Transcription

1 Vol. 26, No. 1 Spring 1993 THE GREAT LAKES ENTOMOLOGIST PUBLISHED BY THE MICHIGAN ENTOMOLOGICAL SOCIETY

2 THE GREAT LAKES ENTOMOLOGIST Published by the Michigan Entomological Society Volume 26 No.1 ISSN TABLE OF CONTENTS A new Eutarsopo/ipus (Acari: Podapolipidae); parasite of Harpa/us herbivagus (Coleoptera: Carabidae) from Michigan Robert W. Husband...,...,... Methods for artificial rearing of solitary eumenid wasps (Hymenoptera: Vespidae: Eumeninae) Charles F. Chilcutt and David p, Cowan,.. "...,.., New distribution records af the tiger math genus Phragmafobia in North America (Lepidoptera: Arctiidae: Arcliinae) Julian P. Donahue..., , Aphid prey of Passa/oecus cuspidatus (Hymenoptera: Sphecidae) John M. Fricke..., , Dytiscidae and Noteridae of Wiscansin (Coleoptera). II. Distribution, habitat, life cycle, and identification of species of Dyliscinae William L Hilsenhoff..., Microctonus pachy/obii (Hymenoptera: Braconidae): New host record from Hylobius radicis (Colebptera: Curculionidae), and additionol notes on its biology George D. Hoffman and Kenneth F. Raffa Field investigations on the American dog tick, Dermacentor variabilis, in northwest Ohio (Acari: Ixodidae) Kelly M. Micher and C. Lee Rockett..., Resistance stability of the secondary tiller of 'Caldwell' wheat ofter the primary culm was infested with virulent Hessian fly (Diptera: Cecidomyiidae) larvae Stanley G. Wellso and Jaime E. Araya,..., New prey families for Crabro advena (Hymenoptera: Sphecidae) Frank E. Kurczewski...,..., Use of communal nest entrances by Osmia simi/lima (Hymenoptera: Megachilidae) Virginia Scoll COVER PHOTOGRAPH Euphydryas phaeton (Drury) larvae (Lepidoptera: Nymphalidae) on Indian paintbrush, Castilleja coccinea, Cheboygan Co., MI. Photograph by Brian Sholtens.

3 THE MICHIGAN ENTOMOLOGICAL SOCIETY President President-Elect Treasurer Secretary Journal Editor Newsletter Editor OFFICERS Fred Stehr Catherine E. Bach M.C. Nielsen Edward Walker Mark F. O'Brien Robert Haack The Michigan Entomological Society traces its origins to the old Detroit Entomological Society and was organized On 4 November 1954 to n promote the science of entomology in all its branches and by all feasible means, and to advance cooperation and good fellowship among persons interested in entomology." The Society attempts to facilitate the exchange of ideas and information in both amateur and professional circles, and encourages the study of insects by youth. Membership in the Society, whicb serves the North Central States and adjacent Canada, is open to all persons interested in entomology. There are four paying classes of membership; Student (to 12th gradel-annual dues $5.00 Active-annual dues $10.00 Institutional-annual dues $25.00 Sustaining-annual contribution $25.00 or more Life - $ Dues are paid on a calendar year basis IJan. I Dec. 3l}. Memberships accepted before July 1 shall begin on the preceding January 1; memberships accepted at a later date shall begin the following January 1 unless the earlier date is requested and the required dues are paid. All members in good standing receive the Newsletter of the Society. published quarterly. All active and sustaining members may vote in Society affairs. All dues and contributions to the Society are deductible for \<'ederal income tax purposes. SUBSCRIPTION INFORMATION Institutions and organizations. as well as individuals not desiring the benefits of membership, may subscribe to The Great Lakes Entomologist at the rate of $20.00 per volume. The journal is published quarterly; subscriptions are accepted only on a volume (4 issues} basis. Single copies of 1'he Great Lakes Entomologist are available at $6.00 each, with a 20 percent discount for 25 or more copies sent to a single address. MICROFILM EDITION: Positive microfilm copies of the current volume of The Great Lakes Entomologist will be available at nominal cost. to members and bona fide subscribers of the paper edition only, at the end of each volume year. Please address all orders and inquiries to University Microfilms, Inc., 300 Zeeb Road, Ann Arbor, Michigan 48106, USA. about back numbers, subscriptions and Society business should be directed to the Executive S",.erAt:n v. Michigan Entomological Society. Department of Entomology, Michigan State University, East Lansing, Michigan l115. USA. Manuscripts and related correspondence should be directed to the Editor (see inside back cover). Copyright The Michigan Entomological Society

4 1993 THE GREAT LAKES ENTOMOLOGIST A NEW EUTARSOPOLIPUS (ACARI: PODAPOLIPIDAE); PARASITE OF HARPALUS HERBIVAGUS (COLEOPTERA: CARABIDAE) FROM MICHIGAN1 Robert W. Husband 2 ABSTRACT Eutarsopolipus porteri n. sp. (Acari: Podapolipidae) is described from Harpa Ius herbivagus and E. elongatus is reported for the first time in North America from Amara aenea (Coleoptera: Carabidae) from Fort Custer State Recreation Area, Kalamazoo County, Michigan. Of the 7 species of Podapolipidae known from American Carabidae, E. elongatus was introduced with Amara aenea and the others are native species. All are parasites restricted to Carabidae. Lindquist (1986) placed Podapolipidae adjacent to Tarsonemidae within the Tarsonemoidea (Heterostigmata). In adaptation to a parasitic mode of life, podapolipids exhibit very unusual characters. These include loss of 1 to 3 pairs of legs in females. migration of the aedeagus to a posterior, middorsal or anterior position and loss of leg setation. Podapolipid mites which are parasites of carabid beetles were considered to be among the most primitive podapolipid mites by Regenfuss (1973), Eickwort (1975) and Husband (1991). Six species of podapolipid mites have been described from North American Carabidae. These species have been assigned to three genera, Ovacarus, Eutarsopo /ipus and Dorsipes. Stannard and Vaishampayan (1971) described Ovacarus clivinae from oviducts of Clivina impressifrons LeConte from Illinois. Husband (1974) described the second species. Ovacarus peellei from the oviducts of Pasimachus elongatus LeConte from North Dakota. Regenfuss (1974) described Eutarsopolipus latus from beneath the elytra of Chlaenius aestivus Say and E. inermis from Evarthrus furtiva LeConte from Georgia. Husband and Swihart (1986) described E. regenfussi from Chlaenius pennsylvanicus Say from Michigan. Husband and Rack (1991) described Dorsipes evarthrusi from Evarthrus americanus Dejean from Georgia. Studies are continuing on undescribed Eutarsopolipus from Canada and the Eastern United States collected by the author or obtained from E. E. Lindquist (Biosystemstics Research Centre, Ottawa, Canada) and G. C. Eickwork (Dept. of Entomology, Cornell Univ., Ithaca, NY). The present paper deals with two species of Eutarsopolipus associated with carabid beetles collected in pitfall traps in Western Michigan by Lee Williams and T. Wayne Porter. Utilizing characters from Regenfuss (1968 and 1974), the mites removed from two host individuals were determined to be E. elongatus (Regenfuss 1968) which had been described from the vicinity of Erlangen, Germany and a lcontribution Number 709, Kellogg Biological Station of Michigan State University, :J!ickory Corners, MI. Biology Dept., Adrian College, Adrian, MI

5 2 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 new species related to E. elongatus and E. pseudopus Regenfuss This species is described and compared with related species. METHODS AND MATERIALS Beetles trapped in cans containing ethylene glycol were individually inspected for mites, then flushed with 70% ethanol to remove remaining mites. More than 20 mites, with all stages represented, were removed from one hind wing base of one beetle. Two mites were removed from a second beetle. Some specimens were placed in lactic acid for clearing before mounting. Long setae are often broken, reflexed and not in horizontal positions in preparations of podapolipid mites. Measurements included in this paper are of entire setae. Mites were examined, drawn and measured with a Wild-Heerbrug phase contrast microscope fitted with a drawing apparatus and stage micrometer. Terminology follows that of Lindquist (1986). All measurements are given in micrometers (/illl). Eutarsopolipus porten new species Female: (Figs. 1,2)-Dimensions as in Table 1. Gnathosoma elongate, length 44-50, width 40-41; dorsal setae 28, ventral setae 12; cheliceral stylets Idiosoma-Length , width Stigmata near anterolateral margin of prodorsal shield, tracheal branches meet medially to form a V under anterior margin of prodorsal plate. Setae VI' Vi vestigial, length at most 112 width of setal socket. Setae SCz 39, extending well beyond posterior border of prodorsal plate. Plate C length 80-85, width ; setae CI 13, cjl microsetae. Plate D length 73, widtb 206. seta d 8. Plate EF length 49, width 180; setae f 3. Venter with apodemes heavily sclerotized, apodemes 1 meet at anterior margin of sternal apodeme. apodemes 2 meet at posterior margin of sternal apodeme. Coxae III separated. Setae hilo, distance between setae hi 12. Setal numbers, including solenidia. for egs I, II, III as in Table 2. Length of solenidion omega on leg I about 2 times width, on leg II omega length nearly equals width. Solenidion phi on leg I 9, adjacent seta k 4; seta k thick. Coxal setae la, 2a 8, lancet-like, setae originate well posterior of apodeme 1. Setae 3a 6, 3b 7. Claw on leg I prominent, no claws on legs II, III. Male: (Figs. 3,4)-Dimensions in Table 1. Gnathosoma length 30-33, width 28-33; dorsal setae 9-11, ventral setae 5-6; cheliceral stylets Idiosoma Len~h , width Prodorsal shield wider than long, setae vl.' V2 vestigial, setae SC2 34. Plates CD fused, setae c, d microsetae. Genital plate length 40-47, width Venter with apodemes 1 meeting at sternal apodeme, apodemes 2 nearly so. Coxae III separated. Legs. Setal numbers as in Table 2. Spine-like setae on tarsi II. III well developed, claws absent. Setae la, 2a 3, both setae thin. Setae 3a, 3b microsetae. Larva: (Figs. 5,6)-Dimensions as in Table 1. Gnathosoma length 35-38, width 29-34, dorsal setae 28, ventral setae 12; cheliceral stylets 30. Idiosoma Length , width Setae v\'. t:.t and C2 microsetae. setae SC Plates C fused with plate D posteriomedially; setae CI 13-18, d 11-13, both setae thick, lancet-shaped. Plate EF wider than long, setae f 9, thin and extending to posterior margin of plate EF. Apodemes 1 and 2 meet sternal apodeme near mid coxae II, coxae III separated. Setae hi 180, h2 25; distal margin of h2 closely apresssed to setae hj' Distance between setae hi 19. Legs. Leg segment setation as in Table 2. Tarsus I solenidion omega 2,

6 1993 THE GREAT LAKES ENTOMOLOGIST 3 -r~ =~ E E (II) o 1 Figure 1. Eutarsopoiipus porteri n. sp., female, dorsal.

7 4 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No mm I ' ~--~~ ~~~ ~- 2 Figure 2a-c. 2a. Eutarsapalipus parten n. sp., female, gnathosoma and legs I, dorsal; ventral aspect of right leg I.; 2b. Eutarsapalipus parten n. sp., female, ventral, coxae I, II.; 2c. Eutarsapolipus parten n. sp., female, ventral, setae hi' tibial solenidion phi 8 with adjacent seta k 3. Coxal setae la 9, 2a 10; setae 3a 10, 3b 8. All coxal setae thick, lancet-shaped. Type data: Holotype larva: Fort Custer Recreation Area, Twp. 2 S, Range 9 W" Sec. 23, Kalamazoo County, Michigan, from Harpalus herbivagus Say (Carabidae), collected 30 July 1991 by Lee Williams. Deposited in the Museum of Zoology, the University of Michigan, Ann Arbor, MI (RWH ). The type host is located in the Museum of Zoology, University of Michigan. Paratypes (3 males, 8 females, 9 larvae)-same data as holotype: 1 male (RWH ), 1 female (RWH ), 1 larva (RWH ) deposited in the Acarology Collection of the Museum of Zoology, University of Michigan, Ann Arbor, Michigan. One male (RWH ), 1 female (RWH ), 1 larva (RWH ) deposited in the Zoologisches Museum, Universitat Hamburg. The balance of paratypes are stored in the Acarology Collection, Biology Department, Adrian College, Adrian, Michigan. Additional specimens are stored in a vial with 70% alcohol in the Acarology Collection at Adrian College. Etymology: The species is named for Dr. T. Wayne Porter in tribute to his

8 1993 THE GREAT LAKES ENTOMOLOGIST 5 Table 1. Selected measurements for Eutarsopolipus spp. All measurements are in micrometers (I'm). Setae designated as vestigial (v) are represented by tiny spots. Setae designated as microsetae (m) are no longer than the diameter of the setal socket. The letter T after a setal measurement indicates that the seta is conspicuously thicker than most setae. The letter t indicates somewhat thicker than most setae.. E. porten E. elongatus E. pseudopus E. regenfussi FEMALES Idiosomal length Idiosomal width Dorsal Gnath. setae Vent. Gnath. setae m Cheliceral stylets Setae vj/v2 V C m m Setae sc Setae f 3 S S m Setae la,2a ST 5/7 514 m LARVAE Idiosomal length Idiosomal width los C heliceral stylets Dorsal Gnath. setae 2S 21 IS 19 Vent. Gnath. setae m Setae vl/v2 v v v 3 Setae Cl 1ST 12t 5T 3 Dist. cjcj Setae 13T lot 5T 3 Distance d-d Setae f Coxal setae la 9T 5T 7T m Coxal setae 2a lot 5T 6T m Coxal setae 3a lot 13t lot 4 Coxal setae 3b ST 6T lot 6 MALE Idiosomallength 17S ISO 136 Idiosomal width Cheliceral stylets Vent Gnath. setae Gen. pl. midwidth Coxal setae la 3 m m Coxal setae 2a 3 m m Table 2. Leg setation for femur, genu, tibia, tarsus for Michigan Eutarsopolipus and E. pseudopus. Solenidia are included. Leg I Leg II Leg III F G Ti Ta F G Ti Ta F G Ti Ta E. porten S E. elongatus S E. pseudopus E. regenfussi S

9 6 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 E E,... o 3 Figure 3. Eutarsopolipus porten n. sp., male, dorsal.

10 1993 THE GREAT LAKES ENTOMOLOGIST 7 4 Figure 4. Eutarsopolipus porteri n. sp. male. ventral.

11 8 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 "y e 1 C!J y 2 E E,... o 5 Figure 5. Eutarsopolipus porteri n. sp., larva, dorsal. helping of students at W. K. Kellogg Biological Station for 37 years and in recognition of his 80th birthday on August 8, Diagnosis: Female E. porten differ from E. elongatus in having coxal setae la and 2a removed from apodemes 1 and 2. These setae are thicker than those illustrated for E. elongatus by Regenfuss (1974) and confirmed by comparison with local E. elongatus. Setae la and 2a are thinner than the respective setae in larval stages. Cheliceral stylets are equal to the width of the gnathosoma in E. porten but are shorter in E. elongatus. Males of E. elonga

12 1993 THE GREAT LAKES ENTOMOLOGIST 9 6 Figure 6. Eutarsapolipus parten n. sp., larva, ventral. tus are unknown. Larval E. porteri have longer coxal setae la, 2a and 3b than are found in E. elongatus but setae 3a are shorter. Setae CI and d are slightly longer and thicker in E. porteri.

13 10 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No. I E E ('W) o Figure 7. Eutarsopolipus elongatus Regenfuss, female, dorsal

14 1993 THE GREAT LAKES ENTOMOLOGIST 11 Figure Sa, h. Sa. Eutarsopolipus elongatus Regenfuss, female, gnathosoma and legs I, dorsal; ventral aspect of right leg I; 8b. Eutarsopolipus elongatus Regenfuss, female, ventral, coxae I, II. Eutarsopolipus elongatus Regenfuss A female and a swollen larva of E. elongatus were removed from Amara aenea DeGeer. a carabid beetle introduced from Europe. A. aenea is the host for E. elongatus in Europe. A female E. elongatus is illustrated by Regenfuss (1974). Both female and larvale. elongatus from Michigan are illustrated here for comparison with E. porten (Figures 7, 8, 9). This is the first record of an introduction of a podapolipid mite from carabid beetles to North America. Since A. aenea is widely distributed in Michigan, the introduced mites may be encountered fairly often and a detailed description is necessary.

15 12 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 A \ ) 9 c Figure 9. a-c. 9a. Eutarsopolipus elongatus Regenfuss. larva, dorsal, propodosoma; 9b. Eutarsopolipus elongatus Regenfuss. larva, ventral. coxae I. II; 9c. Eutarsopolipus elongatus Regenfuss, larva. ventral, coxae III. DISCUSSION Regenfuss (1968) utilized the following characters in assembling five Eutarsopolipus species into a cluster of related species: in females, (1) setae VI and V2 no longer than setal socket, (2) apodeme III lacking, (3) setae SCI absent and (4)ambulacra II and III without claws. In addition, Regenfuss characterized this group by: (1) females with a long, stout femoral I l' seta, (2) larvae with setae hi widely separated and (3) males with genital plate about as broad as long. The single male specimen in this ~oup observed by Regenfuss was the male of E. acanthomus and it was not illustrated. Included in this group were: E. acanthomus, E. alarum, E. assimilis, E. crassisetus and E. elongatus.

16 1993 THE GREAT LAKES ENTOMOLOGIST 13 In 1974, Regenfuss addede.pseudopus to this subset ofthe genus Eutarsopolipus, and E. porten also shares the characteristics of this group. The other North American Eutarsopolipus species belong to different species groups. Regenfuss (1974) placed E. latus and E. inermis, collected in Georgia, with E. desani and E. pterostichi respectively. Husband and Swihart (1986) placed E. regenfussi, collected in Northern Michigan, near E. latus. Within the cluster of mites that includes E. porten, the new species shares the thickened larval coxal setae la and 2a with E. pseudopus, E. elongatus, E. alarum and E. crassisetus. Eutarsopolipus regenfussi is included in Tables 1 and 2 for comparison with the remaining more closely related species. The leg setal patterns for all species are not known. However, species related to E. porten have a seta on genua II and III while this seta is absent in E. regenfussi (Table 2). Femur I in E. pseudopus has 3 setae in contrast to 2 for other species in this discussion. Genu I has 2 setae in E. porten and relatives but none in E. regenfussi. Thus, common setal patterns also support the inclusion of E. porten within the group proposed by Regenfuss. ACKNOWLEDGMENTS I thank Lee Williams for aid in obtaining the beetles, T. W. Porter for help in determining the specific collecting sites, Gary Dunn for identification and information on the host carabid beetles, the staff of the W. K. Kellogg Biological Station of Michigan State University for support during this study. LITERATURE CITED Eickwort, G. C A new species of Chrysomelobia (Acari: Tarsonemina: Podapolipidae) from North America and the taxonomic position of the genus. Canad. Entomol. 107: Husband, Robert W Ovacarus peellei, a new species of mite (Acarina: Podapolipidae) associated with the carabid Pasimachus elongatus. Great Lakes Entomol. 7(1): A preliminary investigation of the phylogeny of Tarsopolipus, a comparison with other early derivative genera. Pp , In: F. Dusbabek and V. Bukva, OOs., Modem Acarology, Vol. 1, SPB Academic Publishing bv., The Hague and Academia, Prague. Husband, Robert W. and Gisela Rack Dorsipes evarthrusi sp.n. (Acari: Podapolipidae), ectoparasite of Evarthrus americanus (Carabidael from Georgia, U.S.A. Entomol. Mitt. zool Mus. Hamburg 10: Husband, Robert W. and Cheryl D. Swihart A new species of mite (Acari: Podapolipidae) from a Michigan carabid beetle, Chlaenius pennsylvanicus. Great Lakes Entomol. 19: Lindquist, E. E The world genera of Tarsonemidae (Acari: Heterostigmata): a morphological, phylogenetic, and systematic revision, with a reclassification of family-group taxa in the Heterostigmata. Mem. Ent. Soc. Canada 136: Ottawa. Regenfuss, H Untersuchungen zur Morphologie, Systematik und Okologie der Podapolipidae (Acarina, Tarsonemini). Z. wiss. Zool. 177: Leipzig BeinrOOuktion und Verlagerung des Koppulationsapparates in der Milbenfamilie Podapolipidae. ein Beispiel fur verhaltensfesteuerte Evolution morphologischer Structuren. Z. zool. Systematik u. Evolutionsforschung 11:

17 14 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No Neue ektoparasitische Arten der Familie PodapoJipidae (Acari: Tarsonemini) von Carabiden. Mitt. Hamburg. Zool Mus. Inst. 71: Hamburg. Stannard, L. J. and S. M. Vaishampayan Ovacarus clivinae, new genus and species (Acarina: Podapolipidae), an endoparasite of the slender seedcorn beetle. Ann. Entomol. Soc. Amer. 64:

18 1993 THE GREAT LAKES ENTOMOLOGIST 15 METHODS FOR ARTIFICIAL REARING OF SOLITARY EUMENID WASPS (HYMENOPTERA: VESPIDAE: EUMENINAE) Charles F. Chilcutt l,2 and David P. Cowan l ABSTRACT Solitary eumenid wasps of the genera Ancistrocerus and Euodynerus can be reared in small cages. Laboratory-reared larvae of the spruce budworm caterpillars, Choristoneura fumiferana (Lepidoptera: Torlricidae) are suitable prey. Many problems in insect biology are difficult or impossible to study in a completely natural setting, thus making artificial rearing methods necessary for further advances. Generally, it is necessary to provide adults with conditions needed for maintenance, an appropriate environment for males and females to mate, proper oviposition stimuli for females, and an adequate diet for immature stages. Fulfilling these conditions for hunting wasps is relatively complex because females require: (1) specific materials (from the environment) for nest construction prior to oviposition; and (2) live arthropod prey for females to paralyze and provision for their offspring. Here we report successfully rearing, under artificial conditions, solitary eumenid wasps of the genera Euodynerus and Ancistocerus. In nature, these wasps nest in hollow twigs and vacant insect borings (Krombein 1967). Within a cavity, a female lays an egg; she then hunts for and paralyzes caterpillars that are placed within the cavity or nest near her egg. When enough prey items have been placed in the cavity with the egg, the female then makes mud by mixing water and dry soil. She uses this mud to seal off a cell that contains her egg and its provisions. This process may be repeated in the same nesting cavity until it is filled with a linear series of cells containing her offspring. Observations by Steiner (1983) and by Veenendaal and Piek (1988) indicate that female eumenid wasps will engage in predatory and nesting behaviors within the confines of small cages. Such behavior makes large-scale artificial rearings possible. MATERIALS AND METHODS Eumenid wasps were obtained by rearing them from trap nests placed out in Kalamazoo and Iron counties, Michigan. Wasps were either removed from trap nests as larvae, or captured as young adults as they emerged from their nests. Males were transferred to a holding cage upon emergence, and two day ldepartment of Biological Sciences, Western Michigan University Kalamazoo, MI 4900~. Current address: Department of Entomology, University of Hawaii Maile Way. Honolulu. HI

19 16 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 post-emergence females were placed in the cage with the males (different ages) for mating. Both mated and unmated females were used in this study. Captive rearings were carried out on the Western Michigan University campus during the summers of Rearing cages were placed in a courtyard surrounded by a 3 story building. This allowed natural weather phenomena, but decreased sun exposure in the morning and evening. We reared wasps in 60 cm X 60 em X 60 em cages constructed of plywood bottoms, with one side and top of glass, and three sides of aluminum screen (18X16 mesh). For access, a 15cm square hole in the comer of a screened side supported a 100cm long cloth sleeve that could be knotted. In 1989, cages were raised 4cm above the ground on boards to prevent rotting. In 1990 and 1991, cages were placed on wooden stands (60cm high), with tangle foot applied to the legs to keep out pests. All outdoor cages contained: (1) a petri dish supplied daily with sugar water, (2) a jar lid of dried clayey mud, and (3) a petri dish supplied daily with water. A microscope slide was placed inside the water dish to allow wasps that fell in the water (a frequent occurrence) an escape route. Generally, the sugar water dried quickly: however, the wasps moistened the crystals with water and ate. Two or four females were placed in each cage, with each cage provided with two nests per wasp, wired to the inside screen or placed on the cage floor. The hole diameters of the nest sticks were the same as the sizes they selected in nature (5.6 mm, 6.4 mm, and 7.1 mm). The day after a nest was completed, as evidenced by a mud plug in the nest-hole entrance, the nest was removed and another nest was provided in its place. To provide wasps with potential prey, diapausing first instars of the spruce budworm, Choristoneura fumiferana (Clemens) were obtained from the Canadian National Forestry Laboratory in Sault Ste. Marie, Ontario, and reared on commercial (Bioserve Corporation) diet, following instructions in Grisdale (1984). Collins and Jennings (1987) found spruce budworms to be a common prey (94% of prey found in trap nests) of Euodynerus and Ancistrocerus species in a spruce-fir forest. By placing budworms on the diet at the time of first-male emergence, prey items of suitable size (> 2mm) were available when female wasps began hunting about 10 days later. As an alternative to the commercial diet, we reared some budworms on a diet of tender buds of white spruce, Picea glauca. Expanding buds were cut early in the growing season and frozen for later use. For rearing budworm larvae, buds were thawed, sprayed with the same fungicide used with the artificial diet, and then placed in 30ml plastic cups. Attempts to rear budworms on mature spruce foliage were unsuccessful. In 1989, 17 adult female Euodynerus foraminatus (Saussure), and one female Parancistrocerus sp. were placed in outdoor cages, with one cage containing four wasps and seven cages containing two wasps apiece, and observed from 23 June until 1 August. In 1990, 18 Euodynerus planitarsis (Bohart) and 8 E. foraminatus were placed in outdoor cages (six cages contained three E. planitarsis apiece and two cages containing 4 E. foraminatus apiece) on 15 June and observed until 21 September. In 1991,13 E. planitarsis, 2 E. foraminatus, 2 Euodynerus hidalgo (Saussure) and 4 Ancistrocerus antilope (Panzer) females were placed in outdoor cages on 1 June and observed until 30 September. Indoor rearings were attempted in Five E. foraminatus females were placed singly in pint jars; four were placed singly in gallon jars. Each jar contained two trap nests, a capful of mud, a capful of water, and white spruce shoots. Jar openings were covered with screening and the wasps were fed on honey smeared on this cover. Honey or sugar water provided needed carbohy

20 1993 THE GREAT LAKES ENTOMOlOGIST 17 drates for the wasps. Due to limited space in the jars, prey were removed from rearing cups and placed on the spruce in the jars. RESULTS AND DISCUSSION During 1989, all 17 female wasps entered nest sticks; however only 3 E. foraminatus hunted, captured prey, and provisioned nests. Beginning on 7 June, these 3 females completed eleven nests that produced adult offspring, 10 females and 9 males During 1990, all 14 E. planitarsis females nested beginning on 30 June. They completed 59 nests that contained 127 provisioned cells, which produced 84 adults. Five of the 8 E. foraminatus females nested and completed 11 nests that contained 46 provisioned cells and which produced 35 adults. All of the offspring reared in 1990 were adult males, indicating that none of the parent females had mated. We reared parental females from larvae placed in vials, and transferred the newly emerged adults to cages with males for a day. We then transferred the females into the rearing cages that contained nesting supplies. Apparently the females were not fully mature and ready to mate when placed in the cage with the males. We avoided this problem in 1989 and 1991 by leaving larvae and pupae in their nests and allowing them to transform to adults and emerge on their own. One day later we placed them in cages with males and watched to be sure they mated. In 1991, the first wasp began nesting in the cages on 7 June, apparently early due to unseasonably warm weather. ThirteenE. planitarsis completed 41 nests which contained 70 provisioned cells. One of 2 E. foraminatus females completed 2 nests with 3 provisioned cells. One of 4 A. antilope females completed 2 nests on the same day, each with one provisioned cell, before escaping from the cage. 1\vo of the 3 A. antilope that did not nest captured prey without placing them in a nest stick. The E. hidalgo females showed no interest in prey, never displaying huntinf5 or searching behaviors. Attempts to rear wasps indoors In jars were less successful. Although 6 of the 9 wasps began nesting, mortality of the females was high. The primary problem was obtaining a b8.1ance between proper light and temperature conditions. Wasps tended to remain in their nest under artificial. g (fluorescent) or when placed in naturally lit areas but not in direct su t. If placed in direct sunlight on warm days, temperatures in the jars soared and wasps died. Placing the jars in direct sunlight with partial shade seemed to help. Nests taken from the jars were damp, and most contained fungal growth. Apparently the heat and water provided for the wasps increased the humidity above safe levels. This problem did not occur outdoors in the better ventilated screen cages. In cramped rearing containers, caterpillars often crawled behind objects where the wasps could not reach. Spruce twigs placed in the rearing jars provided a centralized hunting area where budworms tended to stay. During the normal process of getting water and mud and returning to the nest to build mud partitions, several wasps were seen chewing on the caterpillars' silk and possibly their frass. Perhaps wasps were using something from these materials as additives to the mud partition. All observed wasps spent time searching over the mud provided them before collecting mud from a well chosen spot. Such behaviors indicate that selection of the right kind of mud (apparently cohesive enough for the wasp to mold) is important. The low nesting success in 1989 (only 3 wasps nested) apparently was due to ants and earwigs (Forficula sp.) getting into the cages. Cages placed on boards near the ground were accessible to small insects that could get through

21 18 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 cracks or the screen mesh. Earwigs entered the cages by crawling into the cage sleeve and falling into the cage when the sleeve was used. We later discovered (while opening nests) that earwigs sheltered in the nests, and possibly deterred wasps from nesting. However, several wasps barricaded earwigs in the ends of the nests with mud partitions. Three of 4 nests found this way successfully produced offspring. Ants were the most notable pest problem, because they competed with the wasps for budworm prey. Ants killed and dismembered budworms to remove them through the screen. When ants came upon wasps holding prey (which the wasps often do while waiting for the venom to work, the ants would grab onto the caterpillar and a tug-of-war ensued. The ants always came out the victor (observed > 20 times). In 1989, the cages were placed in an area where they were shaded during the warmer hours of the day (1200 to 1400 h), because we believed that direct sunlight would heat the cages to a deadly temperature. However, the limited nesting success of these wasps prompted us to place the cages in the open (i.e. direct sunlight all day) for subsequent studies in 1990 and Cut evergreen branches were placed on the tops of cages, thus creating shaded and sunny areas within each cage during the day. Only when air temperature exceeded 35 C were wasps observed resting for periods in the shade. One problem encountered by placing the cages in direct sunlight was high spruce budworm mortality. This problem was solved by placing the rearing cups that contained budworm larvae in the area of the cage shaded during mid-day. Our increased success in rearing wasps after 1989, mainly was due to placing the cages on stands with tanglefoot around the legs. Such barriers eliminated all ants and earwigs from the cages, although we still removed an occasional spider. At no time during the rearing studies did we find nest parasites or parasitoids of either wasps or spruce budworms. Hence, we conclude that the increased nesting success from 18% of wasps in the total sample which layed eggs in 1989 to 63% in 1990 primarily was do to exclusion of ants which significantly impact solitary wasps' nesting success. Mortality of E. planitarsis and E. foraminatus immatures reared in 1990 was only 31.2% compared to 45.3% for eumenids observed in nature (Cowan 1981). This difference (14.1 %) is attributable to the absence of nest parasites, parasitoids, and predators. Most of the wasp mortality during our study (> 60%) resulted from our clumsy handling of nests which caused injury to unhatched eggs or delicate immature wasps. Fungi and other unknown pathogens contribute to the remainder of the mortality. We found the following factors most important for successful propagation of wasps: (1) Insure that female wasps have successfully mated; otherwise, only male offspring will result. (2) Supply adequate nesting resources i.e., nest sticks, carbohydrates (sugar or honey), water at all times, dried mud with adequate clay content, and potential prey larvae. (3) Daily exposure to sunlight, with partial shade to avoid overheating, is needed; also, adequate ventilation to control humidity. (4) Measures to exclude pests such as ants improve nesting success. LITERATURE CITED Collins, J.A., and D.T. Jennings Nesting height preferences of eumenid wasps (Hymenoptera: Eumenidae) that prey on spruce budworm (Lepidoptera: Tortricidae). Ann. EntomoL Soc. Am. 80: Cowan, D.P Parental investment in two solitary wasps Ancistrocerus adiabatus and Euodynerus foraminatus (Eumenidae: Hymenoptera). Behav. Ecol. SociobioL 9: Grisdale, D.G A laboratory method of mass rearing the eastern spruce budworm

22 1993 THE GREAT LAKES ENTOMOLOGIST 19 Choristoneura fumiferana. In: E.G. King. and N.C. Leppla (eds.j. Advances and challenges in insect rearing. U.S. Dept. Agric. Tech. Bull. pp Krombein. K.V Trap-nesting Wasps and Bees: Life Histories. Nests. and Associates. Smithsonian Press, Washington, D.C. Steiner, A.L Predatory behavior of solitary wasps V. Stinging of caterpillars by Euodynerus foraminatus (Hymenoptera: Eumenidae). BioI. Behav. 8: Veenendaal. R.L., and T. Piek Predatory behaviour of Discoelius zonalis (Hymenoptera: Eumenidae). Entomol. Ber. (Amsterdam) 48:8-12.

23 1993 THE GREAT LAKES ENTOMOlOGIST 21 NEW DISTRIBUTION RECORDS OF THE TIGER-MOTH GENUS PHRAGMATOBIA IN NORTH AMERICA (LEPIDOPTERA: ARCTIIDAE: ARCTIINAE) Julian P. Donahue] ABSTRACT New distribution records for all three Nearctic species of Phragmatobia include state records (the first records for the states indicated) of P lineata (Maryland, Wisconsin); P fuliginosa rubricosa (Ohio, South Dakota), and P assimilans (Idaho, Montana, Pennsylvania, Wisconsin), all representing southern range extensions at those longitudes except for the Wisconsin records of P lineata, which are northern range extensions. Chelone glabra (Scrophulariaceae) is reported as a larval hostplant of P lineata, and descriptive notes on the larva of this species are included. Midwinter activity of a larva crawling on snow is reported for P fuliginosa rubricosa. The rare original description of Phragmatobia dallii Packard, 1870, is reproduced. Since the late John H. Newman and I reviewed the North American species of Phragmatobia (Donahue and Newman, 1966), I have examined additional material in several collections (see Acknowledgments, where acronyms used to indicate the source of specimens examined are explained), resulting in the discovery of a number of significant new distribution records. I include only first records for a state, records from a region where the species has been infrequently collected and/or may be in danger of extirpation, or records from localities sufficiently distant from the previously documented distribution to be 6f interest. The distribution maps here are those used for the 1966 paper, with the additions and corrections cited in the present paper. In the species accounts I include several corrections and additions to that earlier paper; states and counties within each state are arranged alphabetically. Despite the partial larval descriptions in some of the species accounts, more rearing and comparative larval morphological studies are needed before we have a clear understanding of the larval systematics in this genus. Existing larval descriptions are persistently conflicting, incomplete, and sometimes based on misidentification of the parent female or offspring, or on an assumed identity for a field-collected larva. One such example is Saunders' (1863: 372) report of the larva of P fuliginosa from St. Catherines [sic], Ontario, which clearly refers to the fall webworm, Hyphantria cunea (Drury), predating by five years the first recognized report of this species from Canada [this misidentification will be elaborated upon in a separate paper]. lnatural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, CA

24 22 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 CD:,~ Figure 1. Distribution of Phragmatobia lineata; dots represent localities previously published by Donahue & Newman (1966), stars represent new localities for specimens examined and included in the present paper, triangles represent reliable records published or communicated by others.! ' PHRAGMATOBIA Stephens, 1828 Correction to 1966 paper: Phragmatobia was proposed in 1828 (Stephens, 1828: 55, 73), not 1829 as we stated, although Stephens subsequently proposed the same name twice again in 1829; see Watson et al. (1980: 153) for further details. We mentioned, without further discussion, five Neotropical species that were, or had been, placed in Phragmatobia. Three of them, together with five additional species, are still catalogued inphragmatobia (Watson and Goodger, 1986: 28), but Ferguson (1985: 241) recently proposed the new genus Sonorarctia for two of them [So feruida (Walker) and S. nundar (Dyar)], the latter he reported from the United States for the first time (Huachuca Mts., Cochise Co., Arizona). I expect that further study will demonstrate that Phragmatobia is a strictly Holarctic genus with no Neotropical representatives. PHRAGMATOBIA LINEATA Newman & Donahue, 1966 (fig. 1) Corrections to 1966 paper: I have re-identified as P. lineata the 2 female specimens from ILLINOIS: Peoria Co. (Elmwood) and INDIANA: Lake Co. (Clarke), previously identified and mapped as P. fuliginosa. New Records: MARYLAND [STATE RECORD]: BALTIMORE CO., Stevenson, 27 June 1959, Robert S. Bryant (1 male, RSB); same locality, emerged 14 June 1965 from larva collected on Chelone glabra (Scrophulariaceae) in late May-early June of the same year (1 female, RSB). [This is a new

25 1993 THE GREAT LAKES ENTOMOlOGIST 23 foodplant record for the genus in North America, but not a surprising one, considering the polyphagous nature of the group.] MASSACHUSETTS: PLYMOUTH CO.: East Wareham, Agricultural Experiment Station, 6, 12, & 21 July 1971, W.E. Tomlinson (3 males, FSCA). MICHIGAN: OTSEGO CO.: T29N, R3W, Section 13, mature larva collected 30 March 1968 as it crawled on exposed peat at the margin of a leatherleaf (Chamaedaphne calyculata)-labrador tea (Ledum groenlandicum) bog (both Ericaceae); spun cocoon the same evening, adult moth emerged in laboratory 11 April (1 male, JPD 1/68-6, MSUE). [This record was published without rearing details by Newman and Nielsen (1973: 36).] Because of the locality (in the Northern Lower Peninsula, more than 100 miles north of any previously known Michigan locality for P. lineata), and the uniformly yellow, short, even-length setae, I initially mistook the larva for that of Cycnia oregonensis (Stretch); however, unlike that common species, the skin of this larva was blackish (not yellow). The setal color is in accord with the partial larval description given earlier (Donahue and Newman, 1966: 45), while the black skin, not previously noted, may aid field workers in the discrimination of the similar larvae of these two moths. NEW YORK: McCabe (1990: 8-9,) reports rearing this species from ova obtained from a female collected in the daytime at Browns Tract Pond bog, HAMILTON CO. near the northern limit of known distribution, and figures (p. 29) a larva reared on Spiraea latifolia (Rosaceae). He notes than fewer than 10% (of an unspecified number) of the larvae pupated; the rest diapaused and overwintered, but did not survive (McCabe, pers. comm.). The preserved larva I examined from that rearing is distinctly two toned: the setae of abdominal segments 2-6 are predominantly yellowish. contrasting- sharply with the blackish setae on the anterior and posterior ends; the skm is dusky. but not blackish as I observed in the Michigan specimen noted above, although some fading may have occurred in alcohol. PENNSYLVANIA: SCHUYLKILL CO.: Schuylkill Haven, 27 & 30 June 1969, Wm. Houtz (4 males, LACM); {no city], 16 July 1971, J. Gilbert (2 males, ER). WAYNE CO.: South Sterling, 15 July 1918, Ernest Baylis (1 male, CMP). WISCONSIN {STATE RECORDS]: DANE CO.: T9N, R6E, Sec. 29, 29 June 1981, Les Ferge (1 male, LAF). IOWA CO.: Dodgeville, T7N, R3E, Sec. 31, 27 July 1975 (1 male) & 24 Aug (2 males), Wm. E. Sieker (LACM). KENOSHA CO.: TIN, R23E, Sec. 31, 22 July 1989, Les Ferge (2 males, LAF). WAUKESHA CO.: T5N, R17E, Sec. 16, 2 July 1989, Les Ferge (1 male, LAF). PHRAGMATOBIA FULIGINOSA RUBRICOSA (Harris, 1841) (fig. 2) Corrections to 1966 paper: see the discussion above under P. lineata for the reidentification of 2 specimens from Illinois and Indiana, originally cited as P. fuliginosa rubricosa. Although Phragmatobia fuliginosa is a Holarctic species, said to occur across the entire Palearctic Region from England to Japan (Seitz, 1910: 79), it should be noted that the genitalia figured for this species (from an unspecified locality) by KOda (1988: 9-11, fig. 62) appear to belong to some other species of Phragmatobia; details of the valva and aedeagus differ significantly from those of "true" P. fuliginosa from western Europe and North America. Additional references: Dyar (1899: 130) observed that Phragmatobia fuliginosa "...possibly occurs in Alaska. The National Museum has a specimen collected by Dr. Stejneger on Bering Island, off Kamchatka." We confirmed

26 24 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 Figure 2. Distribution of Phragmatobia fuliginosa rubricosa; symbols as for Fig. 1, scale as in Fig. 3. "Bush" pattern represents approximate northern limit of forests (tree line). the presence of this species in Alaska and cited a number of specimens examined. We did examine the Bering Island specimen in the USNM that Dyar mentioned, but excluded it from consideration in our review because it was extralimital to our study (Bering Island is in the Komandorskiye Islands. Russia), and it did not appear to be referable to any of the Nearctic taxa we were studying; the Palearctic Phragmatobia fauna, beset with many names for few species. remains a taxonomic quagmire to the best of my knowledge. Ferguson (1975: 10) reports the successful rearing of this species on Plantago major (Plantaginaceae) in Nova Scotia, from eggs laid by a female found flying by day on 3 May This reference provides the specific name of the foodplant (on p. 5) and information on parental behavior that were lacking from the label data we published for some of the reared offspring (Donahue and Newman, 1966: 56). Morris (1980: 80) adds one Newfoundland locality (Colinet) to the two previously mapped (Donahue and Newman, 1966: fig. 36), and confirms that it has not yet been reported from Labrador. Curiously, his summary of this species'distribution repeats the statement that it.occurs "south along the

27 1993 THE GREAT LAKES ENTOMOLOGIST 25 Appalachian Mountains into the Carolinas," apparently based on the nearly identical but still unconfirmed statements by Holland (1903: 126) and Seitz (1919: 302) which we noted in our review. Although we first questioned the occurrence of this species in the Carolinas over 25 years ago, I have still not seen any specimens from south of Pennsylvania. It is quite possible that this moth may eventually be found at high elevations in the southern Appalachians, but Holland's reason for believing this to be so remains a mystery. Another perplexing southern record of this moth appeared in Lindroth (1957: 72), who, with a bit more precision than Holland, reported Phragmatobia fuliginosa rubricosa from the state of South Carolina. I can only conclude his record is also based on that of Holland (1903: 126), the only likely source I could find among his references. It is likely that Rothschild's (1910: 116) records of P. fuliginosa rubricosa from "Florida," far outside the range of the species, actually refer to specimens from Florida, Orange County, New York. I have, however, discovered a "Florida" specimen from Florida City [Dade Co.], Florida, 21 April 1934, "CoIl. R.H. Andrews, Lloyd M. Martin" (1 male, LACM). There are two possible explanations for this label: it was applied in error, or the collectors happened to capture a specimen imported (perhaps as an immature) by a tourist from the usual range of this species. I am reasonably confident that this species is not a normal resident of Florida. The bibliography of North American Phragmatobia would not be complete without mentioning a long-overlooked publication by Packard (1870), the subject of a brief note by Banks (1920) that summarized its contents. This paper is so rare that only two copies are known to exist, one in the Museum of Comparative Zoology, Harvard University, the second in the National Agri cultural Library, Beltsville, Maryland (D.C. Ferguson, pers. comm.). On p. 29 of that paper Packard describes Phragmatobia dallii as new, from an unspeci fied number of specimens taken on 15 June [1867?] somewhere in Alaska. Because of the scarcity of this publication, I reproduce here the complete original description: Phragmatobia Dalla n. sp. PI. II, fig. 14. It is rather smaller than P. rubricosa, with the thorax deep reddish brown, being a little darker than in P. rubricosa, while the abdomen is a little darker, with a row of black dots on each side. The fore wings are plain rusty reddish brown, being duller and more tawny than in P. rubricosa, with no markings, except the single black discal dot; hind wings dusky, being almost wholly black, with the inner edge deep red, while the costal edge is pale red, with the apex dusky, and the fringe pale red; discal dot black; beneath uniform dusky tawny, with the dlscal dots prominent; hind wings reddish on the basal half of the costa. and on the inner edge. Legs as in P. rubricosa. Length of the fore wing, 0.50; length of body, 0.50 inch. It differs from P. rubricosa in its smaller size, the uniform tawny reddish brown wings, and in wanting the two dark bands on the fore wings, and in the hind wings being almost wholly dusky. Taken June 15. Packard's reference to P. rubricosa having "two dark bands on the fore wings" indicates that his concept of that taxon actually referred to P. assimilans or, more likely, to the species subsequently named P. lineata, a commonly made mistake (Donahue and Newman, 1966: 41, 45). Packard's description of P. dallii accords well with Alaskan specimens of P. fuliginosa rubricosa, and I agree with Franclemont (1983: 117) that P. dallii is a junior subjective synonym of P. fuliginosa rubricosa. In his entry for this and the other new species of moth Packard described in that paper, Gastropacha alascensis, FrancIemont (1983: 107, 117) questioned the availability of the names, presumably

28 26 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 based on the rarity of the publication and its distribution. It is worth noting that the two new species of wasps Packard described in the same paper, Vespa tripunctata and Vespa alascensis, have been treated as available since at least 1931, as junior synonyms of Vespula austriaca (Panzer) and V. VUlgaris (Linnaeus), respectively (Bequaert, 1931: 90, 106; Miller, 1961: 8, 19; Krombein et ~., 1979: 1519, 1521~ New Records: CANADA: ONTARIO: Riotte (1992: 126) reports P. fujiginosa from the following counties not previously mapped: BRUCE, ESS~X, LAMBTON, MIDDLESEX, NIPISSING, RAINY RIVER, SIMCOE. and WELLINGTON. U.S.A.: COLORADO: DENVER CO.: Denver, 30 April 1892, at light (1 m~e, CSU, Acc. No. 334). LARIMER CO.: Fort Collins, at light, 11 April 1892,17 & 26 July 1897, 19 July [Sept.'!] 1930 (4 m~es, CSU, Acc. No. 1080, 2654, 2670). Rothschild (1910: 116) ~so recorded two m~es each from "Larima" County and Durango, [LA PLATA CO.], Colorado. Ferguson (pers. comm.) reports a specimen from MORGAN CO.: Muir Spring Park and Recreation Area, Fort Morgan, 4300 ft. [1310 m.], 17 July 1987, Terhune S. Dickel (1 m~e, USNM). OHIO [STATE RECORDS]: CUYAHOGA CO.: Hunting Y~ley, Orange Twp., 30 July 1943, B. Quay (1, sex unrecorded, UMMZ). PORTAGE CO.: Ravenna, 28 Dec. 1969, G.S. Ensinger (1 larva, crawling on 14" of snow in field; sky sunny, ambient temperature ca. 30 F; larva preserved, MSUE). WAYNE CO.: Wooster, 14 May & 5 July 1962, 23 July 1963,8 Aug. 1960, A.I. Good (5 m~es, CMP). PENNSYLVANIA: WAYNE CO.: South Sterling, 28 July 1918, Ernest Baylis (1 m~e, CMP). SOUTH DAKOTA [STATE RECORDS]: BUFFALO CO.: Fort Thompson, July 1974, J.M. Cicero (3 m~es, 2 fem~es, LACM); Crow Creek, near Fort Thompson, July 1974, J.M. Cicero (2 m~es, LACM). WISCONSIN: recent intensive survey work in this state by Les Ferge, and specimens in LACM, demonstrate that the species is widely distributed in Wisconsin, as expected. New county records represented in these two collections are: BAYFIELD, DOUGLAS, IOWA, JACKSON, KENOSHA, MARA THON,ONEIDA,OZAUKEE,RUSK,SHEBOYGAN,WAUKESHA,and WALWORTH, with dates of 12 May, 20 June, and 13 July-29 Aug. PHRAGMATOBIA ASSIMILANS W~ker, 1855 (fig. 3) Correction to 1966 paper: the caption for fig. 37 on p. 73 was inadvertently omitted; it should read "distribution of P. assimilans; symbols as in Fig. 36." Additional Reference: Ferguson (1975: 10) reports finding mature larvae crawling on the ground in winter in two Nova Scotia localities (HALIFAX CO.: Waverley; HANTS CO.: Mount Uniacke); they pupated without feeding, producing adult moths on 6 March 1949 and 29 March 1951, respectively. New Records: CANADA: NEW BRUNSWICK: KENT CO.: Kouchibouguac Nation~ Park, 1 May 1977, J.D. Lafontaine Code 5191Q (1 male, CNC); same locality, 13 June 1977, J.D. Lafontaine Code 5279A (1 male, CNC). ONTARIO: RENFREW CO.: La Passe, 23, 24, & 31 May 1980, E.W. Rockburne (3 males, CNC). Riotte (1992: 126) reports P. assimilans from the following Ontario counties not previously mapped: FRONTENAC, LEEDS, RAINY RIVER, and SIMCOE. QUEBEC: RIMOUSKI CO.: 5 km NW St. Guy, elev feet. 14 June 1980, John E. Rawlins (6 m~es. CMP).

29 ::g W 50 -i I m Q ;::0 m ~ :; P. assimilans SCALE MIL ~ LAMBERT'S AZIMUTHAL EQUAL-AREA PROJECTION CD \ 40 '"~ m Z a ~ g ~ Figure 3, Distribution of Phragmatobia assimilans; symbols as for Fig, 1. r-.:l '-I II

30 28 THE GREAT LAKES ENTOMOlOGIST Vol. 26, No.1 U.S.A.: IDAHO [STATE RECORD]: BONNER CO.: Priest Lake, T60N, R4W, Sec. 19, elev ft., 30 May 1972, O.B. Howell (1 male, LACM). MICHIGAN: OTSEGO CO.: T29N, R2W, Section 18, 20 March 1966, J.P. Donahue #66-1: 2 larvae presumed to be this species found on tree trunks, one 7 ft (2.1 m) above the r.0und on the south side of a large aspen (Populus tremuloides ; Salicaceae, the other 2.5 ft (0.8 m) above the ground on an elm (Ulmus sp.; Ulmaceae); ambient temperature ca. 35 F. (1.7 C); sky with a thin overcast. The larvae refused food: 1 spun a thin, flimsy cocoon on 22 March 1966, pupated the next day, but died; the other larva was photographed and preserved. (llarva, 1 pupa, LACM.) MINNESOTA: BECKER CO.: Big Twin Lake, "6-4-71" [4 June 1971]. T.L. McCabe (1 male, LACM). MONTANA [STATE RECORD]: RAVALLI CO.: Hamilton, 23 May 1956 ("cold night"), C.B. Philip (1 male, CAS). This is the southern-most locality known for this species in the Rocky Mountains. NEW YORK: McCabe (1990: 9) discusses and illustrates (p. 30) a larva tentatively identified as this species, collected on Apocynum androsaemifolium (Apocynaceae) at Beaver Meadow. HAMILTON CO., 21 Aug. 1977; the specimen died in the pupal stage. Many other larvae were observed wandering at the time. The larva IS described as having rather stiff, uniform, yellowish hairs. McCabe (pers. conun.) reports that P. assimilans is the most abundant Phragmatobia in the Adirondacks, where he has observed thousands of males but no females. PENNSYLVANIA [STATE RECORDS]: BRADFORD CO.: 8 mi. E. of Canton, near Holcomb Pond, 13 May 1980, John E. Rawlins (4 males, CMP). [This locality in northern Pennsylvania is less than 60 miles (96 km] SSW of Ithaca, New York, the nearest locality from which this species has been previously recorded.] SOMERSET CO: Mt. Davis, elevation 885 meters, 10 May 1986, J.E. Rawlins & S. Thompson (6 males, CMP). This is the southernmost record of the species. The moths were taken in hemlock (Tsuga canadensis; Pinaceae) forest just below the highest point in Pennsylvania, flying with the noctuid Feralia comstocki (Grote); the leaves of the deciduous trees at that site had not yet expanded, indicating an early spring phenology (Rawlins, pers. comm.), the characteristic _period for adult activity of this moth. WISCONSIN [STATE RECORDS]: DOOR CO.: T30N, R28E, Sec. 9, 9 June Les Ferge (1 male, LAF). MARATHON CO.: Nine Mile Swamp Area, 14 May 1976, Les Ferge (2 males, LAF). MARINETTE CO.: T37N, R18E, Sec. 26, 23 May 1981, Les Ferge (1 male, LAF). ONEIDA CO.: Lake Katherine, near Hazelhurst, 23 April to 11 June (the preponderance from May), , H.M. Bower (56 males, LACM). VILAS CO.: T43N, R8E, Sec. 31, 10 June 1983, Les Ferge (1 male, LAF). WAUSHARA CO.: Lake Lucerne, 25 May 1975, Wm. E. Sieker (1 male, LAF). To prevent future misunderstanding about the evidence supporting the belief that P. assimilans is a strictly univoltine spring species, it is pertinent to address here the problem raised by several of Bower's specimens from his Oneida Co. locality that appear to bear erroneous. dates. There are three such specimens in LACM (not cited above) from atherine, dated 1 July 1961 (2 males) and 27 July 1961 (1 male). These specimens are part of a pinned, unspread series with labels printed by and appbed at LACM, after the Bower collection was deposited there in I believe that these July dates, applied by hand to the printed data label, represent errors in transcription of the dates originally on the specimens. In a series of 49 specimens labeled at LACM, dates of collection have been inked onto the labels in three different ways: May 27, 1963; V-27-63; and ; Bower himself apparently used the last system exclusively on specimens he labeled himself. I have been unable to find any Bower specimens with temporary labels, but I believe it is

31 1993 THE GREAT LAKES ENTOMOLOGIST 29 safe to assume that they would have been dated similarly. There is also a series of eight males in the USNM from the same locality and collector, with the months of collection transcribed to the Roman numerals VII and VIII (July and August, respectively). If "May" and "June" are substituted, the USNM specimens would have a more reasonable range of dates between 6 May and 2 June 1960, in concordance not only with most of Bower's other specimens, but with the flight season of the species throughout its range (the latest unquestioned record I have seen is 6 July, in northern Maine). Although Bower may have reared some specimens that emerged prematurely, accounting for the summer label dates, if this were the case one would expect to find in his collection at least one parent or reared female, but no Wisconsin females of P. assimilans are known. Sanders (1991: 58) reports an even narrower window of adult flight activity at Black Sturgeon Lake, in northwestern Ontario near Lake Superior: in nine consecutive years of light trapping ( ), this species was only captured between 25 May and 10 June. ACKNOWLEDGMENTS This paper is dedicated to the memory of my friend and companion, John H. Newman - his Welsh humor and knowledge of moths enriched my years at Michigan State University. I thank J.D. Lafontaine and the late C.L. Hogue for their thoughtful comments on an earlier draft of this manuscript, and D.C. Ferguson for similar useful suggestions and valuable bibliographic assistance. I am grateful to the following individuals, institutions, and curators for the opportunity to examine specimens in their care. The collections and the abbre VIations by which they have been cited above are as follows (with the exception of UMMZ and CSU, acronyms agree with those proposed by Heppner and Lamas, 1982): CAS (California Academy of Sciences, P.H. Arnaud, Jr.); CMP (Carnegie Museum of Natural History, J.E. Rawlins); CNC (Canadian National Collection, J.D. Lafontaine); CSU (Colorado State University, Fort Collins); ER (the late Ed Reid); FSCA (Florida State Collection of Arthropods, H.V. Weems, Jr.); LACM (Natural History Museum of Los Angeles County); LAF (Leslie A. Ferge private collection); MSUE (Michigan State University Entomology Museum, the late R.L. Fischer); RSB (Robert S. Bryant); UMMZ (University of Michigan Museum of Zoology); USNM (U.S. NatIOnal Museum of Natural History, D.C. Ferguson). LITERATURE CITED Banks. N A rare pamphlet (Hym. Lep., Neur.'. Entomol. News 31: 176. Bequaert, J A tentative synopsis of the hornets and yellow-jackets (Vespinae; Hymenoptera, of America. Entomologica Americana 12 (n.s.,: figs. Donahue. J.P. and J.H. Newman The genus Phragmatobia in North America, with the description of a new species (Lepidoptera: Arctiidae,. Mich. Entomol. 1(2,: Dyar, H.G Notes on Alaskan Arctiidae. Entomol. News 10: Ferguson, D.C Host records for Lepidoptera reared in eastern North America. U.S. Dept. Agric.. Tech. Bull No p. Ferguson, D.C Contributions toward reclassification of the world genera of the tribe Arctiini, part 1-introduction and a revision of the Neoarctia-Grammia group (Lepidoptera: Arctiidae; Arctiinae). Entomography 3: Franclemont. J.G Arctiidae, In: Hodges. Ronald W., et al., eds., Check List of the

32 30 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 Lepidoptera of America North of Mexico. E.W. Classey Ltd. & Wedge Entomological Research Foundation. pp Heppner, J.B. and Gerardo Lamas Acronyms for world museum collections of insects, with an emphasis on Neotropical Lepidoptera. Bull. Entomol. Soc. Amer. 28(3): Holland, W.J The Moth Book. A popular guide to a knowledge of the moths of North America. New York: Doubleday, Page & Company. xxiv p., 48 col. pis., 263 figs. KOda, N A generic classification of the subfamily Arctiinae of the Palaearctic and Oriental Regions based on the male and female genitalia (Lepidoptera, Arctiidae). Part II. Tyo Ga 39: Krombein, K.V., P.D. Hurd, Jr., D.R. Smith, and B.D. Burks, eds Catalog of Hymenoptera in America north of Mexico. 3 vols. Washington, D.C.: Smithsonian Institution Press. Lindroth, C.H The Faunal Connections between Europe and North America. 344 p. New York: John Wiley & Sons, Inc. McCabe, T.L. [1990] Atlas of Adirondack Caterpillars with a host list, rearing notes and a selected bibliography of works depicting caterpillars. New York State Mus. Bull. 470: iv p., illustr. Miller, C.D.F Taxonomy and distribution of Nearctic Vespula. Canad. Entomol. Supp!. 22: 52 p., 84 figs. Morris, R.F Butterflies and moths of Newfoundland and Labrador. The Macrolepidoptera. Agriculture Canada, PubL p., 34 col. pis., 40 figs. Newman, J.H. and M.C. Nielsen Moth species new to Michigan. Great Lakes EntomoL 6(2): Packard, A.S Notice of Hymenoptera and Nocturnal Lepidoptera, collected in Alaska, by W.H. Dall, Director Sci. Corps, W.U.T. Exp. Trans. Chicago Acad. Sci. 2: 25-32, plate 2. [Expanded title at the beginning of Article II on p. 25: Notice of Hymenoptera and Nocturnal Lepidoptera collected in Alaska by W.H. Dall, Director of the Scientific Corps of the Western Union Telegraph Company. By A.S. Packard, Jr., M.D., with a list of Neuroptera by P.R. Uhler and Dr. H. Hagen.] Riotte, J.C.E Annotated list of Ontario Lepidoptera. Roy. Ontario Mus. Misc. PubL Life Sci.: viii p., 3 figs., 1 table. Rothschild, W Catalogue of the Arctianae [sic] in the Tring Museum, with notes and descriptions of new species. Novit. Zoo!. 17: 1-85, Sanders, C.J List of the Lepidoptera of Black Sturgeon Lake. northwestern Ontario, and dates of adult occurrence. Great Lakes Entomol. 24: Saunders, W Synopsis of Canadian Arctiadae [sic], including some additional species likely to occur in Canada. Canad. Journal (n.s.) 8 (no. 46): [This paper was apparently re-issued in 1863 or 1864 as a separately-paged publication.] Seitz, A Arctiidae, pp In: Seitz, A., ed., , The Macrolepidoptera of the World. 2: The Palearctic Bombyces and Sphinges. 479 p., 56 pis. Alfred Kernen, Stuttgart. Seitz, A Arctiidae, pp In: Seitz, A., ed., , The Macrolepidoptera of the World. 6: The American Bombyces and Sphinges. 1,452 p., 109 pis. Alfred Kernen, Stuttgart. Stephens, J.F Illustrations of British Entomology... (Haustellata) p. London: Baldwin and Cradock. Watson, A., D.S. Fletcher and I.W.B. Nye In: Nye, I.W.B., ed., The Generic Names of Moths of the World. vol. 2. xiv p. London: British Museum (Natural History). Watson, A. and D.T. Goodger Catalogue of the Neotropical Tiger-moths. Oec. Pap. Syst. Entomol. 1: 1-71.

33 1993 THE GREAT LAKES ENTOMOLOGIST 31 APHID PREY OF PA55ALOECU5 CU5PIDATU5 (HYMENOPTERA: SPHECIDAE) John M. Fricke] ABSTRACT Provisioning activity of Passaloecus cuspidatus extended from 29 May through 6 August Eighty trap-nests contained 281 provisioned cells containing 9,618 aphids. The average number of aphids per cell was 34.2 and the average number of cells provisioned per day was Passaloecus cuspidatus stored the following aphids as provisions: Cinaria sp., Euceraphis sp., Macrosiphum euphorbiae, Myzus sp., Myzus cerasi, Myzus monardae, and Sitobium avenae. The number of aphids provisioned per cell was inversely related to aphid size. The number of aphids provisioned per cell varied significantly (9-74). Passaloecus species provision their nests with aphids (Fye 1965 and Krombein 1955, 1956, , 1961, 1967) and a certain Passaloecus sp. may prey on more than six different aphid species (Fye, 1965). Nesting sites include beetle borings in wooden cow shed walls, artificial borings in elderberry and chinaberry, pine trap-nests, and soda straws. Nests are linear and usually partitioned and closed with pine resin. Prey records of Passaloecus cuspidatus Smith provided by Fye (1965), Krombein (1956,1958,1963,1967), and Vincent (1978) include the following aphids: Cinara, Macrosiphum, Masonaphis, Myzus, and Rhopalosiphum. Aphid predators are more closely associated with particular habitats than they are to particular aphids (Dixon 1973). Passaloecus are flexible in their prey selection. The number of prey provisioned by Passaloecus spp. is extremely varied. Prey provisioned per cell ranged from 7 to 63; prey per nest ranged from 50 to 200. Corbet and Backhouse (1975) suggested that, in favorable conditions, a single female could capture 1500 aphids based upon one cell provisioned per day (30 aphids) and a wasp lifespan of 50 days. The mean number of aphids provisioned per cell has not been reported in the literature. Neither are there reports on whether the species of aphid provisioned influences the number of aphids provisioned. METHODS AND MATERIALS Passloecus cuspidatus Smith was selected for investigation of aphid provisioning because its large size, markings and behavior made field identification possible. P. cuspidatus rings nest openings with resin prior to provision- INatural Science and Mathematics Division, Concordia College, Ann Arbor, MI

34 32 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 Table 1. Aphids provisioned by Passaloecus cuspidatus, Mean Number of Number of Trap-nests Cells /I of Aphids ± S.D ±11.8 sp ± 7.6 Macrosiphum euphorbiae ± 8.2 Euceraphis sp., Myzys sp ± 7.7 Myzus cerasi ± 17.5 Sitobium avenae ± 9.0 ing, facilitating field observations. The study site was located on the campus of Concordia College, Ann Arbor, Michigan during the summer of Paraffin coated, pre-split trap-nests of seven bore diameters ( mm, with 0.8 mm increments) were provided as nesting material. The available frequencies of bore diameters were as follows: 2.4 mm - 211; 3.2 mm - 307; 4.0 mm-307; 4.8 mm-307; 5.6 mm-211; 6.4 mm-211 and 7.2 mm-21i. Trap-nests were removed from the field within one to two days of closure and were replaced with trap-nests of similar bore diameter. Closed trapnests were opened and the contents of each cell were removed, aphids counted and, with the wasp egg or larva, were transferred to a rearing vial. Two aphids from each provisioned cell were retained for identification. Passaloecus cuspidatus was presumably active in the study area prior to site establishment on 29 May Five trap-nests were ringed with resin as of 1 June Trap-nest provisioning was observed on 1 June (4 trips, 2:39 pm-3:20 pm); 3 June (7 trips, 12:31 pm-l:35 pm); 18 June (16 trips, 10:17 am-ll:43 am); and 5 July (8 trips, 4:31 pm to 5:13 pm). Nesting activities continued through the first week of August with no trapnests ringed or closed with resin after 6 August Observations on 8, 14, and 21 August 1987 provided no evidence of additional activity. Passaloecus cuspidatus used trapnests at the following frequencies: 2.4 mm-9; 3.2 mm-37; 4.0 mm-28; 4.8 mm-8; 5.6 mm-l, and 6.4 mm-i. The minimum provisioning period for this wasp population was seventy days. This corresponds well with the range of flight dates (14 June-14 August) for P. cuspidatus material from the Museum of Zoology of the University of Michigan and the Entomology Museum of Michigan State University, and is supported by previous field observations of this species. In 1984, provisioning activity was first observed between 4 and 12 June and terminated between 4 and 13 August. In 1985, emer~ence of P. cuspidatus from a natural nest occurred 10 May and nesting activity ceased 20 August. RESULTS AND DISCUSSION Passaloecus cuspidatus provisioned its cells with Cinaria sp., Euceraphis sp., Macrosiphum euphorbiae (Thomas), Myzus sp., Myzus cerasi (Fabricius), Myzus monardae (Davis), and Sitobium avenae Fabricius (Table 1). Multiple t(ii) tests for differences in number of aphids provisioned were significant for Myzus monardae, Cinaria sp., and Macrosiphum euphorbiae, [Myzus monardae and Cinaria sp., t(li) = , df = 193, p <.001; Cinaria sp. and Macrosiphum euphorbiae, t(ll) , df ::::: 72, p <.001; and Myzus monardae and Macrosiphum euphorbiae, t(ll) = , df = 197,p <.001]. Differences in number of aphids provisioned per cell were inversely related to

35 1993 THE GREAT LAKES ENTOMOLOGIST 33 Table 2. Aphids provisioned by Passaloecus cuspidatus during Summer Provisioning period is divided into 14 5-day intervals beginning 24 May and ending 11 Aug. Upper # is number of aphids; lower # is number of cells. Aphids Ending Dates For 14 Consecutive 5-day Provisioning Periods May June July August Myzus monardae Cinaria sp Macrosiphum euplwrbiae Euceraphis sp and Myzus sp. 6 2 Myzus cerasi Sitobium aveluui 1 5 undetermined aphids aphid size, with Myzus monardae the smallest and Macrosiphum euphorbiae the largest. Two trap-nests had an extraordinarily high number of provisions. One contained 430 aphids (eight cells, aphids per cell) and another contained 334 aphids (five cells, aphids per cell). Unfortunately, aphid samples for identification were not taken from these trap-nests. Table 2 summmarizes data on the seasonal changes in aphid provisioning by P. cuspidatus and relative numbers of aphids provisioned. Passloecus cuspidatus was not restricted to a particular aphid species. Peaks in provisioning rates are assumed to be related to increased aphid numbers. Between 29 May and 7 June, Cinara sp., Euceraphis sp., Myzus sp., and Sitobion avenae Fabricus were used as provisions. From 8 June through 27 July Myzus monardae (Davis) was the preferred prey while Macrosiphum euphorbiae (Thomas) and Myzus cerasi (Fabricius) were provisioned in significant numbers between 28 June and 17 July. Provisioning rates were varied and estimated on the basis of number of cells provisioned and dates of bore ringing and closure. Trap-nests with estimated provisioning periods of one to three days contained from one to ten provisioned cells. Trap-nests with estimated provisioning periods of six to eight days contained two to eight provisioned cells. Trap-nests with provisioning periods of 10 to 21 days contained one to five J:>rovisioned cells. The estimated number of cells provisioned per day ranged from 0 to 5. Several variables could influence provisioning rates. Exceptionally high provisioning rates could be the result of close proximity of aphids, closure materials and the nesting site. Aphids and closure materials somewhat distant from the nesting site could result in low provisioning rates. A possible cause for low provisioning rates is a temporary cessation of provisioning activity. This cessation mif5ht be necessary to allow the development of additional ova following a period of provisioning and oviposition. Other factors including inclement weather, low aphid population levels and disruptive

36 34 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 behavior of parasitoids could influence provisioning rates. Cells (281) were provisioned during a cumulative provisioning period of 381 days. The average provisioning rate per trapnest was 0.73 cells per day. Trap-nests used less than four days had a higher than average provisioning rate and trap-nests used longer than four days had lower than average provisioning rates. ACKNOWLEDGMENTS Aphid specimens were identified by Dr. Manya Stoetzel, USDA. LITERATURE CITED Corbet, S. A., and M. Backhouse Aphid hunting wasps: a field study of Passaloecus. Trans. Roy. Entomol. Soc. London. 127: Dixon, A. F. G Biology of Aphids. The Institute of Biology's Studies in Biology no. 44, 58 pp. Edward Arnold (Publishers) Limited. London. Fricke, J. M Trap-nest bore diameter preferences among sympatric Passaloecus spp. (Hymenoptera: Sphecidae). Great Lakes. Entomol. 24: Fye, R. E The biology of the Vespidae, Pompilidae and Sphecidae from trapnests in northwestern Ontario. Canad. Entomoi. 97: Krombein, K. V Miscellaneous prey records of solitary wasps. I. Bull. Brooklyn Entomol. Soc. 50: Miscellaneous prey records of solitary wasps. II. Bull. Brooklyn EntomoL Soc. 51: Miscellaneous prey records of solitary wasps. III. Proc. BioI. Soc. Washington. 71: Biological notes on some Hymenoptera that nest in sumach pith. Entomoi. News. 71:29-36, _~ Miscellaneous prey records of solitary wasps. IV. Bull. Brooklyn Entomoi. Soc. 56: Trap-nesting Wasps and Bees: Life Histories, Nests, and Associates, ill-vi pp. Smithsonian Press, Washington D.C. Vincent, D. L A revision of the genus Passaloecus (Hymenoptera: Sphecidae) in America north of Mexico. Wasmann J. BioI. 36:

37 1993 THE GREAT LAKES ENTOMOlOGIST 35 DYTISCIDAE AND NOTERIDAE OF WISCONSIN (COLEOPTERA). II. DISTRIBUTION, HABITAT, LIFE CYCLE, AND IDENTIFICATION OF SPECIES OF DYTISCINAEl William L. Hilsenhoff2 ABSTRACT Twenty-one species of Dytiscinae were collected in Wisconsin over the past 30 years, including three species of Acilius, one species of Cybister, eight species of Dytiscus, five species of Graphoderus, two species of Hydaticus, and two species of Thermonectus. Species keys are provided for adults, and except for Dytiscus and Cybister, keys are also provided for larvae. Based on a study of 13,236 adults and 854 larvae, information on the distribution and abundance of each species in Wisconsin is provided along with notes on their habitat, life cycle, and identification. Six genera and 21 species of Dytiscinae were collected in Wisconsin. These include the largest species of Dytiscidae, with adults ranging from nine to 42 mm in length. Because they are large, adults swim rapidly and are difficult to capture with a net. I collected most adult Dytiscinae with bottle traps, and while larvae were more readily captured with a net than adults, they too, were effectively captured with bottle traps (Hilsenhoff 1991). Collecting efforts, measurement of specimens, and general information about life cycles are summarized in Part I of this study (Hilsenhoff 1992). Part I also contains a generic key to adults and a map of Wisconsin with numbered counties that are grouped into nine areas; this map is often referred to below. Based on adults and 853 larvae that could be identified to species, the general distribution and relative abundance of species of Dytiscinae is summarized in Table 1 for the nine areas of Wisconsin. Totals for collections that resulted from intensive studies in McKenna Pond (Hilsenhoff 1992) and the Leopold Memorial Reserve (Sauk County) are included separately in this table because of the large number of beetles that were collected. The latter includes monthly (March-October) net and trap collections from 16 ponds between May 1989 and May 1992, which are part of an ecological study of pond insects by Leonard Huebner. Dytiscidae collected by Huebner were not included in part I of this study (Hilsenhoff 1992), hut will be included in this and suhsequent parts. The 658 adult Dytiscinae that were collected with bottle traps from Horicon Marsh by Kevin Kenow during the summer months from 1983 through 1985 are reflected in totals for the south-central area (Table 1); most were Graphoderus and Hydaticus. He did not collect heetles during other months and no larvae were saved from his collections. 1Research supported by the College of Agricultural and Life Sciences and the Grad?,ate School at the University of Wisconsin-Madison. Department of Entomology, University of Wisconsin. Madison, WI

38 36 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 Table 1. Numbers of Dytiscinae adults (A) and larvae (L) from nine areas of Wisconsin (Hilsenhoff 1992), McKenna Pond (McK), and Leopold Memorial Reserve (LMR) collected between 1962 and NW Ne NE we e EC SW se SE MeK LMR TOTAL Acilius mediatus A A. mediatus L A. semisulcatus A A semisulcatus L A. sylvanus A A. sylvanus L Cybister fimbriolatus A C. fimbriolatus L Dytiscus alaskan us A D. carolinus A D. cordieri A D. dauricus A D. fasciventris A D. harrisii A D. hybridus A D. uerticalis A Graphodems fascicollis A G. libems A G. liberos L G. manitobensis A G. manitobensis L G. occidentajis A G. perplexus A G. perplexus L Hydaticus aruspex A H aruspex L H piceus A H piceus L Themwnectus basilj.o:ris A T. omaticollis A T.omaticollisL Below, under each genus, is a key to adults of species that occur or may occur in Wisconsin; a size-range for adults from Wisconsin is included for each species. A key to third instar larvae (larvae having lateral spiracles) is included, except for Cybister and Dytiscus. Usually second instar larvae can also be identified with these keys, but first instar larvae are structurally very different. Following the keys is information on distribution and abundance in Wisconsin, general range in North America, habitat, and identification. County records are represented by numbers on the map in part I (Hilsenhoff 1992); records based only on larvae are marked with an asterisk. Acilius Leach, 1817 Four species occur in northeastern North America (Hilsenhoff 1975); only three were found in Wisconsin as a result of this study. It is possible that Acilius {raternus fraternus may also occur in Wisconsin, since it was reported by wallis from four localities in Minnesota (Larson, 1973); however, none of these specimens could be located to verify Wallis' identifications. A male adult A. fraternus fraternus in the University of Wisconsin Insect Research Collection is labeled only "Wis." and "Collection of W.S. Marshall". This specimen, which was probably collected before 1925, may not have been collected in

39 1993 THE GREAT LAKES ENTOMOLOGIST 37 Wisconsin. Larvae of species collected from Wisconsin were identified by using available descriptions and circumstantial evidence such as relative abundance of larvae and adults. their distribution, and especially collections of adults and larvae from the same site. Inclusion of the larva of A. fratemus fratemus is based upon the description and figures of Wolfe (1980). All species normally have a univoltine life cycle in Wisconsin. Adults overwinter in deeper ponds and margins of lakes and streams, but often fly to a wide variety of other habitats in early spring where they apparently mate and oviposit in late March and April, and then die. Most larvae develop in May (first and second instars) and June (third instar), and pupate in June. Most adults emerge in June and early July. Key to Species of Adult AciJjus in Wisconsin 1. Testaceous ventrally; mm long....sylvanus Black, or mostly black ventrally...2 2(1). Smaller, mm long; metatibia and tarsus black; males without tufts of hairs on inner ventral margin of 3 basal mesotarsal segments; females without elytral sulci and with a distinct M-mark on dorsum of head....mediatus Larger, >12.5 mm long; metatibia and tarsus testaceous to rufous; males with tufts of golden hairs on inner ventral margin of 3 basal mesotarsal segments; females with elytral sulci or without a distinct M-mark on head...3 3(2). Metafemur testaceous with a small basal infuscation; second visible abdominal sternum usually with a pale lateral spot on each side; anterior pro- and mesotarsal claws of male longer and thicker than.rosterior claws; female elytra sulcate, longest sulci extending to basal seventh of elytra; mm long....semisulcatus Metafemur rufous to piceous, lighter apically; second visible abdominal sternum without pale lateral spots; pro- and mesotarsal claws of male not modified; female elytra, if sulcate, with longest sulci extending only to basal fourth of elytra; mm long (Indiana specimens)....fraternus fraternus Key to Species of Larval AciNus in Wisconsin 1. Anterior of frontoclypeus not darker than posterior; ligula of labium forked before basal third or past apical third, rarely near middle..2 Anterior of frontoclypeus darker than posterior; ligula of labium forked near middle...3 2(1). Ligula of labium usually forked past apical third, apical spines inconspicuous and about as wide as long....semisulcatus Ligula of labium forked before basal third. with a pair of distinct, elongate spines at apex of each fork....sylvanus 3(1). Ligula short and stout, width of stem 3/4 length past fork mediatus Ligula elongate, width of stem < 112 length past fork fraternus fraternus

40 38 THE GREAT lakes ENTOMOlOGIST Vol. 26, No.1 AciHus fraternus fraternus (Harris, 1828) Distribution and Abundance: Range: IA-MA-GA-AR. A. fraternus disma Ius Matta and Michael, 1976 occurs farther south. Habitat: Michael and Matta (1977) most often collected this species in shaded ponds and pools with some leaf litter. Life Cycle: Unknown, but likely univoltine in the northern part of its range. Identification: Adults are similar to A. semisulcatus, but are readily separated from that species by characters in the key. The M -mark on the head is very obscure; often it cannot be seen. The larva of A. f fraternus, as described by Wolfe (1980), is similar to the larva of A. mediatus because the ligula is forked near the middle. However, the ligula in Wolfe's figure is more slender than in A. mediatus and the frontoclypeus of A. f fraternus does not have the distinct, dark, anterior rectangle of A. mediatus. AciliuB mediatub (Say, 1823) Distribution and Abundance: Uncommon statewide (Table 1). County records: 10, 12-13, 15, 18*, 19, 30-31, 57. Range: MN-NB-NC-MO. Habitat: All adults and larvae were collected from swamps or small, softwater ponds in areas above precambrian formations. Life Cycle: A second ins tar larva was collected in early June; a third instar larva and a teneral adult were collected in late June; a slightly teneral adult was found August 10 in the extreme north. Most adults were collected in April, or from July through October. This indicates a univoltine life cycle as described above for the genus. Identification: Adults can be recognized in the field by their smaller size, dark coloration, and bold black and yellow fasciae across the apical third of the elytra. The second instar larva was collected from a site where 15 adults were collected on six dates and no other species of Acilius has been found. The ligula and markings on the head differed markedly from other Acilius larvae that I collected in Wisconsin; therefor I conclude it must be the larva of A. mediatus. Subsequently a third instar larva was collected from a Sphagnum habitat, which is typical for adults of this species. The head capsule width of the third instar larva was less than 0.5 mm, and narrower than in third instar larvae of A. semisukatus and A. sylvanus. Acilius semisulcatus Aube, 1838 Distribution and Abundance: Very common in northern two-thirds, common in southern third (Table 1). County records: 1-27, 29-61, Range: AK-NF-NJ-IL-AB Habitat: Adults and larvae were collected most frequently from permanent ponds, but they also occurred in deeper marshes, margins of swamps, and vernal ponds and marshes. Life Cycle: Most adults were collected in March and April. and from late June to November; they were uncommon in May and early June. Second instar larvae were found from May 7 to June 27, third instar larvae from May 5 to July 24. and teneral adults from June 1 to August 21. Two larvae collected August from cold northern swamps probably resulted from delayed development. The univoltine life cycle described for the genus is typical for this species. Two larvae were collected from a warm central-wisconsin marsh in mid-august and two more from a flooded field in October; teneral adults were also collected in October. These records probably represent a partial second generation, which may occur in some years. Identification: The testaceous metafemora with a small basal infuscation

41 1993 THE GREAT LAKES ENTOMOlOGIST 39 immediately distinguishes adults from other species with dark venters. Larvae are readily identified by the elongate ligula, which usually is branched between the apical third and apical sixth. I have two larvae from northern Wisconsin with their ligula branched near the middle, but their heads lack the dark anterior marks found in A. mediatus and A. f fraternus and I believe they are A. semisulcatus. The brief larval description by Watts (1970) describes this species and not A. sylvanus, which in 1970 was considered to be a color variant of A. semisulcatus. Wilson's (1923) larval description is erroneous; he describes a Graphoderus larva. AciJius sylvanus Hilsenhoff, 1975 Distribution and Abundance: Common statewide (Table 1). County records: 2-6, 8-11, 13, 16-25, 27-44, 45*, 46-52, 54-63, 66, Range: MB-PQ-ME-NJ-IL Habitat: The habitat is similar to that of A. semisulcatus, except adults and larvae were infrequently found in acid waters. Often adults of A. semisulcatus and A. sylvanus occurred together. Life Cycle: Second instar larvae were found from May 11 to July 13, third instar larvae from May 8 to July 13, and teneral adults from June 21 to August 10. Almost no adults were collected from late April to mid-june. This suggests a life cycle like that described for the genus. Identification: The testaceous ventral surface of adults and lack of a distinct sub-apical fascia on the elytra are distinctive. The elongate branches of the ligula with distinct apical and sub-apical spines readily separate larvae from other species. Cybister Curtis, 1827 A single species of this southern genus occurs in the northern United States and Canada. Cybister fimbriolstus (Say, 1823) Distribution and Abundance: Uncommon in central and southern Wisconsin (Table 1). County Records: 26, 46, 48*, 51, 57-58, 60-61, 67. Range: MB NS-FL-TX. Habitat: Larvae and adults inhabit deeper water of open permanent ponds. Life Cycle: Adults were collected from April to November. Second instar larvae were found from June 5 to 21, third instars from June 23 to July 15, and teneral adults from August 17 to September 24. Adults overwinter, probably in deeper ponds, and oviposit in spring. Larvae develop in late spring ana summer, and apparently pupate from late July into September. Adults emerge in late summer and early autumn to complete a univoltine life cycle. Identification: Adults resemble small Dytiscus verticalis or large D. h'lbridus, but are distinctly widened at the apical third and have numerous tilly, green spots on the elytra. They lack a pale sub-apical fascia on the elytra. which occurs in D. verticalis, and almost always have a green prosternal process. The protarsal disc of males is angulate on the anterior margin, giving It a triangular shape when not expanded. Females have elongate, longitudinal aciculations laterally on the elytra. Larvae resemble those of Dytiscus because second and third instars are large and have a setal fringe on the last two abdominal segments. They are readily identified, however, because their urogomphi are vestigial and they possess long teeth on the labroclypeus. Wilson (1923) described the larva.

42 40 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 IJytiscusLinnaeus, 1758 R ey (1990) revised the world fauna, providing a wealth of information 0 fication, distribution, habitat, life history, and phylogeny. Part I of his revision deals only with adults; part II, which has not yet been published, will contain keys and descriptions of larvae. I was unable to identify larvae of some species, so my study ofijytiscus in Wisconsin is based only on adults. Eight species were collected and are keyed below along withd. circumcinctus, which also may occur in Wisconsin. Because they are large, adults were measured with a caliper instead of an ocular micrometer. The length of Wisconsin specimens was generally longer than reported by Roughley (1990). Aquarium and field observations, along with collection records and date of occurrence of teneral specimens, indicate all species have a univoltine life cycle similar to that reported by Aiken and Wilkinson (1985) for D. alaskanus. Adults of Dytiscus overwinter in a variety of deeper aquatic habitats and disperse in spring as soon as these habitats become free of ice and air temperatures are sufficiently warm to permit flight. Adults mate very early in the spring and oviposit when their breeding habitat becomes free of ice, which may be from late March to early May; ding on habitat and latitude. Most adults apparently die after mating oviposition, but some survive well into summer. Eggs of most species probably hatch from late April to late May, depending on water temperatures and date of oviposition. Larval development is completed from early June to early August, with much variation among and between species. At least 60% of adult Dytiscus were collected in March and April with bottle traps, often from overwintering habitats still partially covered with ice. The remaining adults were collected mostly from June through August; few were collected in May and after September. The scarcity of adults after September is probably due to movement into deeper water to overwinter and decreased activity, which makes them less vulnerable to trapping. Key to Species of Adult Dytiscus in Wisconsin 1. Metacoxal process rounded, never spinose; venter pale or dark...2 Metacoxal process distinctly spinose; venter mostly pale...7 2(1). Abdominal sterna testaceous; mm long....cordieri Abdominal sterna rufous, piceous, or with conspicuous piceous fasciae (2). Larger, > 33 mm long; elytra with a sub-apical pale fascia....4 Smaller, < 29 mm long; elytra without a sub-apical pale fascia...5 4(3). Metacoxal plate and first visible abdominal sternum dark rufous, same color as rest of metasternum and venter; posterior margin of pronotum without a pale band; mm long....verticalis Anterior of metacoxal plate and first visible abdominal sternum testaceous, much lighter than rest of metasternum and venter; posterior margin of pronotum with a pale band; mm long...harrlsii 5(3). Mesotarsus of male with median glabrous area dividing suckers; female without elytral sulci; venter dark rufous, without black fasciae; mm long....hybridus Mesotarsus of male without glabrous area dividing suckers; female with elytral sulci; venter with black fasciae...6 6(5). Metacoxal plate and first visible abdominal sternum pale, much lighter than infuscate mid-metasternum; anterior of pronotum with pale border; mm long....fasciventris Metacoxal plates and first visible abdominal sternum dark, similar in

43 1993 THE GREAT LAKES ENTOMOLOGIST 41 color to metasternum; anterior of pronotum without a pale border; mm long... "..carolinus 7(1). Head narrowly pale along inner margin of eye; piceous marks on basal abdominal sterna absent or very narrow basal lines; mm long (Roughley 1990)....circumcinctus Head not pale along inner margin of eye; piceous marks on basal abdominal sterna usually expanded posteriorly...8 8(7). Larger, mm long; basolateral piceous marks extend at least half distance to posterior margin on visible abdominal sterna 2 and 3....dauricus Smaller, mm long; basolateral piceous marks usually do not extend half distance to posterior margin on visible abdominal sterna 2 and 3....alaskanus Dytiscus alaskanus J. Balfour-Browne, 1944 Distribution and Abundance: Very rare in west-central area (Table 1). County records: 25, 29. Range: AK-NF-NH-MN-WA+WY+CO. Habitat: Wisconsin sl!ecimens were collected from emergent vegetation in large permanent ponds. This species is more abundant west and north of Wisconsin, where it inhabits permanent ponds and lakes (Roughley 1990). Life Cycle: Aiken and Wilkinson (1985) studied the bionomics in northcentral Alberta. They reported a univoltine life cycle with overwintering adults, mating in April, and oviposition as soon as the lake was free of ice. First instar larvae appeared.from mid-to late May, and peak numbers of third instar larvae occurred throughout July. In Wisconsin this sequence probably occurs earlier because of a warmer climate. Identification: Adults could be confused with D. dauricus, but in Wisconsin they are distinctly smaller. In males, the penis (median lobe of aedeagus) in dorsal view is broad and evenly tapered to the apex (Fig. 1), while in D. dauricus (Fig. 2) and D. circumcinctus it is narrower and subapically sinuate to form an apical knob. In sulcate females. ridges on elytral intervals 8 and 9 meet to form a "V". These ridges do not meet in D. dauricus or D. circumcinctus. The size of black basolateral marks on visible abdominal sterna 2 and 3 varies in the three Wisconsin specimens from narrow to expanded almost halfway to the posterior margin, but the marks are distinct on the last visible sternum in all specimens. Dytiscus circumcinctus adults lack a black mark on the last visible abdominal sternum. Dytiscus carolinus Aube 1838 Distribution and Abundance: Rare in southern third (Table 1). County records: 52, 55, 57, 60, 72. Range: WI-MA-GA-AR. Habitat: All except one adult were collected from ponds and sloughs associated with large rivers. Life Cycle: Six of the seven collections were overwintering adults collected from March 17 to April 4. The seventh collection contained four adults, which were trapped June in LaCrosse County; two were teneral, which indicates oviposition in early spring and completion of larval development by mid June. Identification: Adults are most similar to D. fasciventris, but can be separated from that species by characters in the key. Abdominal sterna are rufous with black fasciae; in D. fasciventris they are usually testaceous with black fasciae and when darker the first abdominal sternum and the metacoxal plate are always paler.

44 42 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 2 Figures Apical 2 mm of penis (dorsal view). 1. Dytiscus alaskanus. 2. D. dauricus. Dytiscus circumcinctus Ahrens, 1811 Distribution and Abundance: Not yet found in Wisconsin. Range: AK-PQ ME-NC-IA-ND-WA. Habitat: Roughley (1990) reported that adults in western North America were found in a wide range of aquatic habitats. Life Cycle: The life cycle is al.lparently univoltine, with adults overwintering in both terrestrial and aquatic habitats (Rou~hley 1990). Identification: The lack of black abdolillnal markings, except for extremely narrow basal lines on visible abdominal sterna 1-3 (Roughley 1990), separates this species from D. alaskan us and D. dauricus in Wisconsin. Dytiscus cordieri Aube, 1838 Distribution and Abundance: Fairly common statewide (Table 1). County records: 3-6,8,10,12,14-15,17,19-25,33-35,37-39,41-48, 50-51, 57-59, 61, 64, 67-68, Range: AK-NB-NC-CO-CA. Habitat: Adults were collected from open ponds and marshes. Life Cycle: The life cycle is typical of that described above for the genus. It is completed somewhat earlier than in most other species, probably due to warmer water in its open breeding habitat. Teneral adults, along with many mature adults, were collected from June 23 to August 2. Identification: This is the only Dytiscus in Wisconsin in which adults are almost entirely testaceous ventrally. They lack the spinose metacoxallobes of other medium to large Dytiscus that are mostly testaceous ventrally. Dytiscus dauricus Gebler, 1832 Distribution and Abundance: Uncommon in northern third (Table 1). County records: 1-2, 4-5, 9-13,15,17-18, Range: AK-LB-NH-SD-CO AZ-CA.

45 1993 THE GREAT LAKES ENTOMOlOGIST 43 Habitat: Adults were collected from bogs, swamps, and a variety of ponds. Life Cycle: Most adults were captured in traps in April and August; the remainder (five) were collected in June and July. Two teneral adults were found in August, indicating a typical univoltine Dytiscus life cycle. Identification: The very large size readily separates adults from those of D. alaskanus and most D. circumcinctus. All Wisconsin adults had basolateral marks on visible abdominal sterna 2 and 3 extending posteriorly to the middle of the segment or beyond, which would separate them from D. circumcinctus and most D. alaskanus. Other differences are discussed under D. alaskanus. Dytiscus fasciventris Say, 1824 Distribution and Abundance: Common statewide (Table 1). County records: 2-6, 9-27, 29-42, 44-52, 54-61, 64, 66-68, Range: YK-LB-NJ IL-MN-MT-BC. Habitat: Adults were collected from a variety of ponds and marshes, especially those with sedges (Carex, Eleochans). Life Cycle: The univoltine life cycle is typical of Dytiscus. Teneral adults were trapped from June 6 to August 10, especially from late June through July. Identification: The combination of small size and testaceous venter with black basal fasciae on each segment is distinctive. Dytiscus harrisii Kirby, 1837 Distribution and Abundance: Uncommon in northern two-thirds, rare in southern third (Table 1). County records: 2-13,15-17,20,27,29-39,42,44,46, 48, 59, 61, 65. Range: AK-NF-PA-NE-ND-BC. Habitat: Most adults were collected from a variety of ponds and marshes; two were collected from swamps, and two from the margin of rivers in October. All teneral adults (five from four sites) were trapped in sedge marshes or shallow ponds with sedge margins; I believe this is the larval habitat. Other than the two overwintering adults collected in October, I did not notice an association with streams as mentioned by Roughley (1990). Life Cycle: Teneral adults were collected in central Wisconsin between June 23 and July 2, accounting for most adults collected in June and July. Because of the shallow, warm habitat in which larvae apparently develop, they complete development earlier than most other Dytiscus. Identification: Because of their large size, a subapical pale fascia on the elytra, and a mostly dark venter, adults can be confused only with D. verticalis. The pale first visible abdominal sternum, light mark on the anterior of each metacoxal plate, and broadly pale basal band on the pronotum easily separate adult D. harnsii from that species. Dytiscus hybridus Aube Distribution and Abundance: Common in southern two-thirds, fairly common in northern third. County records: 1, 3-6, 8, 12-16, 20, 25-26, 28-32, 34-41,44-58, Range: AB-NB-SC-CO+OR Habitat: Overwintering adults were collected from a variety of ponds and deeper marshes, but most teneral adults were found in shallow cattail (Typha) marshes, which is likely their breeding habitat. Life Cycle: The life cycle is typical of Dytiscus, with teneral adults being collected between June 23 and August 1.

46 44 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 Identification: The small size and uniformly dark venter of adults is a distinctive combination. Dytiseus vertiealis Say, 1823 Distribution and Abundance: Common statewide, especially so in north west. County records: 1-23, 25-27, 29-40,44-51,53-55,57-61,65-68, Range: MB-PE-VA-IN-MN Habitat: Adults were collected from a variety of ponds, and less frequently from marshes and bogs. Most teneral adults were collected from shallow ponds with cattails along the margin; one was found in a sedge marsh. Life Cycle: The relatively large number of adults collected in June suggests that overwintering individuals live longer than most other species and may continue to oviposit in late spring. Larval development was apparently later than in other species. All teneral adults were collected July 13 to August 20, except for one trapped in a sedge marsh on July 2. Identification: The dark venter, large size, and sub-apical pale fascia on the elytra separate adults from all other dytiscids, except D. harnsii. Their separation is discussed under D. harrisii. Graphoderus Dejean, 1833 All five North American species were collected. Wallis (1939) provided a key to male adults of North American species, which he concluded were distinct from species found in Europe. but he did not delineate their distribution. Lacking a recent revision, ranges of species within the United States are inadequately defined. Both adults and larvae of G. liberus differ significantly in many respects from the other four species, which are similar to one another. Male adults of these four species can be readily identified by the number of palettes on the meso tarsi; a table for separation of females was prepared by Tracy and Hilsenhoff (1982). The distinctive larva of G. liberus was described by Barman (1972) and that of G. perplexus (as zonatus) by Watts (1970), but the latter description was not helpful in separating G. perplex us larvae from those of the three similar species, which are undescribed. A study of Wisconsin larvae revealed characters by which third instar larvae of most species can be separated, but I was unable to separate larvae of G. fascicollis and G. occidentalis. A provisional key to larvae is provided below. Larvae (except G. liberus) were associated with adults by circumstantial evidence such as relative abundance, distribution within Wisconsin, date of occurrence, and presence of larvae and adults in the same site. Adults of G. fascicollis are most similar to those of G. occidentalis, while adults of G. manitobensis and G. perplexus are similar to each other. These relationships apparently also hold for the larvae. Numerous adults (359) of all species except G. liberus were collected in bottle traps from Horicon Marsh by Kevin Kenow from June tbrough August, accounting for unusually high totals in the south-central area (Table 1). Although all species of Graphoderus are predominantly univoltine, with adults overwintering in aquatic habitats. substantial differences in life cycles seem to exist between species. Overwintering adults of G. fascicollis and G. occidentalis apparently do not become active until later in the spring then G. manitobensis and G. perplexus, and thus oviposition and larval development is delayed. All of the species may have a partial second generation in some years.

47 1993 THE GREAT LAKES ENTOMOLOGIST 45 Key to Species of Adult Graphoderus in Wisconsin 1. Head and pronotum without distinct black markings; smaller, mm long....liberus Head and pronotum with distinct black markings; larger, > 12.7 mm long...2 2(1). Width of testaceous area of metasternal wing 0.40 mm or less at narrowest point adjacent to mesocoxa; male mesotarsus with palettes, protarsus with palettes...3 Width of testaceous area of metasternal wing 0.48 mm or more at narrowest point adjacent to mesocoxa; male mesotarsus with 0 or 12 palettes, protarsus with palettes (2). Anterior and posterior pronotal black bands separated from margin by testaceous bands, the basal testaceous band being narrow and occasionally absent mesally; male mesotarsus with palettes; female pronotum with moderate corrugated macrosculpture laterally (Fig. 3); mm long... '"....perplexus Anterior and posterior pronotal black bands not separated from margin by a testaceous band; male mesotarsus with palettes; female pronotum with pronounced corrugated macrosculpture laterally (Fig. 4); mm long...,..manitobensis 4(2). Anterior black band on pronotum separated from margin by rufopiceous band; mesotarsus of male dilated, with 12 palettes in 2 rows; pronotum of female with moderate corrugated macrosculpture laterally (Fig. 3); mm long...,....fascicollis Anterior black band on pronotum not separated from margin; mesotarsus of male not dilated, and without palettes; pronotum of female with lateral corrugated macrosculpture indistinct (Fig. 5); mm long....occidentalis Key to Species of Larval Graphoderus in Wisconsin 1. Head without temporal spines; thoracic and abdominal terga with a longitudinal pale strife bordered by dark stripes....liberus Head with a row temporal spines; thoracic and abdominal terga without longitudinal stripes...2 2(1). Labium parallel-sided in basal half; length of spines on ligula subequal to width of ligula at mid-length; anterior margin of frontociypeus narrowly black...3 Sides of labium diverging from base; length of spines on ligula > times width of ligula at mid-length; anterior margin of frontociypeus brown or fuscous....fascicollis or occi!kntalis 3(2). Inner margin of stipes rapidly widened in apical half, with apical fourth much wider and slightly rounded before apex (Fig. 6)....perplexus Inner margin of stipes gradually widened in apical half, with apical fourth slightly widened and not rounded (Fig. 7)....manitobensis Graphoderus fascicollis (Harris, 1828) Distribution and Abundance: Fairly common statewide (Table 1). County records: 3-7, 9, 12, 15, 18-20, 25, 29,34,37-38,40,46-48, 50-51, 57-59, 61, 67-70, 72. Range: MN-PQ-ME-IN. Habitat: Most adults were collected from small ponds containing cattails along at least one margin. Almost half of all adults were collected from Horicon Marsh, a very large marsh with numerous stands of cattails.

48 46 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 Figures SEM of lateral fourth of female pronotum. 3. Graphoderus perplexus. 4. G. manitobensis. 5. G. occidentalis. 6 Figures Right stipes of larva (ventral view). 6. Graphoderus perplexus. 7. G. manitobensis. 8. G. fascicollis.

49 1993 THE GREAT LAKES ENTOMOlOGIST 47 Life Cycle: Almost all third instar larvae believed to be G. fascicollis or G. occidentalis were collected June 7 to July 20; teneral adults were found June 25 to August 25, three-fourths of them in July. While adults were collected from March to October, 77% were found from June through August (excludin15 summer only collections from Horicon Marsh). I believe that most overwmtering adults do not mate and oviposit until late Mayor June, with larval development being completed mostly in late June and July. One third instar larva that I believe is this species was collected October 5 in a flooded area; it probably represents a partial second generation. Identification: Adults can be readily identified by the key. The area separating the anterior pronotal black band from the margin rarely may be rufotestaceous, but the posterior band is never separated from the margin as in G. perplexus. The three large and small palettes on the protarsi and 12 palettes on the mesotarsi of males are diagnostic. In addition to characters in the key, larvae of this species and G. occidentalis have the inner apical margin of the stipes truncated (Fig. 8), which differs from the rounded inner apical margin in G. perplexus (Fig. 6). Graphoderus liberus (Say, 1825) Distribution and Abundance: Common statewide, especially so in northern third (Table 1). County records: }-20, 22-23, 25,27,30-31,33-35,38,42, 44,57-58,61,66,68-70,72. Range: NT-NF-FL-MN. Habitat: Adults and larvae were collected mostly from margins of deeper ponds and small lakes. Larvae were found most frequently, but not exclusively, in acidic habitats. Life Cycle: Almost all third instar larvae were collected from June 7 to August 16; one was collected from McKenna Pond on September 15. Only 1% of adults were collected before May, but substantial numbers were collected from May through September. Unlike other Dytiscinae and other Graphoderus, adults were more readily collected with a net than with bottle traps (HiIsenhoff 1987). Reliance on bottle traps for collections in March and April and prolonged ice cover in the normal deepwater habitat probably account for low numbers found in early spring. This species apparently has a univoltine life cycle; adults overwinter in ponds and lakes, mate in early spring, and oviposit when the ice has thawed. Presence of third instar larvae in McKenna Pond from June 7 to August 6, 1976 suggests staggered oviposition, which may also account for the single larva found that year on September 15. Identification: The lon~tudinal stripes on the larvae are distinctive. Adults can be easily recogruzed in the field by their shape, distinctive color, and very active behavior; they tend to "jump" when out of water and disturbed. Graphoderus manitobensis Wallis, 1933 Distribution and Abundance: Uncommon in southern half (Table 1). County records: 39-40, 46-47, 49, 52, 54, 57, 59, 61. Range: MB-WIIA. Habitat: Large sedge and cattail marshes or ponds in open areas. Life Cycle: Twenty-one adults were collected from McKenna Pond April 3 to May 24; 20 were collected from Horicon Marsh June 13 to July 28 (only summer collecting). The remaining 13 adults were collected from various sites between April and early July. All adults, except one, were collected with bottle traps; none were found after July 28. Eight third instar larvae were collected June lo-30, with one second instar also present on the latter date. Another larva was found on September 10; it probably represents a partial second generation. Teneral adults occurred June 23 to July 14. Adults apparently

50 48 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 overwinter, probably in the breeding site, mate in early spring, and oviposit soon thereafter. Larvae develop in May and June and pupate; adults emerge from mid-june to mid-july to complete a univoltine life cycle. Identification: The palettes on mesotarsi of adult males is diagnostic; three large and small palettes occur on protarsi. Adult females, which were described by Tracy and Hilsenhoff (1982), have a more pronounced corrugated macrosculpture on the pronotum (Fig. 4) than other similar species. They most resemble females of G. occidentalis because both anterior and posterior black pronotal bands are contiguous with the margin, but in G. occidentalis the corrugated macrosculpture is obsolete and the metasternal wing is much wider. The metasternal wing in G. manitobensis is very narrow (0.33 mm), narrower than in almost all G. perplexus and much narrower than in G. fascicollis and G. occidentalis. Larvae resemble those of G. perplexus, but the inner margin of the stipes is gradually widened to the apex (Fig. 7) and not abruptly widened as in the other three species of Graphoderus with similar larvae (Figs. 6, 8). The sclerite on the larval prosternum is rectangular, and not narrowed anteriorly as in G. perplexus. In mature larvae it is slightly longer than wide, while in mature G. perplexus larvae it is about twice as long as wide, but this character varies somewhat with the age of the larva. Graphoderus occidentaus Horn, 1883 Distribution and Abundance: Uncommon statewide; 81 % of adults collected from Horicon Marsh and McKenna Pond (Table 1). County records: 3, 6-7,19-20,24-25,29,37,43-44,46,48,50,57-59,61, 67, Range: YK PQ-NY-WI-ND-UT. Habitat: Adults were collected from a variety of ponds. The fact that half of the adults were collected from Horicon Marsh by Kevin Kenow suggests that this large, predominantly cattail marsh is a preferred habitat. Unfortunately, larvae from Horicon Marsh were not saved. Life Cycle: Teneral adults were collected June 28 to August 15, which suggests a life cycle like that described for G. fascicollis. One teneral adult was collected October 3, indicating a partial second generation in some years. Identification: The 2 large and small palettes on protarsi and lack of palettes on mesotarsi of males, and the obscure corrugated sculpturing on the pronotum of females (Fig. 5), separate adults of this species from those of similar Graphoderus. Because the black anterior band on the pronotum is contiguous with the anterior margin (in pinned specimens the area near the margin may appear rufopiceous), adults most resemble those of G. manito bensis; separation of females is discussed under that species. Two larvae that may be this species have 40 or fewer spines on the last abdominal sternum compared with 46 or more in larvae beheved to be G. fascicollis, but this character IS variable and no other differences between these larvae and those believed to be G. fascicollis were found. Graphoderus perplexus Sharp, 1882 Distribution and Abundance: Common statewide (Table 1). County records: 1-8, 11-15, 17-21,23,25,27-29,31,34-44,46-47,48*,50,52,54, 57-59,61,64,66-67, Range: AK-NF-AR-UT Habitat: Adults and larvae were found in a variety of ponds, most of them being relatively shallow and vegetated with cattails, bur-reed (Sparganium), and sedges. About 75% of all adults were collected from McKenna Pond (Table 1), a relatively shallow 0.8 h pond containing cattails and bur-reed. Life Cycle: Seventy percent of adults were collected from March through May. Adults obviously overwinter in ponds. mate in early spring, and oviposit

51 1993 THE GREAT LAKES ENTOMOLOGIST 49 in April. In McKenna Pond, first instar larvae were collected May 1, and small third instar larvae were found as early as May 5, with large third instar larvae appearing by May 15. Peak numbers of third instar larvae were collected in late May and early June; a few were found as late as July 21 in the north. Third instar larvae of other species of Graphoderus were not found before June 7. Teneral adults were collected from June 7 to July 23, to complete a univoltine life cycle. In 1977 (after a drought in 1976), McKenna Pond was completely dry by July, but flooded July after heavy rain; on August 4 several second instar and two third instar larvae were collected. In 1980 and 1981 single third instar larvae were collected from McKenna Pond in early September and a teneral adult was found November 5, Teneral adults were collected elsewhere on August 15 and September 21. These lateoccurring larvae and the teneral adults probably represent a partial second generation. Identification: Adults can be readily distinguished by characters in the key. The corrugation of the pronotum of females, while readily seen (Fig 3), is not pronounced as it is in G. manitobensis (Fig. 4). The 3 large and small palettes on protarsi and palettes on mesotarsi of males is diagnostic. Larvae are most similar to those of G. manitobensis, but the shape of the stipes differs as described in the key. Larvae of G. perplexus and G. manitobensis can be readily separated from those of G. fascicollis and G. occidentalis by couplet 2 of the key and by conspicuous darkening at the base of spines on the last abdominal sternum, which is not evident in the latter two species. Hydaticus Leach, 1817 Roughley and Pengelly's (1981) study of Hydaticus in North America provides the most recent key and descriptions for adults of the five species. Two species occur in the western Great Lakes region and are commonly found in Wisconsin. Larvae resemble smaller larvae of Dytiscus, but lack lateral fringes on the urogomphi and have two projecting lobes on the labium (Figs. 9, 10). Although only the larva of H. aruspex has been described (Watts 1970, as H stagnalis), a study of larvae collected in Wisconsin enabled me to identify both species and to develop a key to third instar larvae. The life cycle differs from other Dytiscinae, except Thermonectus, because adults overwinter in terrestrial habitats. Galewski (1964) reported that Hydaticus was one of four genera of European dytiscids that overwinters as adults in forest litter. Although I found four adults in aquatic habitats as early as late March, substantial numbers were not collected before the last 10 days of April. Temperatures sufficiently warm to thaw overwintering sites and permit flight frequently occur in Wisconsin as early as the last half of March. Key to Species of Adult Hydaticus in Wisconsin 1. Elytra black with rufotestaceous lateral margins, or vittate in many females; pronotum rufotestaceous with basal black mark that is continuous with black on elytra; mm long....aruspex Elytra rather uniformly rufopiceous, sli~htly darker on disc; pronotum lighter than elytra and without markings; mm long piceus

52 50 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No. I Key to Species of Larval Hydaticus in Wisconsin 1. Anterior projections of labium pronounced, about a long as distance between their tips, and separated by au-shaped emargination (Fig. 9); 4-8 small spmes on genae on each side of gular area....aruspex Anterior projections of labium short, length much less than half distance between their tips, and separated by a shallow emargination (Fig. 10); no groups of small spines on genae....piceus Hydaticus aruspex Clark, 1864 (= H. modestus Sharp, 1882) Distribution and Abundance: Very common statewide (Table 1). County records: 1-45,46*, 47-59, 61, 63-64, 65*, 66-68, 70-71, 72*. Range: AK NF NJ MO-CA. Habitat: Adults and larvae were collected from Sphagnum bogs and swamps, sedge and cattail marshes, and from a wide variety of temporary and permanent ponds. Life Cycle: Overwintering adults fly from terrestrial sites to breeding habitats in early spring, with mating and oviposition occurring throughout April. Most larvae develop in May and June, with 70% of larvae having been collected in June. Third instar larvae were collected May 11 to October 8, with significant numbers found in August and September. This suggests at least a partial second generation in late summer. Less than 2% of the adults were collected from aquatic habitats after August, which indicates that they fly to terrestrial overwintering sites in late summer or early autumn. Identification: Adults and larvae of the two species are readily identified by characters in the key. Adults of H. aruspex, which have 3 elytral patterns, can be readily recognized in the field. Male adults of H aruspex have fasciate and non-fasciate elytra. In non fasciate males, elytra are black with a lateral yellow border; in fasciate males, the pattern is similar except that there is a pale sub-basal fascia. In addition to these patterns, female adults often have about six longitudinal yellow vittae on each elytron. All patterns occur in Wisconsin, with the fasciate pattern in males and vittate pattern in females predominating. Hydaticus piceus LeConte Distribution and Abundance: Common in southern third to rare in northern third (Table 1). County records: 1, 3, 6, 21, 24, 26*,38-41,44-45,47-52, 54-61, 64, 66-68, Range: AB-NS NJ-MO. Habitat: Adults and larvae were found in permanent ponds and marshes, especially those with cattails and bur-reed. Life Cycle: Adults were collected from April 8 to August 18; most were collected before July. Most larvae were found June 5-30; two were found August Overwintering adults apparently enter ponds in April, mate, and oviposit. Larvae develop in May and June, with apparently at least a partial second generation in late July and August. Identification: In size. shape, and general coloration adults superficially resemble Rhantus sinuatus, Agabus erichsoni, and several species of Ilybius, but they can be immediately recognized by their solid rufopiceous color with a somewhat lighter head and pronotum.

53 1993 THE GREAT LAKES ENTOMOLOGIST 51 Figures Labium of larva (ventral view). 9. Hydaticus aruspex. 10. H. pice us. Thermonectus Dejean Dejean initially named this genus Thermonetus in 1833, but later (1837) changed the spelling (Nilsson et ali989). In anticipation that the earlier spelling ultimately will be suppressed by the International Commission of Zoological Nomenclature. I will continue to use Thermonectus. The genus was revised in North America by McWilliams (1969), but the revision has not been published. except for descriptions of two new species in the southwestern United States and Mexico (Goodhue-McWilliams 1981). Only two species occur in the Great Lakes region; both were found in Wisconsin. Larvae of both species were described by Wilson (1923); his description of T. basillaris was relied upon for its inclusion in the larval key. The life cycle is apparently univoltine, with adults overwintering in terrestrial habitats, which they do not leave until late spring or early summer. Larval development is rapid, as documented below for T. ornaticollis. Key to Species of Adult Thermonectus in Wisconsin l. Elytra yellow with black irrorations and a black fascia in apical third; anterior and posterior black marks on pronotum separated from margins; larger, mm long....ornaticollis Elytra black on disc, with yellow spots at base and black irrorations laterally and in apical third; anterior and posterior black marks on pronotum extendmg to anterior and posterior margins and enclosing a narrow yellow line on disc; smaller, mm long...basillaris Key to Species of Larval Thermonectus in Wisconsin l. Labium with a pair of long, spine-like setae ventrally at base of ligula; pair of spines at apex of ligula less than 112 length of ligula basillaris Labium without spine-like setae ventrally at base of ligula; pair of spines at apex of ligula subequal to length of ligula....ornaticollis

54 52 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 Thermonectus bssillsris (Harris, 1829) Distribution and Abundance: Very rare in west-central area and southern third (Table U. County records: 31, 61, 67, 70. Range: WI-MA-FL-CA-MO. Habitat: Two adults were collected from McKenna Pond when it was a permanent pond; another was found in a temporary, cattail-sedge pond, and a fourth occurred in the margin of a spring. All were collected between August 3 and October 19. Life Cycle: The life cycle is probably similar to that of T. ornaticollis. Identification: Adults of the two species are readily separated by the key. The only larvae I have seen are those of T. ornaticollis; they differ significantly from the larva described and figured by Wilson (1923) for T. basillaris. Thermonectus ornsticollis (Aube, 1838) Distribution and Abundance: Uncommon in southern two-thirds (Table 1/. County records: 25-26, 28-29, 48, 51, 53-54, 57-58, 60-61, 71 *, 72. Range: ND-VT-FL-AZ-WY. Habitat: Adults were collected from ponds in open areas, which contained a variety of vegetation or lacked vegetation. Larvae were collected from recently flooded areas that had been dry. Life Cycle: Adults were collected June 14 through October 6, third instar larvae July 1 through August 18, and teneral adults August 18 to September 30. Since all t pes of habitats, including temporary ones, were sampled extensively in and May with traps and nets, I assume that adults were absent from these habitats before June. In McKenna Pond, which was sampled at weekly intervals after ice-out in 1980 and 1981, adults were first collected June 14, 1980 and June 16, 1981; 20 (3/4 c) were collected on the latter date. Adults apparently oviposit in recently flooded areas, and larvae complete development within a month in late spring and summer. Rapid development was documented in McKenna Pond, which was dry in July 1977 until flooded by a six-inch rainfall the night of July On July 21 one female adult was collected, and on August 4 nine male adults, one second instar larva, and 23 third instar larvae were collected. On August 18, three teneral and 51 mature adults were collected alon~ with one third instar larva. I believe that adults overwinter in terrestrial SItes where they remain until warm late spring or summer rains inundate temporary len tic habitats and margins of more permanent ones. Adults then fly to these habitats and oviposit. Eggs hatch within a few days (Wilson 1923) and larvae complete development within a few weeks. LITERATURE CITED Aiken, R.B., and C.W. Wilkinson Bionomics of Dytiscus alaskan us J. Balfour Browne (Coleoptera: Dytiscidae) in a central Alberta lake. Can. J. Zool. 63: Barman, E.H., Jr The biology and immature stages of selected species of Dytiscidae (Coleoptera) of central New York State. Ph.D. Diss., Cornell Univ. v pp. Galewski, K The hibernation of the adults of the European species of Dytiscidae (Coleoptera) out of water. Polskie Pismo Entomol. 34: Goodhue-McWilliams, KL Two new species of Thermonectus (Coleoptera: Dytiscidae) from southwestern U.S.A. and western Mexico. Coleopts. Bull. 35: Hilsenhoff, W.L Notes on Nearctic Acilius (Dytiscidae), with the description of a new species. Ann. EntomoL Soc. Amer. 68: Effectiveness of bottle traps for collecting Dytiscidae (Coleoptera). Coleopts. Bull. 41: Comparison of bottle traps with a D-frame net for collecting adults and larvae of Dytiscidae and Hydrophilidae (Coleoptera). Coleopts. Bull. 45:

55 1993 THE GREAT LAKES ENTOMOLOGIST Dytiscidae and Noteridae of Wisconsin (Coleoptera). I. Introduction, key to genera of adults, and distribution, habitat, life cycle, and identification of species of Agabetinae, Laccophilinae and Noteridae. Great Lakes Entomol. 25: Larson, D.J An annotated list of the Hydroadephaga (Coleoptera: Insecta) of Manitoba and Minnesota by J.B. Wallis. Quaest. Entomol. 9: McWilliams, KL A taxonomic revision of the North American species of the genus Thermonectus Dejean (Coleoptera: Dytiscidae). Ph.D. Diss., Indiana Univ. 220 pp. Michael, A.G., and J.F. Matta The insects of Virginia No. 12. The Dytiscidae of Virginia (Coleoptera: Adephaga) (Subfamilies: Laccophilinae, Colymbetinae, Dytiscinae, Hydaticinae and Cybistrinae). Res. Div. Bull. 124, V.P.I. and State Univ., Blacksburg, VA. 53 pp. Nilsson, A.N., R.E. Rougbley, and M. Brancucci A review of the genus and family-group names of the family Dytiscidae Leach (Coleoptera). Entomol. Scandinavica 20: Rougbley, R.E A systematic revision of species ofdytiscus Linnaeus (Coleoptera: Dytiscidae). Part 1. Classification based on adult stage. Quaest. Entomol. 26: and D.H. Pengelly Classification, phylogeny, and zoogeography of Hydati cus Leach (Coleoptera: Dytiscidae) of North America. Quaest. Entomol. 17: Tracy, B.H., and W.L. Hilsenhoff The female of Graphoderus manitobensis with notes on identification of female Graphoderus {Coleoptera: Dytiscidael. Great Lakes Entomol. 15: Wallis, J.B 'The genus Graphoderus Aube in North America (north of Mexico) (Coleoptera). Can. Entomol Watts, C.H.S The larvae of some Dytiscidae (Coleoptera) from Delta, Manitoba. Can. Entomol. 102: Wilson, C.B Water beetles in relation to pondfish culture, with life histories of those found in fishponds at Fairport, Iowa. Bull U.S. Bur. Fish. 34: Wolfe, G.W The larva and pupa of Acilius fratemus fratemus (Coleoptera: (Dytiscidae) from the Great Smoky Mountains, Tennessee. Coleopt. Bull. 34:

56 1993 THE GREAT LAKES ENTOMOLOGIST 55 MICROCTONUS PACHYLOBII (HYMENOPTERA: BRACONIDAE): NEW HOST RECORD FROM HYLOBIUS RADICIS (COLEOPTERA: CURCULIONIDAE), AND ADDITIONAL NOTES ON ITS BIOLOGY George D. Hoffman l and Kenneth F. Raffa l ABSTRACT The endoparasite Microctonuspachylobii was discovered parasitizing a new weevil host, Hylobius radicis. Thirteen of the 154 H. radids adults collected were parasitized (8.5%). The median numbers of parasites per weevil were 26 (x = 22.5) during the period April through June, and 4 (x =9.4) during August and September. The median male:female sex ratio was 0.91 (x = 0.65). Males emerged approximately 1 day earlier than females. Median parasite mortality while in the cocoon was 10.2% per parasitized weevil (x = 11.8%). Microctonus pachylobii was not found parasitizing two previously recorded weevil hosts from field samples, Hylobius rhizophagus and H. pales, and a laboratory study suggests that the parasite may have difficulty parasitizing the latter species. Microctonus pachylobii Muesebeck was originally described as a parasite of the pitch-eating weevil, Pachylobius picivorus (Germar) (Muesebeck 1961). Its reported host range and geographical distribution were long confined to P. picivorus collected from southeastern United States (Krombein et al. 1979). Recently M. pachylobii was found parasitizing the pine root tip weevil, Hylobius rhizophagus Millers, Benjamin and Warner, and the pales weevil, Hylobius pales (Herbst), collected from central and northwest Wisconsin (Rieske et al. 1989). These two weevil species, plus P. picivorus and the root collar weevil, Hylobius radids Buchanan, are pests in Christmas tree plantations in the Great Lakes region. The discovery of M. pachylobii in Wisconsin has led to questions regarding its potential as a biological control agent against these weevil pests. There is currently little information in the literature on the biology of this parasite. We recently discovered M. pachylobii parasitizing a new weevil host, H. radicis, during the course of a study at the site in central Wisconsin where Rieske et al. (1989) had.previously found M. pachylobii. Data were collected on the incidence of parasitism of the three Hylobius species, the number of parasite progeny per parasitized weevil, differences between female and male parasite pupal development time, parasite mortality during the pupal period, and the parasitization rates of the three Hylobius species under laboratory conditions. IDepartment of Entomology, University of Wisconsin, Madison, WI,

57 56 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 MATERIALS & METHODS Endoparasitic larvae were initially recovered from H. radicis during routine dissections of adults of the three Hylobius species. The weevils were collected in 1991 as part of a study on weevil reproductive biology. The study site was a red pine, Pinus resinosa, and jack pine, P. banksiana, plantation 10 miles south of Milston, Jackson County, Wisconsin. Living weevils were collected from either pitfall traps baited with turpentine and ethanol, or screen traps wrapped around tree trunks. After the initial discovery of the parasite larvae a portion of the Hylobius weevils were held alive for parasite emergence. These individuals were not a random sample of all the collected weevils, but were from weekly collections in which parasitized weevils were found during dissections. The adult weevils were placed in individual diet cups containing a section of fresh red pine twig (weevil food) and peat moss (pupation medium for emerging parasite larvae). They were held at 22 C and 16:8 L:D photoperiod. The diet cups were checked for the presence of parasite cocoons every three to four days for two months. When cocoons were found they were separated from the peat moss, placed individually in gelatin capsules, and held at the same temperature and photoperiod. The peat moss was placed in a petri dish and observed for emergence of parasites from any undiscovered cocoons. The adult weevil, usually dead, was dissected and any unemerged parasite larvae recorded. Cocoons were checked daily for adult parasite eclosion. Cocoons from which parasites did not emerge were opened to determine the developmental stage at which the parasite died. Because M. pachylobii was not found parasitizing field collected H. pales and H. rhizophagus during this study, we investigated the ability of the parasite to successfully parasitize the three Hylobius species under laboratory conditions. Parasites that emerged in the laboratory were mated when two days old as adults. Four females were placed with four males in a diet cup containing a honey and water source. Two days later, individual females were placed with individual H. radicis (n ;: 8), H. rhizophagus (n = 4), and H. pales (n = 10) under the above conditions. The former two weevil species had been collected in the field and held in the laboratory for 3 months to assure they were not already parasitized. The H. pales adults were from a laboratory colony. Twenty four hours later the parasites were removed and the weevils held for 3 months while observed twice weekly fortarasite emergence. Weevils that died during this period were dissected an checked for parasite larvae. Weevils still alive after three months time were dissected and checked for the presence of diapausing first instar parasite larvae. Microctonus species diapause as first instar larvae within the host (Loan 1967). RESULTS Microctonus pachylobii larvae were found in 7 of the 118 dissected H. radicis, and larvae emerged from 6 of the 36 isolated weevils. The total parasitization rate was 8.5%. No parasites were found in the 52 dissected and 16 isolated H. rhizophagus, or the 77 dissected and 10 isolated H. pales weevils. The data on the number of parasites per parasitized H. radicis are separated into two groups based on the date the weevils were collected. We report results using the median and range for data from small sample sizes that appear to be non-normally distributed. The conventional mean and its standard error are also reported. The first group is comprised of weevils collected during the early peak of weevil activity (April through June). The second group consists of root collar weevils collected during the second peak of activity (August and September). Weevil activity is very low from mid-july to early

58 1993 THE GREAT LAKES ENTOMOLOGIST 57 August (Raffa and Hall 1988). The median number of parasites (larvae or cocoons plus unemerged larvae) in the April-June group (6 female and 2 male weevils) was 26, with a range of 9 to 32 (x= 22.5 ± 2.9). The median number in the August-September group (2 female and 3 male weevils) was 4, with a range of 1 to 22 (x = 9.4 ± 4.2). A few parasite larvae did not successfully emerged from their hosts. Two of the isolated weevils, from which 23 and 14 larvae emerged and pupated, contained 2 and 1 unemerged parasite larvae, respectfully. The median male:female sex ratio of the brood irrespective of weevil collection date was 0.91, with a range of 100% female to 1.14 (n 6) (x = 0.65 ± 0.18). The median parasite mortality while in the cocoons was 10.2% per weevil (n =6), with a range of 4.3% to 23.1 % (x = 11.8% ± 2.5). Five of the 14 dead parasites died as larvae or shortly after spinning the cocoon, three died during the pupal stage, and six of the dead were completely formed adults. Some of these adults appeared to have been unable to completely free themselves from the pupal skin. Because isolated weevils were not:checked every day for parasite emergence, the exact length of the pupal period is unknown for most of the parasites. Our data suggests that on a per weevil basis (n = 5), male parasites (n = 43) edose 0.82 days earlier (median) than females (n = 55). Parasite larvae were seen emerging from a laboratory parasitized H. rhizophagus. In this incident the average length of the pupal period was 11.6 ±.12 days for the all male brood (n = 17). Four of the 8 H. radicis weevils confined with female M. pachylobii were parasitized. One of the parasitized weevils died, possibly due to its food drying out, and the parasite larvae were discovered upon dissection of the cadaver. All male parasite progeny emerged from the other three weevils. Three of the four H. rhizophagus weevils were parasitized. Parasite larvae were found in one moribund weevil, and the other two weevils gave rise to all male parasites. None of the H. pales weevils were parasitized (n = 10). Dissections of the nonparasitized weevils of all three species revealed no diapausing first instar parasites. In this laboratory parasitism test the mean duration of the egg plus larval stage of M. pachylobii in H. radicis and H. rhizophagus was 41.0 ± 1.14 days (n = 5). The oviposition behavior of M. pachylobii during the first five to 10 minutes after contact with H. radicis and H. rhizophagus did not seem conducive to successful parasitization. The parasites were highly attracted to feeding weevils, and less so to feeding scars. Many of the parasites would repeatedly thrust their ovipositor at the mouthparts of feeding weevils. Weevils which were not movin~ were less attractive to parasites than feeding scars. Parasites would routmely walk over stationary weevils without making attempts to oviposit. We saw a few parasites chase after walking weevils, but no complete ovipositional thrusts were made, even after the weevil had stopped walking. Two parasites were seen to hang from the lower abdomen of a weevil and thrust their ovipositor forward. The ovipositors did not appear to penetrate the intersegmental membrane, but we had a poor view of the parasites. Weevil behavior did not appear to be influenced by the presence of the parasite. We did not observe the behavior of parasites placed with H. pales. DISCUSSION Finnegan (1962) and Shenefelt & Millers (1960) each reported the rare occurrence of a larval parasite of H. radicis. This study is the frrst to identify a Hymenopteran parasite of adult H. radicis. The expansion of the host range of

59 58 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 M. pachylobii to include H. radicis is not surprising given that the parasite's range includes two weevil genera, and the fact that H. radicis and H. rhizophagus are sibling species. What is surprising is that we did not find parasitized H. pales and H. rhizophagus in our field collections. Four years earlier, Rieske et al. (1989) found parasitized H. pales andh. rhizophagus at this site, but not parasitized H. radicis. The laboratory parasitism test showed that M. pachylobii can parasitize H. rhizophagus as readily as H. radicis, but may have difficulty parasitizing H. pales. The 8.5% parasitism rate of H. radicis is a pooled value combining all collections over the summer. Depending upon the life history of the host, parasitism rates expressed in this manner can misrepresent the total loss to parasitism for a life stage over a generation (Van Driesche et al. 1991). The long adult life and seasonal history ofh. radicis (Wilson and Millers 1983) will reduce the magnitude of this error. The time at which H. radicis weevils were collected from the field appears to affect the number of larvae found in both female and male weevils. The April-June peak of weevils was primarily composed of reproductive individuals. Female weevils were filled with eggs in May and June, and ovipositing parasites may be able to detect a host with substantial nutritional reserves. Females collected in August-September were almost all postreproductive or newly emerged adults. Conversely, males collected during both peaks were mostly sexually mature with fully enlarged gonads (G.D. Hoffman unpublished data). This suggests that differences in nutritional reserves between weevils collected during the April-J une and August-September peaks may not explain the difference in the number of parasites per weevil during these two periods. It is not known how many generations M. pachylobii has each year, but the long juvenile development period, and length of time over which parasitized weevils were found, suggests there are two generations a year. The all male M. pachylobii progeny that resulted from the laboratory parasitism test suggests that female parasites were not fertilized during the two days they were confined with males. We observed male and female parasites coupling soon after the two sexes were placed in the same vial. We did not investigate the duration of copulation or note behaviors of the parasites which could have disrupted fertilization. Alternative explanations are that the parasites can control the sex ratio of their. progeny, or that the male parasites were infertile. The life histories of H. rhizophagus (Kearby and Benjamin 1969) and H. pales (Finnegan 1959, Bliss and Kearby 1970) are similar to that of H. radicis (Wilson and Millers 1983); and same is true for the annual pattern of ovary and teste development (G.D. Hoffman unpublished data). This suggests that constraints imposed on the parasite's biology by the life history and physiology of the three weevil species should be similar. The data from this and other studies suggest that M. pachylobii will not be an effective biological control agent of Hylobius weevils in the Great Lakes region. We recorded low parasitism rates of H. radicis, and found that the parasite does not consistently parasitize a given weevil species at a single location. Of particular interest is that during the six-year period in which our laboratory has studied these four weevil pests, M. pachylobii was never found parasitizing P. picivorus, despite high local densities of this weevil. Nor has M. pachylobii been found to parasitize H. pales in the southern United States (Lynch 1984).

60 1993 THE GREAT LAKES ENTOMOLOGIST 59 ACKNOWLEDGMENTS We thank Steven Krauth, University of Wisconsin, and Paul Marsh, Systematic Entomology Lahoratory, USDA, for the identification of M. pachylobu. Richard Hofstetter and Todd Patton assisted with the weevil collections. We thank Dan Mahr and Michael Strand for reviewing the manuscript. This study was supported by The University of WisconsinMadison Graduate School, the University of Wisconsin-Madison College of Agricultural and Life Sciences Center for Integrated Agricultural Systems, The Wisconsin Christmas Tree Producers Association, and USDA Competitive Grants 86 CRCR , and 86-FSTY LITERATURE CITED Bliss, M. J. & W. H. Kearby Notes on the life history of the pales weevil and northern pine weevil in Central Pennsylvania. Ann. Entomol. Soc. Am. 63: Finnegan, R. J The pales weevil, Hylobius pales (Hbst), in southern Ontario. Can. EntomoL 91: The pine root collar weevil, Hylobius radicis Buch., in southern Ontario. Can. Entomol. 94: Kearby, W. H. & D. M. Benjamin Life history and damage of the pine root tip weevil Hylobius rhizophagus, in Wisconsin. Ann. Entomol. Soc. Amer. 62: Krombein, K. V. J. P. D. Hund, D. R. Smith. & B. D. Burks Catalog of Hymenoptera in America north of Mexico, Vol 1: Symphyta and Apocrita (Parasitical p. Smithsonian Inst. Press, Washington, D.C. Loan, C. C Studies on the taxonomy and biology of the Euphorinae (Hymenoptera: Braconidae). II. Host relations of six Microctonus species. Ann. Entomol. Soc. Am. 60: Lynch. A. M The pales weevil, Hylobius pales (Herbst): a synthesis of the literature. J. Ga. Entomol. Soc. 19:1-34. Muesebeck, C. F. W A new Opius and two new species of Microctonus (Hymenoptera: Braconidae). Bull. Brooklyn Entomol. Soc. 56: Raffa, K. F. & D. H. Hall Seasonal occurrence of pine root collar weevil, Hylobius radicis (Coleoptera: Curculionidae), in red pine stands undergoing decline. Great Lakes Entomol. 21: Rieske, L. K., D. W. A. Hunt & K. F. Raffa Microctonus pachylobii (Hymenoptera: Braconidae) parasitizes Hylobius weevils in Wisconsin: new host genus and geographic records. Entomol. News. 100: Shenefelt, R. D. & I. Millers A new species of Eracon from the pine root collar weevil. Can. EntomoI. 92: Van Driesche, R. G., T. S. Bellows, Jr., J. S. Elkinton, J. R. Gould, and D. N. Ferro The meaning of percentage parasitism revisited: solutions to the problem of accurately estimating total losses from parasitism. Wilson, L. F. & 1. Millers Pine root collar weevil-its ecology and management. U.S.D.A. For. Servo Tech. Bull. No

61 1993 THE GREAT LAKES ENTOMOLOGIST 61 FIELD INVESTIGATIONS ON THE AMERICAN DOG TICK, DERMACENTOR VARIABILlS, IN NORTHWEST OHIO (ACARI: IXODIDAE) Kelly M. Micher! and C. Lee Rockett! ABSTRACT Ecological investigations on the American dog tick, Dermacentor variabilis, were conducted in two metroparks located in Lucas County, Ohio. Adult tick surveys were conducted in 1989 and For both years, adult tick activity began in late April, and adult ticks were most abundant from early May to mid-june. Observed activity had ceased by early August, producing a unimodal pattern of activity. Sunny days with temperatures between 24 and 32 C were most conducive to adult tick activity, and adult ticks were most abundant on grass trails and in meadows. Using nest boxes inhabited by white-footed mice, immature tick surveys were conducted in 1989 and For both years, larval abundance peaked in early May and nymphal abundance peaked in late June or early July. The American dog tick, Dermacentor variabilis (Say), is established in large areas of eastern and central North America, and in addition, there are several disjunct populations which include populations in the Great Lakes region and Canada (Sonenshine 1979). Ecological investigations of D. variabilis have taken place in Virginia (Sonenshine 1979). Tennessee (Zimmerman et al. 1987, 1988), Georgia (Newhouse 1983), Nova Scotia (Campbell 1979) and Ohio (Conlon and Rockett 1982, Harlan 1986). The importance of D. variabilis as a vector of Rocky Mountain spotted fever (RMSF) has provided much of the incentive behind such studies (Mount and Haile 1989). Much remains to be learned of the distribution, abundance, seasonal activity, host and vegetative parameters, and environmental aspects of D. variabilis. The American dog tick has a life cycle of egg, larva, nymph, and adult. Campbell and MacKay (1979j reported the importance of the whitefooted mouse, Peromyscus leucopus (Rafinesque); the meadow vole, Microtus pennsylvanicus Ord; and the red-backed vole, Clethrionomys gapperi Vigors, as hosts for larval dog ticks. The meadow vole, M pennsylvanicus, was listed as an important host for nymphal dog ticks. Grey squirrels, Sciurus carolinensis Gmelin; raccoons, Procyon lotor (L.); eastern chipmunks, Tamias striatus (L.); and red squirrels, Tamiasciurus hudsonicus Erxleben, also were reported to be important nymphal hosts (Garvie et al. 1978, Campbell and MacKay 1979, Zimmerman et al. 1988). The adult ticks tend to parasitize larger mammals such as raccoons, skunks, Mephitis mephitis (Schreber); woodchucks, Mal' mota monax (L.); opossums, Didelphis virginiana Kerr; horses, dogs, and man IDepartment of Biological Sciences, Bowling Green State University, Bowling Green, OH

62 62 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 (Zimmerman et al. 1988, Fish and Dowler 1989). Tick seasonal activity patterns vary with regional temperature regimes. In the northern part of the dog tick's range such as Canada and Massachusetts, overwintering larvae and adults begin activity in the spring with peak activity occurring in early summer (McEnroe 1978, Campbell 1979). Tick activity in these areas normally ceases by mid-autumn (Campbell 1979). Ecological studies of D. vanabilis have provided information on environmental factors that may influence tick seasonal activity and survival. Important controlling factors appear to be ambient temperature, annual temperature regime, solar radiation received at ground level, and moisture (Atwood and Sonenshine 1967, Conlon and Rockett 1982, McEnroe 1978,1984). Differences in relative abundance and distribution of ticks with respect to vegetation have also been noted. The purpose of this study was to obtain information on the life cycle, seasonal dynamics, vegetative and host associations, and other environmental parameters of the American dog tick in two heavily used park areas of Lucas County, Ohio. In 1979 and 1980, after extensive surveys, Conlon and Rockett (1982) reported extremely low number of collected dog ticks in one of these two park areas. This low number of collected ticks limited the significance of the overall ecological investigation necessitating additional studies. Despite an apparent low number of ticks, Lucas County, located in northwest Ohio, accounted for more than 21 percent (107/491) of the RMSF cases reported from Ohio between the years 1964 through 1990 (Fontaine 1991). MATERIALS AND METHODS Secor and Oak Openings, located in southwestern Lucas County, maintain breeding populations of dog ticks. Secor Park, a 243 ha preserve is located approximately 24 km northeast of the much larger (1,484 hal Oak Openings Preserve; both parks are open to visitors daily. From the beginning of March through the middle of October in 1989, Secor Park was surveyed weekly for male and female adult ticks. Sex ratios were noted. Both Secor and Oak Openings were surveyed in 1990 using the same date parameters as used for Secor Park in Again, sex ratios were noted. Adult dog ticks were collected with a 3 m 2 white flannel drag as described by Smith et al. (1946). Four different vegetative areas of Secor Park and five different vegetative areas of Oak Openings were each "dragged" for a ten-minute period every week during the survey. The flannel cloth was dragged over vegetation for one minute at a time at 50 paces per minute and any attached ticks were removed. At that point, dragging would then continue. Dragging time did not include the time taken to remove ticks from the cloth, and no areas received repetitive drags during any weekly survey. In both parks a grass path and grassy roadside, each approximately 800 m long and containing primarily domestic yard grass, were surveyed for ticks. A meadow with assorted grasses and weeds plus a second growth, mixed, deciduous woodlot with numerous oak trees were also surveyed in each park. In addition, a pine woodlot containing mainly red and white pines, was examined in Oak Openings. All meadows and woodlots were approximately m 2 in area. Comparisons of habitat to number of ticks collected were analyzed using the Kruskal Wallis ANOVA (Zar 1984), which accounted for the unequal variances at each site. The air temperature and the solar radiation condition were recorded each week. The solar radiation condition was rated as being sunny, cloudy, or mixed sun and clouds. A comparison of ticks collected to daily air temperature was accomplished with simple regression analysis (Zar

63 1993 THE GREAT LAKES ENTOMOLOGIST 63 Table 1. Numbers of adult ticks collected according to park, year, sex, and solar radiation condition on day of capture. Number of Ticks Collected Park and Year Male Female Ticks/Minute Sunny and Clouds Cloudy 1989 Secor Oak Openings Total ). The significance of different solar conditions to collected tick numbers was determined with chi-squared analysis (Zar 1984). Since immature dog ticks are rarely captured by "dragging" the vegetation (Smith et al. 1946), hosts and their nests were examined directly for ticks using mice nest boxes. Sherman-type animal traps were not used due to certain disadvantages such as not having permission to trap in certain areas of the public parks and possible vandalism to traps. In 1989, twenty mouse nest boxes were constructed of plywood, using a modified design described by Mitchell and Micher (1989). Nesting material (polyester fiber filling) was placed inside the boxes and each was baited weekly with a handful of sunflower seeds. The boxes were nailed to trees at a height of 0.5 to 1 m above the ground. Five boxes were attached to trees within each of four areas of Oak Openings Park. These areas consisted of the same deciduous and coniferous woodlots and meadow used for the adult tick survey, plus a different meadow containing a few apple trees. The boxes were checked weekly for ticks and mice from 14 April-13 October and 3 March-13 October for 1989 and 1990, respectively. Collected mice were anesthetized with ether, ticks were removed, and the mice were subsequently released back into their home area. The number of ticks and developmental stage were recorded for each box every week. Comparisons of immature tick number to habitat were analyzed using the Kruskal Wallis ANOVA (Zar 1984), which accounted for unequal variances at each site. RESULTS During the two year study period, 1174 adult Dermacentor variabilis were collected (Table 1). A marked difference in the number of dog ticks collected was noted between Secor Park (1142 ticks, 97%) and Oak Openings (32 ticks, 3%). In Secor Park, the capture rate decreased from a seasonal average of 1,45 ticks collected per minute in 1989 to a seasonal average of 0.31 ticks collected per minute in During the 1990 season of adult tick activity in Oak Openings. the average capture rate was 0.04 ticks collected per minute. Adult ticks were collected more often on sunny days than on days with mixed sunny and cloudy days with high air temperatures (Le., above 25 C). For example, in 1989 with 16 collection dates, one sunny day with the air temperature below 15 C accounted for 5.2% of the total ticks captured, while three cloudy days and two mixed sunny and cloudy days each with air temper atures above 25 C accounted for only 4.5% and 2.3%, respectively, of the total tick captures. During the 1989 season, seven sunny days accounted for 73.3% of the total tick captures within Secor Park. Four mixed sunny and cloudy days accounted for 18.3% of the total tick captures and five cloudy days produced only 8.5% of the ticks. During the 1990 season, ten sunny days

64 64 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No III II! u f= :5 140 ~ ' <I.l 100..Q E ::I 80 Z t,..-. Secor \ \ +--+ Secor \,,'-... Oak Openings ,J,,,,, \ 1,,'", J, ",, ". I ", I ",,,, 1, I April May June July August Week of Collection Figure 1. Seasonal distribution of adult Dermacentor vanabilis in Secor Park and Oak Openings Park, Lucas Co., Ohio. accounted for 67.1% of the total adult ticks collected within both parks combined, and five cloudy days produced 21.8% of the ticks. The level of adult tick activity increased significantly as the air temperature increased (P < 0.001). Adult ticks were most commonly collected (58%) at ambient temperatures between 24 C and 32 C. Adult ticks were not collected at ambient temperatures below 12 C. During this study, no temperatures higher than 32 C were recorded. Infestation levels were significantly different among habitats within Secor Park (P ::$; ). Eight hundred and twenty adult ticks (71.8%) were collected from the grass path habitat. Two hundred thirtyseven ticks (20.8%) were collected from the meadow and 78 ticks (6.8%) were collected from the grassy roadside. Only seven adult ticks (0.6%) were collected from the mixed deciduous woodlot in Secor Park. Infestation levels also were significantly different among habitats within Oak Openings (P < ). As with Secor Park, the majority of adult ticks (17 ticks or 53.1 %) were collected from the grass path habitat. Seven (21.9%) and six adult ticks (18.75%) were captured from the meadow and roadside, respectively. Two ticks (6.25%) were collected in the coniferous woodlot and no ticks were collected in the mixed deciduous woodlot within Oak Openings Park. Adult tick seasonal activity, as observed by the authors, is depicted in Figure 1. The first adult ticks were collected on 21 April 1989 and 22 April In Secor Park, adult ticks were most abundant from early May to mid June for both years of study. Even during this period, observed tick activity

65 1993 THE GREAT LAKES ENTOMOLOGIST 65 til..::l ~ ~ ::::I rti E E '0... ~ z 24, , 22 I<- --,t. larvae Nymphs larvae Nymphs March April May June July August Sept. Oct. Week of Collection Figure 2. Seasonal distribution of immature Dennacentor variabilis in Oak Openings Park, Lucas Co., Ohio. was decreased and atypical during rainy periods or conditions when air temperatures were below 15 C. This is reflected in Figure 1 (i.e., week 2, May 1989; week 3, June 1989; week 4, May 1990). In Oak Openings, adult ticks maintained a constant low level of observed activity throughout the season. Apparent adult tick activity had ceased by early August for both years and both parks producing a unimodal pattern of distribution. A majority (54%) of ticks collected was male. A total of 185 immature dog ticks was collected from Oak Openings duro ing the two-year study. One hundred and thirty six larvae and 24 nymphs were collected from inside the nest boxes and nesting material while 24 larvae and one nymph were collected directly from mice found in the nest boxes. Most (77%) of immature ticks collected were engorged larvae. In 1989, larval collections increased to a peak in May and then decreased thereafter (Figure 2). Similarly, larval activity peaked in May of No larvae were caji>tured after the first week of July 1989, however, two larvae were collected m late September of In 1989, the majority of nymphs (64%) was collected in early July and apparent activity ceased by late August. In 1990, peak activity occurred in late June when 78% of the nymphs were collected. Apparent activity ceased by late August. Immature tick density with respect to nest box location was significantly different for larvae (P < ), but was not significantly different for nymphs. One hundred and two larvae (63.75%) were collected from boxes or mice found in the boxes placed in the first meadow habitat. Thirty five larvae

66 66 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 (21.88%) were collected from the pine woodlot. Nineteen larvae (11.88%) and four larvae (2.5%) were collected from the mixed deciduous woodlot and second meadow habitat (with a few, scattered apple trees), respectively. Eleven (44%) and nine (30%) nymphs were collected from the mixed deciduous and coniferous woodlots, respectively. Three nymphs (12%) were collected from the first meadow habitat while two (8%) were captured from boxes placed in the second meadow containing apple trees. White-footed mice were the only mammals captured from the nest boxes. Twenty-two and 19 mice were captured from nest boxes in 1989 and 1990, respectively. For both years, the majority of captured mice were from the coniferous woodlot-1989 (51 %) and 1990 (53%). DISCUSSION Considerably more ticks were collifcted from Secor Park than from Oak Openings. This is interesting, given the close Qroximity of the parks (less than 24 kilometers aqart). In a previous study, Conlon and Rockett (1982) reported low population levels of dog ticks in Oak Openings. Unlike Oak Openings, Secor Park is a small metropark that is located in the middle of an urbanized area. Trails or paths in Secor Park are subject to greater and more frequent host traffic, particularly dogs and humans, than Oak Openings. It is tempting to speculate that enhanced host availability would be a factor in increasing tick densities along the path areas in Secor Park. Smith et al. (1946) reported that even in heavily infested regions, abundance of D. vanabilis is localized and sporadic. Soil surveys have determined that Oak Openings has many more sand dunes, beaches and ridges compared to Secor Park which has more low lying areas (Stone et al. 1980). This should produce a much drier situation in Oak Openings as the sand hills will allow better drainage of water while the low lying areas of Secor Park are subject to ponding. Greater moisture content of the soils in Secor Park may have contributed to the survival success of dog ticks simply because the ticks were not put under as much risk to desiccation. Dodds et al. (1969) reported h' r tick abundance in wetter areas of the Tobeatic Game Sanctuary in Nova tia when compared to tick surveys in drier areas in the sanctuary. The above authors concluded that the presence of considerable moisture with accompanying grass or sedge is needed to maintain high tick densities. Adult dog tick abundance decreased markedly in Secor Park during the second year of the study; over four times as many ticks were captured in 1989 than in This decrease may be explained partially by differences in known winter weather conditions, particularly December 1988 and According to the National Oceanic and Atmospheric Administration (NOAA 1988,1989). the mean temperature in December, 1988, was only slightly below normal (0.2 C below normal). but the mean temperature of December. 1989, was well below normal (6.2 C below normal). The seasonal snowfall was below normal for both winters (NOAA) 1988, 1989). The low tick abundance observed in 1990 may be due partially to lowered overwintering success caused by the harsh December temperature (monthly mean of -8.4 C) of 1989 coupled with inadequate snow cover. McEnroe (1977) reported that when winter temperature means fall below O C the overwintering ticks are subject to mortality from desiccation as the water maintenance pump works less adequately under extreme temperatures. Conlon and Rockett (1982) and McEnroe (1984) reported that snow cover might help to insulate ticks from colder winter temperatures within their overwinterinl? microhabitat and help maintain soil and leaf-litter moisture above the ticks critical humidity equilibrium.

67 THE GREAT LAKES ENTOMOlOGIST 67 Solar radiation condition data denoted higher adult tick activity on sunny days than on days with mixed sun and clouds or cloudy days. This supports Atwood and Sonenshine's (1967) finding that questing activity in adult ticks is positively correlated to the amount of solar radiation received at ground level. New house (1983) observed that cloudy days negatively affected tick activity, and Conlon and Rockett (1982) reported higher capture rates on sunny days. Temperature data from Secor Park and Oak Openings indicated that adult tick activity also increased as ambient air temperatures increased. Newhouse (1983) reported a positive correlation between tick activity and ambient temperature. Smith et al. (1946) reported the importance of ambient temperature to the incubation period of eggs, molting time for immature ticks, tick activity, and longevity in ticks. In the present study, adult ticks were not captured at air temperatures below 12"C or above 32 C. Hall and McKiel (1961) and McEnroe and McEnroe (1973) reported a lower temperature threshold of 5 C for adult tick questing activity and a curbing of questing at higher temperatures (40 C). During 1989 and 1990, adult tick activity began in late April, peaked in mid-may to early June, and declined thereafter. Larval activity began in late March and p~aked in May. Nymphal activity was most evident in late June to early July. This pattern of seasonal tick activity suggested a two-year life cycle for D. variabilis. The spring adults would produce eggs that hatch into larvae in the summer. These larvae would overwinter until the following spring. In the spring, the overwintered larvae would become active and then the engorged larvae would molt into nymphs which quest for hosts in June and July. The engorged nymphs would molt into adults in late summer. Most of these late-summer adults would not actively quest, but instead, would overwinter and become active in late April the following year for a two-year cycle. Some late summer adult ticks will quest, but, McEnroe (1974) and Smith et al. (1946) noted that latesummer, questing adults usually do not contribute to the following generation because eggs cannot successfully overwinter in cold climates. In Lucas County, the last annual freeze is typically between April. Initial adult activity began in both Secor Park and Oak Openings just after these dates (21 April 1989 and 22 April 1990), the same dates reported by Conlon and Rockett (1982). Adults were not collected by "dragging" in late summer of 1989 and Comparable seasonal activity patterns for D. variabilis were reported by Campbell (1979), Garvie et al. (1978), and Smith et al. (1946). In contrast to the present study, Conlon and Rockett (1982) and McEnroe (1974) reported bimodal activity for adult dog ticks with a second small peak of activity observed in late summer. This bimodal activity pattern may be attributed to the maintenance of a few individuals with aberrant behavior (Wilkinson 1979). The individuals may quest for hosts in late summer or fall if environmental conditions are suitable and produce the second small peak of activity sometimes observed. Adult dog ticks were most often captured on grassy paths. This conformed with the results of other authors such as Newhouse (1983) who observed high tick abundance on grass {laths and trails in Georgia. Sonenshine et al. (1972) reported that D. variabllis is often associated with edges of trails, roads, meadows and other clearings. The scent of hosts may be more concentrated on grass paths and trails in Georgia. Sonenshine et al. (1972) reported that D. variabilis is often associated with edges of trails, roads, meadows and other clearings. The scent of hosts may be more concentrated on grass paths and thus in turn could attract more dog ticks to an area. Dukes and Rodriguez (1976) reported that nymphal dog ticks were more attracted to dog and rabbit hair than sterile cotton. Dog hair was preferred over rabbit hair. Many (244) adult ticks were collected from the meadow habitats in the

68 68 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 parks. Smith et al. (1946) noted that tick infestations were usually associated with an abundance of grassy cover in which meadow mice could thrive. In this study, trees were scattered throughout the meadows and the meadow habitat bordered second growth woodlots. The majority of the meadow captured ticks were collected from the borders and around trees in the meadow. As stated previously, a majority of ticks were caught on sunny days. Presumably, the trees would provide shade from direct sunlight, therefore ticks could quest for longer periods and not risk quick desiccation (Conlon and Rockett 1982). In addition to the benefit of shade, the dog ticks may have aggregated near trees in the meadows because host scent may have been concentrated there, as many mammal burrows were seen in these areas. A considerable number (78) of adult ticks were collected from grassy roadsides within the parks. Smith et al. (1946) partially attributed the high density of dog ticks observed on road sides to the possible concentrated scent of their hosts. Furthermore, carbon dioxide has been determined to be an attractant to dog ticks (Garcia 1962) and some carbon dioxide is released from automobile traffic in the parks. Very few adult ticks (9) were encountered within the mixed deciduous and coniferous woodlands of Secor and Oak Openings Parks. This finding was not unexpected as this trend was observed by Conlon and Rockett (1982), Dodds et al. (1969), and Sonenshine and Levy (1972). Dodds et al. (1969) suggested that a well-developed canopy prevents the growth of adequate understory needed to maintain large numbers of small mammal hosts for the dog ticks. Interestingly, more male dog ticks (54%) were collected than females. Conlon and Rockett (1982) also captured more male ticks by dragging. Sonenshine et al. (1966) found that males tend to appear earlier than females. It is unknown whether this is a reflection of a more vigorous questing behavior in males or more rapid acclimation to spring weather conditions after overwintering. In contrast to the above, Garvie et al. (1978) found female ticks outnumbered male ticks on a study site in Nova Scotia, but did not speculate as to why females were more numerous. Moderate numbers of larvae (160) and low numbers of nymphs (25) were collected from mice nesting boxes placed throughout Oak Openings in 1989 and Mitchell and Micher (1989) reported the capture of 143 larvae and 8 nymphs from Secor Park in one season of study utilizing 24 nest boxes. Jackson and DeFoliart (1976) collected 7706 immatures in four seasons of study from 337 of 851 nests of white-footed mice in Wisconsin. However, a direct comparison cannot be made with the present study because Jackson and DeFoliart (1976) used both natural nest and nest boxes and did not distinguish between numbers of each. The natural nests may be more attractive to mice than wooden boxes. Interestingly, Jackson and DeFoliart (1976) also reported low numbers of nymphs in nests were at least partially the result of sampling without replacement, i.e., removal of engorged larvae. This could help to explain the low numbers of nymphs captured from nest boxes in the present study. Also, if medium-sized mammals were serving as major nymphal hosts, the population would certainly be much larger than the nymphal capture rate reported from the relatively small nest boxes. In a previous unpublished study, we noted the presence of potential nymphal hosts such as chipmunks, ground squirrels, red squirrels, raccoons and woodchucks, in Secor and Oak Openings. The American dog tick remains an important pest and vector of disease (RMSF) in Lucas County. Much remains to be learned of the bionomics of D. vanabilis. Studies are in progress to further detail the seasonal dynamics, host-specific aspects, and vegetative parameters of D. vanabilis in northwest Ohio.

69 1993 THE GREAT LAKES ENTOMOLOGIST 69 LITERATURE CITED Atwood, E. L., and D. E. Sonenshine Activity of the American dog tick, Dermacentor uariabilis (Acarina: Ixodidae), in relation to solar energy changes. Ann. Entomol. Soc. Am. 60: Campbell, A Ecology of the American dog tick, Dermacentor uariabilis (Say), in Southwestern Nova Scotia. pp In: J. J. Rodriguez. ed. Recent Advances in Acarology. Vol. 2. Academic Press, London. Campbell, A., and P. R. MacKay Distribution of the American dog tick. Dermacentor uariabilis (Say), and its small-mammal hosts in relation to vegetation types in a study area of Nova Scotia. Can. J. Zool. 57: Conlon, J. M., and C. L. Rockett Ecological investigations of the American dog tick. Dermacentor variabilis (Say). in Northwest Ohio (Acari: Ixodidae). Internat. J. Acarol. 8: Dodds, D. G. A. M. Martell, and R. E. Yescott Ecology of the American dog tick, Dermacentor uariabilis (Say). in Nova Scotia. Can. J. Zool. 47: Dukes, J. C.. and J. H. Rodriguez A bioassay for host-seeking responses of tick nymphs. J. Kans. Entomol. Soc. 49: Fish, D. and R. C. Dowler Host associations of ticks (Acari: Ixodidae) parasitizing medium-sized mammals in a Lyme disease endemic area of southern New York. J. Med. Entomol. 26: Fontaine, R. E. (Editor) Society for Vector Ecology (SOVE). Newsletter. Santa Ana, CA, 15 pp. Garcia, R Carbon dioxide as an attractant for certain ticks (Acarina: Argasidae and Ixodidae). Ann. Entomol. Soc. Am. 55: Garvie. M. B., J. A. McKiel, D. E. Sonenshine. and A. Campbell Seasonal dynamics of the American dog tick Dermacentor uariabilis (Say). populations in southwestern Nova Scotia. Can. J. Zool. 56: Hall. R. R., and J. A. McKiel Occurrence of the American dog tick, D. uariabilis (Say) in western Nova Scotia. Canadian Entomol. 93: Harlan. H. J Temperature effects on host-seeking by larval American dog ticks, Dermacentor uariabilis (Say). Ohio J. Sci. 86: Jackson, J. 0., and G. R. DeFoliart Relationship of immature Dermacentor uariabilis (Acari: Ixodidae) with the white-footed mouse, Peromyscus leucopus in southwestern Wisconsin. J. Med. Entomol. 12: McEnroe, W. D.. and M. A. McEnroe Questing behavior in the adult American dog tick. Dermacentor uariabilis (Say). (Acarina: Ixodidae). Acarologia 15: McEnroe, W. D The regulation of the adult American dog tick, Dermacentor variabilis (Say), seasonal activity and breeding potential. Acarologia 16: Winter survival microhabitat and constant density regulation of Dermacentor uariabilis (Say). Acarologia 19: The effect of the temperature regime on Dermacentor uariabilis (Say) populations in eastern North America. Acarologia 20: The effect of snow cover on an American dog tick, Dermacentor uariabilis (Say), population under a harsh winter environment. Z. Ang. Entomol. 97: Mitchell, L. and K. Micher Use of mouse nest boxes to survey for immature American dog ticks. Proc. Ohio Mosquito Control Assoc. 19: Mount, G. A. and O. G. Haile Computer simulation of population dynamics of the American dog tick (Acari: Ixodidae). J. Med. Entomol. 26: National Oceanic and Atmospheric Administration (NOAA) Climatological data of Ohio. Annual Summary, National Climatic Data Center, Asheville, NC, 94(13):34 pp Climatological data of Ohio. Annual Summary, National Climatic Data Center, Asheville, NC, 95(13):34 pp Climatological data of Ohio. Annual Summary, National Climatic Data Center, Asheville, NC. 96(13):34 pp.

70 70 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 Newhouse, V. F Variations in population density and rickettsial infection rates in a local population of Dermacentor variabilis (Acarina: Ixodidae) ticks in the Piedmont of Georgia. Environ. Entomol. 12: Smith, C. N., M. M. Cole, and H. K. Gouck Biology and Control of the American dog tick. USDA Tech. Bul. 905:74 pp. Sonenshine, D. E., E. L. Atwood, and J. T. Lamb, Jr The ecology of ticks trans mitting Rocky Mountain spotted fever in Virginia. Ann. Entomol. Soc. Am. 59: Sonenshine, D. E.. and G. F. Levy Ecology of the American dog tick, Dermacentor variabilis (Say), in a study area in Virginia. 2. Distribution in relation to vegetative types. Ann. Entomol. Soc. Am. 65: Sonenshine, D. E., A. H. Peters,and G. F. Levy Rocky Mountain spotted fever in relation to vegetation in the eastern United States, Am. J. Epidemiology 96: Sonenshine, D. E Zoogeography of the American dog tick, Dermacentor variabi lis. pp In: J. J. Rodriguez, ed. Recent Advances in Acarology. Vol. 2. Aca demic Press. London. Stone, K. L., E. H. McConoughey, G. D. Botrell, and D. J. Crowner, Soil Conservation Service Soil survey of Lucas County, Ohio. USDA, ODNR, U.S. Government Printing Office, 139 pp. Wilkinson, P. R Early achievements, recent advances, and future prospects in the ecology of the Rocky Mountain wood tick. pp , In: J. J. Rodriguez. ed. Recent Advances in Acarology. Vol. 2. Academic Press, London. Zar, J. H Biostatistical Analysis. Prentice Hall, Inc., Englewood Cliffs, N.J. 718 pp. Zimmerman, R. H., G. R. McWherter, and S. R. Bloemer Role of small mammals in population dynamics and dissemination of Amblyomma americanum and Derma centor variabilis (Acari: Ixodidae) at Land Between the Lakes, Tennessee. J. Med. Entomol. 24: Medium sized mammal host of Amblyomma americanum and Dermacentor variabilis (Acari: Ixodidae) at Land Between the Lakes, Tennessee, and effects of integrated tick management on host infestations. J. Med. EntomoL 25:

71 1993 THE GREAT LAKES ENTOMOLOGIST 71 RESISTANCE STABILITY OF THE SECONDARY TILLER OF 'CALDWELL' WHEAT AFTER THE PRIMARY CULM WAS INFESTED WITH VIRULENT HESSIAN FLY (DIPTERA: CECIDOMYIIDAE) LARVAE Stanley G. Wellso 1 and Jaime E. Araya 2 ABSTRACT Secondary tiller resistance of 'Caldwell' wheat, Triticum aestivum, with the H6 gene for larval resistance to Hessian fly, Mayetiola destructor, was maintained, after the primary culm had been previously infested with virulent larvae. Earlier studies showed that a primary culm infested initially with a virulent larva allowed subsequent normally avirulent larvae to survive on that cultivar; however, in our study the resistance of secondary tillers was maintained even though the primary culm was infested earlier with virulent Hessian fly larvae. This gene stability for resistance is important for optimizing wheat yield of those cultivars that possess genes resistant to the Hessian fly that are tillering and infested with different biotypes. Wheat. Triticum aestivum. is a very adaptable crop being grown across a wide range of environments. and leads in production and acreage of all crops (Briggle and Curtis 1987). It is also quite tolerant of insect attack due to its ability to produce secondary tillers. Wheat usually has eight tiller buds, but generally only three or four of these develop fully (Williams and Langer 1975). Thus, an insect may feed upon and destroy the primary culm. while laterdeveloping tillers may not be damaged and often produce seeds. Three major components affect wheat yield: density of fertile heads per unit area, number of seeds Iler head. and seed weight (often expressed as the weight per 1000 seeds) (Schlehuber and Tucker 1967). At maximum yield levels, a substantial increase in one yield component generally results in a decrease in one or both of the others. Biotic and abiotic conditions influence tiilering. Most studies of tillering in Grarnineae have focused on cultivar differences and the effects of a wide range of environmental factors. The physiology of tillering has been investigated by studying the effects caused by various growth substances and inhibitors (Leopold Williams et al Williams and Langer 1975). Little information. however. is available about the impact of insect numbers on wheat tillering. Multi-tillering wheat varieties were able to tolerate heavier infestations of the wheat bulb fly, Delia coarctata (Fallen), but this had the dis~dvantage of greater pest survival the following season (Oakley 1980). 1Insect and Weed Control Research Unit. U.S. Department of Agriculture. Agriculture Research Service. Department of Entomology, Purdue University. West Lafayette. Ind Department of Entomology, Purdue University, West Lafayette, Ind Current Address: Departmento de Sanidad Vegetal. Facultad de Ciencias Agrarias y Forestales. Universidad de Chile, Casilla 1004, Santiago. Chile.

72 72 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 Successful Hessian fly, Mayetiola destructor (Say), infestation of the primary culm usually kills it, and may result in the production of tillers; however, young plants do not tiller under severe infestation; abundantlytillering wheat cultivars appear to survive Hessian fly infestations better (Barnes 1956). Buntin and Raymer (1989) noted that low to moderate Hessian fly damage levels reduced forage yield primarily by reducing tiller size and weight. rather than tiller density. Tillering of 'Arthur 71' (H3H5 genes for resistance), 'Monon' (H3), and 'Seneca' (H7Hs) increased when infested with Hessian fly biotypes B, D, or GP at higher temperatures (Sosa and Foster 1976). Tillering of 'Knox 62' (Hs and perhaps H7Hs) was relatively stable when infested with biotypes B, C, D, or GP, and biotype C did not cause an increase in tiller numbers in any of the four cultivars. 'Monon' and 'Newton' wheat cultivars differ in their tillering response when infested by Hessian fly (unpublished data, Wellso and Hoxie). Although 'Newton' tillered less than 'Monon' at the 0 infestation level, infestations of 1-3 puparia (indicative of the number of infesting larvae per seedling) per primary culm resulted in a greater number of tillers. 'Monon' had the same number of tillers at 0-3 puparia per primary culm; however, both cultivars tillered less at 4 or more puparia than at 1-3 puparia per primary culm. Thus, the Hessian fly mayor may not increase tillering under light infestations; however, higher infestation levels lessened tillering, perhaps due to the depletion of soluble carbohydrates (Wellso et al The main resistance mechanism of wheat to the Hessian fly is larval antibiosis, resulting in the death of young larvae due to their inability to maintain sustained feeding (Gallun 1965, Shukle et al. 1990) and the resistant plant continues to grow with little evidence of insect feeding. Grover et al. (1989) noted that infestation by a single virulent larva resulted in an alteration of resistance of the plant, allowin~ survival of all normally avirulent larvae that concurrently or subse ntly mfested the plant. Their research dealt only with the primary culm. used four cultivars ('Abe', Purdue line 6549, 'Caldwell' and 'Monon') with various combination of infestation with four Hessian fly biotypes (B, C, D, and L). Little information is available about the effectiveness of Hessian fly resistant wheat genes when the primary culm is infested with a biotype that is either virulent or avirulent and a secondary tiller is subsequently infested with the same or another biotype. The objective of this study was to evaluate the stability of resistance in the primary culm and secondary tiller of 'Caldwell' wheat infested by virulent or avirulent Hessian fly biotype combinations. MATERIALS AND METHODS Test Insects. Hessian fly biotypes Band L, originally isolated in Indiana approximately 21 and 13 years ago, respectively, have subsequently been maintained in culture at the Insects and Weed Control Research Unit, U.S. Dept. of Agriculture, Purdue University, Ind. To maintain biotype cultures for these studies, mated females oviposited on susceptible wheat (e.g., 'Newton' CI 17715, Ho gene for resistance) maintained in a growth chamber at 21.1 C, a 16:8 (L:D) photoperiod, light intensity of 300,uEinsteins/m 2 /sec, and 65% R.H. Biotype purity was verified periodically with wheat cultivars having known resistance genes (Sosa and Gallun 1973). Test Plants. Three 'Caldwell' wheat (CI 17897; Hs gene; resistant to Hessian fly biotype B, but susceptible to biotype L) seeds, obtained from Purdue University's wheat breeding program, were soaked for 1 h, planted in a mix

73 1993 THE GREAT LAKES ENTOMOLOGIST 73 ture of soil-vermiculite in forty 10 cm diameter plastic pots (with 4 replicates), and later thinned to one seedling per pot (24 seedlings of each cultivar per treatment). Plants were maintained in a growth chamber set at 21.1 C, 65 ± 10% RH and 14:10 (L:D), and provided with Hoagland's solution biweekly and watered as needed. Treatments. 'Caldwell' seedlings were exposed to ovipositing Hessian fly as follows: Control, not infested; Primary culm infested 1 wk after planting with biotype B; Primary culm not infested; secondary tiller infested 3 wk after planting with biotype B; Primary culm infested 1 wk after planting with biotype B; secondary tiller of the same seedling infested 3 wk after planting with biotype B; Primary culm infested 1 wk after planting with biotype L; secondary tiller of the same seedling infested 3 wk later with biotype B. Adult Hessian fly of the desired biotype were released in a growth chamber to oviposit on the first or second leaf of the primary culm, and oviposition was verified within 24 h. Leaves without eggs were exposed to adults until eggs were found on each seedling. Secondary tiller leaves were infested by confining a mated female in a cylindrical screen cage (2.5 x 12 cm) with plastic foam plugs supported by a metal rod enclosing a normal leaf [as opposed to dark green leaves on a primary culm that previously had been infested successfully by virulent larvate)]. The cage was removed after oviposition was confirmed. The plants were scored 6 wk after planting relative to the number of tillers. plant length from the crown to the apex of the longest leaf, above ground fresh weight, live Hessian fly larvae or puparia and dead larvae (usually small red larvae about 0_5 mm in length). The experiment was replicated on four dates. Data Analysis: After analysis of variance (ANOVA, SAS Institute 1985) using a split-plot design, significantly different means ( P S 0.05) were separated by Student-Newman-Keuls test (Steel and Tonie 1980). RESULTS AND DISCUSSION There was a significantly higher (P=0.05) level of infestation of primary culms of 'Caldwell' seedlings by virulent biotype L than by avirulent biotype B larvae (Table 1). Secondary tillers also were resistant to biotype B larvae, including those from seedlings on which the primary culm was infested by virulent biotype L larvae. The mean numbers of tillers (stems) per plant was not significantly different among the Hessian fly treatments (Table 1/. Plant length and fresh weight did not differ significantly between control plants infested by avirulent biotype B larvae on either the primary or secondary tiller, however, plant length was significantly less when both the primary culm and secondary tiller were infested with biotype B larvae. Virulent biotype L larvae had the greatest effect on plant parameters. Infestation of the primary culm by biotype L larvae resulted in a significant decrease in plant weight, length and number of leaves per plant. The greatest impact to the plant by biotype L larvae was reduction in fresh plant weight, which decreased 50% of that of the control. Wellso et al. ( ) reported that uninfested plants of 'Winoka' and 'Monon', respectively, had significantly greater plant length, weight. and number of leaves than plants infested with virulent larvae. The 50% reduction in fresh plant weight of 'Caldwell' was less than the 65 to 89% reduction in

74 ~ -I Table 1. Effect of virulent (biotype L) and avirulent (biotype B) larval infestation of the Hessian fly (FH) on the primary culm and/or second;ary ::J: tillers of 'Caldwell' winter wheat in the laboratory" m Gl Treatmentsb % HF infestation±sd HFc per tiller ± SD ;0 Stems Leaves Plant Fresh m Primary Primary Secondary Primary Secondary per plant per plant length plant wt ~ culm tiller culm tiller culm tiller ±SD ±SD (cm±sd) (g±sd) B 2L9± 2.6b 3.7± ±1.1a 16.8±3.3a 46.3±2.3a 4.8± 1.4a ~ m B 18.8±12.5a 5.8± ±1.4a 18.2±4.1a 45.4±2.9a 6.0± 1.5a (/1 B B 15.6± 12.0b 12.5 ± 1O.2a 1l.8± ± ±1.2a 16.3±3.7ab 43.2±4.1b 4.4± 1.5a m L B 75.0±49.9a 9.4± 1.2a 8.2± ± ± 1.5a 13.6±1.5a 36.0±6.1c 2.4± 1.3b Z 3.6±O.9v. 15.9±3.2b 46.0±3.0a 4.8±1.3a o s: ameans in the same column followed by the same letter are not significantly different (P:5 0.05), according to a Fisher's Protected LSD test. for o % HF infestation of BB and LB (df = 1; F = 21.7), stems per plant (df = 4; F = 1.1), leaves per plant (df = 4,5; F = 5.9), plant length (df = 4; 5 F = 38.9), and fresh plant weight (df = 4; F = 28.9). This experiment included four replicates and 8 plants per replicate. Gl bhf biotype on primary colm and/or a secondary tiller. clarvae and puparia per tiller. ~ 'ti,~ z 9

75 1993 THE GREAT LAKES ENTOMOLOGIST 75 weight of Winoka' infested with 1 to 3 or 7 to 9 larvae, respectively, for 8 wk old plants (Wellso et al. 1989). The greater loss in weight of 'Winoka' plants may have been due, in part. to environmental differences between the test parameters and to the longer duration of the study. The adverse impact (stunting) of infestation by virulent biotype L larvae on development of the primary culm of susceptible 'Caldwell' seedlings was clearly demonstrated in this study. Of significance, however, was the fact that secondary tillers which developed from stunted primary culms (susceptible to biotype L), retained resistance to the avirulent biotype B larvae. This is in contrast to the finding that resistance of the primary culm to normally avirulent larvae is lost following feeding by a virulent larva (Grover et al1989). The response of tillers observed in this test would be beneficial where different biotypes are present in the same field, because tillers would be capable of surviving and producing grain, even after the primary culm was destroyed by a virulent biotype. ACKNOWLEDGMENTS We thank Wyman Nyquist and Judith Santini for statistical assistance, and Clifford Papsdorf for technical assistance. Purdue Univ. Agr. Exp. Stn. Journal Paper No.. LITERATURE CITED Barnes, H. G Gali midges of economic importance. VII. Gall midges of cereal crops, the Hessian fly, pp Crosby Lockwood and Sons, LTD. London. Briggle. L. W. and B. C. Curtis Wheat worldwide, pp In: E. G. Heyne (ed.), Wheat and wheat improvement. Agron. Monogr. No. 13. ASACCSA-SSA, Madison, Wisconsin. Buntin, G. D. and P. L. Raymer Hessian fly (Diptera: Cecidomyiidae) damage and forage production of winter wheat. J. Econ. Entomol. 82: Galiun, R. L The Hessian fly, and how to control it. USDA Leaflet Genetic basis of Hessian fly epidemics. Ann. N. Y. Acad. Sci. 287: Grover, P. B., Jr., R. H. Shukle and J. E. Foster Interactions of Hessian fly (Diptera: Cecidomyiidae) biotypes on resistant wheat. Environ. Entomol. 18: Leopold, A. D Plant growth and development. McGraw-Hill, New York. Oakley, J. N An integrated control strategy for the wheat bulb fly. ADAS Quart. Rev. 38: SAS Institute SAS user's guide: statistics. 5th ed. SAS Institute, Cary, N.C. Schlehuber, A. M. and B. B. Tucker Culture of wheat, pp In: K. S. Quisenberry and L. P. Reitz (ed.) Wheat and wheat improvement. Agron. Monogr. No. 13. ASA-CCSA-SSA, Madison, Wisconsin. ShukIe, R. H., P. B. Grover, Jr. and J. E. Foster Feeding of Hessian fly (Diptera: Cecidomyiidae) larvae on resistant and susceptible wheat. Environ. Entomol. 19: Sosa, 0., Jr. and J. E. Foster Temperature and the expression of resistance in wheat to the Hessian fly. Environ. Entomol. 5: Sasa, 0., Jr. and R. L. Galiun Purification of races B and C of the Hessian fly by genetic manipulation. Ann. Entomol. Soc. Am. 66: Steel, R. G. D. and J. H. Tome Principles and procedures of statistics, with special reference to the biological sciences. McGraw Hill, New York. Wellso, S. G., R. P. Hoxie and C. R. Olien Effects of Hessian fly (Diptera: Cecido